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HARVARD UNIVERSITY

Library of the

Museum of

Comparative Zoology

€ GREAT BASIN NATURALIST

January 31, 1983

Brigham Young University

iviUS. COMP. ZOOL
LIBRARY

UMiVEJRSiTY

GREAT BASIN NATURALIST

Editor. Stephen L. Wood, Department of Zoology, 290 Life Science Museum, Brigham Young

University, Provo, Utah 84602.
Editorial Board. Kimball T. Harper, Chairman, Botany; James R. Barnes, Zoology; Hal L.
Black, Zoology; Stanley L. Welsh, Botany; Clayton M. White, Zoology. All are at Brig-
ham Young University, Provo, Utah 84602.
Ex Officio Editorial Board Members. Bruce N. Smith, Dean, College of Biological and Agricul-
tural Sciences; Norman A. Darais, University Editor, University Publications.
Subject Area Associate Editors.
Dr. Noel H. Holmgren, New York Botanical Garden, Bronx, New York 10458 (Plant

Taxonomy).
Dr. James A. MacMahon, Utah State University, Department of Biology, UMC 53, Lo-
gan, Utah 84322 (Vertebrate Zoology).
Dr. G. Wayne Minshall, Department of Biology, Idaho State University, Pocatello,

Idaho 83201 (Aquatic Biology).
Dr. Ned K. Johnson, Museum of Vertebrate Zoology and Department of Zoology, Uni-
versity of California, Berkeley, California 94720 (Ornithology).
Dr. E. Philip Pister, Associate Fishery Biologist, California Department of Fish and

Game, 407 West Line Street, Bishop, California 93514 (Fish Biology).
Dr. Wayne N. Mathis, Chairman, Department of Entomology, National Museum of

Natural History, Smithsonian Institution, Washington, D.C. 20560 (Entomology).
Dr. Theodore W. Weaver III, Department of Botany, Montana State University, Boze-
man, Montana 59715 (Plant Ecology).
The Great Basin Naturalist was founded in 1939 and has been published from one to four
times a year since then by Brigham Young University. Previously unpublished manuscripts in
English of less than 100 printed pages in length and pertaining to the biological natural his-
tory of western North America are accepted. Western North America is considered to be west
of the Mississippi River from Alaska to Panama. The Great Basin Naturalist Memoirs was es-
tablished in 1976 for scholarly works in biological natural history longer than can be accom-
modated in the parent publication. The Memoirs appears irregularly and bears no geographi-
cal restriction in subject matter. Manuscripts are subject to the approval of the editor.

Subscriptions. The annual subscription to the Great Basin Naturalist for private individuals
is $16.00; for institutions, $24.00 (outside the United States, $18.00 and $26.00); and for stu-
dent subscriptions, $10.00. The price of single issues is $6.00 each. All back issues are in print
and are available for sale. All matters pertaining to subscriptions, back issues, or other busi-
ness should be directed to Brigham Young University, Great Basin Naturalist, 290 Life Sci-
ence Museum, Provo, Utah 84602. The Great Basin Naturalist Memoirs may be purchased
from the same office at the rate indicated on the inside of the back cover of either journal.

Scholarly Exchanges. Libraries or other organizations interested in obtaining either journal
through a continuing exchange of scholarly publications should contact the Brigham Young
University Exchange Librarian, Harold B. Lee Library, Provo, Utah 84602.
Manuscripts. See Notice to Contributors on the inside back cover.

3-83 650 64094

ISSN 017-3614

TABLE OF CONTENTS

Volume 43
Number 1 - January 31, 1983

Life history of the Lahontan cutthroat trout, Salmo clarki henshawi, in Pyramid Lake,
Nevada. WilHam F. Sigler, William T. Helm, Paul A. Kucera, Steven Vigg, and

Gar W. Workman 1

A review of the genus Soliperla (Plecoptera: Peltoperlidae). Bill P. Stark 30

A bibliography of Colorado vegetation description. William L. Baker 45

Evaluation of a program to control hydatid disease in central Utah. Ferron L. Andersen,

John R. Crellin, Craig R. Nichols, and Peter M. Schantz 65

Influence of cryptogamic crusts on moisture relationships of soils in Navajo National

Monimient, Arizona. Jack D. Brotherson and Samuel R. Rushforth 73

A vascular flora of the San Rafael Swell, Utah. James G. Harris 79

Pronghom responses to hunting coyotes. Timothy D. Reynolds 88

Floristics of the upper Walker River, California and Nevada. Matt Lavin 93

Agrapyron arizonicum (Gramineae: Triticeae) and a natural hybrid from Arizona. Grant

L. Pyrah 131

Species composition, distribution, and phytosociology of Kalsow Prairie, a mesic

tallgrass prairie in Iowa. Jack D. Brotherson 137

Deer mouse, Peromyscus maniculatus, and associated rodent fleas (Siphonaptera) in the
arctic-alpine life zone of Rocky Mountain National Park, Colorado. R. B. Eads and

E. G. Campos 168

Food of larval Tui chubs, Gila bicolor, in Pyramid Lake, Nevada. David L. Galat and

Nancy Vucinich 175

Number 2 - April 30, 1983

Utah flora: Compositae (Asteraceae). Stanley L. Welsh 179

Haplopappus crispus and H. zionis (Asteraceae): new species from Utah. Loran C.

Anderson 358

A bouquet of daisies {Erigeron, Compositae). Stanley L. Welsh 365

New taxa in Thelesperma and Townsendia (Compositae) from Utah. Stanley L. Welsh .... 369

New Haplopappus variety in Utah (Compositae). Stanley L. Welsh and Frank J. Smith ... 371
New species of Hymenoxys and Perityle (Compositae) from Utah. Stanley L. Welsh and

Elizabeth Neese 373

New variety of Stephanomeria tenuifolia (Compositae) from Utah. Sherel Goodrich and

Stanley L. Welsh 375

Number 3 - July 31, 1983

A re-evaluation of the postglacial vegetation of the Laramie Basin, Wyoming-Colorado.

Deborah L. Elliott-Fisk, Betty S. Adkins, and Jeanine L. Spaulding 377

Comparative life history and floral characteristics of desert and mountain floras in Utah.

Patrick D. Collins, Kimball T. Harper, and Burton K. Pendleton 385

Flora of the Lower Cretaceous Cedar Mountain Formation of Utah and Colorado, part

I. Paraphyllanthoxylon utahense. G. F. Thayne, W. D. Tidwell, and W. L. Stokes .. 394

Aculeata Hymenoptera of Sand Mountain and Blow Sand Mountains, Nevada. R. W.

Rust, L. M. Hanks, and R. C. Bechtel 403

Status and life history notes on the native fishes of the Alvord Basin, Oregon and

Nevada. Jack E. Williams and Carl E. Bond 409

Kramer Palouse natural area. Del W. Despain and Grant A. Harris 421

Winter food habits of cougars from northeastern Oregon. Chris Maser and Ronald S.

Rohweder 425

A new species of Penstemon (Scrophulariaceae) from the Uinta Basin, Utah. Elizabeth

Neese and Stanley L. Welsh 429

A comparative study of coyote food habits on two Utah deer herds. Jordan C. Pederson

and R. Cary Tuckfield 432

A small carnivore survey technique. Tim W. Clark and Thomas M. Campbell III 438

Evaluation of Draba oligosperma, D. pectinipila, and D. juniperina complex

(Cruciferae). Robert W. Lichvar 441

Presence of maxillary canine teeth in mule deer in Utah. Jordan C. Pederson 445

Comparative successional roles of trembling aspen and lodgepole pine in the Southern

Rocky Mountains. Albert J. Parker and Kathleen C. Parker 447

Differential utilization of bat boxes by house wrens {Troglodytes aedon). Hal L. Black 456

Plant and soil relationships in two hydrothermally altered areas of the Great Basin. N.

M. Milton and T. L. Purdy 457

Plasticity and polymorphism in seed germination of Mimulus guttatus

(Scrophulariaceae). Robert K. Vickery, Jr 470

Predatory behavior of larval Ambystoma tigrinurn nebulosiim on Limnephilus

(Trichoptera) larvae. Joseph R. Holomuzki 475

Notes on reproduction of the side-blotched lizard Uta stansburiana stansbiiriana in

southwest Idaho. George Bakewell, Joseph M. Chopek, and Gary L. Burkholder .... 477
Observations on alpine vegetation near Schoolroom Glacier, Teton Range, Wyoming.

John R. Spence and Richard J. Shaw 483

Winter stomach contents of South Dakota badgers. E. Blake Hart and Michael Trumbo . 492
A list of Utah spiders, with their localities. Dorald M. Allred and B. J. Kaston 494

Number 4 - October 31, 1983

Alpine and subalpine wetland plant communities of the Uinta Mountains, Utah. George

M. Briggs and James A. MacMahon 523

Utah flora: Salicaceae. Sherel Goodrich 531

A mammalian humerus from the Upper Jurassic of Colorado. Donald R. Prothero and

James A. Jensen 551

Bats of the Colorado oil shale region. Robert B. Finley, Jr., William Caire, and Dallas E.

Wilhelm 554

New generic concepts in the Triticeae of the Intermountain Region: keys and

comments. Mary E. Barkworth, Douglas R. Dewey, and Riley J. Atkins 561

Reproductive attributes of some Rocky Mountain subalpine herbs in successional

context. David J. Schimpf and Robert L. Bayn, Jr 573

Applicability of the universal soil loss equation for southeastern Idaho wildlands. Mark

E. Jensen 579

Winter stoneflies (Plecoptera) of New Mexico. Gerald Z. Jacobi and Richard W.

Baumann 585

Daily and yearly movement of the Devil's Hole pupfish Cyprinodon diabolis Wales in

Devil's Hole, Nevada. Thomas M. Baugh and James E. Deacon 592

A revision of the genus Microrhopala (Coleoptera: Chrysomelidae) in America north of

Mexico. Shawn M. Clark 597

Flora of the Stansbury Mountains, Utah. Alan C. Taye 619

New synonymy and new species of American bark beetles (Coleoptera: Scolytidae), part

IX. Stephen L. Wood 647

Plant commimity variability on a small area in southeastern Montana. James G.

MacCracken, Daniel W. Uresk, and Richard M. Hansen 660

New leafhopper species of Coelidia with a revised key and notes on homonymy and

distribution (Homoptera: Cicadellidae, Coelidiinae). Mervin W. Nielson 669

Eye fluke {Diplostomum spathaceum) of fishes from the upper Salmon River near

Obsidian, Idaho. Richard Heckmann 675

Evaluation of varieties in Stanleya pinnata (Cruciferae). Robert W. Lichvar 684

Some aspects of the presettlement vegetation of the Piceance Basin, Colorado. William

L.Baker 687

New variety of Opuntia hasilaris (Cactaceae) from Utah. Stanley L. Welsh and

Elizabeth Neese 700

Vegetative types and endemic plants of the Bryce Canyon Breaks. Robert A. Graybosch

and Hayle Buchanan 701

Seasonal growth of the Tui chub, Gila bicolor, in Pyramid Lake, Nevada. Joseph L.

Kennedy 713

Bird distributional and breeding records for southeastern Idaho, Utah, and adjacent

regions. Clayton M. White, Herbert H. Frost, Dennis L. Shirley, G. Merrill Webb,

and Richard D. Porter 717

Birds of southwestern Idaho. Daniel A. Stephens and Timothy D. Reynolds 728

New records for the vascular flora of Wyoming and Montana. Robert W. Lichvar,

Robert D. Dom, and Erwin F. Evert 739

Range extensions for two dwarf mistletoes (Arceuthobium spp.) in the southwest. Robert

L. Mathiasen and Kenneth H. Jones 741

First nest records for the Plain Titmouse and Blue-gray Gnatcatcher in Wyoming. Scott

L. Findholt 747

A new species of Thelesperma (Asteraceae) from Wyoming. Robert D. Dorn 749

Index 751

The Great Basin Naturalist

Published at Provo, Utah, by
Brigham Young University

ISSN 0017-3614

Volume 43 January 31, 1983 No. 1

LIFE HISTORY OF THE LAHONTAN CUTTHROAT TROUT,
SALMO CLARKI HENSHAWI, IN PYRAMID LAKE, NEVADA

William F. Sigler', William T. Helm^ Paul A. Kucera\ Steven Vigg*, and Gar W. Workman'

Abstract.— The Pyramid Lake Lahontan cutthroat trout (Salmo clarki henshawi) population was sampled on a
monthly basis from November 1975 through December 1977. A subsample of 676 trout, stratified by fish size and
lake habitat, provided biological data. The entire population is presently derived from hatchery production, stocked
at lengths of approximately 75 to 300 mm. Peak annulus formation occurs in March and April, followed by the peri-
od of maximum growth. Scale patterns illustrate a variable growing season. Maximum growth in length is in the first
three years of life; after that males begin to grow faster than females. Males attained a greater age in our sample; i.e.,
the oldest male was seven years old compared to six years for females. The Pyramid Lake Lahontan cutthroat trout
exhibit nearly isometric growth.

The legal sport fishery removed <20,000 adult fish in 1977 (>380 mm); other decimating factors are poorly un-
derstood. No evidence of the following diseases or pathogens was found in the Pyramid Lake population, presuming
a carrier incidence of 2 percent at the 95 percent confidence level: infectious pancreatic necrosis, infectious hemato-
poietic necrosis, viral hemorrhagic septicema, bacterial kidney disease, enteric redmouth, furunculosis, whirling dis-
ease, blood fluke; however, 7 of 235 (=:;3 percent) adults sampled at the Marble Bluff fish way were positive for
furunculosis.

Small trout feed primarily on zooplankton and benthic invertebrates; cutthroat trout >300 mm are piscivorous,
feeding almost exclusively on tui chub {Gila bicolor). The spawning migration of Pyramid Lake cutthroat trout to the
Marble Bluff egg taking facility in spring 1976 and 1977 peaked in April and May. Females mature at three or four
years (352-484 mm), and males mature at two or three years (299-445 mm). Mean diameter of mature eggs is 4.51
mm; both ovum size and fecundity are a function of fish size. Fecundity ranges from 1241 to 7963 eggs, with a mean
of 3815.

Lahontan cutthroat trout comprise <2 percent of the numerical relative abundance and <7 percent of the total
fish biomass. Distribution patterns vary on a seasonal basis, with maximum activity during late fall and winter. Man-
agement objectives are presented and recommendations are discussed.

The Lahontan cutthroat trout (Salmo clarki The dechne and ultimate extinction of the

henshawi) is unique in its abihty to withstand original strain of cutthroat trout in Pyramid

the alkaline-saline waters of remnant Great Lake was caused primarily by degradation of

Basin lakes. Coevolution of Pyramid Lake spawning habitat associated with diversion of

Lahontan cutthroat trout in a continuous lake water out of the Truckee River-Pyramid

environment for 50,000-100,000 years with Lake ecosystem (Trelease 1953). The Pyra-

an abundant prey species (tui chub, Gila mid Lake trout fishery has been reestablished

bicolor) resulted in a unique predator— the via hatchery propagation of Heenan, Walker,

world's largest cutthroat trout (18.6 kg). and Summit lake strains of Lahontan cut-

'W. F. Sigler & Associates Inc., P.O. Box 1350, Logan, Utah 84322.

'Utah State University, Logan, Utah 84322.

'Nez Perce Tribe, Lapwai, Idaho 83540.

'Desert Research Institute, University of Nevada System, Reno, Nevada 89507.

'Utah State UniveRity, Logan, Utah 84322.

Great Basin Naturalist

Vol. 43, No. 1

Fig. 1. The largest post-1943 Lahontan cutthroat
trout (12.7 kg), captured by Ralston Fillmore from Pyra-
mid Lake, Nevada, April 1976. Photograph courtesy of
Alan Ruger.

throat trout, and, in the past, of cutthroat-
rainbow trout {Salmo gairdneri) hybrids. The
subspecies S. c. henshawi currently has
"threatened" status (Deacon et al. 1979).

Pyramid Lake presently supports a trophy
sport fishery; the average trout retained by
fishermen is 500 mm in length and weighs 1 .2
kg. In 1976, a Paiute Indian, Ralston Fill-
more, captured a 12.7 kg Lahontan cutthroat
trout that represents a record for the post-
1943 fishery (Fig. 1). Recent catches are evi-
dence that the environment of Pyramid Lake
is capable of supporting at least a limited
valuable and unique fishery. However, hu-
man demands on limited Truckee River wa-
ter and recent droughts have jeopardized the
trout in Pyramid Lake. The ethics and prior-
ities of our society, as a whole, may ulti-
mately decide the fate of the Pyramid Lake
Lahontan cutthroat trout.

Nine now discrete basins once conjoined to
form vast (area 22,300 km^; maximum depth
270 m) Lake Lahontan in the northwestern
Great Basin (Hubbs and Miller 1948). Pyra-
mid Lake is the deepest remnant of this once
great lake system that experienced several
cycles of water level fluctuations during the
Pleistocene Epoch (Houghton 1976). Great
Basin lakes have desiccated to the present
state since the last pluvial period some 10-12
thousand years before present (BP). Benson
(1978) concludes via sediment analyses that
Pyramid Lake was greatly reduced in size
9-5 thousand years BP, but did not become
dry and had subsequently been rising until
the cultural impacts of the past century.

Pyramid is a graben lake approximately 40
km long and 6.5 to 16 km wide, with a north-
south axis (Figure 2). At the mean 1976 ele-
vation of 1157 m (United States Geological
Survey 1977), Pyramid Lake has a surface
area of 446.4 km^, a volume of 26.4 km^, a
mean depth of 59 m and a maximum depth
of 103 m (Harris 1970). Pyramid is the deep-
est and most voluminous saline terminal lake
in the western hemisphere (Galat et al. 1981).

Pyramid Lake, located entirely within the
Pyramid Lake Paiute Indian Reservation, is
the terminal water body of the endorheic
Truckee River system originating 193 river
km upstream at oligotrophic Lake Tahoe.
The evaporation loss is about 1.2 m annually.
Due to transbasin diversion of the Truckee
River, the lake level declined 23 m between
1905 and 1979; this amounts to a 30 percent
reduction in lake volume. The lake water is
highly ionic (Na+ > K+ > Mg2+ > Ca2+;
CI- > HCO3 > CO32- > SO42-), with a pH
of 9.2. The 1976 total dissolved solids (TDS)
concentration was 5235 mg/1 at elevation
1157 m. On a worldwide perspective, 71 per-
cent of some 350 saline lakes listed by
McCarraher (1972) are more saline than
Pyramid, but, compared to USA saline lakes.
Pyramid is in the moderate range (Galat et
al. 1981).

During 1976 and 1977 mean surface tem-
perature ranged from 6.1 to 23.1 C. As winds
subside and surface water temperature in-
creases, a thermocline is formed in June and
lasts through December at 16 to 22 m. The
lake is monomictic (Hutchinson 1957); turn-
over begins in early winter and mixing ex-
tends to spring.

January 1983

SiGLER ET AL.: LaHONTAN CuTTHROAT TrOUT

Fox Valley

Needles

Cormorant
Rock

Helk
Kitchen

Anderson
Bay

True
North

PopcortT"

Fig. 2. Bathymetric map of Pyramid Lake, Nevada; depth contours are in meters.

Physical changes, including out-of-basin
and inbasin water diversions, channelization,
and destruction of riparian habitat, have ad-
versely affected the ecology of the Truckee
River-Pyramid Lake ecosystem. Historically
the Lahontan cutthroat trout moved out of

Pyramid and Winnemucca lakes, Nevada,
and spawned in the entire Truckee River and
its tributaries, a total length of 525 km. They
also moved into Lake Tahoe, Nevada-Califor-
nia, and spawned in its tributary streams.
Derby Dam, completed in 1905, 62 km

Great Basin Naturalist

Vol. 43, No. 1

above Pyramid Lake, effected a transbasin
diversion of much of the lower Truckee Riv-
er flow. This obstacle reduced river spawning
to the area below the dam. The dam not only
had the direct effect of reducing flows in the
lower river, but indirectly caused the buildup
of the delta at the mouth. The numbers of
trout diminished steadily imtil 1930, which
was the last successful spawning year for the
original population of Pyramid Lake Lahon-
tan cutthroat trout (Sumner 1939). The U.S.
Bureau of Fisheries (now U.S. Fish and Wild-
life Service) stocked limited numbers in the
lake in 1931 and 1932. None was seen in
Pyramid Lake after 1943 and very few after
1938. However, before that time millions of
eggs had been taken from the Pyramid Lake
cutthroat and stocked elsewhere (Townley
1980).

In 1976 the Marble Bluff complex, con-
sisting of a dam and impoundment, a build-
ing, and a 5.6 km fishway ending at the lake,
was completed. This facility was built so that
spawning fish could migrate upriver when
there was not enough water in the river del-
ta. The fishway, operating at 0.85 to 1.27
mVsec, allows fish to move upstream via four
step-up ladders. At the upper end of the fish-
way the fish may be shunted on upstream or
into the building.

Importance

The Pyramid Lake Lahontan cutthroat
trout is potentially of substantial economic
and social importance to the Pyramid Lake
Paiute Indian Tribe. Its adaptation to the
highly saline waters of the lake make it a
unique natural history entity.

In 1977 an estimated 27,241 people spent
276,532 hours fishing for Lahontan cutthroat
trout in Pyramid Lake. They landed 43,841
fish, of which 19,930 or 46 percent were le-
gal size (381 mm). This is at the rate of 0.16
fish per hour landed and 0.07 kept.

It has been estimated that the historic an-
nual production of Lahontan cutthroat trout
was at least 454,000 kg (Behnke 1974). The
Pyramid Lake trout was the mainstay in the
diet of the Pyramid Lake Paiutes and many
other Indian tribes. They were also shipped
to mining camps and other markets as far
away as San Francisco.

Range and Distrirution

An ancestoral cutthroat trout probably in-
vaded ancient Lake Lahontan from the Co-
lumbia River Basin and developed into what
is now known as the Lahontan cutthroat
trout (Behnke and Zarn 1976). When the
great lake desiccated, two populations of La-
hontans evolved, one best adapted to lakes
and the other to streams. The major lake
populations of Lahontan cutthroat trout were
then in Pyramid Lake, Walker Lake, Donner
Lake, Independence Lake, and Lake Tahoe
(Miller 1951). The trout in some of these
lakes, which held an abundance of forage
fish, became predatory at an early age, grew
fast and large, and were moderately long
lived.

Currently, the largest population of lake-
dwelling Lahontan cutthroat trout is in Pyra-
mid Lake. Walker Lake supports a small
population that has no opportunity to repro-
duce. Summit Lake and Independence Lake
may contain the most nearly pure strain of
the Lahontan, but both lakes are small and
have few forage fish (Behnke and Zarn 1976).
A number of western lakes support reproduc-
ing populations of Lahontan cutthroat trout.
Recently a small stream-dwelling population
of Lahontan cutthroat trout, believed to have
been transplanted from Pyramid Lake before
1930, was discovered near Pilot's Peak, north
of Wendover, Utah-Nevada (Hickman and
Duff 1978, Hickman and Behnke 1979).
There are a number of stream-dwelling La-
hontan cutthroat trout populations in the
Great Basin.

In 1950, the Nevada Fish and Game De-
partment initiated a small-scale stocking pro-
gram of Lahontan cutthroat and other trout
in Pyramid Lake (Trelease 1969). The pro-
gram has now grown to 2.2 million 75-300
mm fish per year, supplied by two hatcheries
of the Pyramid Lake Indian Tribal Enter-
prises (PLITE) and the Lahontan National
Fish Hatchery at Gardnerville, Nevada.

Morphology and Genetics

Despite the diverse evolutionary histories
of western trout (genus Saltno), some species
are related closely enough to interbreed free-
ly and produce fertile hybrids. It is this po-
tential presence of all degrees of hybrids

January 1983

SiGLER ET AL.: LaHONTAN CuTTHROAT TrOUT

within a habitat that complicates identi-
fication and evaluation of pure stock (Behnke
and Zam 1976). The original stock of Lahon-
tan cutthroat trout was apparently resistant
to hybridization due to its long isolation in
the Lahontan basin. The present subspecies
does not share this characteristic. The isola-
tion also encouraged a high degree of adapt-
ability for lake habitat.

The following are typical meristics of the
Lahontan cutthroat trout (Behnke and Zam
1976):

Scale counts
lateral series
two rows above

lateral line 150-180

above lateral line
(origin of dorsal
fin to lateral

line) 33-43

Vertebrae 61-63

Gillrakers 21-28

Pyloric caecae 40-75

Basibranchial teeth Numerous and

well developed

The number of pyloric caecae is higher in the
Lahontan cutthroat than in other subspecies
of cutthroat. The large, round, rather dull
reddish spots that appear on the head as well
as on the caudal peduncle and occasionally
ventrally are the best field characteristic.

The following data were collected as part
of a study contracted between W. F. Sigler &
Associates Inc. and the United States. This
study was to provide an ecological evaluation
of Pyramid Lake and its fishery resources and
habitat.

Procedures

Fish life history data were taken by month-
ly nettings from November 1975 through De-
cember 1977. Fish were sampled by bottom
set variable-mesh gill nets, vertical set gill
nets, beach seines, fyke nets, and trawls. Fish
were measured to the nearest millimeter in
fork (FL), standard (SL), and total (TL)
lengths and weighed to the nearest gram (Sig-
ler and Kennedy 1978).

Scale samples were collected from the left
side in the region above the lateral line and
midway between the posterior edge of the

operculum and the origin of the dorsal fin.
Five scales per fish were selected and impres-
sions made of them on plastic slides with the
use of a roller press (Smith 1963).

The length-weight relationship is expressed
by the formula W = aL'' (Sigler 1951), where
W = weight in grams, L = fork length in
cm, and a and b are constants. A log transfor-
mation of W produces a linear equation. The
constants a and b are calculated by the meth-
od of least squares.

Validity of the scale method was deter-
mined by criteria suggested by Van Oosten
(1923, 1929, 1944) and Hile (1941). To avoid
possible bias, scales were first read without
knowledge of the size of the fish. The scales
were read at least three times. Further
checks for accuracy of age assignment includ-
ed comparisons with known age and tagged
fish, Peterson's method, and use of year
marks on other bony parts. All scales were
examined with an Eberbach microprojector
at a magnification of SOX.

The body-scale relationship was calculated
according to Tesch (1971). The condition fac-
tor K = WxlOVL^ was calculated according
to Carlander (1969), where W = weight in
grams and L = fork length in mm. Calcu-
lations were accomplished using an age-
growth computer program (Nelson 1976).

Creel census information was collected
from January 1977 to April 1978. Four week-
days and six weekend days were randomly se-
lected each month for censusing, with holi-
days treated as weekend days. On each
selected day a check station was manned on
the principal highway leading to Pyramid
Lake and three aerial counts were made.
Check stations were in operation from noon
until dark, where all pertinent information
was collected from fishermen. Aerial surveys
were conducted by dividing a day into three
equal time segments and an aerial count was
made at the midpoint of each segment (Fig.
3). Inclement weather caused cancellation of
5 percent of the flights (Kennedy 1978).
Shore fishermen and boats were counted on
each flight, with the number of boat fish-
ermen obtained by multiplying the number
of boats by the average number of fishermen
per boat on the day of the count (Johnson
and Wroblewski 1962). When less than 10
boats were checked, the yearly mean number

Great Basin Naturalist

Vol. 43, No. 1

Needles

Hells Kitchen

Anderson Bay

''aREAN- Dago Bay

EIGHT s^

Depth contours (m]

To Reno

To Nixon

Truckee River

Fig. 3. Lake areas used for creel census data collec-
tion and location of creel check stations.

of fishermen per boat was used. Rate of suc-
cess, effort, and harvest were calculated by
computer program (David Wheaton, pers.
comm. 1977). All calculations were expanded
to a 30-day month. Mean lengths and weights
of fish caught were also calculated.

Food habits were determined by examining
the stomachs of five fish per size group from
each net catch. Food habit analyses were
conducted by percent of frequency of occur-
rence and percent of total volume.

Fecundities were determined by actual egg
counts (Kucera and Kennedy 1977). Criteria
described by Nikolsky (1963) were used for
determining stage of maturity; only ripe fe-
males and fresh ovaries were used for fecun-
dity studies. Linear and logio (Y+1) regres-
sions between fecundity, fork length, weight,
age, ovum diameter, ovary weight, and net
weight (body weight minus ovary weight)
were used to examine the interrelationships
between these variables.

Age and Growth

Appearance of Scales and
Formation of Annuli

Fork length at the time of scale formation
for hatchery-reared Lahontan cutthroat plan-
ted in Pyramid Lake is 25.8 mm. Calhoun

(1942) lists 40 mm (FL) as the size of Lahon-
tan cutthroat from Blue Lake, California, at
time of scale formation. Yellowstone cut-
throat develop scales when they are between
40.3 mm and 42.8 mm (FL) (Brown and Bail-
ey 1952). Laakso and Cope (1956) report 39.3
mm (FL) as the size of cutthroat trout at the
time of scale formation. Cutthroat trout sam-
pled in Montana had formed scales at 41.2 to
63.2 mm (FL) (Brown and Bailey 1952). Ir-
ving (1953) reports 23.9 mm (FL) for cut-
throat trout in Henry's Lake, Idaho. Rob-
ertson (1947) found considerable variation in
the size of cutthroat trout at the time of scale
formation.

Nearly all scales examined from Pyramid
Lake fish showed early growth patterns char-
acteristic of hatchery rearing. Scales also
showed crowded circuli beginning in late
September 1975 and 1976 and in early No-
vember 1977. Nearly all scales aged showed
winter bands of thin and closely spaced cir-
culi. Summer growth bands appear as thick
and widely spaced circuli. The beginning of
growth, the first appearance of summer
banding, is assumed to correspond with for-
mation of the annulus. The period of annulus
formation extended from about February
through May, peaking in late April 1976 and
late March 1977.

All annuli were readily visible. Check
marks that appeared throughout all fields
were prominent during the first year's
growth for the majority of fish examined.
These stress conditions that resulted in
growth interruptions existed during the first
(0 age) year for fish from the National Fish
Hatchery (Lahontan National Fish Hatchery
personnel, pers. comm. 1976). This situation
presumably does not occur every year.

Seasonal Growth

Our analyses of age and growth for Lahon-
tan cutthroat trout from Pyramid Lake is
based on scale samples from 676 specimens
taken almost exclusively with nets from No-
vember 1975 through November 1977. The
general shapes of the 1976 and 1977 growth
curves (Fig. 4) were the same, but during
1977 growth was more rapid and extended
over a longer period of time than for com-
parably aged fish in 1976. Increments of

January 1983

SiGLER ET AL.: LaHONTAN CuTTHROAT TrOUT

growth declined with increasing age of fish in
1976, but increased in 1977. Growth in-
creased sharply in spring, slowed in late sum-
mer, and ceased during fall and winter.

Annual Growth

Annual growth (back-calculated lengths)
values were derived from the body length-
scale radius relationship FL = A -I- B(SR);
FL = fork length in mm and SR = anterior
scale radius (Table 1). Body-scale regression
equations, based on data collected over the
entire study, were used to calculate the
lengths. The results for 676 fish are: for fe-
males FL = 155.881 + 3.5364 (SR), for
males FL = 79.176 + 4.2599 (SR), for in-
determinates FL = 112.872 + 4.0834 (SR),
and for combined FL = 132.952 + 3.8079
(SR).

Young-of-the-year Lahontan cutthroat
trout sampled from Lahontan National Fish
Hatchery averaged 152 mm in length at age
eight months. By the end of Year I, hatchery
trout are approximately 203 mm FL (Lahon-
tan National Fish Hatchery personnel, pers.
comm. 1976). These data demonstrate that
the back-calculated lengths for age I Pyramid
Lake trout are accurate.

Growth in length is nearly isometric from
the end of the first through the seventh years
of life. Variation by sex is evident in the
growth rates of certain age groups. Annual
increments of growth in length for males are
greater than for females from age II on. Ac-
cording to Irving (1953), male trout from
Henry's Lake, Idaho, grow faster than fe-
males, and Bulkley (1961) reports male trout
outlive females. Pyramid Lake Lahontan cut-
throat trout appear to follow these patterns.
Others have reported no difference between
the sexes in growth rates (Drummond 1966,
Snyder and Tanner 1960).

The oldest male and female aged from
Pyramid Lake were in their seventh and sixth
year, respectively. This longevity is some-
what less than historical data. Sumner (1939)
found the oldest age groups of trout in Pyra-
mid Lake were the seven- to nine-year-olds.
Studies in smaller high altitude lakes. Upper
Blue Lake (Calhoun 1944), and Topaz Lake
(Johnson 1958), indicate that few trout live
past their sixth year (Table 2).

500 ■

J. 300

i£ 200

100 ■

AgeV

-il91_
Days

= 273.

Days

1976

1977

AJ AOD FA JAOD
Month

Fig. 4. Seasonal growth curves, 1976 and 1977, for
Lahontan cutthroat trout age groups I through V. Fish
were collected from Pyramid Lake, Nevada, from No-
vember 1975 through November 1977. The graph ab-
scissa is divided into bimonthly intervals beginning with
April.

Lahontan cutthroat trout greater than 787
mm were not sampled with our nets and thus
do not appear in our age and growth studies;
however, larger ones were taken by anglers.
In the 1977 creel sample, which was about
three times that of the net sample, 22.5 per-
cent of the 1916 trout exceeded 600 mm in
length; the longest one was 990 mm FL. The
average size was 505 mm (S.D. = 107.0). Ei-
ther these large fish grew faster than the av-
erage in our studies, or they were older than
the maximum ages of our net-caught fish. In
April 1976, an Indian angler, Ralston Fill-
more, captured a 12.7 kg Lahontan cutthroat
trout, the largest recorded since December
1925, when another Indian, John Skim-
merhorn, caught an 18.6 kg trout, the world
record cutthroat (Wheeler 1974). There are
numerous unconfirmed reports of ones larger
than this being marketed around the turn of
the century.

LeCren (1951) states the length-weight
relationship equation, in addition to provid-
ing a method of converting length to weight,
also indicates taxonomic differences and
events in the life history. The value of the
constant "B" will equal 3.0 where growth is
symmetrical or isometrical (Ricker 1971).
Values less than 3.0 demonstrate linear
growth is taking place faster than growth in

Great Basin Naturalist

Vol. 43, No. 1

weight. Values greater than 3.0 demonstrate
the reverse; both are allometric growth.

The length-weight equations calculated for
Lahontan cutthroat trout show growth
slightly exceeds the cubic relationship; this
represents allometric growth. We combined
all years, drawing on data from 561 trout.
Sizes ranged from 189 mm (36 g) to 787 mm
(6163 g) (Table 3, Fig. 5). Tesch (1971) notes
allometric growth in some salmonid stocks.

The length-weight curve of the Lahontan
cutthroat trout shows the importance of for-
age fish. In Pyramid Lake cutthroat trout
weight gains tend to exceed the increases in
length when the diet shifts from invertebrates
to fish. They are in their third year and sim-
ilar to cutthroat trout from Independence
Lake, California (Lea 1963). Lea states the
rate of growth for the Independence Lake
cutthroat trout population is only slightly

Table 1. Summary of the mean calculated fork lengths and increments of growth for Lahontan cutthroat trout
collected from Pyramid Lake, Nevada, from November 1975 through November 1976.

Age group

Number
of fish

Calculated fork lengths (mm) at end of each year of life
2 3 4 5 6

(Female)

III

42
68
80
25
23
15

238
236
233
236
236
233

306
305
304
303
306

374
376

374
374

442
442
447

500
508

580

Grand average
Increments of growth
Number of fish

235
235
253

305

211

374

143

443
68
63

503
59

580
72
15

(Male)

III

VII

178

168

255

169

253

341

171

254

344

439

168

254

339

437

516

167

250

341

439

518

165

250

345

435

521

588
575

664

Grand average
Increments of growth
Number of fish

169
169
113

253

109

342
89
95

438
96
69

517
79
44

585
66
20

664

(Indeterminate)

217

206

286

III

110

203

285

359

202

281

353

416

200

274

343

407

475

197

264

334

407

471

VII

199

274

335

384

460

(Combined)

225

133

219

294

III

216

293

365

121

217

290

362

429

215

287

358

429

494

213

286

360

440

506

VII

212

286

362

433

508

536
538

574
565

578

Grand average

204

282

353

412

474

537

578

Increments of growth

204

Number of fish

310

284

233

123

629

Grand average
Increments of growth
Number of fish

217
217
676

291

75
605

362

72
472

431

256

499

135

573
66
48

629

January 1983

SiGLER ET AL.: LaHONTAN CuTTHROAT TrOUT

curvilinear until age III, at which time the
relative weight increase accelerates greatly.
Lea also reports that, for Independence Lake
cutthroat trout less than 225 mm, forage fish
are of minor significance, but for those over
300 mm, fish become the major forage item.
Hazzard and Madsen (1933) report cutthroat
trout from Jackson Lake, Wyoming, also
show a definite change in diet from Crustacea
to fish at a length of approximately 300 mm.

The condition factor (K = W X 10 VL^) is
used as an index of well-being or relative ro-
bustness. The average K of 561 Pyramid
Lake Lahontan cutthroat trout, sexes and age
groups combined, was 1.00. A slight sexual
dimorphism is noted for condition factor,
with males having a slightly higher K value
than females (Table 4). This is in agreement
with results from other studies. Fleener
(1952) and Madsen (1940) also report higher
K values for male over female cutthroat
trout. However, the extent of the sexual di-
morphism may vary with fish size and season;

i.e., gravid females weigh significantly more
just before spawning season than after. Thus
the K of females is more variable than males
on a seasonal basis.

A direct relationship between size and
condition factor of Pyramid Lake Lahontan
cutthroat trout is evident (Table 5). Lea
(1963) reports a trend of increasing condition
factor with increasing length for Independ-
ence Lake Lahontan cutthroat trout. Fleener
(1952), however, states condition factor de-
creases with length for cutthroat trout from
Beaver Creek, a small tributary of the Logan
River, Utah. Irving (1953) says size, age, and
sex are not related to condition factor for
Henry's Lake cutthroat trout. It seems logical
that condition factor would be directly re-
lated to fish size in lake environments where
large fish have a predatory advantage. This
situation occurs in Pyramid Lake since, at the
critical size of about 300 mm, Lahontan cut-
throat trout are able to utilize the huge for-
age base of tui chubs.

Table 2. Growth of cutthroat trout from 14 Western lakes.

Calculated mean

total length

(mm) and increments at each

annulus

Location

Number

III

VII

Pyramid Lake, NV^

676

217

291

362

431

499

573

629

(217)

(75)

(72)

(70)

(66)

(64)

Lower No Name Lake, WY

102

145

190

221

231

(Robertson 1947)

(102)

(43)

(45)

(31)

(10)

Priest Lake, ID

135

211

287

348

371

(Bjomn 1957)

(81)

(54)

(76)

(61)

(23)

Upper Priest Lake, ID

142

216

292

338

391

(Bjomn 1957)

(94)

(48)

(74)

(76)

(46)

(53)

White Rock and Ted's Lake,

UT 22

130

185

201

221

(Sigler and Low 1950)

(130)

(55)

(16)

(20)

Granby Reservoir, CO

109

196

251

290

(Finnell 1966)

(109)

(87)

(55)

(39)

Yellowstone Lake, WY

5057

130

224

312

394

442

486

(Bulkley 1961)

(46)

(84)

(94)

(88)

(82)

(48)

(44)

Montana Lake, MT

2158

163

241

307

384

(Peters 1964)

(76)

(87)

(78)

(66)

(77)

Island Lake, UT

157

211

249

300

343

(Sigler and Low 1950)

(157)

(54)

(38)

(54)

(43)

Thompson Lake, MT

130

198

262

318

(Echo 1955)

(130)

(68)

(64)

(56)

Upper No Name Lake, WY

112

178

274

381

421

478

(Robertson 1947)

(112)

(66)

(96)

(107)

(40)

(57)

Blue Lake, CA

419

180

307

378

361

(Calhoun 1942)

(66)

(114)

(127)

(71)

Heenan Lake, CA

117

216

330

445

(Calhoun 1942)

(97)

(119)

(114)

(115)

Henry's Lake, ID

356

170

325

437

503

551

594

(Irving 1953)

(170)

(155)

(112)

(66)

(48)

"Fork lengths converted to total length by factors of 1.07 (189 mm - 490 mm), 1.05 (500 mm - 590 mm), and 1.03 (> 590 mm).

Great Basin Naturalist

Vol. 43, No. 1

6163

4938 +

3712 +

? 2487

Graph symbol

No. points represented

W= .0027L33271*
n = 561

• 23456 789 A BC D
1 2 3 4 5 6 7 8 9 10 11 12 13

Age
VII

. Age
VI

Age
V

Age
IV

1261 +

Age

/..•■

•4 3 » 2

J 2 . ».

-Age

Age I

+ 2*26S4*3*

189*

2 . .3«'5
I /3. .

> ••2*32 •42**

• ••7847 •*
il 2 I * 2^23489664222 •

, I •37AC6943 2 •

I 4 5638 37087^ 233 ••• 2

44494S7S732I^^2* •
26528848A94522** •

309"

428

548

677

787

Fork length (mm)

Fig. 5. Length-weight relationship of combined data for Lahontan cutthroat trout from Pyramid Lake, Nevada,
November 1975 through November 1977, with mean length-weight values by age groups. FL in cm.

Conversion Factors

Factors for converting TL to FL and SL to
FL for cutthroat trout from Pyramid Lake,
Nevada are:

TL = 1.07 FL (189 - 490 mm)
TL = 1.05 FL (500 - 590 mm)
TL = 1.03FL(>.590mm)
SL = .888 FL (189 - 300 mm)
SL = .893 FL (301 - 500 mm)
SL = .895FL(>501 mm)
FL = .935 TL (202 - 524 mm)
FL = .953 TL (525 - 620 mm)
FL = .970TL(>608mm)
FL = 1.13 SL (168 - 266 mm)
FL = 1.12 SL (268 -447 mm)
FL = 1.12SL(>447mm)

The ratios vary with size, necessitating more
than one set of conversion factors. Con-
version factors for cutthroat trout have been
reviewed by Cope (1953).

Hatchery-reared Lahontan Cutthroat Trout,

Length-weight Relationships and

Condition Factors

In 1976, 612 fingerlings were taken from
the Lahontan National Fish Hatchery, Gard-
nerville, Nevada, to determine length-weight
relationships. The fingerlings represent wild
Summit Lake stock ranging in fojk length
from 57.9 mm to 125 mm (Table 6) and La-
hontan fifth-generation domestic stock, origi-
nally from Summit Lake, ranging in fork
length from 40 mm to 250 mm (Table 7). The
length-weight relationships are calculated as:
Summit Lake brood W = .00001L2-8749

Lahontan brood W = .000007L30588

The K-factors for domestic stock range
from 1.01 to 1.21, the exponent indicating
slightly faster growth in weight than length.

January 1983

SiGLER ET AL.: LaHONTAN CuTTHROAT TrOUT

The K-factors for wild stock show sUghtly de-
creasing trends; the exponent is less than 3.0,
indicating these fish are getting slimmer as
they grow in length.

The Lahontan brood attain greater weight
per length than do Summit Lake brood. The
weight differences can be attributed to brood
spawned from wild stock being more active
(wild) and domestic stock being more passive
(Lahontan National Fish Hatchery Manager,
Charles R. Messier, pers. comm. 1976).

Mortality and Morbidity

Various factors may cause mortality of
Pyramid Lake's trout population, including
angling, death during stocking of hatchery
recruits, predation on juveniles, spawning-re-
lated deaths of adults, and disease. When
other decimating factors are inoperative or
eliminated, senility must ultimately cause
death. Among 676 Lahontan cutthroat trout
sampled from Pyramid Lake during this
study, no females and only four males
reached age VII. Sumner's (1939) data in-
dicates very few of the original population of
Lahontan cutthroat trout lived beyond eight
years, although a few may have lived to be
11.

Chemical constituents of the aquatic habi-
tat are rarely neutral in their effects on the
biota. Toxic substances often first express
themselves as growth suppressants, reproduc-
tive inhibitors, increased vulnerability to dis-
ease, or destroyers of the most sensitive link
in the food chain. Increased levels of TDS
could be detrimental to the Pyramid Lake

fishery. Walker Lake's now extinct Sacra-
mento perch population reached its limit of
"alkalinity" tolerance when it could no long-
er reproduce in the early 1950s. At that time,
the total alkalinity was approximately 2500
mg/1 as HCO3 (Cooper 1978). In 1952, the
TDS of Walker Lake was 6790 mg/1 (Koch
et al. 1979). Rainbow trout are stressed by
and grow poorly in Pyramid Lake water;
they do not survive in more concentrated al-
kaline waters such as Walker Lake (Knoll et
al. 1979) and Omak Lake (Paul A. Kucera,
pers. comm. 1982). Taylor (1972) notes that
carbonate and bicarbonate salts are more tox-
ic to Lahontan cutthroat trout at elevated
TDS levels than sodium chloride alone. Ele-
vated temperatures may have a synergistic
effect on salinity toxicity and vice versa
(Vigg and Koch 1980).

Mortalities range widely among fish cap-
tured and released. Hooking mortality of
lure-caught cutthroat trout in Yellowstone
Lake was relatively low (<6.5 percent);
however, the combination of natural baits
and high water temperature resulted in sig-
nificantly higher mortality (Mamell 1969,
Marnell and Hunsaker 1970). In Pyramid
Lake, where fish <483 mm TL are illegal (as
of 1 July 1982), all types of artificial lures are
used. The losses from hook and release in
Pyramid Lake have not been established. Le-
gal sport fishing removed < 20,000 fish in
1977, not a seriously decimating factor for
the Pyramid Lake population.

Infectious disease is a potential threat to
wild and cultured fish alike. Rational man-

Table 3. Length-weight relationships (linear and curvilinear) for Lahontan cutthroat trout from Pyramid Lake,
Nevada, from November 1975 through November 1977.*

Class

Log-log transformation
(linear)

Exponential
(curvilinear)

(Female)

(Male)

(Indeterminate)

(Combined)

logioW = -2.6218 + 3.3754 logioL
(i^ = .95, n = 224)
F = 602.4

logioW = -2.8052 -I- 3.4778 logioL
(1^ = .94, n = 73)
F = 1089.9

logioW = -2.3690 -I- 3.2023 logioL
(r2 = .92, n = 262)
F = 3017.2

logioW = -2.5531 + 3.3271 logjoL
(i^ = .94, n = 561)
F = 9293.3

W = .0023L3-3754

W = .0016L3'«778

W = .0042L3-2023

W = .0027L3-3271

^Equations were calculated using fork length in centimeters.

Great Basin Naturalist

Vol. 43, No. 1

agement and utilization of any fisheries must
incorporate a realistic understanding of the
serious pathogens extant in the system. A fish
population undergoing environmental deg-
radation is subjected to environmental factors
that may predispose the population to dis-
ease. Various interstate and international
regulations have been formulated to restrict
movement of serious fish pathogens.

PLITE has a program to determine the
presence of pathogens in Pyramid Lake
trout. Particular attention is given to those
pathogens included in restrictive lists. A
series of inspections begun in 1976 followed
proper procedure as specified by the Fish
Health Section of the American Fisheries So-
ciety and The Colorado River Wildlife Coun-
cil-Fish Disease Policy (American Fisheries
Society: Fish Health Section 1975), Con-
sultation and confirmation of procedure were
reviewed by Ron Goede, fish pathologist of
the Utah Division of Wildlife Resources, and
by Dennis E. Anderson, U.S. Fish and Wild-
life Service, Fort Morgan, Colorado.

The inspections in 1977 included the fol-
lowing diseases and/or the pathogens in-
ducing the diseases:

Viral: IPN (Infectious pancreatic necrosis) (1976 &

1977)
IHN (Infectious hematopoietic necrosis)
VHS (Viral hemorrhagic septicemia)

Bacterial: Bacterial kidney disease {Renebacteriiirn
salmoninarum)
Enteric redmouth {Yersinia ruckerii)
Funmculosis {Aeromonas sahnonicida)

Parasitic: Whirling disease {Myxosoma cerebralis)
Blood fluke {Sanguinicola sp.)

Pyramid Lake and lower Truckee River
fish populations were sampled by hook and
line, gill net, and electrofishing. Fish were
also collected from the Marble Bluff fishway
and the Dunn Hatchery, Sutcliffe, Nevada.

The inspection in 1976 was conducted by
Biometrics Inc., Tacoma, Washington. In

1977, the Fish Disease Control Center, U.S.
Fish and Wildlife Service, Fort Morgan, Col-
orado, conducted inspections.

Attribute sampling for IPN in 1976 pre-
sumed a 5 percent carrier incidence and
achieved 95 percent confidence limits. No
evidence of any pathogen was detected in
1976 (Ferjancic 1976). Sampling in 1977 pre-
sumed a carrier incidence of 2 percent and
achieved 95 percent confidence. Inspections
conducted in 1977 (Ruger 1977) detected no
evidence of IPN, IHN, VHS, bacterial kidney
disease, enteric redmouth, furunculosis,
whirling disease, or blood fluke, except 7 of
235 adults sampled at the Marble Bluff fish-
way were positive for furunculosis.

Sample sizes were sufficiently large to ex-
tend confidence beyond original required
sampling presumption. Regulation and pro-
tocol require assumption of a 2 percent car-
rier incidence for 95 percent confidence in
wild populations. Sample sizes in this study
are sufficient at the 1 percent carrier in-
cidence to permit 95 percent confidence in
detecting IPN and at the 2 percent carrier in-
cidence to permit 95 percent confidence in
detecting all other listed diseases.

Food and Feeding Habits

Lahontan cutthroat trout in Pyramid Lake
are largely piscivorous after they reach a size
of approximately 300 mm. They then feed al-
most exclusively on tui chubs, but they may
feed opportunistically on other fish and they
feed to some extent on aquatic insects. Small
trout feed on zooplankton and benthic in-
vertebrates. From January through Decem-
ber 1976, 192 Lahontan cutthroat trout were
examined for food habits; 35 had not recently
fed. The highest percentage of the 35 non-
feeders occurred during the winter and early
spring months when v/ater temperatures and.

Table 4. Coefficient of condition for Lahontan cut- Table 5. Coefficient of condition based on increasing

throat trout from Pyramid Lake, Nevada, fork length in
mm, November 1975 through November 1977.

fork length (mm) for Lahontan cutthroat trout from
Pyramid Lake, Nevada, November 1975 through No-
vember 1977.

Male

74
1.08
.634-1.416

Female

225
1.03
.857-1.212

Sexes
combined*

562
1.00
.634-1.416

Mean

Range

Fork length (mm)

150-350
351-550
551-700

Number of fish

200

273
89

(K)

.831

.994

1.160

^Includes fish in which sex was undetermined.

January 1983

SiGLER ET AL.: LaHONTAN CuTTHROAT TrOUT

therefore, trout metabolism and feeding ac-
tivity were low.

The piscivorous nature of Lahontan cut-
throat trout was predictable. Fish, the most
frequent food item, was eaten by 62.4 per-
cent of the trout (Table 8). Fish also account-
ed for the largest volume of food (84.5 per-
cent). Snyder (1917) found adults in lakes
feed largely on minnows, with one fish from
Pyramid Lake described as containing three
large minnows. In Johnson's (1958) food anal-
yses of 20 Pyramid Lake cutthroat trout, fish
were dominant. Insects, zooplankton, and
amphipods appear in descending order of im-
portance. Invertebrates rather than fish are
the major source of food for Lahontan cut-
throat trout in two Sierra lakes, presumably
because the trout occur in different areas of
the lake than other species of fish (Calhoun
1942).

A diet succession from invertebrates to fish
is apparent for Lahontan cutthroat trout as
they increase in size (Table 9). Invertebrates
make up 51.2 percent and fish 38.3 percent
of the volume of food eaten by trout 198-300
mm FL. The volume of invertebrates eaten
decreases with increasing trout size. Larger
trout, 300-399 mm, consume 32.8 percent in-
vertebrates and 60.8 percent fish. This is also
true for Utah cutthroat trout (Sigler 1962,
Sigler and Miller 1963).

Chironomids, the second most important
food item, are consumed by 41.4 percent of
the trout, but account for only 4.0 percent of

the volume. Chironomid pupae are eaten
nearly twice as frequently as larvae. This is
also true for Lahontan cutthroat in Blue
Lake, California (Calhoun 1944) and sea-
sonally in Omak Lake, Washington (Paul A.
Kucera, unpubl. data 1981). Platts (1959b) re-
ports chironomidae pupae are the most im-
portant forage item for cutthroat trout in
Strawberry Reservoir, Utah.

The remaining food items in the cutthroat
diet were of relatively minor importance
compared to consumption of fish and chi-
ronomids. Some of the items, however, can
be of significant value seasonally or during
certain life stages, such as zooplankton and
smaller invertebrates for young-of-the-year
trout. Other food items consumed in order of
percent of frequency of occurrence were:
terrestrial insects (10.2); amphipods, both
Hyallela and Gammarus (9); algae (7.6); zoo-
plankton (7.6); bottom substrate (4.5); He-
mipterans (3.2); odonates (1.9); vascular
plants (1.9); coleopterans (.6); and hydracari-
nads (.6).

Reproduction

The Lahontan cutthroat trout spawning
migration into the Truckee River and Marble
Bluff fishway peaked in April and May of
1976 and 1977 at river water temperatures
varying from 8 to 16 C (Fig. 6). Although
only 563 fish were captured in 1976, the run
was reported to be the largest in recent years

Table 6. Expected Lahontan cutthroat trout measurements (FL) based on measurements of 200 Lahontan cut-
throat trout, Summit Lake brood. Lahontan National Fish Hatchery, Gardnerville, Nevada, 1976.

Length (L)

Weight (W)

Grams per

"K"

Fish per

in inches

in millimeters

in grams

centimeter

factor

kilogram

2.16

1.66

.30

.99

602.94

2.36

2.16

.36

.99

463.37

2.55

2.64

.40

.96

379.11

2.75

3.60

.51

1.04

278.02

2.95

4.56

.60

1.08

219.49

3.14

4.88

.61

.95

205.10

3.34

6.21

.73

1.01

161.16

3.54

7.17

.79

.98

139.57

3.74

8.50

.89

.99

117.75

3.93

100

10.41

1.04

96.14

4.13

105

11.74

1.11

1.01

80.84

4.33

110

13.34

1.21

1.00

75.02

4.52

115

14.99

1.30

.98

66.76

4.72

120

16.91

1.40

.97

59.17

4.92

125

18.24

1.45

.93

54.87

5.11

130

20.16

1.55

.91

49.63

Great Basin Naturalist

Vol. 43, No. 1

(U.S. Fish and Wildlife Service pers. comm.
1976). In comparison with the number of fish
in the lake large enough to reproduce, this
number is amazingly small. The 1977 creel
census (Kennedy 1978) produced an esti-
mated sport fishing catch of < 20,000 Lahon-
tan cutthroat trout >380 mm TL. This size is
a reasonable approximation of the average
length at reproductive maturity. It is obvious
that the population of mature fish must be
considerably greater than 563, the number
that were captured. Netting did not indicate

an unusual concentration of cutthroat trout
in the lake near the Truckee River mouth at
this time. However, large numbers of trout
congregated around the Sutcliffe area where
hatchery reared cutthroat are planted. As
Ball (1955) postulates, since these planted fish
were not imprinted on an inflowing stream,
they may be milling about the area where
they were originally planted.

Limited data on the maturation of female
Lahontan cutthroat trout in Pyramid Lake
suggest that consecutive-year spawning does

Table 7. Expected Lahontan cutthroat trout measurements (FL) based on measurements of 412 Lahontan cut-
throat trout, Lahontan domestic brood. Lahontan National Fish Hatchery, Gardnerville, Nevada, 1976.

Length (L)

Weight (W)

Grams per

"K"

Fish per

in inches

in milhmeters

in grams

centimeter

factor

kilogram

1.57

.65

.16

1.01

1539.84

1.77

.93

.20

1.02

1076.23

1.96

1.29

.25

1.03

775.87

2.16

1.73

.31

1.03

578.54

2.36

2.27

.37

1.05

440.92

2.55

2.91

.44

1.05

343.94

2.75

3.67

.52

1.06

272.71

2.95

4.54

.60

1.07

220.46

3.14

5.55

.69

1.08

178.57

3.34

6.70

.78

1.09

149.39

3.54

8.00

.88

1.09

125.11

3.74

9.46

.99

1.10

105.80

3.93

100

11.09

1.10

90.24

4.13

105

12.90

1.22

1.11

77.58

4.33

110

14.90

1.35

67.15

4.52

115

17.11

1.48

1.12

58.99

4.72

120

19.52

1.62

L12

51.26

4.92

125

22.15

1.77

1.13

45.17

5.11

130

25.02

1.92

1.13

39.99

5.31

135

28.12

2.08

1.14

35.58

5.51

140

31.48

2.24

1.14

31.79

5.70

145

35.10

2.42

1.15

28.51

5.90

150

38.99

2.59

1.15

25.66

6.10

155

43.16

2.78

1.15

23.17

6.29

160

47.62

2.97

1.16

21.01

6.49

165

52.39

3.17

L17

19.09

6.69

170

57.47

3.38

Lie

17.39

6.88

175

62.88

3.59

L17

15.92

7.08

180

68.61

3.81

1.17

14.57

7.28

185

74.70

4.03

1.17

13.38

7.48

190

81.13

4.27

1.18

12.32

7.67

195

87.94

4.50

1.18

11.38

7.87

200

95.12

4.75

1.18

10.52

8.07

205

102.69

5.00

1.19

9.74

8.26

210

110.65

5.26

1.19

9.03

8.46

215

119.03

5.53

1.19

8.40

8.66

220

127.82

5.81

1.20

7.83

8.85

225

137.04

6.09

1.20

7.30

9.05

230

146.70

6.37

1.20

6.81

9.25

235

156.82

6.67

1.20

6.37

9.44

240

167.39

6.97

1.21

5.97

9.64

245

178.44

7.28

1.21

5.60

9.84

250

189.98

7.59

1.21

5.25

January 1983

SiGLER ET AL.: LaHONTAN CuTTHROAT TrOUT

not occur. Judging by ovary development,
several females of age groups IV and V col-
lected prior to the spawning season were not
going to reproduce the current year but
would the following spring. Similar situations
have been reported elsewhere. Some 1-15
percent of the cutthroat in Arnica Creek,
Yellowstone National Park, spawn each year
after reaching maturity, 10-26 percent are
alternate-year spawners, and 46 percent skip
two years (Ball and Cope 1961). Seven per-
cent of the female Lahontan cutthroat trout
in Blue Lake, California, spawned in con-
secutive years, and 10.5 percent of the origi-
nally marked females were in the run two
years later (Calhoun 1942).

Female cutthroat trout in Pyramid Lake
mature at age III or IV, when they are 352 to
484 mm FL; males mature at ages II or III,
when 299 to 445 mm FL. Lahontan cutthroat
trout in Independence Lake, California, ma-
ture at ages III or IV (Lea 1963). Lea attri-
butes the presence of small numbers of ma-
ture three-year-olds to a precocial element of
the population. Rankel (1976) reports spawn-
ing runs of Lahontan cutthroat trout from
Summit Lake, Nevada, consist mainly of

four-year-old fish. If an alternate-year spawn-
ing pattern is typical, then most female La-
hontan cutthroat trout in Pyramid Lake will
spawn a maximum of twice in their lifetime.
If they mature at age IV, they may live to
spawn only once. Some of those that mature
at age III may spawn again at age V.

The sex ratio of Lahontan cutthroat trout
in our net catches was 1 male:4.23 females
(n = 455). This is not representative of the
population in the lake. The ratio of males to
females in the spawning runs was variable,
i.e., 1.06:1 in 1976 and 1:2.35 in 1977. The
ratios of Summit Lake Lahontan cutthroat
trout spawning runs, from 1968 to 1975, var-
ied from 1:1.3 to 1:2.2 and averaged 1:1.6
males to females (Rankel 1976). Angling is
male-selective in Pyramid Lake. This is also
true in Omak Lake (Paul A. Kucera, pers.
comm. 1981).

Female Lahontan cutthroat trout spawn
after attaining an average gonadal somatic
index (percent gonad weight to total body
weight) of 11 percent. The progression in go-
nadal somatic indices, observed from October
through December 1977, indicates a steady
increase in germinal growth through the

Table 8. Food of 157 Lahontan cutthroat trout. Pyramid Lake, Nevada, 1976.

Percentage

of total

Percentage

volume

frequency

Volume

excluding

total

digested

Food item

Frequency

occurrence

Rank

volume

Rank

matter

Fish

62.42

1225.7

82.89

84.50

Chironomidae

14.01

11.6

0.78

0.80

larvae

Chironomidae

27.39

46.8

3.16

3.23

pupae

Odonata

1.91

2.7

0.18

0.19

Coleoptera

0.64

0.7

Trace*

Hemiptera

3.18

1.0

Trace*

Terrestrial

10.19

19.6

1.33

1.35

insects

Amphipoda

8.92

9.2

0.62

0.63

Ostracoda

1.91

1.4

Trace*

0.10

Zooplankton

7.64

10.9

0.74

0.75

Algae

7.64

93.1

6.30

6.42

Vascular

1.91

2.0

0.14

plants

Hydracarina

0.64

Trace*

Bottom substrate

4.46

25.8

1.74

1.78

Digested matter

11.46

28.2

1.91

—

Total

1478.7

99.98

99.89

*Trace = less than 0.1

Great Basin Naturalist

Vol. 43, No. 1

winter. The largest increase involves a shift
from an average value of 5.9 percent in No-
vember to 8.1 percent in December.

The diameter of mature eggs of Pyramid
Lake Lahontan cutthroat trout range from
2.76 to 5.09 mm and average 4.51 mm.

Monthly progression in egg sizes from Octo-
ber through December 1977 indicated a con-
sistent, gradual increase in ovum size. The
mean egg diameter in December was 4.11
mm, with some eggs as large as 4.60 mm.
Some females may be able to spawn in

Table 9. Percentage of total volume and frequency of occurrence of food items consumed by Lahontan cutthroat
trout from Pyramid Lake, Nevada, in relation to size. Trout were captured from January through December 1976
with bottom-set gill nets.

Food item

Volume

Percent
of total
volume

Frequency

by
occurrence

Frequency of
occurrence
by percent

Size group 99-198 mm (n =

= 1)

100.0

Fish

1.0

100.0

Fork length = 184 mm

Weight = 54

Size group 198-300 mm (n = 22)

Fish

Benthic invertebrates

Terrestrial insects

Zooplankton

Digested matter

Vascular plants

12.1

38.3

50.0

12.2

38.6

54.5

4.0

12.7

9.5

0.5

1.6

4.8

2.0

6.3

9.5

0.8

2.5

Mean fork length = 251 mm
Range = 203-297 mm

Mean weight = 136 g
Range = 54-240 g

Size group 300-399 mm (n = 63)

Fish

Benthic invertebrates
Terrestrial insects
Zooplankton
Periphyton
Vascular plants
Bottom substrate
Digested matter

67.1

60.8

66.7

27.6

25.0

39.7

7.3

6.6

11.1

1.3

1.2

3.2

2.6

2.4

3.2

1.0

0.9

1.6

Trace*

Trace^

1.6

3.4

3.1

4.8

Mean fork length = 356 mm
Range = 305-398 mm

Mean weight = 408 g
Range = 218-730 g

Size group 399-498 mm (n = 60)

Fish

Benthic invertebrates
Terrestrial insects
Zooplankton
Periphyton
Vascular plants
Bottom substrate
Digested matter

530.7

77.8

58.3

30.0

4.4

41.7

8.3

1.2

13.3

9.1

1.3

13.3

57.5

8.4

15.0

0.2

Trace^

1.7

23.5

3.4

10.0

22.8

3.3

15.0

Mean fork length = 434 mm
Range = 400-498 mm

Mean weight = 789 g
Range = 476-1496 g

Size group 500 + mm (n = 11)

Fish

Benthic invertebrates

Periphyton

Bottom substrate

615.8

94.1

90.9

3.6

0.5

9.1

33.0

5.0

9.1

2.3

0.4

9.1

Mean fork length = 531 mm
Range = 500-596 mm

Mean weight = 1741 g
Range = 1161-2753 g

*Trace = less than 0.1

January 1983

SiGLER ET AL.: LaHONTAN CuTTHROAT TroUT

J F
1977
Month and year

Fig. 6. Mean monthly Pyramid Lake cutthroat trout catches (15 gill net sets per month) from November 1976
through December 1977, in comparison with the spawning runs at the Marble Bluff fish passage facility.

December or January. For example, as early
as January, 1981 and 1982, mature cutthroat
were running up a small stream south of Sut-
cliffe, Nevada, artificially produced by
pumping about .057 m^/sec of lake water
through rearing ponds and allowing it to run
back into the lake. Egg size is positively cor-
related with fish length (r = .48; P<0.05)
and weight (r = .51; P<0.05), indicating
that egg size increases with fish size.

The number of eggs produced by a female
Lahontan cutthroat trout is significantly re-
lated to age, fork length, and weight
(P<0.05). The above relationships have sig-
nificant linear fits with and without logio
transformation of data. Fork length provides
the best predictor for fecundity, followed by
weight and age. Increases in fecundity corre-
spond to increases in length and weight. The
log 10 equation for fork length and fecundity is
logioF = 2.83 (logioFL) - 4.16, and for
weight and fecundity is logioF = .81
(logioWT) + .92. These fish range in fecun-

dity from 1241 to 7963 and average 3815
eggs per female. Lea (1963) reports fecun-
dities of Lahontan cutthroat trout in Inde-
pendence Lake, California, vary from 669 to
2080 eggs and average 1191 eggs per female.

HUbitat and Ecology

The most characteristic feature of the
Pyramid Lake environment is the high level
of salts; TDS concentration was about 5350
mg/1 during 1976-1977 (Sigler and Kennedy
1978). Although sodium chloride is the domi-
nant salt (over 70 percent), alkalinity may be
the most important constituent. The mean
pH is 9.2. The historic increase in TDS levels
was associated with the decline in lake level
(Fig. 7). Since the baseload of salts is relative-
ly constant, TDS varies inversely with the
volume of the lake. Various studies, although
preliminary in nature, have demonstrated
that NaCl is relatively irmocuous, but alka-
linity (HCO3 + CO3J is toxic to sahnonids

Great Basin Naturalist

Vol. 43, No. 1

1186

1183 ^

1180

1176 -

1173 -

1170 -

-^ 1167 -
a>

I 1164 ^

1161 -

1158 -

1155 -

1152

_l 1 I 1 1 L.

X
I \
X \
/ \

x->

\" \

• • • • TDS

X — X Fragmentary record

More complete record
(September altitudes are shown

5500

5000

4500 H

- 4000

- 3500

3000

— I 1 1 1 1 1 1 1 1 1 I

1860 1870 1880 1890 1900 1910 1920 1930 1940 1950 1960 1970 1980

Year

Fig. 7. Water level and total dissolved solids fluctuations in Pyramid Lake, Nevada, 1867-1979 (From Galat et al.
1981).

(Beatty 1959, Mitchum 1960, Taylor 1972,
Knoll et al. 1979).

The five major species of fish in Pyramid
Lake, in order of relative abundance are: tui
chub, Tahoe sucker, Lahontan cutthroat
trout, cui-ui (Chasmistes cujus), and Sacra-
mento perch {Archoplites interruptus). Al-
though numerous species have been in-
troduced, the current species composition is
almost exclusively represented by the original
fish species, the only exception being the Sac-
ramento perch (Vigg 1981). This fact is prob-
ably due to the harsh environmental condi-
tions of Pyramid Lake, specifically the TDS
levels. In contrast, exotic fish introductions
have nearly extirpated the native fish fauna
of oligotrophic (low TDS) Lake Tahoe at the
upper end of the Truckee River (Miller
1951).

The maximum surface (0-1 m) water tem-
perature in Pyramid Lake was 21.4 and

23.1 C in July 1976 and August 1977, respec-
tively. The lake is thermally stratified from
June through December; wind-generated
mixing occurs from January through May.
The thermocline forms at a depth ranging
from 16 to 22 m. The euphotic depth aver-
aged 11 m for 1976 and 1977, which resulted
in a trophogenic zone of about 4.67 km^ (Ga-
lat et al. 1981).

Surface-dissolved oxygen (DO) is above 8
mg/1, and thus not limiting to fish. Meta- and
hypolimnetic DO depletion occurs beginning
in July following stratification and algal de-
composition; maximum DO deficits occur in
the profundal zone just prior to late fall mix-
ing. Hypolimnetic DO deficits in stratified
lakes are generally associated with decompo-
sition of organic matter, which is generated
by primary production in surface water, and
gradually sink to the bottom. During Decem-
ber, the mean DO level is <4 mg/1 at depths

January 1983

SiGLER ET AL.: LaHONTAN CuTTHROAT TrOUT

>61 m and <0.2 mg/1 at depths >92 m
(Vigg 1980). In contrast to the anoxic condi-
tions, which are very hmited on a temporal
and spatial basis in Pyramid Lake, Walker
Lake exhibits extensive DO depletions that
severely restrict fish distribution (Cooper
1978, Koch et al. 1979).

Diatoms {Cyclotella sp. and Stephanodiscus
spp.) dominate the phytoplankton commu-
nity during winter; but the most abundant
chlorophyte, Crucigenia sp., attains max-
imum abundance in spring (Sigler and Ken-
nedy 1978). Blue-green algae are by far the
dominant form in Pyramid Lake, comprising
>74 percent. Nodularia spumigena is the
most abundant species; blooms begin as early
as July and may last as late as October. Tem-
poral nutrient dynamics inversely relate to
phytoplankton abundance. Following vernal
increases of algal growth, orthophosphate
and nitrate are depleted and remain at low
levels during the summer period of maximum
primary production. Silica, in addition to ni-
trate, probably limits diatom production in
Pyramid Lake (Galat et al. 1981).

Benthic macroinvertebrates, periphyton,
and zooplankton all are important energy
sources for juvenile fish in Pyramid Lake. Di-
atom domination of the periphyton commu-
nity is demonstrated by sampling with glass
slides (<99 percent). The chlorophyte,
Cladophora glomerata, was the dominant
epilithophyton in Pyramid Lake during May
and June in 1976 and 1977 (Sigler and Ken-
nedy 1978). Chironomids are the lake's most
abundant macroinvertebrates (63 percent),
followed by Oligochaetes (33 percent), which
are especially abundant in the profundal
zone (Robertson 1978). Two euryhaline am-
phipods, Gammarus lacustris and Hyallela
azteca, are associated with tufa and rocks.
The zooplankton community is composed of
five cladocerans, three copepods, and four
rotifers (Lider and Langdon 1978). The cla-
doceran, Diaptomus sicilis, is a perennial spe-
cies and the most abundant zooplankter
throughout the year.

The Lahontan cutthroat trout is the third
most numerous fish in Pyramid Lake. Com-
pared to the more abundant tui chubs and
Tahoe suckers, the trout population is numer-
ically small, about 1.3 percent (Vigg 1981).

However, the relative biomass of the Lahon-
tan cutthroat trout population is estimated at
6.4 percent. Theoretically, the biomass of a
primary piscivore such as cutthroat trout
may be as much as 20 percent of the biomass
of the fish forage (McConnell et al. 1978).
The trout population at present is far below
its theoretical maximum.

Activity of the Lahontan cutthroat trout
population is at a maximum from December
through March. Peak spawning migrations
occurred during April and May in 1976 and
1977 (Fig. 6). In 1978 the run was from
March 8 to June 13 (Wolcott 1978). The
greatest trout activity observed in our study
corresponds very closely to the historical
spawning period of the winter race of Pyra-
mid Lake Lahontan cutthroat trout (Snyder
1917). Snyder observed that the spawning
migrations of Lahontan cutthroat occurred in
two distinct periods. The larger winter run of
trout out of Pyramid and Winnemucca lakes
began following the rise in river flows Octo-
ber-December; the spawning migration ex-
tended through March. As the winter run
waned, the spring nm of the smaller, darker,
and more heavily spotted trout commenced.
This migration peaked in April and extended
to May.

The sport fishing catch was highest in win-
ter, corresponding to the high catches in the
1975-1977 net sampling program (Figure 8).
The proportion of large trout was greatest
during winter. The high level of winter activ-
ity of the Pyramid Lake population of La-
hontan cutthroat trout is apparently a mani-
festation of innate spawning-related
behavior. The larger and older spawners in
the cutthroat trout population in Yellowstone
Lake are predominant in the early part of
each spawning run, with the smaller spawn-
ers comprising the latter part of the runs
(Bulkley and Benson 1962).

The differences between the 1976-1977
Marble Bluff spawning runs and the activity
patterns of the lake population of cutthroat
trout may be explained by three factors: (1)
few Pyramid Lake Lahontan cutthroat trout
were apparently imprinted on the Truckee
River; (2) the spawning runs were composed
of a disproportionately large number of cut-
throat-rainbow hybrids that were raised (thus
imprinted) in the Truckee River watershed;

Great Basin Naturalist

Vol. 43, No. 1

-S 38-
>

-Q 36 ■
o

■S 34 ■

g- 32

T 30-

"I 28

-i 26

24 -
22-
20

u> 16

3 14

I 12-

i 10 ■

I 8

-5 6
o

i 4^

S 2

'^ 0

i ^

^;^

152

30 35

Fig. 8. The
Pyramid Lake,

DJ FMAMJ JASONDJ FMAMJ JASON
1976 1977

Month and year

proportion of large (>550 mm) cutthroat trout taken in the monthly bottom-set gill net catches in
Nevada, November 1975 through 1977.

and (3) Lahontan cutthroat trout are prob-
ably genetically programmed for winter as
well as spring spawning, but early winter
peak flows are now diminished or eliminated
by diversions. For example, during 1976
flows in the lower river peaked in March and
steadily decreased to the lowest annual level
in December (U.S. Geological Survey 1977).

Spawning cutthroat trout instinctively re-
turn to the stream in which they were born
(Ball 1955, Platts 1959a, McCleave 1967,
Jahn 1969). Their olfactory development pat-
terns indicate cutthroat trout are capable of
imprinting on a home stream odor at a very
early age (Jahn 1972). Moreover, mature cut-
throat are able to return to home streams
even when deprived of their sense of vision
or smell, indicating an inborn "compass"
homing mechanism.

Ball (1955) says, since predetermination of
spawning site is established by early life

stream association, streams can be lost as nur-
sery areas if no natural reproduction occurs,
even though mature adults are present in the
lake. Ball postulates that nonimprinted fish,
such as gravid adults, may randomly move
throughout a lake or mill about in the area
where they were planted.

On an annual basis the majority of Lahon-
tan cutthroat trout in Pyramid Lake occur at
depths <61 m (Fig. 9). Compared to other
species, its depth preference is intermediate,
and most closely associated with that of tui
chub. There is a differential seasonal depth
distribution of cutthroat trout (predator) and
tui chub (prey) in Pyramid Lake, i.e., inshore
versus 46 m (Fig. 10). Cutthroat trout appar-
ently prefer inshore areas during all seasons
except summer, when shallow water temper-
atures are high. Tui chub are generally in-
shore during spring and summer, inter-
mediate during autumn, and offshore during

January 1983

SiGLER ET AL.: LaHONTAN CuTTHROAT TroUT

Percent species composition

" -^ Tui chub

Mean catch
■■ ■■ Cutthroat trout

^ -A Tui chub

• • Cui-ui

o o Tahoe sucker

Depth (m)

Fig. 9. Percent species composition of tui chub and percent of the mean catch of cutthroat trout, tui chub, cui-ui,
and Tahoe sucker by depth in Pyramid Lake, Nevada. Data are derived from the total catch of 108 bottom gill net
sets on a quarterly basis (September, December, March, and June) during 1976-1977 (Vigg 1980).

winter. Maximum overlap of the two popu-
lations occurs during spring, the period of
maximum cutthroat trout growth. The two
species are opposite with respect to depth
distribution during winter, when trout meta-
bolism and feeding are low.

Changes in net catch/effort and benthic
depth distribution of cutthroat trout occurred
on a seasonal basis in 1976-1977 (Vigg 1978).
During late fall and winter, when the total
catch rate of cutthroat trout was about 1.5
times that of other seasons, they inhabited
predominantly inshore areas. As surface wa-
ter temperatures increase from 10 to 16 C
during late spring, the trout population

moves into cooler, deeper waters (Fig. 11).
This temperature relationship clearly illus-
trates the habitat preference of Lahontan
cutthroat trout for cooler waters than their
prey the tui chub. The lowest summer den-
sity of cutthroat trout occurs in littoral ben-
thic and inshore surface waters, and the high-
est density in benthic waters in or below the
thermocline. During the summer months,
cutthroat are well represented at depths of
20-60 m in benthic areas, while avoiding the
surface waters of the offshore limnetic zone.
From June to October the majority of the
limnetic trout are at depths of 15-28 m with
negligible ntmibers at greater depths (Vigg

Great Basin Naturalist

Vol. 43, No. 1

100
90
80
70
60
50
40
30
20
10
0

n Cutthroat trout
^ Tuichub

n J

Fall Winter Spring Summer Fall Winter Spring Sumnner Fall
(N-D) (J-M) (A-J) (J-S) (O-D) (J-M) (A-J) (JS) (OD)
1975 1976 1977

Season, month and year

Fig. 10. Percent of the total seasonal variable mesh bottom-set gill net catches of cutthroat trout and tui chub
taken at inshore (versus 46 m) sampling stations in Pyramid Lake, Nevada, from November 1975 through December
1977.

1980). As surface temperatures cool below 16
C in the fall, trout return to surface waters
and inshore areas.

The profundal zone of Pyramid Lake (>61
m), which constitutes about half of the bot-
tom area and 20 percent of the volume, is
nearly devoid of cutthroat trout in summer.
Temperatures at these depths are less than 7
C during all seasons, and oxygen is low dur-
ing the fall and early winter. Cutthroat trout
densities are slightly higher in the profundal
zone during winter, but this deep area is not
an important habitat for trout (Sigler and
Kennedy 1978).

Management

Management of any fishery should ensure
that biological, social, economic and political
values are given appropriate consideration so
as to produce maximum benefits to society
from a given stock of fish. Although the pri-
mary beneficiary of the Pyramid Lake fishery
is the Pyramid Lake Paiute Indian Tribe, so-

ciety, the secondary beneficiary, must also be
satisfied if the goal of the tribe is to be
reached. This goal, as articulated by the U.S.
Justice Department, is to produce a viable
fishery in Pyramid Lake. The above state-
ment may be assumed to be synonymous with
or an extended definition of viable fishery.

As far as can be determined, there is vir-
tually no natural reproduction of Lahontan
cutthroat trout in the Truckee River at pres-
ent. An important aspect of the management
program should be to reestablish successful
spawning runs in the Truckee River (Innis et
al. 1981). This will require rehabilitation to
stabilize stream banks and provide shade to
reduce water temperatures, installation of
fish ladders to permit spawning adults to mi-
grate upriver, fish screens to keep down-
stream migrants from entering irrigation
canals, and augmented stream flow during
critical seasons. Sufficient water will be re-
quired for adults to migrate in winter and
early spring; to keep temperatures below
13.3 C through the fry stage; and below

January 1983

SiGLER ET AL.: LaHONTAN CuTTHROAT TrOUT

Fig. 11. Percent of the catches of cutthroat trout and tui chub taken on the surface from surface and bottom gill
nets (adjusted to unit of net area) at the 23 m depth in Pyramid Lake, Nevada, from February through November
1977.

21.8 C (Vigg and Koch 1980) during juvenile
residence. Since it will take years to restore
river habitat, the Lahontan cutthroat trout
population must be sustained by stocking.

The catch rate of legal size (>380 mm TL)
Lahontan cutthroat trout in 1977 was ap-
proximately one fish per 14 hours of effort.
Under the minimum legal size of 457 mm the
catch rate was one fish per 18.9 hours (Alan
Ruger, pers. comm. 1982). Catch rates should
be increased five- to tenfold to fall within ac-
ceptable limits. This will require substantially
increased recruitment rates and the reduction
of incidental causes of mortality. Since ma-
ture fish appear to congregate in the vicinity
of stocking sites in late fall, winter, and early
spring, during the time when sport fishing is
best, some of the stocking effort should be di-
rected to the vicinity of popular fishing areas
and access points (Table 10).

Unless Lahontan cutthroat trout have been
imprinted on Truckee River water, they are
disinclined to attempt to ascend the river to
spawn. Some hatchery fish should be stocked
in the lower river to initiate a spawning run.
The trout reared in the PLITE Numana
Hatchery may be imprinted on the Truckee
River because the hatchery outfall runs into
the river. The reasons for the very substantial
spawning runs of Lahontan cutthroat trout
up the Sutcliffe outflow (>9000 in 1982) ap-
pears to contradict some long-held beliefs
and raises more questions than answers. Alan
Ruger (pers. comm. 9 June 1982) thinks the
fish are returning to hatchery odors, such as
fish feed and juvenile fish. The modified an-
nual temperature regime in the Truckee Riv-
er, due to the regulation in flow that will be
necessary to provide for all uses, must be
considered in planning for the reestablish-

Great Basin Naturalist

Vol. 43, No. 1

ment of natural reproduction in the river.
Temperatures acceptable for spawning and
incubation of eggs exist during winter and
early spring; thus successful natural repro-
duction in the river will depend in part on an
early spawning migration. Behnke (1979) sug-
gests selecting maximum-size fish at first
maturity for breeding stock. He also recom-
mends using the genetic diversity in remnant
stocks to produce the best-adapted strain to
Pyramid Lake conditions. As pointed out by
Snyder (1917), Pyramid Lake Paiute Indians
early in the century recognized two spawn-
ing runs of Lahontan cutthroat trout: one in-
volved large fish in late November, Decem-
ber, and January; the other occurred in the
spring when the smaller fish spawned. The
spring-spawning fish always faced the haz-
ards of high water temperatures, but not at
the level that exists today. Competition for
river flows is much more intense in spring
than in winter. It is generally agreed that
some remnant of the original gene pool of
Lahontan cutthroat trout persists. Therefore,

some of the present population should and
apparently do tend to spawn in winter.
When hatchery brood stock are to be used as
a source of eggs, a program of selective
breeding utilizing early-spawning fish should
be initiated. Stocking in the river and lake
should be limited to Lahontan cutthroat
trout; hybrids should not be utilized.

Benthic invertebrates are the major food
source of Lahontan cutthroat trout until they
exceed 300 mm FL. Survival of smaller
stocked fish may be limited by the avail-
ability of benthic invertebrates. The feasibil-
ity of stocking fish as large as 300-330 mm,
and their survival and costs in comparison to
the size conventionally stocked, should be
evaluated. In view of the larger number of
trout that must be stocked, introducing larger
fish could eliminate benthic invertebrate
abundance as a limiting factor and thus in-
crease survival rates of stocked fish.

Currently all trout <483 mm TL that are
landed must be released. The present catch
rate of undersized fish is much greater than

Table 10. Comparison of gill netting catch rate, surface water temperature, and trout fishing success during 1977
at Pyramid Lake, Nevada.

Gill net data

Fish per i

net

Percent

(inshore

(inshore)

Monthly

Two-month X

Monthly

Two-month X

January

33.5

February

March

April

19.5

May

22.5

June

July

10.5

August

September

21.5

October

24.5

November

44.5

December

January

February

March

January 1983

SiGLER ET AL.: LaHONTAN CuTTHROAT TroUT

those fish longer than 483 mm. The reason
for releasing a fish is the assumption that it
will survive to spawn and/ or be caught later.
This assumption should be tested, and the
size limit implemented accordingly.

The Lahontan cutthroat trout now in Pyra-
mid Lake probably is physiologically capable
of hybridizing with rainbow trout. A popu-
lation of mature rainbow trout in the middle
and upper Truckee River where Lahontan
cutthroat trout spawn would, therefore, po-
tentially threaten the maintenance of the
lake strain. In addition, brown trout residents
in the upper Truckee River will compete
with and prey on young Lahontan cutthroat
trout. Since a large part of the Truckee River
is managed by the Fish and Game Depart-
ment of California and the Nevada Wildlife
Department, the decision is theirs to imple-
ment reduction in the nonnative resident
populations of the river fish.

Nonfishing recreation on Pyramid Lake
represents approximately 500,000 hours of
use annually. This use is concentrated during

the summer months, but lasts from May
through November. Nonfishing recreationists
currently represent a significant segment of
the lake users, almost twice the use of fishing
effort (Fig. 12).

Summary

The lake form of Lahontan cutthroat trout
is the largest of all cutthroat. Its ancestors in-
vaded ancient Lake Lahontan from the Co-
lumbia River drainage about 70,000 years
BP. Before the coming of white men, the La-
hontan cutthroat trout was a staple in the
diet and an item of trade for the Paiute In-
dians of Pyramid Lake. Later, both white
men and Indians commercialized the trout
fishery in markets as far away as San Fran-
cisco. At one time the annual production
may have been as much as 454,000 kg. In
1943 the last of the Lahontan cutthroat trout
disappeared from Pyramid Lake. Very few
had been seen after 1938. Lahontan cutthroat
and other trout were stocked in the lake

Table 10 continued.

Total
catch

Fish
kept

Percent
fish
kept

Trout

per hour

Water
temperature

Total
catch
rate

Catch
rate

of fish
kept

size

surface

keeper

6.81

4381

3180

.0837

.0608

523

6.60

4993

2495

.0956

.0471

538

6.60

7266

3193

.1558

.0685

521

8.73

1926

820

.0946

.0403

529

10.28

301

101

.0458

.0154

478

16.32

439

164

.0849

.0317

437

21.28

.0073

23.12

.0130

21.21

2234

750

.3471

.1165

459

16.51

5846

1992

.3246

.1106

455

14.50

9164

3495

.3452

.1317

498

11.50
7.80
7.10

10.00

7205
4970
2997
4941

3654

2050

969

1924

51
41
32

.2250
.1211
.0755
.1513

.1141
.0499
.0244
.0589

488
556
565
482

Great Basin Naturalist

Vol. 43, No. 1

130,
120
110
100
90

Nonfishing recreation
Ibtol hsnermen

Fig. 12.

J F
1977

Month and year

Hours per month of fishing and nonfishing recreation at Pyramid Lake, Nevada, in 1977-1978.

Starting in 1950. Today, there is virtually no
natural reproduction.

Lahontan cutthroat trout in Pyramid Lake
live six to seven years. They start maturing at
age three to four, some as early as December
or as late as April, May, or June. There may
be what amoimts to two potential spawning
runs. An average-size female produces about
3815 eggs and is mature when the gonadal
somatic index reaches 11 percent. None of
the eight potential disease organisms ex-
plored in 1976-1977 were considered a haz-
ard. The fish are most active in the lake from
December through March, a time of most
fishing effort. Cutthroat <300 mm feed pri-
marily on invertebrates; after that size they
feed heavily on fish. Five species of fish con-
stitute >99 percent of the population. They
are, in order of abundance, tui chub, Tahoe
sucker, Lahontan cutthroat trout, cui-ui, and
Sacramento perch.

Pyramid Lake, entirely within the Pyramid
Lake Paiute Indian Reservation, is the termi-
nus of the Truckee River, which is its only
source of water except for a few desert show-
ers. The average annual loss to evaporation is
1,2 m. The lake is 40 km long, 6.5 to 16 km

wide, covers an area of 446.4 km^, and has a
mean depth of 59 m and a maximum of 103
m. Derby Dam, completed in 1905, effects a
transbasin diversion of part of the Truckee
River flow. The TDS of Pyramid Lake in
1977, at an altitude of 1157 m, was 5235
mg/1. The base load of TDS is reasonably
stable. Pyramid Lake stratifies into three
well-defined layers in June-July. It destra-
tifies in December-January. It is a midlevel
productivity lake. Pyramid Lake, a remnant
of Lake Lahontan, has a pH of 9.2 and is high
in carbonates and bicarbonates. Summer sur-
face temperatures are 21-23 C. There is
ample dissolved oxygen in the epilimnion and
thermocline at all times. Nodularia, a blue-
green alga, dominates much of the lake from
late summer to early fall. Diatoms dominate
the periphyton communities. Chironomids
are the most abundant macroinvertebrates.

Conclusions

The Lahontan cutthroat trout fishery in
Pyramid Lake is currently not a viable one.
The annual catch was < 20,000; the rate, one
fish for > 14 hours effort, when the minimum

January 1983

SiGLER ET AL.: LaHONTAN CuTTHROAT TrOUT

legal size was 381 mm TL. Fishing success
should be increased in the magnitude of
five-ten times. Any adverse changes in the
lake ecology may stress the fish that will in
turn make them more susceptible to disease.
The 1905 diversion of the Truckee River,
which in dry years may take most of the
flow, reduced available stream spawning area
for the cutthroat from >500 km to <62 km
of substandard stream. Derby Dam, over a
period of 25 years, doomed the historical cut-
throat fishery. Successful reproduction in the
lower river demands stable riparian habitat
and water temperatures < 13.3 C, until after
spawning-hatching-fry emergence and <21.8
C thereafter. Brood stock or wild egg-pro-
ducing fish for hatcheries should be selected
for large size at first maturity and for winter
or early spring maturing. Part of the matur-
ing fish in the lake should be imprinted on
the Truckee River. Since the effluent from
Numana Hatchery flows into the Truckee
River, it may be these trout will be im-
printed. The biological implications of the
large lam of cutthroat into the Sutcliffe flow
should be explored in depth.

Since the base load of TDS is constant in
Pyramid Lake, the concentration varies in-
versely to lake volume. Any significant in-
crease in TDS may prove harmful to key or-
ganisms in the food chain and to the trout.
The median level of productivity that Pyra-
mid Lake currently enjoys is considered more
desirable than a higher level for Pyramid
Lake Lahontan cutthroat trout.

Acknowledgments

This work was performed under Bureau of
Indian Affairs contract H50C 14209487. As-
sistance and cooperation was provided by
employees of W. F. Sigler & Associates Inc.
(WFSAI), members of the Pyramid Lake
Paiute Indian Tribe, and the U.S. Fish and
Wildlife Service, Fisheries Assistance Office,
Reno, Nevada. Denise Robertson and Roy
Whaley, formerly of WFSAI, were respon-
sible for the two sections on age and growth,
and food habits, respectively. The manuscript
was reviewed by Alan Ruger, fisheries direc-
tor. Pyramid Lake Indian Tribal Enterprises,
Sutcliffe, Nevada.

Literature Cited

American Fisheries Society: Fish Health Section.
1975. Suggested procedures for detection and
identification of certain infectious diseases of
fishes. Amer. Fish. Soc, Washington, D.C.

Ball, O. P. 1955. Some aspects of homing in cutthroat
trout. Proc. Utah Acad. Sci., Arts, Lett. 32:75-80.

Ball, O. P., and O. B. Cope. 1961. Mortality studies on
cutthroat trout in Yellowstone Lake. U.S. Dept.
Inter., Fish Wildl. Serv. Res. Rep. 55. 62 pp.

Beatty, D. B. 1959. An experimental study of the tox-
icity of sodium bicarbonate, sodium chloride, and
sodium sulfate to rainbow trout. Unpublished
thesis. Univ. of Wyoming.

Behnke, R. J. 1974. The effects of the Newlands project
on the Pyramid Lake fishery. File Rep. 15 pp.

1979. Monograph of the native trouts of the

genus Salmo of western North America. U.S. For.
Serv., U.S. Fish Wildl. Serv., U.S. Bur. Land Mgt.
Washington, D.C.

Behnke, R. J., and M. Zarn. 1976. Biology and manage-
ment of threatened and endangered western
trout. USDA, For. Serv. Gen. Tech. Rep., Fort
Collins, Colorado. 45 pp.

Benson, L. V. 1978. Fluctuations in the level of pluvial
Lake Lahontan for the past 40,000 years. Quar-
temary Res. 9:300-318.

BjoRNN, T. C. 1957. A survey of the fishery resources of
Priest and Upper Priest lakes and their tribu-
taries. Dingell-Johnson Completion Rep. Idaho
Fish Game Dep., Boise, Idaho F-24-R. 176 pp.

Brown, C. J. D., and J. E. Bailey. 1952. Time and pat-
tern of scale formation in Yellowstone cutthroat
trout {Salmo clarkii lewisii). Trans. Amer. Mi-
crosc. Soc. 71(2): 120-124.

BuLKLEY, R. V. 1961. Fluctuations in age composition
and growth rate of cutthroat trout in Yellowstone
Lake. U.S. Fish Wildl. Serv. Res. Rep. 54. 31 pp.

BuLKLEY, R. v., and N. G. Benson. 1962. Predicting
year-class abundance of Yellowstone Lake cut-
throat trout. U.S. Fish Wildl. Serv. Res. Rep.
50:1-20.

Calhoun, A. J. 1942. The biology of the black-spotted
trout {Salmo clarkii henshawi [Gill and Jordan])
in two Sierra Lakes. Unpublished dissertation.
Stanford University. 218 pp.

1944. The food of the black-spotted trout {Salmo

clarkii henshawi [Gill and Jordan]) in two Sierra
Nevada lakes. California Fish Game Dep.
30(2): 80-85.

Carlander, K. D. 1969. Handbook of freshwater fish-
eries biology, Iowa State Univ. Press, Ames,
Iowa. 1:1-752.

Cooper, J. J. 1978. Contributions to the life history of
the Lahontan tui chub, Gila bicolor obesa (Gi-
rard), in Walker Lake, Nevada. Unpublished
thesis. Univ. of Nevada, Reno, Nevada.

Cope, O. B. 1953. Length measurements of Yellowstone
Lake trout. U.S. Fish Wildl. Serv. Spec. Sci. Rep.
Fish. 102. 17 pp.
Deacon, J. £., G. Kobetich, J. D. Willl\ms, and S.
Contreras. 1979. Fishes of North America en-
dangered, threatened, or of special concern. Fish.
Bui. Amer. Fish. Soc. 4(2):29-44.

Great Basin Naturalist

Vol. 43, No. 1

Drummond, R. a. 1966. Techniques in the collection
and mounting of trout scales. Prog. Fish Cult.
28(2):113-116.

Echo, J. B. 1955. Some ecological relationships between
yellow perch and cutthroat trout in Thompson
Lakes, Montana. Trans. Amer. Fish. Soc.
84:239-249.

Ferjancic, K. p. 1976. The results of the 1976 I.P.N. (In-
fectious pancreatic necrosis) survey conducted on
Lahontan cutthroat trout (Salmo clarki henshawi)
at Pyramid Lake, Nevada. Pyramid Lake Indian
Tribal Enterprises, Sutcliffe, Nevada Rept. 4 pp.

FiNNELL, L. M. 1966. Granby Reservoir Studies. Colo-
rado Div. Wildl., Fisheries Res. No. 3:4-6.

Fleener, G. G. 1952. Life history of the cutthroat trout
{Salmo clarkii Richardson), in Logan River, Utah.
Trans. Amer. Fish. Soc. 81:235-248.

Galat, D. L., E. L. Lider, S. Vigg, and S. R.
Robertson. 1981. Limnology of a large, deep
North American terminal lake. Pyramid Lake,
Nevada. USA. Hydrobiologia. 82:281-317.

Harris, E. E. 1970. Reconnaissance bathymetry of Pyra-
mid Lake, Washoe County, Nevada. Hydrologic
Investigation Atlas HA-379, U.S. Geol. Surv.

Hazzard, a. S., and M. J. Madsen. 1933. Studies of the
food of the cutthroat trout. Trans. Amer. Fish.
Soc. 63:198-207.

Hickman, T. J., and D. A. Duff. 1978. Status of cut-
throat subspecies in the western Bonneville Ba-
sin. Great Basin Nat. 38(2): 193-202.

Hickman, T. J., and R. J. Behnke. 1979. Probable dis-
covery of the original Pyramid Lake cutthroat
trout. Prog. Fish Cult. 41:135-137.

Hile, R. 1941. Age and growth of rock bass (Ambloplites
rupestris Rafinesque) in Nebesh Lake, Wisconsin.
Trans. Wisconsin Acad. Sci., Arts, Lett.
33:189-337.

HuBBS, C. L., and R. R. Miller. 1948. The Great Basin
with emphasis on glacial and postglacial times.
n. The zoological evidence. Univ. of Utah. Bull.
38(20): 17-166.

Houghton, S. G. 1976. A trace of desert waters. The
Great Basin story. Arthur H. Clark Co., Glendale,
California. 287 pp.

Hutchinson, G. E. 1957. Treatise on limnology: geogra-
phy, physics, and chemistry. Vol. 1. John Wiley
and Sons, New York. 1015 pp.

Innis, G. S., D. F. Hanson, and J. W. Haefner. 1981. A
simulation model of management alternatives in
a freshwater fishery. Elsevier Scientific Publ. Co.
Amsterdam, The Netherlands. 12:267-280.

Irving, R. B. 1953. Ecology of the cutthroat trout {Salmo
clarkii Richardson) in Henry's Lake, Idaho. Un-
published thesis. Utah State Agricultural College.
101 pp.

Jahn, L. a. 1969. Movements and homing of cutthroat
trout {Salmo clarki) from open-water areas of Yel-
lowstone Lake. J. Fish. Res. Bd. Can.
26:1243-1261.

1972. Development of the olfactory apparatus of

the cutthroat trout. Trans. Amer. Fish. Soc.
101(2):284-289.

Johnson, M. W., and L. Wroblewski. 1962. Errors as-
sociated with a systematic sampling creel census.
Trans. Amer. Fish. Soc. 91(2):201-207.

Johnson, V. K. 1958. Fisheries management report.
Lakes— Pyramid, Walker, and Tahoe. Dingell-
Johnson Completion Rep. Nevada Fish and
Game Dept. FAF-4-R. 47 pp.

Kennedy, J. L. 1978. A creel survey of Pyramid Lake,
Nevada. Pages 414-437 in W. F. Sigler and J. L.
Kennedy, eds.. Pyramid Lake ecological study.
W. F. Sigler & Associates Inc., Logan, Utah.

Knoll, J., D. L. Koch, R. Knoll, J. Sommer, L.
Hoffman, and S. Lintz. 1979. Physiological ad-
aptations of salmonid fishes {Salmo clarki hen-
shawi, Salmo gairdneri and Oncorhynchus
kisutch) to alkaline saline waters and their toxic
effects. Report to Office of Water Research and
Technology. Desert Res. Inst. Bioresources Cen-
ter Piibl. 50009. Reno, Nevada. 82 pp.

Koch, D. L., J. J. Cooper, E. L. Lider, R. L. Jacobson,
and R. J. Spencer. 1979. Investigations of Walker
Lake, Nevada: Dynamic ecological relationships.
Desert Res. Inst. Bioresources Center Publ. Reno,
Nv. No. 50010. 191 pp.

Kucera, p. a., and J. L. Kennedy. 1977. Evaluation of a
sphere volume method for estimating fish fecun-
dity. Prog. Fish Cult. 39(3):115-117.

Laakso, M., and O. B. Cope. 1956. Age determination
in Yellowstone cutthroat trout by the scale meth-
od. J. Wildl. Mgt. 20(2): 138-153.

Lea, R. N. 1963. Ecology of the Lahontan cutthroat
trout {Salmo clarki henshawi) in Independence
Lake, California. Unpublished thesis. Univ. of
California, Berkeley. 95 pp.

LeCren, E. D. 1951. The length-weight relationship and
seasonal cycle in gonad weight and condition in
perch {Perca fluviatilis). J. Anim. Ecol.
20(2):201-219.

Lider, E. L., and R. W. Langdon. 1978. Plankton ecolo-
gy. Pages 241-293 in W. F. Sigler and J. L. Ken-
nedy, eds., Pyramid Lake ecological study. W. F.
Sigler & Associates Inc., Logan, Utah.

Madsen, M. J. 1940. Report on age and growth of cut-
throat trout {Salmo lewisii) of Yellowstone Lake,
Wyoming. U.S. Fish Wildl. Serv. Res. Rept. 12
pp.

Marnell, L. F. 1969. Hooking mortality of cutthroat
trout. Unpublished dissertation, Colorado State
Univ. 99 pp.

Marnell, N. E., and D. Hunsaker 11. 1970. Hooking
mortality of lure-caught cutthroat trout {Salmo
clarki) in relation to water temperature, fatigue
and reproductive maturity of released fish. Trans.
Amer. Fish. Soc. 99(4): 684-688.

McCarraher, D. B. 1972. A preliminary bibliography
and lake index of the inland mineral waters of
the world. FAO. Fish. Cir. 146.

McCleave, J. D. 1967. Homing and orientation of cut-
throat trout {Salmo clarki) in Yellowstone Lake
with special reference to olfaction and vision. J.
Fish. Res. Bd. Can. 24(10):201 1-2044.

McCoNNELL, W. J., D. L. Galat, and K. Hamilton-
Galat. 1978. Potential fish production of Pyra-
mid Lake based on organic matter contributions.
Colorado Coop. Fish. Res. Unit. Fort Collins,
Colorado. 84 pp.

January 1983

SiGLER ET AL.: LaHONTAN CuTTHROAT TrOUT

Miller, R. G. 1951. The natural history of Lake Tahoe
fishes. Unpubhshed dissertation. Stanford Univ.
160 pp.

MiTCHUM, D. L. 1960. An experimental study of the tox-
icity of calcium carbonate, calcium sulphate,
magnesium carbonate, and magnesium sulphate
to rainbow trout. Unpublished thesis. Univ. of
Wyoming.

Nelson, L. 1976. SHAD II. A model for analysis of fish-
eries age and growth data. Wildl. Sci. Dep., Utah
State Univ., Logan, Utah. 15 pp.

NiKOLSKY, C. B. 1963. The ecology of fishes. Acad. Press,
New York. ,393 pp.

Peters, J. C, ed. 1964. Summary of calculated growth
data on Montana fishes, 1948-1961. Dingell-John-
son Completion Rep., Montana Fish and Game
Dept., Helena, Montana. F-23-R. 76 pp.

Platts, W. S. 1959a. Homing, movements, and mor-
tality of wild cutthroat trout {Salmo clarki)
spawned artificially. Prog. Fish Cult. 1959
(January):36-38.

1959b. Food habits of the cutthroat trout in

Strawberry Reservoir, Utah. Proc. Utah Acad.
Sci., Arts, Lett. 36:119-121.

Rankel, G. L. 1976. Fishery management program,
1976, Summit Lake Indian Reservation, Hum-
boldt County, Nevada. Manuscript, U.S Fish
Wildl. Serv., Reno, Nevada. 35 pp.

RiCKER, W. E., ed. 1971. Methods for assessment of fish
production in freshwaters. Blackwell Sci. Publ.,
Oxford. 348 pp.

Robertson, O. H. 1947. An ecological study of two high
mountain trout lakes in the Wind River Range,
Wyoming. Ecology 28(2):87-112.

Robertson, S. R. 1978. The distribution and relative
abundance of benthic macroinvertebrates in
Pyramid Lake, Nevada. Unpublished thesis,
Univ. of Nevada.

RuGER, A. W. 1977. The results of the 1977 disease sur-
vey conducted on Lahontan cutthroat trout (Sal-
mo clarki henshawi) at Pyramid Lake, Nevada.
Manuscript, Pyramid Lake Indian Tribal Enter-
prises, Sutcliffe, Nevada.

SiGLER, W. F. 1951. The life history and management of
the mountain whitefish (Prosopium williamsoni
(Girard)) in Logan River, Utah. Utah State Agri-
cultural College Bull. 347. 36 pp.

1962. Bear Lake and its future, 26th Fac. Assn.

Honor Lecture. Utah State Univ., Logan, Utah.
23 pp.

SiGLER, W. F., AND J. L. KENNEDY, eds. 1978. Pyramid
Lake ecological study. W. F. Sigler & Associates
Inc., Logan, Utah. 545 pp.

SiGLER, W. F., AND J. B. Low. 1950. Age composition
and growth of fish and fishermen success in
Utah's high Uinta lakes. Utah Acad. Sci., Arts,
Lett. 27:32-36.

Sigler, W. F., and R. R. Miller. 1963. Fishes of Utah.
Utah Dept. Fish and Game, Salt Lake City, Utah.
203 pp.

Smith, S. H. 1963. Making plastic impressions of fish
scales with a roller press. Prog. Fish Cult.
16(2): 75-78.

Snyder, G. R., and H. A. Tanner. 1960. Cutthroat trout
reproduction in the inlets to Trappers Lake. Col-
orado Div. Wildl. Tech. Bull. 7:1-85.

Snyder, J. O. 1917. The fishes of the Lahontan system of
Nevada and northeastern California. U.S. Bur.
Fish. Bui. 1915-16(35:31-86).

Sumner, F. H. 1939. The decline of Pyramid Lake fish-
ery. Trans. Amer. Fish. Soc. 69:216-224.

Taylor, R. E. L. 1972. The effects of increasing salinity
on the Pyramid Lake fishery. Agric. Expt. Sta.,
University of Nevada. 8 pp.

Tesch, F. W. 1971. Age and growth. Pages 98-131 in
W. E. Ricker, ed., Methods of assessment of fish
production in freshwaters. Blackwell Sci. Publ.,
Oxford.

TowNLEY, J. M. 1980. The Truckee Basin fishery,
1844-1944. Nevada Historical Society in coopera-
tion with Desert Research Institute, Univ. of Ne-
vada, Reno.

Trelease, T. J. 1953. The death of a lake. Field and
Stream. February.

1969. The rebirth of a lake. Nevada Outdoors

and Wildlife Review. Nevada Dept. Fish and
Game 4:3.

United States Geological Survey. 1977. Water re-
sources data for Nevada, water year 1976. U.S.
Dept. Inter., Carson City, Nevada. 344 pp.

Van Oosten, J. 1923. The whitefishes [Coregonus
clupeaformis). A study of the scales of whitefishes
of known ages. Zoologica 11(17):380-412.

1929. Life history of the lake herring (Coregonus

artedii LeSueur) of Lake Huron as revealed by its
scales, with a critique of the scale method. U.S.
Bur. Fish. 44(1928):265-428.

1944. Factors affecting the growth of fish.

Trans. Ninth N. Amer. Wildl. Conf. 9:177-183.

Vigg, S. 1978. Vertical distribution of adult fish in Pyra-
mid Lake, Nevada. Great Basin Nat.
38(4):417-428.

1980. Seasonal benthic distribution of adult fish

in Pyramid Lake, Nevada. California Fish and
Game J. 66(l):49-58.

1981. Species composition and relative abun-
dance of adult fish in Pyramid Lake, Nevada.
Great Basin Nat. 41(4):395-408.

Vigg, S., and D. L. Koch. 1980. Upper lethal temper-
ature range of Lahontan cutthroat trout in waters
of different ionic concentration. Trans. Amer.
Fish. Soc. 109:336-339.

Wheeler, S. S. 1974. The Desert Lake: the story of Ne-
vada's Pyramid Lake. Caxton Printers, Ltd.,
Caldwell, Idaho. 133 pp.

Wolcott, Roger S. C, Jr. 1978. Evaluation of the 1978
Lahontan cutthroat trout run to the Pyramid
Lake fishway and comparison with past year.
U.S. Fish Wildl. Serv., Fish. Asst. Office, Reno,
Nevada. 30 pp.

A REVIEW OF THE GENUS SOLIPERLA (PLECOPTERA: PELTOPERLIDAE)

Bill P. Stark'

Abstract.— The western Nearctic stonefly genus Soliperla is reviewed and six species are recognized. Soliperla
sierra (Calif.) and S. tillamook (Ore.) are described as new to science and illustrations of diagnostic features are pre-
sented for all species. Males and nymphs are keyed and a phylogeny for the group is proposed.

Soliperla was proposed by Ricker (1952) as
a monotypic subgenus of Peltoperla to con-
tain P. thyra Needham & Smith. At that time
the species was known only from the male
holotype but Jewett (1954) described the fe-
male along with males and females of two
additional species, P. campanula and P.
quadrispinula; the nymph of P. campanula
was also described at this time. Jewett (1955)
described the fourth member of the group, P.
fenderi, from a single male. These species
have remained poorly known since their dis-
covery, with only synoptic notes and regional
keys (Jewett 1959, 1960) appearing until
Stark and Stewart (1981) gave additional
characters that supported Illies's (1966) ele-
vation of the group to generic status.

During recent field work with colleagues
in Washington, Oregon, and California, Soli
perla nymphs were common in splash zones
of small streams and springs. Through this
work, nymphs were associated for the four
known species, and the additional material
collected along with specimens obtained

from museums permits the first com-
prehensive treatment of Soliperla. Methods
were given by Stark and Stewart (1981).

Soliperla Ricker

Peltoperla (Soliperla) Ricker 1952: 157. Type-species of
subgenus: Peltoperla thyra Needham & Smith.
Monotypic.
Soliperla lilies 1966:26.

Adults and nymphs of Soliperla are unusual
among Nearctic Peltoperlidae in displaying
distinctive pigmentation patterns. Adults are
typified by a dark mesal pronotal stripe that
contrasts sharply with the light yellow back-
ground (Figs. 23, 31), and nymphs have con-
spicuous white areas on the abdominal terga
that contrast with the dark background (Figs.
4, 15). Monophyly for the group is asserted
on the basis of the distinctive epiproct, with
recurved crenulate apex and the membranous
pair of lobes associated with the epiproct
(Figs. 5, 13). The genus is currently known
from the western Nearctic region from cen-
tral California to Washington (Fig. 1).

Keys to Soliperla males

1. Meso ventral area of aedeagus with two irregular longitudinal rows of short,

thick setae (Figs. 3, 30) 2

— Mesoventral area of aedeagus with short, thick setae, if present, not in

longitudinal rows 3

2(1). Lateral aedeagal lobes terminating in sclerotized spine with 2-4 small

subapical setae (Fig. 29) sierra

'Department of Biology, Mississippi College, Clinton, Mississippi 39058.

January 1983

Stark: Review of Soliperla

Fig. 1. Distribution of Soliperla species. S. campanula = open circles, S. fenderi = closed circles, S. quad-
rispinula = open squares, S. sierra = triangles; S. thyra = closed squares, S. tillamook = x.

32 Great Basin Naturalist Vol. 43, No. 1

— Lateral aedeagal lobes membranous apically with 2-4 small subapical setae
(Fig. 2) campanula

3(1). Ventral aedeagal lobes with large sclerotized spine or a single long, thick seta

(Figs. 19,35) 4

— Ventral aedeagal lobes membranous with numerous scattered setae
(Figs. 12,45) 5

4(3). Ventral aedeagal lobes with sclerotized bilobed process (Fig. 35) thyra

— Ventral aedeagal lobes with a terminal long, thick seta (Fig. 19) quadrispinula

5(3). Ventral aedeagal lobes with irregular row of long, slender setae along apical

margin (Fig. 12) fenderi

— Ventral aedeagal lobes with scattered short setae along apical margins
(Fig. 45) tillamook

Preliminary key to nymphs
(sierra and tillamook unknown)

1. Some long setae in abdominal tergum 9 posterior fringe bent (Fig. 39);

abdominal tergum 5 typically with lateral pale spots (Fig. 22) 2

— Long setae in abdominal tergum 9 posterior fringe straight (Fig. 6); abdominal
tergum 5 typically without pale spots (Fig. 4) 3

2(1). Mesal area of abdominal tergum 8 posterior fringe with ca 20 clavate setae be-
tween long setae (Fig. 40); mesal pale spots on abdominal terga 5 and 6
rounded (Fig. 38) thyra

— Mesal area of abdominal tergum 8 posterior fringe with ca 5-7 clavate setae
between long setae; mesal pale spots on abdominal terga 5 and 6 irregularly
linear to triangular (Fig. 22) quadrispinula

3(1). Lateral pale spots on abdominal tergum 4 conspicuously larger than mesal spot

(Fig. 15); known from Mt. Rainier, Washington fenderi

— Lateral pale spots on abdominal tergum 4 subequal to mesal spot in size (Fig.

4); widely distributed in northern Oregon campanula

Soliperla campanula (Jewett) Females cannot be distinguished with cer-
tainty from several related species, and the

Peltaperla {Soliperla) campanula Jewett 1954: 167. Holo- ,^. - r.. , , i features

type $ (CAS), Oxbow Springs, Hood River Co., ^§8^ ^^^g^' /' ^> ^^^° f ^"^ /° *^^ , teatures

Oregon th^t would distinguish them from other mem-

Jewett (1954) detailed the major diagnostic bers of the genus. The species is currently

features of this species. Males are distin- known only from northern Oregon (Fig. 1).

guished from other Soliperla by the structure „ Material EXAMiNED.-Oregon: Cl^ckarna. Co Mt.

°c 1 ■ 1 1 4.1 1 Hood, near Timberline Lodge, 31-V-77, K. W. Stewart,

of the epiproct and aedeagus. The anterodor- ^ ^ Szczytko, 2 s (reared) (NTSU); same location, 20-

sal face of the epiproct is about twice as wide vn-67, S. G. Jewett, l ? (USNM); Mt. Hood, Still Crk.

as the stalk, and the lateral margins are Cmp. Gnd., 12-VII-79, B. Stark, K. W. Stewart, 2 s

curved inward near the crenulate antero-ven- (^P^); trib. Still Crk, Mt. Hood, 17-VI-67, S. G. Jewett, 1

.. 1 _r /IT- tr\ A .. 1 in 1^ ■ S (USNM); Mt. Hood, 1.2 mi N Hwy 26, 20- vn-67, J.

tral surface (Fig. 5). Approximately 12-14 ir- ^^\j ^^^j^^y^ ^^.^^ S^,^^^ ^.^ ^^ ^^^^ 15-vn-54, S.

regular teeth are present along this surface. g. Jewett 1 S (USNM). Hood River Co., Oxbow Springs,
The ventral aspect of the aedeagus includes a 26-V-40, S. G. Jewett, 15,1? (OSU); Iron Crk, W of
large mesal lobe and two small lateral lobes. Bennetts Pass, ll-VII-68, E. Evans, 13,2? (USNM).

The mesal lobe has two irregular rows of ^^ ?"' ^f f f'o^nJn^M'^Trf ?t ?V h'^""

, ~ ^r, 1. 1-1 111 78, B. Frost, 1^,1? (OSU); Mack Crk, H. J. Andrews

about 5-10 short peglike setae, and the later- g^p p^r., 25-VI-74, N. H. Anderson, l s (OSU); 12.5

al lobes have 2-4 subapical peglike setae mi NE Blue Riv, H. J. Andrews Exp. For., 19-VII-78, B.

(Fig. 3). Frost (OSU). Linn Co., Ice Cap Crk, 7-VII-66, J. Bedea,

January 1983

Stark: Review of Soliperla

Figs. 2-6. S. campanula. Fig. 2. Aedeagus, lateral. Fig. 3. Aedeagus, A = dorsal, B = ventral. Fig. 4 Nymphal ab-
domen, dorsal. Fig. 5. Epiproct, anterodorsal. Fig. 6. Nymphal abdominal tergum 8, posterior fringe.

Great Basin Naturalist

Vol. 43, No. 1

Figs. 7-10. Soliperla eggs. Fig. 7. S. campanula, 280X. Fig. 8. S. campanula, lOOOX. Fig. 9. S. fenderi, 300X. Fig.
10. S. fenderi, lOOOX.

2 3 (OSU); 14 mi NE Blue Riv, H. J. Andrews Expt.
For., 6-VII-78, B. Frost 1 <? (OSU). Multnomah Co.,
Wahkeena Falls, 3-V-47, S. G. Jewett, 2 3,2$ (ROM)
(OSU); same location, 4-V-82, R. W. Baumann & S. G.
Jewett, 1 $ (MLB).

Soliperla fenderi (Jewett)

Peltoperla (Soliperla) fenderi Jewett 1955: 145. Holotype
<5 (SGJ), Saint Andrews Crk, Mt. Rainier Natl.
Pk., Washington

January 1983

Stark: Review of Soliperla

m-i^^iii<^'

Figs. 11-17. S. fenderi. Fig. 11. Aedeagiis, lateral. Fig. 12. Aedeagus, A = dorsal, B = ventral. Fig. 13. Epiproct,
anterodorsal. Fig. 14. Epiproct; lateral. Fig. 15. Nymphal abdomen, dorsal. Fig. 16. Female sterna 8 and 9. Fig. 17.
Nymphal abdominal tergum 8, posterior fringe.

Great Basin Naturalist

Vol. 43, No. 1

Jewett's (1955) description of this species
from a single male included diagnostic fea-
tures of the epiproct and aedeagus. The an-
terodorsal face of the epiproct is about the
same width as the shaft and is armed by
about 5-6 teeth along the anteroventral sur-
face (Fig. 13). The apical aedeagal section
has two large membranous ventral lobes that
have long slender setae in an irregular row
along the distal margins (Fig. 12).

Females are distinguished on the basis of
the shallow V-shaped notch on the subgenital
plate (Fig. 16) and the eggs (Figs. 9, 10) are
typical of the genus. This species is presently
known only from Mt. Rainier National Park,
but a single nymph collected by R. W. Bau-
mann near Snoqualmie Pass could be this
species (Fig. 1).

Material examined.— Washington: Pierce Co., Mt.
Rainier Natl. Pk. spring seeps along St. Andrews Crk,
13-VII-79, B. Stark, K. W. Stewart, 1 ? (reared) 7
nymphs (BPS); same location, 29-VI-81, K. W. Stewart,
W. Shephard, 10 S , 6 ? (reared); small stream at Re-
flection Lk, 14-VII-79, B. Stark, K. W. Stewart 1 S
(reared) (NTSU). Seeps along Puyallap Riv, 29-VI-81, K.
W. Stewart, W. Shephard, 1 <5 , 2 ? (reared) (NTSU).
Christina Falls, lS-VI-69, R. W. Baumann, 1 $ (MLB).

Soliperla quadrispinula (Jewett)

Peltoperla (Soliperla) quadrispinula Jewett 1954: 169.

Holotype S (CAS), Wrangle Gap Camp, Jackson

Co., Oregon
Jewett's (1954) description suggested that
the aedeagus of this species, as the name im-
plies, has four prominent spines. I have found
considerable variation in this character even
within individuals taken from the same local-
ity. However, all males examined had a single
long, thick seta on each ventral lobe (Fig.
19); the variation occurs in the number of
these setae on the dorsal lobes. The range of
variation seen in this character extends from
no setae (found on one male from Oak Crk,
Benton Co., Oregon) to 4 setae (found on sev-
eral individuals from Jackson Co., Oregon).
The epiproct, as Jewett (1954) indicated, is
similar to that of S. campanula, but the dor-
sal carina on the anterodorsal face is not de-
veloped mesally and the lateral margins near
the anteroventral surface curve outward (Fig.
20).

The female and egg (Figs. 25, 26) are in-
distinguishable from several others in the
genus. The species has the largest known
range for any member of the genus. Records

are presently from northern Oregon (Clatsop
Co.) to northern California (Humboldt and
Trinity Cos.). Adults reared from Fieldbrook,
California, mated readily in captivity but did
not mate with S. thyra specimens from Napa
Co., California.

Material examined.— California: Humboldt Co.,
Grassy Crk, Fieldbrook, 22-V-82, B. Stark, D. Ziegler, 6

3,3? (reared), 54 nymphs (BPS) (NTSU). Oregon:
Benton Co., Parker Crk, l-VII-71, G. Steyskal, 1 $
(USNM); Oak Crk, 8-VI-79, P. Hammond, 3 $ (BK);
same location, 10-13-V-69, C. Kerst, 1 S (OSU); same
location, 4-7-VI-68, 1 ? (OSU). Clatsop Co., Osweg Crk,
2 mi E Elsie, 30-V-64, S. G. Jewett, 1 <? (ROM). Jackson
Co., 1.5 mi N Wrangle Cmp., 8-V1I-79, B. Stark, K. W.
Stewart, 19 <5 , 4 ? , 4 nymphs (BPS) (NTSU); 15 mi S
Talent, 8-VII-79, B. Stark, K. W. Stewart, 6 3 , 3 $ , 6
nymphs (BPS) (NTSU). Josephine Co., Lake Crk, Oregon
Caves Natl. Mon., 9-VII-79, B. Stark, K. W. Stewart, 1

? (reared) (BPS). Yamhill Co., 5 mi E Hack Crk Rd-
Hwy 5 jet, 13-V-82, K. W. Stewart, D. Ziegler, 1 ?
(reared) (NTSU).

Soliperla sierra, n. sp.

Male.— Body length, 10 mm; fore wing
length, 12 mm. General color yellow pat-
terned with light brown. Head without
brown markings; pronotum with long light
spot in mesal dark stripe (Fig. 31). Antero-
dorsal face of epiproct about 2x wide as
stalk, lateral margins curved to anteroventral
margin; mesal teeth widely separated (Fig.
32). Ventral lobe of aedeagus with 2 irregular
mesal rows of about 5-8 short, thick setae;
lateral lobes with a large conical terminal
spine and about 2-4 subapical short, thick
setae (Fig. 30).

Female.— Body length, 12 mm; forewing
length, 14 mm. General color similar to male.
Subgenital plate large, parabolic, reaching
posterior margin of sternum 9.

Egg.— Typical of the genus (Figs. 27, 28)
but micropylar orifices are irregular rather
than smooth.

Types.— Holotype ( S ), allotype ( $ ), and
5 $ and 2 ? paratypes from Plumas Co.,
California, French Crk, 1 mi N Caribou, Butt
Reservoir Rd, 25-VI-80, R. W. Baumann and
J. Stanger, deposited in the U.S. National
Museum (100090). Additional Paratypes: Cal-
ifornia: Sierra Co., Big Springs, Hwy 49, N
Sierra City, 24-VI-80, R. W. Baumann, J.
Stanger, 13 5 , 4 ? (MLB).

Etymology.— The specific name sierra re-
fers to the mountain range in which it was
collected.

January 1983

Stark: Review of Soliperla

Figs. 18-24. S. quadrispinula. Fig. 18. Aedeagus, lateral. Fig. 19. Aedeagus, A = dorsal, B = ventral. Fig. 20. Epi-
proct, anterodorsal. Fig. 21. Epiproct, lateral. Fig. 22. Nymphal abdomen, dorsal. Fig. 23. Female head and pro-
notum. Fig. 24. Nymphal abdominal tergum 8, posterior fringe.

Great Basin Naturalist

Vol. 43, No. 1

Figs. 25-28. Soliperla eggs. Fig. 25. S. qiiadrispimila, 240X. Fig. 26. S. quadrispinula, lOOOX. Fig. 27. S. sierra,
240X. Fig. 28. S. sierra, lOOOX.

Soliperla thyra (Needham & Smith)

Peltaperla thyra Needham & Smith 1916: 87. Holotype

(5 (Cornell Univ.), Nevada.
Peltoperla (Soliperla) thyra: Ricker 1952: 157.
Peltaperla (Soliperla) thyra: Jewett 1954: 167.

Jewett (1954) described the sahent features
of male and female genitalia and discussed
the holotype specimen. After examining the
holotype, I concur with Jewett in his usage of
thyra. The anterodorsal face of the epiproct

January 1983

Stark: Review of Soliperla

Figs. 29-33. S. sierra. Fig. 29. Aedeagus, lateral. Fig. 30. Aedeagus, A = dorsal, B = ventral. Fig. 31. Female
head and pronotum. Fig. 32. Epiproct, anterodorsal. Fig. 33. Epiproct, lateral.

is slightly wider than the stalk and about 10
irregular teeth are located along the antero-
ventral surface (Fig. 36). The ventral aedea-
gal lobes bear a partially sclerotized spinelike
bilobed process; the ventral portion of the

process is less heavily sclerotized and bears
about 6-8 short, thick setae, including an
apical cluster (Fig. 35). The dorsal mem-
branous lobes bear several scattered short,
thick setae (Fig. 35).

Great Basin Naturalist

Vol. 43, No. 1

\ i

Jlf^^~T'J' 'i^'^- ^'^- ^/^^«fg"^' •^t^'-'^l- Fig- 35. Aedeagus, A = dorsal, B = ventral. Fig. 36. Epiproct,
^terodorsal. F.g. 37^ Epiproct, lateral. Fig. 38. Nymphal abdomen, dorsal. Fig. 39. Nymphal abdominal tergSm 9
l-ig. 40. Nymphalabdommaltergum 8, posterior fringe.

The female and egg appear in-
distinguishable from several other members
of the genus. The species is known from scat-
tered localities around the San Francisco
area, but no specimens have been taken in
Nevada or eastern California except the holo-
type(Fig.l).

Material examined.— California: Marin Co. Cascade
Crk, Mill Valley, 25-IV-57, H. B. Leech, 1 <5 (ROM);
same location, 20-V-55, H. B. Leech, 1 ? (ROM); trickle
nr. Alpine Lk, l-V-55, S. W. Hitchcock, 1 <J , 2 nymphs
(OSU); Mt. Tamalpais, 25-V-74, D. G. Denning, 1 <J
(MLB). Mendocino Co., sm. stream at Leggett, 22- V-
1982, B. Stark, D. Ziegler, 1 $ ,1 ? , 2 nymphs (NTSU).
Napa Co., 9 mi N Calistoga, Hwy 29, 21-V-82, B. Stark,
D. Ziegler, 4 ^ , 10 ? , 4 nymphs (2 <? , 6 ? reared)

January 1983

Stark: Review of Soliperla

Figs. 41-43. S. thyra nymphal setae. Fig. 41. Bent setae, posterior fringe tergum 9, 700X. Fig. 42. Straight and cla-
vate setae, posterior fringe tergum 8, 600X. Fig. 43. Clavate setae, posterior fringe tergum 8, 28(X)X.

(BPS); Angwin, IV-76, D. Ashley, 1 ? (BPS). Santa Clara
Co., Uras Canyon, 25-V-74, D. G. Denning, 1 $ (MLB).
Nevada: no additional data (Holotype <5 ) (Cornell).

Soliperla tiHamook, n. sp.

Male.— Body length, 11 mm; forewing
length, 13 mm. General color yellow pat-
terned with brown. Anterodorsal face of epi-
proct slightly wider than stalk; antero ventral
surface with about 7-9 teeth (Fig. 46). Ven-
tral and lateral lobes of aedeagus mem-
branous with scattered apical short, thin
setae (Fig. 45).

Female.— Body length, 13 mm; forewing
length, 15 mm. Subgenital plate large, para-
bolic, reaching to posterior margin of ster-
num 9.

Types.— Holotype ( $ ) and 1 $ paratype
from Clatsop Co., Oregon, Osweg Crk, 2 mi
E Elsie, 13-VI-64, S. G. Jewett. Holotype de-
posited in the Oregon State University mu-
seum. Allotype ( ? ) and 1 $ paratype from
Clatsop Co., Oregon, trib. Big Crk, Salmon
Hatchery, 28-V-49, S. G. Jewett (OSU).

Etymology.— The specific name honors
the Tillamook Indians, who inhabited the
area near the type locality.

Discussion

In this study several characters suitable for
making phylogenetic inferences were ob-

served. Most of these have been selected
from male genitalic structures since the egg,
nymph, and female show little apparent vari-
ation. A taxonomic out group (TOG) con-
sisting of Yoraperla and Sierraperla was
formed to assist in inferring polarity of se-
lected characters (Watrous and Wheeler
1981).

1. Aedeagal setae. Two variations of setae
occurring on the ventral lobes were noted in
the TOG and TIG (taxonomic in group). In
Yoraperla and Sierraperla these setae are not
appreciably thickened. Fine setae also occur
in Soliperla fenderi and S. tillamook (Figs. 12,
45), but in other Soliperla species these setae
are distinctly thickened (Figs. 3, 19). This is
most conspicuous in S. quadrispinula, in
which the setae are also longer than usual.
Hence, thickened setae are regarded as
apomorphic.

2. Epiproct tip. Within Soliperla several
species have the epiproct tip expanded into a
process that is much wider than the epiproct
base (Figs. 5, 20), but in other species the epi-
proct tip is about as wide as the base (Fig.
13). Since Yoraperla and Sierraperla lack a
developed epiproct, the TOG was expanded
to include other Nearctic peltoperlid genera
{Tallaperla, Peltoperla, and Viehoperla). In
this group, both Peltoperla and Viehoperla
have narrow epiproct tips, and in Tallaperla
the epiproct is poorly developed (Stark and
Stewart 1981). Hence, the expanded epiproct

Great Basin Naturalist

Vol. 43, No. 1

Figs. 44-46. S. tillamook. Fig. 44. Aedeagus, lateral. Fig. 45. Aedeagus, A = dorsal, B = ventral. Fig. 46. Epi-
proct, anterodorsal.

tip of S. campanula, S. quadrispinula, S.
sierra, and S. thyra is regarded as
apomorphic.

3. Subgenital plate margin. The apex of
the female subgenital plate is emarginate
(Sierraperla) or notched (Yoraperla) in the
TOG, but most Soliperla have a large para-
bolic plate that is entire along the margin.
Only S. fenderi has a notched plate, but this
notching is considered nonhomologous to
that in the TOG and the notched plate is re-
garded as apomorphic.

4. Outline of ventral aedeagal lobe. In
Yoraperla, Sierraperla, and several Solipera
species, the ventral aedeagal lobe is divided
longitudinally by a cleft into lateral lobes
(Figs. 12, 35). In two Soliperla species (S.
campanula and S. sierra) the apical margin of
this lobe is entire; hence, this is considered to
be the apomorphic feature.

5. Arrangement of aedeagal setae. Typi-
cally in Yoraperla, Sierraperla, and Soliperla
species the setae on the ventral aedeagal lobe
are scattered, but in two species (S. camp-

anula and S. sierra) these setae form two ir-
regular mesal rows (Figs. 3, 30). This latter
variation is considered apomorphic.

6. Sclerotization of ventral aedeagal lobe.
The TOG and most Soliperla species have en-
tirely membranous ventral aedeagal lobes,
but in S. thyra and S. quadrispinula these
lobes are sclerotized (Figs. 19, 35). This latter
modification is considered apomorphic.

These six characters were used to derive
the cladogram shown in Fig. 47. This analysis
must be regarded as tentative due to the lim-
ited number of characters utilized. The posi-
tion of S. tillamook could not be determined
since it exhibits the plesiomorphic condition
for each of the six characters.

It seems probable, given the apparent high
degree of endemicity in the genus, that addi-
tional species await discovery from the
Olympic Peninsula, Mt. Baker, and other iso-
lated areas of the Pacific Northwest. It is
hoped material from these areas and a larger
sample of S. tillamook will permit rigorous
testing of this cladogram.

January 1983

Stark: Review of Soliperla

0)
•H
W

■H

•H

r— 1
rH
•H

-p

+->

- - 6

_- 5

O.
D.
D-

col

■Q

n -Aponorphic
O -Plesionorphic

Fig. 47. Proposed phylogeny for Soliperla species. See text for explanation.

Acknowledgments

Several individuals and museums cooper-
ated in providing material for study. These
include R. W. Baumann (Monte L. Bean Mu-
seum, Brigham Young University), O. S. Flint
(United States National Museum), B. Kon-
dratieff, J. Lattin (Oregon State University
Museum), B. Mather, L. L. Pechuman (Cor-
nell University Museum), K. W. Stewart
(North Texas State University Museum) and
G. Wiggins (Royal Ontario Museujn). K. W.

Stewart and D. Ziegler gave valuable assis-
tance in collecting and rearing efforts. S. W.
Szczytko (University of Wisconsin, Stevens
Point) and S. Faison (University of Mississippi
Dental School) assisted in preparing SEM mi-
crographs. This study was supported, in part,
by NSF grant DEB 78-12565.

Literature Cited

Illies, J. 1966. Katalog der rezenten Plecoptera. Das
Tierreich, 82. Walter de Gruyter and Co. 632 pp.

Great Basin Naturalist

Vol. 43, No. 1

Jewett, S. G. 1954. New stoneflies from California and
Oregon (Plecoptera). Pan-Pac. Entomol.
30:167-179.

1955. Notes and descriptions concerning west-
em North American stoneflies (Plecoptera). Was-
mann Jour. Biol. 13:145-155.

1959. The stoneflies (Plecoptera) of the Pacific

Northwest. Oregon State Monographs, Studies
Entomol. 3:1-95.

1960. The stoneflies (Plecoptera) of California.

Bull. Calif. Insect Surv. 6:125-177.

Needham, J. C, AND L. W. Smith. 1916. The stoneflies
of the genus Peltoperla. Canadian Entomol.
48:80-88.

RicKER, W. E. 1952. Systematic studies in Plecoptera.
Indiana Univ. Pub. Sci. Ser. 18:1-200.

Stark, B. P., and K. W. Stewart. 1981. The Nearctic
genera of Peltoperlidae (Plecoptera). J. Kans. En-
tomol. Soc. 54:285-311.

Watrous, L. E., and Q. D. Wheeler. 1981. The out-
group comparison method of character analysis.
Syst. Zool. 30:1-11.

A BIBLIOGRAPHY OF COLORADO VEGETATION DESCRIPTION

William L. Baker'

Abstract.— A list of 658 references to Colorado vegetation description is presented, along with county and sub-
ject indexes to the list.

This bibliography includes published and
selected unpublished references through
1981. Included are references to vegetation
description, including studies that contain
only qualitative description, along with more
detailed quantitative studies. Also included
are selected references to age /size structure,
fire, succession, floristics, vegetation history
(primarily palynology), phenology, plant ge-
ography, vegetation zonation, and the tim-
berline. An index is included to subjects other
than vegetation description. Coverage of
these tangential subjects may not be com-
prehensive. Autecological and environmental
studies are generally excluded, as are studies
pertaining only to nonvascular vegetation.

References are included here if at least
part of the study area is in Colorado. Ex-
cluded are the many works from adjoining
states that may have relevance, particularly
to the margins of Colorado, but have no part
of their study areas inside Colorado. Re-
searchers should consult bibliographies from
adjoining states for these references.

Included below is an index to the bibliog-
raphy by county. A single reference may per-
tain to more than one county. Also included
are a list of studies pertaining to the whole
state, and a list of "regional studies" that per-
tain to an undefined, or poorly demarcated
part of the state. Researchers seeking com-
plete coverage of a particular county should
also check references in these lists.

Subject Index

Age/Size Structure: 286, 291, 302, 330, 373, 488, 489,

547, 558, 559, 624, 628, 630
Bibliographies: 3, 4, 7, 214, 658

Fire: 2, 3, 4, 5, 13, 19, 24, 25, 37, 38, 49, 70, 92, 93, 98,
113, 114, 150, 152, 153, 171, 211, 228, 234, 235, 249,
306, 395, 396, 417, 490, 556, 557, 655, 656, 658, 659

Floristics: 10, 23, 30, 45, 60, 101, 102, 126, 164, 203, 215,
216, 298, 335, 372, 397, 402, 403, 404, 493, 496, 497,
500, 531, 561, 599, 616

Palynology/Vegetation History: 11, 307, 336, 337, 338,
340, 425, 426, 427, 428, 617

Phenology: 30, 42, 155, 217, 218, 219, 220, 221, 222, 240,
246, 249, 368, 369, 443, 506, 564, 619, 620, 621

Plant Geography: 179, 215, 216, 249, 287, 288, 298, 326,
372, 375, 419, 421, 424, 443, 454, 493, 497, 500, 611,
613, 614, 615, 619, 620, 621

Succession: 5, 8, 13, 15, 21, 24, 25, 26, 32, 37, 38, 49, 69,
70, 73, 83, 84, 92, 93, 98, 105, 112, 113, 114, 125, 128,
131, 140, 141, 146, 150, 152, 153, 161, 171, 178, 182,
183, 184, 185, 188, 211, 228, 229, 234, 239, 242, 255,
258, 261, 263, 286, 299, 301, 311, 312, 318, 319, 321,
327, 343, 345, 346, 347, 352, 354, 360, 375, 384, 395,
396, 400, 401, 407, 415, 417, 418, 420, 429, 448, 466,
472, 478, 483, 490, 504, 507, 510, 517, 518, 519, 520,
524, 525, 537, 542, 545, 546, 547, 548, 556, 557, 558,
559, 580, 611, 624, 625, 626, 627, 630, 632, 633, 635,
636, 637, 638, 639, 640, 641, 646, 653, 656, 658, 659

Timberline: 49, 62, 167, 192, 227, 466, 492, 547, 606

Vegetation Maps: 5, 6, 12, 17, 31, 48, 68, 70, 82, 86, 94,
95, 103, 108, 137, 139, 140, 169, 173, 181, 186, 225,
239, 243, 262, 265, 266, 267, 269, 271b, 290, 291, 292,
293, 294, 299, 301, 322, 323, 329, 336, 337, 350, 351,
353, 361, 369, 376, 388, 400, 401, 403, 404, 409, 461,
468, 479, 486, 487, 512, 521, 522, 533, 538, 544, 547,
562, 563, 568, 582, 586, 588, 589, 597, 623, 631, 647,
660

Vegetation Zonation: 23, 76, 78, 79, 81, 86, 88, 107, 126,
128, 129, 135, 138, 177, 181, 193, 230, 233, 239, 298,
326, 336, 337, 380, 421, 436, 438, 439, 440, 449, 482,
495, 498

County Index

Adams: 351, 447, 592, 593

Alamosa: 138, 193, 450, 451, 457, 458

Arapahoe: 351

Archuleta: 508, 542, 648

Baca: 10, 125, 263, 486, 487, 504, 511, 573

Bent: 309, 310, 311, 312, 316

'Colorado Natural Heritage Inventory, 1550 Lincoln Street, Suite 110, Denver, Colorado 80203.

Great Basin Naturalist

Vol. 43, No. 1

Boulder: 8, 14, 20, ,33, 36, 37, 38, 45, 48, 49, 51, 74, 75,
84, 94, 95, 104, 126, 140, 141, 144, 148, 155, 165, 169,
189, 209, 216, 228, 229, 230, 231, 232, 239, 240, 246,
250, 251, 252, 265, 275, 276, 277, 278, 286, 287, 288,
289, 290, 291, 302, 307, 325, 332, .333, 334, 338, 341,
342, ,343, 344, 345, .346, 347, 348, .351, 367, 368, 369,
370, 377, .381, 382, 391, .392, 393, 397, 403, 406, 407,
409, 417, 418, 419, 420, 421, 422, 425, 447, 448, 459,
464, 466, 478, 479, 484, 513, 526, 5.35, .545, 546, 554,
574, 582, 592, 593, 594, 595, 596, 606, 611, 612, 616,
656

Chaffee: 52, 53, 56, 234, .321, 522, 560

Cheyenne: no references

Clear Creek: 49, 52, 53, 56, 68, 74, 75, 112, 206, 209,
215, 216, 2.39, 291, 421, 446, 448, 594, 595, 596, 616

Conejos: 138, 139, 193, 450, 457, 458

Costilla: 138, 139, 193, 450, 457, 458, 650, 651, 652

Crowley: no references

Custer: 101, 421

Delta: 70, 191, 467, 468, 542, 557, 577, 578

Denver: 351

Dolores: 86, 136, 336, 337, 355, 533, 542

Douglas: 40, 234, 351, ,361, 452, 575, 594, 595, ,596, 616

Eagle: 49, 133, 210, 2,39, 303, 350

Elbert: 161, 351, 361, 619, 620, 621

El Paso: 46, 52, 53, 56, 99, 100, 161, 171, 215, 216, 234,
263, 279, 280, 318, 319, 320, 361, 421, 504, 509, 510,
514, 516, 517, 524, 525, 560, 561, 594, 595, 596, 616,
631, 643, 644, 645, 647

Fremont: 61, 160, 576, 650, 651, 652

Garfield: 58, 59, 70, 82, 133, 157, 158, 177, 204, 208, 213,
239, 253, 266, 267, 269, 270, 271a, 352, 353, 355, 373,
374, 375, 480, 507, 556, ,557, 563, 568/569, 570, 571,
597

Gilpin: 72, 112, 168, 209, 215, 216, 239, 308, 354, 441,
442, 443, 444, 447, 448, 460, 469, 594, 595, 596, 616

Grand: 47, 74, 75, 77, 78, 84, 112, 173, 202, 206, 207,
209, 210, 213, 228, 229, 230, 231, 232, 239, 253, 254,
287, 288, 289, 354, 397, 398, 421, 448, 470, 535, 537,
545, 546, 552, 564, 624, 625, 626, 627, 628, 629, 630,
646

Gunnison: 9, 21, 22, 23, .30, 50, 70, 71, 73, HI, 127, 143,
163, 164, 239, 243, 244, 245, 272, 295, 296, 297, 298,
299, ,300, 301, 321, 327, 328, 329, 340, 385, 413, 414,
415, 506, 537, 542, 557, 572, 598, 600, 649

Hinsdale: 11, 23, 52, 53, 56, 69, 139, 148, 249, 534, 611

Huerfano: 291, 421, 550, 594, 595, 596, 650, 651, 652

Jackson: 2, 74, 75, 77, 78, 209, 213, 228, 229, 2,30, 231,
232, 239, 253, 254, 355, 485, 532, 537, 562, 6,32

Jefferson: 63, 64, 65, 94, 95, 189, 234, 239, 264, 273, 351,

404, 452, 594, 595, 596, 610
Kiowa: 125

Kit Carson: 100, 399, 573
Lake: 215, 216, 359
La Plata: 18, 136, 201, 242, 336, 337, 379, 542, 558, 559,

648
Larimer: 2, 13, 24, 25, 31, 34, 35, 36, 37, 38, 41, 42, 43,
44, 52, 53, 56, 57, 68, 83, 84, 92, 93, 98, 104, 127, 145,
146, 168, 172, 175, 182, 183, 184, 185, 187, 188, 194,
195, 196, 197, 198, 199, 200, 202, 206, 207, 209, 212,
215, 216, 218, 219, 220, 221, 222, 228, 229, 230, 231,
232, 239, 275, 276, 277, 278, 304, 305, 306, 322, 323,
335, 356, 362, 395, 396, 397, 417, 418, 419, 420, 421,
422, 429, 437, 447, 461, 463, 465, 473, 488, 489, 507,
529, 535, 537, 538, 539, 541, 545, 546, 547, 581, 594,

595, ,596, 601, 602, 603, 616, 618, 6,33, 634, 635, 636,

637, 638, 639, 640, 641, 650, 651, 652, 654, 657, 660
Las Animas: 311, 312, 486, 487, 594, 595, 596, 619, 620,

621
Lincoln: 125

Logan: .36, 89, 90, 91, 116, 117, 118, 119, 120, ,364, 505
Mesa: 66, 70, 86, 147, 191, 208, 210, 266, 324, 507, 537,

542, 557, 577, 578, 588, 589
Mineral: 136, 1,39, 542, 548
Moffat: 60, 103, 121, 1,32, 177, 208, 213, 239, 352, ,353,

357, ,358, 659
Montezuma: 16, 19, 134, 150, 151, 152, 153, 355, 426,

427, 428, 542, 617
Montrose: 50, 70, 71, 86, 163, 164, 208, 295, 355, 413,

414, 415, 467, 468, 542, 572, 600, 649
Morgan: 36, 89, 90, 91, 125, 162, 236, 237, 238, 312, 313,

364, 505
Otero:309, 310, 311,312, 316
Ouray: 70
Park: 234, 291, 400, 401, 402, 474, 515, 522, 523, 537,

560, 583, 584, 585, 599, 616
Phillips: no references
Pitkin: 52, 53, 56, 70, 210, 239, 247, 557
Prowers: 309, 310, 311, 312, 316
Pueblo: 160, ,309, 310, 311, 312, 316, 594, 595, 596
Rio Blanco: 58, 59, 82, 158, 177, 204, 208, 210, 213, 239,

253, 254, 268, 271b, 281, 282, 352, ,353, 355, 373, 374,

375, 480, 507, 556, 563, 570, 571, 597, 604, 605, 623
Rio Grande: 52, 53, 56, 138, 139, 193, 450, 457, 458
Routt: 77, 78, 79, 80, 213, 239, 248, 355, 553, 556, 632
Saguache: 23, 32, 127, 138, 139, 193, 291, 339, 450, 457,

458, 522, 650, 651, 652
San Juan: 11, 148, 3,36, ,337, 372, 534, 537, 611
San Miguel: 86, 208, 355, 371, 507, 542
Sedgewick: 89, 90, 91, 312, 313, 364, 399, 505, 573
Summit: 49, 52, 53, 56, 123, 167, 173, 174, 209, 234, 239,

285, 359, 360, 400, 401, 402, 537
Teller: 171, 234, 255, 256, 257, 258, 259, 260, 261, 361,

435, 509, 510, 530, 560, 616, 631
Washington: 124, 154, 180, 491, 503, 504, 519, 527
Weld: 15, 36, 54, 55, 84, 89, 90, 91, 105, 162, 166, 176,

223, 224, 236, 237, 238, 241, 283, 284, 312, 313, 314,

330, 331, 364, 366, 378, 383, 384, 399, 405, 447, 454,

455, 456, 471, 472, 477, 505, 521, 527, 555, 592, 593,

594, 653
Yuma: 36, 125, 154, 389, 390, 399
Statewide Studies: 1, 3, 4, 7, 12, 88, 102, 106, 107, 109,

128, 129, 135, 142, 149, 170, 178, 179, 181, 214, 262,

294, 326, 365, 376, 380, 387, 388, 424, 438, 439, 445,

447, 449, 481, 482, 492, 493, 494, 495, 496, 497, 498,

499, 500, 501, 502, 512, 520, 531, 540, 565, 566, 586,

587, 607, 613, 614, 615
Regional Studies:

Plains: 39, 87, 110, 115, 205, 235, 274, 433, 454, 549,
608, 609

Mountains: 5, 6, 17, 19, 26, 28, 29, 76, 96, 108, 122,
139, 211, 227, 349, 386, 412, 416, 433, 580

Western Colorado Plateaus: 225, 226, 416, 480, 622,
658

Bibliography

1. Aldous, A. E., and H. L. Shantz. 1924. Types of
vegetation in the semiarid portion of the United States
and their economic significance. J. Agr. Res. 28: 99-128.

January 1983

Baker: Colorado Bibliography

2. Alexander, M. E. 1980. Four fire scar records on
lodgepole pine (Pinus contorta Dougl.) in north-central
Colorado. S. W. Natur. 25:432-434.

3. Alexander, M. E., and D. V. Sandberg. 1976. Fire
ecology and historical fire occurrence in the forest and
range ecosystems of Colorado: a bibliography. Colo. St.
Univ., Dept. of For. and Wood Sci., Occas. Report. 17
pp.

4. Alexander, M. E., and D. V. Sandberg. 1979. Fire
ecology and historical fire occurrence in the forest and
range ecosystems of Colorado: a bibliography, supple-
ment No. 1. Colo. St. Univ., Dept. of For. and Wood
Sci., Occas. Report. 4 pp.

5. Alexander, R. R. 1974a. Silviculture of central and
southern Rocky Mountain forests: a summary of the sta-
tus of our knowledge by timber types. USDA For. Serv.
Res. Paper RM-120, Rocky Mt. For. and Range Expt.
Stn., Fort Collins, Colo. 36 pp.

6. Alexander, R. R. 1974b. Silviculture of subalpine
forests in the central and southern Rocky Mountains: the
status of our knowledge. USDA For. Serv. Res. Paper
RM-121, Rocky Mt. For. and Range Expt. Stn., Fort
Collins, Colo. 88 pp.

7. Allison, E. M. 1908. Bibliography and history of
Colorado botany. Univ. of Colo. Studies 6: 51-76.

8. Amundsen, C. C. 1967. The subalpine forest of
Wild Basin, Front Range, Colorado. Unpubl. dis-
sertation, Univ. of Colo., Boulder. 129 pp.

9. Andersen, D. C, R. S. Hoffman, and K. B. Armi-
tage. 1979. Aboveground productivity and floristic struc-
ture of a high subalpine herbaceous meadow. Arct. and
Alp. Res. 11:467-476.

10. Anderson, J. M., Jr. 1950. A flora of Baca County,
Colorado. Unpubl. thesis, Univ. of Colo., Boulder. 39 pp.

11. Andrews, J. T., P. E. Carrara, F. B. King, and R.
Stuckenrath. 1975. Holocene environmental changes in
the alpine zone, northern San Juan Mountains, Colo-
rado: evidence from bog stratigraphy and palynology.
Quat. Res. 5:173-197.

12. Armstrong, D. M. 1972. Distribution of Mammals
in Colorado. Mus. Nat. Hist., Univ. of Kansas, Monogr.
No. 3. [Vegetation: p. 26-31].

13. Ashton, R. E. 1930. Preliminary observations on
revegetation of the Twinsisters Bum in Rocky Mountain
National Park. J. Colo.-Wyom. Acad. Sci. 1(2): 13
(Abstract).

14. Bachrach, J. H. 1951. The plant communities of a
valley in the montane forest of the Colorado Front
Range. Unpubl. thesis, Univ. of Colo., Boulder. 68 pp.

15. Badaracco, R. J. 1971. An interpretive resource
analysis of Pawnee Buttes, Colorado. Unpubl. dis-
sertation, Colo. St. Univ., Fort Collins. 341 pp.

16. Bader, E. H. 1932. The vegetation of the Mesa
Verde National Park, Colorado. Unpubl. thesis, Univ. of
Colo., Boulder. 64 pp.

17. Bailey, D. K. 1970. Phytogeography and taxonomy
of Pinus subsection Balfourianae. Ann. Missouri Bot.
Card. 57:210-249.

18. Baker, C. F. 1898. Botanizing in the La Plata
Mountains. Plant World 2:29-32.

19. Baker, F. S. 1925. Aspen in the central Rocky
Mountain region. USDA Bull. 1291. 46 pp.

20. Barber, R. A. 1935. An ecological study of a small,
montane, spring-fed, stream area in the University of
Colorado camp, on Mount Niwot. Unpubl. thesis, Univ.
of Colo., Boulder.

21. Barclay, H. G. 1941a. Plant communities of the
Gothic area and their successional relations. Bull. Ecol.
Soc. Am. 22:34 (Abstract).

22. Barclay, H. G. 1941b. A subalpine climax grass-
land in west central Colorado. Bull. Ecol. Soc. Am.
22:35 (Abstract).

23. Barren, J. 1969. Flora of the Gunnison Basin:
Gunnison, Saguache, and Hinsdale counties. Natur.
Land Inst., Rockford, 111.

24. Barrows, J. S., E. W. Mogren, K. Rowdabaugh,
and R. Yancik. 1977. The role of fire in ponderosa pine
and mixed conifer ecosystems. Final report, Co-op
Agreem. with U.S. Nat. Park Serv. (CX-1200-7-8011) and
Rocky Mt. For. and Range Exp. Stn. (16-542-CA), on file
at Rocky Mt. For. and Range Expt. Stn., Fort Collins,
Colo. 101 pp.

25. Barth, R. C. 1970. Revegetation after a subalpine
wildfire. Unpubl. thesis, Colo. St. Univ., Fort Collins.
142 pp.

26. Bates, C. G. 1917. Forest succession in the central
Rocky Mountains. J. For. 15:587-592.

27. Bates, C. G. 1923. The transect of a mountain val-
ley. Ecology 4: 54-62.

28. Bates, C. G. 1924. Forest types in the central
Rocky Mountains as affected by climate and soil. USDA
Bull. No. 1233. 152 pp.

29. Bates, C. G., F. B. Notestein, and P. Keplinger.
1914. Climatic characteristics of forest types in the cen-
tral Rocky Mountains. Proc. Soc. Am. For. 9:78-94.

30. Bathke, D. M. 1968. The flora of the subalpine
and alpine zones in Robinson Basin, Colorado. Unpubl.
thesis. Western St. College, Gunnison, Colo. 92 pp.

31. Baumann, T. G. 1978. Winter ecology of bighorn
sheep in the Mummy Range, Colorado. Unpubl. thesis,
Colo. St. Univ., Fort Collins.

32. Beck, R. F. 1965. Pocket gopher habitat relations
in mountain rangelands in Colorado. Unpubl. thesis,
Colo. St. Univ., Fort Collins. 61 pp.

33. Beidleman, R. G. 1948. The vertebrate ecology of
a Colorado plains cottonwood river bottom. Unpubl.
thesis, Univ. of Colo., Boulder.

34. Beidleman, R. G. 1952. The ponderosa-grassland
ecotone in north central Colorado. J. Colo.-Wyom.
Acad. Sci. 4(4):70 (Abstract).

35. Beidleman, R. G. 1953. The island of pines. Liv-
ing Wilderness 46:7-10.

36. Beidleman, R. G. 1954. The cottonwood river-bot-
tom community as a vertebrate habitat. Unpubl. dis-
sertation, Univ. of Colo., Boulder. 358 pp.

37. Beidleman, R. G. 1957. Fire bum succession stud-
ies in the northern Colorado Rockies. J. Colo.-Wyom.
Acad. Sci. 4(9):39 (Abstract).

38. Beidleman, R. G. 1967. Twenty-year successional
studies in northern Colorado. J. Colo.-Wyom. Acad. Sci.
5(8):63-64 (Abstract).

39. Beidleman, R. G. 1978. The cottonwood-willow
riparian ecosystem as a vertebrate habitat, with particu-
lar reference to birds. Pages 192-195 in W. D. Graul and
S. J. Bissell, eds.. Lowland river and stream habitat in
Colorado: a symposium. (Greeley, Colo. 4-5 Oct. 1978).
Colo. Chap. Wildl. Soc. & Colo. Audubon Council.

40. Beidleman, R. G., and W. Fischer. 1979. Environ-
mental reconnaissance study of Perry Park Ranch,
Douglas County, Colorado. Wright-Ingram Institute,
Colorado Springs, Colo.

Great Basin Naturalist

Vol. 43, No. 1

41. Bell, K. L. 1974a. Autecology of Kobresia bel-
lardii: why winter snow accumulation patterns affect lo-
cal distribution. Unpubl. dissertation, Univ. of Alberta,
Edmonton. 167 pp.

42. Bell, K. L. 1974b. Autumn, winter, and spring
phenology of some Colorado alpine plants. Am. Midi.
Natur. 91:460-464.

43. Bell, K. L. 1977. Autecology of Kobresia beUardii:
why winter snow accumulation affects local distribution.
Bull. Ecol. Soc. Am. 58(2):41 (Abstract).

44. Bell, K. L., and L. C. Bliss. 1979. Autecology of
Kobresia bellardii: why winter snow accumulation limits
local distribution. Ecol. Monogr. 49: 377-402.

45. Benedict, J. B. 1977. A master plan for the Bunker
Hill Placer Preserve, Boulder County, Colorado. Un-
publ. report on file at The Nature Conservancy, Denver,
Colo. 61 pp.

46. Bichel, M. A. 1959. Investigations of a Nebraska
and a Colorado prairie and their impact on the relict
concept. Unpubl. dissertation, Univ. of Nebraska, Lin-
coln. 180 pp.

47. Bierly, K. F. 1972. Meadow and fen vegetation in
Big Meadows, Rocky Mountain National Park. Unpubl.
thesis, Colo. St. Univ., Fort Collins, Colo. 102 pp.

48. Biggins, J. and M. H. Dodson. 1970. Marshall
Mesa natural area study. Unpubl. report, Dept. Geogr.,
Univ. of Colo., Boulder. 54 pp.

49. Bollinger, W. H. 1973. The vegetation patterns af-
ter fire at the alpine forest-tundra ecotone in the Colo-
rado Front Range. Unpubl. dissertation, Univ. of Colo.,
Boulder. 2 vols.

50. Bond, H. L. 1959. Revegetation and disintegration
of pocket gopher mounds on Black Mesa, Colorado. Un-
publ. thesis, Colo. St. Univ., Fort Collins.

51. Bonde, E. K. 1967. Pinus aristata (Pinaceae) in
Boulder County, Colorado: northward extension of its
known Colorado range. S.W. Natur. 12:474-475.

52. Bonham, C. D. 1966. An ordination of alpine hair-
grass (Deschampsia caespitosa Beauv.) meadows. Un-
publ. dissertation, Colo. St. Univ., Fort Collins. 124 pp.

53. Bonham, C. D. 1972. Vegetation analysis of
grazed and ungrazed alpine hairgrass meadows. J. Range
Mgmt. 25: 276-279.

54. Bonham, C. D., and J. S. Hannan. 1978. Blue
grama and buffalograss patterns in and near a prairie
dog town. J. Range Mgmt. 31:63-65.

55. Bonham, C. D., and A. Lerwick. 1976. Vegetation
changes induced by prairie dogs on shortgrass range. J.
Range Mgmt. 29: 221-225.

56. Bonham, C. D., and R. T. Ward. 1970. Phytoso-
ciological relationships in alpine tufted hairgrass {Des-
chanvpsia caespitosa (L.) Beauv.) meadows. Arct. and
Alp. Res. 2:267-275.

57. Bowes, A. L. 1958. Ecological analysis of mule
deer winter range. Cache la Poudre canyon, Colorado.
Unpubl. thesis, Colo. St. Univ., Fort Collins.

58. Boyce, D. A. 1976. A preliminary description of
the ecosystems adjacent to the South Fork of the White
River, Rio Blanco, and Garfield counties, Colorado. J.
Colo.-Wyom. Acad. Sci. 8(1):45 (Abstract).

59. Boyce, D. A. 1977. Vegetation of the South Fork
of the White River Valley, Colorado. Unpubl. dis-
sertation, Univ. of Colo., Boulder.

60. Bradley, R. A. W. 1950. The vascular flora of Mof-
fat County, Colorado. Unpubl. thesis, Univ. of Colo.,
Boulder. 71 pp.

61. Brandegee, T. S. 1878. Notes from Colorado. Bot.
Gaz. 3:166.

62. Brandegee, T. S. 1880. Timber line in the
Sawatch Range. Bot. Gaz. 5:125-126.

63. Branson, F. A., R. F. Miller, and I. S. McQueen.
1961. Soil-water availability and use by grasslands on ad-
jacent stony and shale-derived soils in Colorado. Short
papers in the geologic and hydrologic sciences. USGS
Prof. Paper 424C:251-253.

64. Branson, F. A., R. F. Miller, and I. S. McQueen.

1964. Effects of two kinds of geologic materials on plant
communities and soil moisture. Am. Soc. Agron. Spec.
Publ. No. 5:165-175.

65. Branson, F. A., R. F. Miller, and I. S. McQueen.

1965. Plant communities and soil moisture relationships
near Denver, Colorado. Ecology 46:311-319.

66. Branson, F. A., R. F. Miller, and L S. McQueen.
1976. Moisture relationships in twelve northern desert
shrub communities near Grand Junction, Colorado.
Ecology 57:1104-1124.

67. Branson, F. A., and J. B. Owen. 1970. Plant cover,
runoff, and sediment yield relationships on Mancos shale
in western Colorado. Water Resources Res. 6:783-790.

68. Braun, C. E. 1969. Population dynamics, habitat,
and movements of white-tailed ptarmigan in Colorado.
Unpubl. dissertation, Colo. St. Univ., Fort Collins.

69. Brewster, J. F. 1968. Ecological factors in the
revegetation of SlumguUion Mudflow near Lake City,
Colorado. Unpubl. thesis, Univ. of Illinois, Urbana. 94
pp.

70. Brown, H. E. 1958. Gambel oak in west-central
Colorado. Ecology 39:317-327.

71. Brown, H. E., and J. R. Thompson. 1965. Summer
water use by aspen, spruce, and grassland in western
Colorado. J. For. 63:756-760.

72. Bruderlin, K. 1911. A study of the lodgepole-pine
forests of Boulder Park (Tolland, Colorado). Univ. of
Colo. Studies 8:265-275.

73. Buck, P. 1960. Vegetational succession in sub-
alpine ponds in the Rockies. Proc. Okla. Acad. Sci.
40:2-6.

74. Buckner, D. L. 1976. Occurrence and mainte-
nance of ribbon forest in the Colorado Rocky Moun-
tains. J. Colo.-Wyom. Acad. Sci. 8(1):46 (Abstract).

75. Buckner, D. L. 1977. Ribbon forest development
and maintenance in the central Rocky Mountains of
Colorado. Unpubl. dissertation, Univ. of Colo., Boulder.
224 pp.

76. Bujakiewicz, A. 1975. Vegetational zonation in
the Front Range of the Rocky Mountains, Colorado.
USA [in Polish] Frag. Florist, et Geobot. 21:99-142.

77. Bunin, J. E. 1974. The vegetation of the west
slope of the Park Range, Colorado. J. Colo.-Wyom.
Acad. Sci. 7(5):30 (Abstract).

78. Bunin, J. E. 1975a. The vegetation of the west
slope of the Park Range, Colorado. Unpubl. dissertation,
Univ. of Colo., Boulder. 2 pp.

79. Bunin, J. E. 1975b. Aspen forests of the west slope
of the Park Range, north-central Colorado. Paper pre-
sented at the 1975 AIBS/ESA Meetings, 17-22 Aug.
1975, Oregon St. Univ., Corvallis. 22 pp.

80. Bunin, J. E. 1975c. Aspen forests of the west slope
of the Park Range, northcentral Colorado. Bull. Ecol.
Soc. Am. 56(2):48 (Abstract).

January 1983

Baker: Colorado Bibliography

81. Bums, S. F. 1980. A comparison of mountain life
zones of the Swiss Alps with the Colorado Front Range.
J. Colo.-Wyom. Acad. Sci. 12(1):31 (Abstract).

82. Butler, J. R. and J. L. England. 1979. Vegetation
map of the southeastern Uinta Basin, Utah and Colo-
rado, uses Misc. Investig. Series, Map 1-1141.

83. Butter)^', R. F. 1955. Range conditions and trends
resulting from winter concentrations of elk in Rocky
Mountain National Park, Colorado. Unpubl. thesis,
Colo. St. Univ., Fort Collins. 117 pp.

84. Cacek, T. L. 1973. An ecological interpretation of
north central Colorado. Unpubl. dissertation, Colorado
St. Univ., Fort Collins. 261 pp.

85. Cancalossi, J. 1979. Biological evaluation for Coal
Creek and Crescent damsites, Colorado. Unpubl. report
prepared for USDl, Bur. of Reel., Denver, Colo.

86. Cannon, H. L. 1960. The development of bo-
tanical methods of prospecting for uranium on the Colo-
rado Plateau. USGS Bull. 1085-A. 50p.

87. Carpenter, J. R. 1940. The grassland biome. Ecol.
Monogr. 10:617-684.

88. Cary, M. 1911. A biological survey of Colorado.
USDA Bur. Biol. Survey, No. Am. Fauna No. 33. 256 pp.

89. Christy, S. J. 1972a. Woody vegetation along the
South Platte River in northeastern Colorado. J. Colo.-
Wyom. Acad. Sci. 7(3): 106 (Abstract).

90. Christy, S. J. 1972b. Plant communities of the
South Platte River floodplain in Colorado. J. Colo.-
Wyom. Acad. Sci. 7(2):31 (Abstract).

91. Christy, S. 1973. An analysis of the woody vegeta-
tion on the South Platte River flood plain in north-
eastern Colorado. Unpubl. thesis, Univ. of N. Colo.,
Greeley. 82 pp.

92. Clagg, H. B. 1975. Fire ecology in high-elevation
forests in Colorado. Unpubl. thesis, Colo. St. Univ., Fort
Collins. 137 pp.

93. Clagg, H. R. and D. R. Stevens. 1974. Fire man-
agement in Rocky Mountain National Park. Tall Tim-
bers Fire Ecol. Conf. No. 14:77-86.

94. Clark, S. J. V. 1977. The vegetation of Rocky
Flats, Colorado. Unpubl. thesis, Univ. of Colo., Boulder.

95. Clark, S. V., P. J. Webber, V. Komarkova, and W.
A. Weber. 1980. Map of mixed prairie grassland vegeta-
tion-Rocky Flats, Colorado. Univ. of Colo., Inst, of Arct.
and Alp. Res. Occas. Paper No. 35. 66 pp.

96. Clary, W. P. 1978. Arizona fescue mountain
rangelands. Pages 205-207 in D. N. Hyder, ed., Proc. 1st
Intemat. Rangeland Congr., Denver, Colo., USA— 14-18
Aug. 1978, Soc. for Range Mgmt., Denver.

97. Clements, F. E. 1904. Formation and succession
herbaria. Univ. of Nebr., Univ. Studies IV(4):329-355.

98. Clements, F. E. 1910. The life history of lodge-
pole bum forests. USDA For. Serv. Bull. No. 79. 56 pp.

99. Clements, F. E., and G. W. Goldsmith. 1924. The
phytometer method in ecology: the plant and commu-
nity as instruments. Camegie Inst, of Wash. Publ. No.
356 [Climaxes and climates of the Pike's Peak region,
pp. 14-16].

100. Clements, F. E., and J. E. Weaver. 1924. Experi-
mental vegetation: the relation of climaxes to climates.
Camegie Inst, of Wash. Pub. No. 355. 172 pp.

101. Cockerell, T. D. A. 1891. Notes on the flora of
high altitudes in Custer County, Colorado. Bull. Torr.
Bot. Club 18:167-174.

102. Cockerell, T. D. A. 1906. The alpine flora of Col-
orado. Am. Nat. 40:861-873.

103. Colorado River Water Conservation District.
1980. Application for license: project No. 2757, Juniper-
Cross Mountain Project, Exhibit W-Environmental
Report.

104. Cooper, W. S. 1908. Alpine vegetation in the vi-
cinity of Long's Peak, Colorado. Bot. Gaz. 45:319-337.

105. Costello, D. F. 1944a. Natural revegetation of
abandoned plowed land in the mixed prairie association
of northeastern Colorado. Ecology 25:312-326.

106. Costello, D. F. 1944b. Important species of the
major forage types in Colorado and Wyoming. Ecol.
Monogr. 14:107-134.

107. Costello, D. F. 1954. Vegetation zones in Colo-
rado. Pages iii-x in H. D. Harrington, ed.. Manual of the
plants of Colorado. Sage Books, Denver, Colo.

108. Costello, D. F., and H. E. Schwan. 1946. Condi-
tions and trends on ponderosa pine ranges in Colorado.
USDA For. Serv. Mimeo. 33 pp.

109. Costello, D. F., and G. T. Turner. 1941. Vegeta-
tion changes following exclusion of livestock from
grazed ranges. J. For. 39:310-315.

110. Costello, D. F., and G. T. Turner. 1944. Judging
condition and utilization of short-grass ranges on the
central Great Plains. USDA Farmer's Bull. No. 1949. 21
pp.

111. Cox, B. J. 1968. A vegetational comparison of the
Gothic and Galena Mountain area. Trans. Mo. Acad.
Sci. 2:72-83.

112. Cox, C. F. 1933. Alpine plant succession on
James Peak, Colorado. Ecol. Monogr. 3:299-372.

113. Crandall, C. S. 1897a. Natural reforestation on
the mountains of northern Colorado. I. Garden and for-
est 10 (506): 437.

114. Crandall, C. S. 1897b. Natural reforestation on
the mountains of northern Colorado. II. Garden and for-
est 10 (507):446-447.

115. Cringan, A. T., and R. L. Dix. 1975. Ecological
theme analysis and inventory of potential natural land-
marks of the Great Plains Natural Region. Report to the
National Park Service, Contract No. 14-10-9-900-248
with Colo. St. Univ., Dept. of Fishery and Wildlife Biol-
ogy, and Dept. of Botany and Plant Pathology. 465 pp.

116. Crouch, G. L. 1961a. Inventory and analysis of
wildlife populations and habitat, South Platte River Val-
ley. Final report. Fed. Aid in Wildlife Restor., Project
W-104-R-1-2, Colo. Game & Fish Dept. 68 pp.

117. Crouch, G. L. 1961b. Wildlife populations and
habitat conditions on grazed and ungrazed bottomlands
in Logan County, Colorado. Unpubl. thesis, Colo. St.
Univ. Fort Collins, Colo.

118. Crouch, G. L. 1978. Effects of protection from
livestock grazing on a bottomland wildlife habitat in
northeastern Colorado. Pages 118-125 in W. D. Graul
and S. J. Bissell, eds.. Lowland river and stream habitat
in Colorado: a symposium. (Greeley, Colo., 4-5 Oct.
1978). Colo. Chapt. Wildl. Soc. and Colo. Audubon
Council.

119. Crouch, G. L. 1979a. Long-term changes in cot-
tonwoods on a grazed and an ungrazed plains bot-
tomland in northeastern Colorado. USDA For. Serv. Res.
Note RM-370. Rocky Mt. For. & Range Exp. Stn., Fort
Collins, Colo. 4 pp.

Great Basin Naturalist

Vol. 43, No. 1

120. Crouch, G. L. 1979b. Changes in the vegetation
complex of a cottonwood ecosystem on the South Platte
River. Pages 19-22 in Riparian and wetland habitats of
the Great Plains: proceeding of the 31st annual meeting.
Great Plains Agric. Council, Forestry Committee, Gt.
Plains Agric. Counc. Publ. No. 91, Rocky Mt. For. &
Range Expt. Stn., Fort Collins, Colo.

121. Cunningham, H. 1971. Soil-vegetation relation-
ships of a bitterbrush-sagebrush association in north-
western Colorado. Unpubl. thesis, Colo. St. Univ., Fort
Collins. 94 pp.

122. Currie, P. O. 1975. Grazing management of pon-
derosa pine-bunchgrass ranges of the central Rocky
Mountains: the status of our knowledge. USDA For.
Serv. Res. Paper RM-159, Rocky Mt. For. & Range Expt.
Stn., Fort Collins. 24 pp.

123. Curry, R. R. 1962. Geobotanical correlations in
the alpine and subalpine regions of the Tenmile Range,
Summit County, Colorado. Unpubl. thesis, Univ. of
Colo., Roulder. 123 pp -I- map.

124. Dahl, B., and J. J. Norris. 1965. The effect of
grazing intensity on native sandhill ranges. Eastern Col-
orado Range Station. Colo. Agric. Expt. Sta. Prog. Rep.
162. 2 pp.

125. Daley, R. H. 1972. The native sand sage vegeta-
tion of eastern Colorado. Unpubl. thesis, Colo. St. Univ.,
Fort Collins. 62 pp.

126. Daniels, F. P. 1911. The flora of Boulder, Colo-
rado, and vicinity. Univ. Missouri Studies, Sci. Series
11(2). [Vegetation: p. 8-48].

127. Dart, D. J. 1977. AUelopathic effects of Artemisia
tridentata ssp. vaseyana (Rydb.) Beetle and aspects of its
ecology in Colorado. Unpubl. thesis, Univ. of Colo.,
Boulder. 85 pp.

128. Daubenmire, R. F. 1943a. Vegetational zonation
in the Rocky Mountains. Bot. Rev. 9:325-393.

129. Daubenmire, R. F. 1943b. Soil temperature ver-
sus drought as a factor determining lower altitudinal
limits of trees in the Rocky Mountains. Bot. Gaz.
105:1-13.

130. Davis, M. L. 1981. Habitat diversity and its ef-
fects on the genetic and ecological structures of two suc-
cessional populations of Douglas-fir. Unpubl. dis-
sertation, Univ. of Colo., Boulder.

131. Dawson, R. E., E. Hotton, I. Takahashi, and C.
Doty. 1979. Natural revegetation of a mixed-grass
prairie. J. Colo.-Wyom. Acad. Sci. 11(1):38 (Abstract).

132. Dean, J. S., and D. O. Bowden, III. 1976. Den-
drochronological analysis of wood samples from living
trees in the Skull Creek Basin, Colorado. Final report
prepared for the Craig, Colorado District Office, Bureau
of Land Management by Laboratory of Tree-Ring Re-
search, Univ. of Arizona, Tucson, Ariz.

133. De Leuw, Gather and Company. 1977. Vegeta-
tion sampling (Glenwood Canyon). Unpubl. report pre-
pared for Colo. Div. of Highways, Glenwood Springs,
Colo., by De Leuw, Gather, and Company, Consulting
Engineers and Planners, Chicago, 111.

134. Dillinger, K. C. 1970. Evaluation of mule deer
habitat in Mesa Verde National Park. Unpubl. thesis,
Colo. St. Univ., Fort Collins. 40 pp.

135. Dix, R. L. 1974. Regional ecological systems of
Colorado. Pages 7-17 in P. O. Foss, ed.. Environment
and Colorado: a handbook. Env. Resources Center,
Colo. St. Univ., Fort Collins.

136. Dix, R. L., and J. D. Richards. 1976. Possible
changes in species structure of the subalpine forest in-
duced by increased snowpack. Pages 311-322 in H. W.
Steinhoff and J. D. Ives, eds.. Ecological impacts of
snowpack augmentation in the San Juan Mountains,
Colorado. Final Report, San Juan Ecology Project, Colo.
St. Univ., Fort Collins, to the USDI Bur. Reel., Div. of
Atmos. Wat. Res. Mgmt., Denver, Colo.

137. Dixon, H. N. 1969. The growth of lodgepole pine
in the Colorado Front Range as related to environment.
Unpublished dissertation, Univ. of Colorado, Boulder.
120 pp.

138. Dixon, H. N. 1971. Flora of the San Luis Valley.
Pages 133-136 in New Mexico Geol. Soc. 22nd Field
Conference Guidebook.

139. Dortignac, E. J. 1956. Watershed resources and
problems of the Upper Rio Grande Basin. Unpubl. re-
port on file at the Rocky Mt. For. & Range Expt. Stn.,
Fort Collins, Colo.

140. Douglass, M. M. 1954a. Ecology of forest succes-
sion on a ridge in the subalpine forest of northern Colo-
rado. Unpubl. thesis, Univ. of Colo., Boulder. 134 pp.

141. Douglass, M. M. 1954b. Subalpine fir succession
on a ridge in Boulder County, Colorado. J. Colo.-Wyom.
Acad. Sci. 4(6):45-46 (Abstract).

142. Douglass, M. M., and J. R. Douglass. 1955. The
distribution and growth of the limber pine in Colorado.
J. Colo.-Wyom. Acad. Sci. 4(7):46-47 (Abstract).

143. Dunlap, N. J. 1962. A comparison of vegetation
at Crested Butte and Gunnison, Colorado. Unpubl.
thesis. Western State College, Gunnison, Colo.

144. Ebert, D. C. 1972. Primary productivity as a
function of vegetation composition in an alpine tundra
environment. J. Colo.-Wyom. Acad. Sci. 7(2):27-28
(Abstract).

145. Eddleman, L. E. 1963. Abrupt transitions in al-
pine communities. J. Colo.-Wyom. Acad. Sci. 5(4):39-40
(Abstract).

146. Eddleman, L. E. 1967. A study of phyto-edaphic
relationships in alpine tundra of northern Colorado. Un-
publ. dissertation, Colo. St. Univ., Fort Collins. 148 pp.

147. Ellis, S. L., T. Shoemaker, and R. Sanz. 1979. In-
ventories of plants, birds, mammals, reptiles, and am-
phibians of the Unaweep Canyon Springs, Mesa County,
Colorado. Unpubl. report prepared for the Colorado
Natural Areas Program, Department of Natural Re-
sources, State of Colorado, Denver. 25 pp.

148. Emerick, J. C. 1976. Effects of artificially in-
creased winter snow cover on plant canopy architecture
and primary production in selected areas of Colorado al-
pine tundra. Unpubl. dissertation, Univ. of Colo., Boul-
der. 193 pp.

149. Ensign, E. T. 1888. Report on the forest condi-
tions of the Rocky Mountains. USDA, Forestry Div.,
Bull. No. 2 [Colorado: p. 115-137].

150. Erdman, J. A. 1962. Ecology of the pinyon-juni-
per woodland of Wetherill Mesa, Mesa Verde National
Park, Colorado. Unpubl. thesis, Univ. of Colo., Boulder.
109 pp.

151. Erdman, J. A. 1964. Climatic differences of the
Mesa Verde, and their effects on its vegetation. Bull.
Ecol. Soc. Am. 45(3):89 (Abstract).

152. Erdman, J. A. 1969. Pinyon-juniper succession
after fires on residual soils of the Mesa Verde, Colorado.
Unpubl. dissertation, Univ. of Colo., Boulder. 81 pp.

January 1983

Baker: Colorado Bibliography

153. Erdman, J. A. 1970. Pinyon-juniper succession
after natural fires on residual soils of Mesa Verde, Colo-
rado. Brigham Young Univ. Sci. Bull., Biol. Ser. 11(2):
1-26.

154. Evans, K. E. 1964. Habitat evaluation of the
Greater Prairie Chicken in Colorado. Unpubl. thesis,
Colo. St. Univ., Fort Collins. 98 pp.

. 155. Fareed, M., and M. M. Caldwell. 1975. Pheno-
logical patterns of two alpine tundra plant populations
on Niwot Ridge, Colorado. N.W. Sci. 49:17-23.

156. Ferchau, H. A. 1973a. Vegetation. In G. L.
Hayes, ed.. Environmental analysis and potential im-
pacts of proposed developments at Marble Sid Area. Un-
publ. report, Thome Ecol. Inst., Boulder, Colo.
■ 157. Ferchau, H. A. 1973b. Vegetation inventory
analysis and impact study of the Parachute Creek area,
Garfield County, Colorado. Part II, Vol. I, Chap.
VI: 1-77 in Colony environmental study. Unpubl. report
prepared for Colony Development Operation, Atlantic
Richfield Co., Operator, Denver, Colo, by Thome Ecol.
Inst., Boulder, Colo.

158. Ferchau, H. A. 1974. An ecological analysis of
the vegetation, Piceance Creek Basin, Rio Blanco and
Garfield counties, Colorado. V. 14. 19 pp. In Regional
oil shale study. Unpubl. report to the State of Colo, by
Thome Ecol. Inst., Boulder, Colo.

159. Ferchau, H. A., and B. McLarty, 1980. O.C.
Mine Environmental Report. Unpubl. report prepared
for O.C. Mine, Gunnison, Colo. 24 pp.

160. Fink, B. 1907. A round trip between Iowa and
Puget Sound. I. Westward bound. Plant World
10:49-58.

161. Fish, E. B. 1966. Secondary succession on upper
Kiowa Creek watershed. Unpubl. thesis, Colo. St. Univ.,
Fort Collins, Colo. 102 pp.

162. Fitzgerald, J. P. 1978. Vertebrate associations in
plant communities along the South Platte River in
northeastern Colorado. Pages 73-88 in W. D. Graul and
S. J. Bissell, eds.. Lowland river and stream habitat in
Colorado: a symposium. (Greeley, Colo., 4-5 Oct. 1978).
Colo. Chap. Wildl. Soc. & Colo. Audubon Council.

163. Flowers, S. 1962. Vegetation of Morrow Point
and Blue Mesa Reservoir basins of the upper Gunnison
River, Colorado. Pages 12-46 in A. M. Woodbury, ed..
Ecological studies of the flora and fauna of the Cure-
canti Reservoir basins, western Colorado. Univ. of Utah,
Dept. of Anthropol., Anthropol. Papers No. 59 (Upper
Colo. Ser. No. 8).

164. Flowers, S., H. H. Hall, and R. N. Reynolds.
1962. Plants found in Curecanti Reservoir basins, 1961.
Pages 47-102 in A. M. Woodbury, ed.. Ecological stud-
ies of the flora and fauna of the Curecanti Reservoir ba-
sins, western Colorado. Univ. of Utah, Dept. of Anthro-
pol., Anthropol. Papers No. 59 (Upper Colo. Series No.
8).

165. Forsyth, J. K. 1980. Principal components analy-
sis of subalpine vegetation on the east slope of the Colo-
rado Rocky Mountains. Unpubl. dissertation, Univ. of
Colo., Boulder. 101 pp.

166. Fraley, L., Jr. 1971. Response of shortgrass plains
vegetation to chronic and seasonally administered gam-
ma radiation. Unpubl. dissertation, Colo. St. Univ., Fort
Collins.

167. Fritz, R. J. 1981. Alpine vegetational patterns
around isolated tree islands on the eastern and western
slopes of the Tenmile Range, Summit County, Colorado.
Unpubl. thesis, Univ. of Colo., Boulder. 233 pp.

168. Fuller, G. D. 1930. A comparison of certain
Rocky Mountain grasslands with the prairie of Illinois.
Trans. 111. Acad. Sci. 8:121-130.

169. Gable, D. J. 1978. Vegetation map of the Ward
Quadrangle, Boulder County, Colorado. USGS Misc.
Field Studies Map MF-940.

170. Gannett, H. 1878. Grazing lands of Colorado.
Ch. 5 of "Report on the arable and pasture lands of Col-
orado." In F. V. Hayden, ed.. Tenth Ann. Rep. of the
U.S. Geol. and Geogr. Surv. of the Terr., embracing Col-
orado and parts of adjacent territories, U.S. Govt. Print.
Off., Washington, D.C.

171. Gardner, W. J. 1905. Results of a Rocky Moun-
tain forest fire studied fifty years after its occurrence.
Proc. Soc. Am. For. 1:102-109.

172. Geist, J. M. 1965. Site-vegetation relationships in
the montane zone of northern Colorado. Unpubl. thesis,
Colo. St. Univ., Fort Collins. 104 pp.

173. Giese, T. G. 1975. The ecology of the Middle
Blue River Valley, Summit County, Colorado, with an
analysis of modifications due to powerline construction.
Unpubl. thesis, Univ. of Colo., Boulder. 109 pp.

174. Giese, T. G., and J. W. Marr. 1973. Shale barrens
of Summit County, Colorado. J. Colo.-Wyoming Acad.
Sci. 7(4):26-27 (Abstract).

175. Giunta, B. C. 1968. Coactions of bitterbmsh,
ponderosa pine, and herbaceous vegetation. Unpubl.
thesis, Colo. St. Univ., Fort Collins.

176. Goodding, L. N. 1923. An interesting area of lim-
ber pine extending into southwestern Nebraska. J. For.
21:175-176.

177. Graham, E. H. 1937. Botanical studies in the
Uinta Basin of Utah and Colorado. Ann. Camegie Mu-
seum 26:1-432.

178. Graul, W. D., and S. J. Bissell (tech. coord.)
1978. Lowland river and stream habitat in Colorado: a
symposium (Greeley, Colo., 4-5 Oct. 1978). Colo. Chapt.
Wildl. Soc. and Colo. Audubon Council. 195 pp.

179. Gray, A., and J. D. Hooker. 1880. The vegetation
of the Rocky Mountain region and a comparison with
that of other parts of the world. Bull. U.S. Geol. and
Geogr. Surv. Terr. VI(l):l-77.

180. Green, N. E. 1969. Occurrence of small mam-
mals on sandhill rangelands in eastern Colorado. Un-
publ. thesis, Colo. St. Univ., Fort Collins. 39 pp.

181. Gregg, R. E. 1963. The ants of Colorado. Univ.
of Colo. Press, Boulder. 74-119 pp -I- map.

182. Greller, A. M. 1974. Vegetation of roadcut slopes
in the tundra of Rocky Mountain National Park, Colo-
rado. Biol. Conserv. 6:84-93.

183. Greller, A. M., M. Goldstein, and L. Marcus.
1974. Snowmobile impact on three alpine tundra plant
communities. Environ. Conserv. 1:101-110.

184. Griggs, R. F. 1955. Competition and succession
on the fellfield of Rocky Mountain National Park. Bull.
Ecol. Soc. Am. 36(3):76 (Abstract).

185. Griggs, R. F. 1956. Competition and succession
on a Rocky Mountain fellfield. Ecol. 37:8-20.

186. Groeneveld, D. P., and P. V. Krebs. 1974. Forest
cover-type mapping using color infrared aerial photogra-
phy. J. Colo.-Wyom. Acad. Sci. 7(5):46 (Abstract).

Great Basin Naturalist

Vol. 43, No. 1

187. Gross, J. E. 1955. A vegetation survey of the
Poudre bighorn sheep range. Paper for course WM-
190D, Colo. St. Univ., Fort Collins, Colo. 65 pp.

188. Gysel, L. W. 1960. An ecological study of the
winter range of elk and mule deer in the Rocky Moun-
tain National Park. J. For. 58:696-703.

189. Hadley, R. F., and F. A. Branson. 1965. Surficial
geology and microclimatic effects on vegetation, soils,
and geomorphology in the Denver, Colorado area. Pages
56-63 in C. B. Schulz and H. T. Smith, eds.. Guidebook
for one-day field conferences-Boulder area, Colorado.
Vllth Int. Assoc, for Quat. Res. Congress. Nebraska
Acad. Sci., Lincoln.

190. Hammerson, G. A. 1979. Structure and repro-
duction of "tree island" populations of Engelmann
spruce (Picea engelnuinnii) and subalpine fir (Abies lasio-
carpa) in the lower 'alpine tundra of Colorado. J. Colo.-
Wyom. Acad. Sci. ll(l):23-24 (Abstract).

191. Hansen, R. M., and A. L. Ward. 1966. Some rela-
tions of pocket gophers to rangelands on Grand Mesa,
Colorado. Colo. Agric. Expt. Stn. Tech. Bull. No. 88. 22
pp.

192. Hansen-Bristow, K. J. 1981. Environmental con-
trols influencing the altitude and form of the forest-al-
pine tundra ecotone, Colorado Front Range. Unpubl.
dissertation, Univ. of Colo., Boulder. 274 pp.

193. Hanson, H. C. 1929. Range resources of the San
Luis Valley. Colo. Agric. Expt. Stn. Bull. No. 335. 61 pp.

194. Hanson, H. C. 1955. Characteristics of the Stipa
comata-Bouteloua gracilis-Bouteloua curtipendula associ-
ation in northern Colorado. Ecol. 36:269-280.

195. Hanson, H. C. 1957. The use of basic principles
in the classification of range vegetation. J. Range Mgmt.
10:26-33.

196. Hanson, H. C. 1959. Variation within a pon-
derosa pine forest in Colorado. Bull. Ecol. Soc. Am.
40(3):71-72 (Abstract).

197. Hanson, H. C, and W. S. Ball. 1928. An appli-
cation of Raunkiaer's law of frequence to grazing stud-
ies. Ecology 9:467-473.

198. Hanson, H. C. and E. Dahl. 1957. Some grass-
land communities in the mountain-front zone in north-
em Colorado. Vegetatio 7:249-270.

199. Hanson, H. C, L. D. Love, and M. S. Morris.
1931. Effects of different systems of grazing by cattle
upon a western wheat-grass type of range near Fort Col-
lins, Colorado. Colo. Agric. Expt. Sta. Bull. No. 377. 82
pp.

200. Hanson, H. C, and F. B. Smith. 1928. Some
types of vegetation in relation to the soil profile in
northern Colorado. J. Am. Soc. Agron. 20:142-151.

201. Harmon, W. E. 1980. Survey of the flora and
vegetation of the Bodo Wildlife Management Area. Un-
publ. report prepared for The Nature Conservancy,
Denver, Colo. 40 pp.

202. Harrington, F. A., Jr. 1978. Ecological segrega-
tion of ungulates in alpine and subalpine communities.
Unpubl. dissertation, Colo. St. Univ., Fort Collins.

203. Harrington, H. D. 1964. Manual of the plants of
Colorado. Sage Books, Swallow Press, Chicago. 2d ed.
666 pp.

204. Harris, J. T. 1958. Analysis of elk winter range.
South Fork of the White River, Colorado. Unpubl.
thesis, Colo. St. Univ., Fort Collins. 136 pp.

205. Heerwagen, A. 1958. Management as related to
range site in the central plains of eastern Colorado. J.
Range Mgmt. 11:5-9.

206. Heifner, M. A. 1974. Ecological studies in Colo-
rado alpine willow marshes. Unpubl. thesis, Colo. St.
Univ., Fort Collins. 66 pp.

207. Helm, D. J. 1977. Variations in alpine snowfield
vegetation. Unpubl. thesis, Colo. St. Univ., Fort Collins.
95 pp.

208. Helm, D. J. 1981. Vegetation diversity indexes in
several vegetation types of western Colorado. Unpubl.
dissertation, Colo. St. Univ., Fort Collins. 113 pp.

209. Hess, K. 1981. Phyto-edaphic study of habitat
types of the Arapaho-Roosevelt National Forest, Colo-
rado. Unpubl. dissertation, Colo. St. Univ., Fort Collins.
558 pp.

210. Hess, K., and C. H. Wasser. 1982. Habitat type
classification of a selected part of the White River-Ara-
paho National Forest. Unpubl. report on file at the U.S.
For. Serv., Region 2, Range and Wildlife Management,
Denver, Colo.

211. Hodson, E. R., and J. H. Foster. 1910. Engel-
mann spruce in the Rocky Mountains. USDA For. Serv.
Circ. 170. 23 pp.

212. Hoff, C. C. 1957. A comparison of soil, climate,
and biota of conifer and aspen communities in the cen-
tral Rocky Mountains. Am. Midi. Nat. 58:115-140.

213. Hoffman, G. R., and R. A. Alexander. 1980. For-
est vegetation of the Routt National Forest in north-
western Colorado: a habitat type classification. USDA
For. Serv. Res. Paper RM-221, Rocky Mt. For. and
Range Exp. Stn., Fort Collins, Colo. 41 pp.

214. Hoffman, G. R., and G. J. Tomlinson. 1966. A
bibliography of vegetational studies of Colorado. S.W.
Natur. 11:228-237.

215. Holm, T. 1908. Studies in the Gramineae. IX.
The Gramineae of the alpine region of the Rocky Moun-
tains in Colorado. Bot. Gaz. 46:422-444.

216. Holm, T. 1927. The vegetation of the alpine re-
gion of the Rocky Mountains in Colorado. Nat. Acad.
Sci. Mem. 19(3):l-45 + 7 plates.

217. Holway, J. G. 1960. Phenological observations on
the major alpine species in the vicinity of False Mummy
Pass, Rocky Mountain National Park, Colorado. Unpubl.
thesis, Colo. St. Univ., Fort Collins. 84 pp.

218. Holway, J. G. 1961. Phenological observations on
alpine species. J. Colo.-Wyom. Acad. Sci. 5(2);38-39
(Abstract).

219. Holway, J. G. 1962a. Phenology of Colorado al-
pine plants. Unpubl. dissertation, Colo. St. Univ., Fort
Collins. 154 pp.

220. Holway, J. G. 1962b. Effects of snow and di-
verted melt-water in the alpine tundra. J. Colo.-Wyom.
Acad. Sci. 5(3):33 (Abstract).

221. Holway, J. G., and R. T. Ward. 1963. Snow and
meltwater effects in an area of Colorado alpine. Am.
Midi. Nat. 69:189-197.

222. Holway, J. G., and R. T. Ward. 1965. Phenology
of alpine plants in northern Colorado. Ecology 46:73-83.

223. Hyder, D. N., R. E. Bement, E. E. Remmenga,
and D. F. Hervey. 1975. Ecological responses of native
plants and guidelines for management of shortgrass
range. USDA Tech. Bull. No. 1503. 87 pp.

January 1983

Baker: Colorado Bibliography

224. Hyder, D. N., R. E. Bement, E. E. Remmenga,
and C. Terwilliger, Jr. 1966. Vegetation-soils and vegeta-
tion-grazing relations from frequency data. J. Range
Mgmt. 19:11-17.

225. loms, W. V., C. H. Hembree, and G. L. Oak-
land. 1965. Water resources of the Upper Colorado Riv-
er Basin-Technical Report. USGS Prof. Paper 441.
(Plates 5 & 9— vegetation maps).

226. Isaacson, H. E. 1967. Ecological provinces with-
in the pinyon-juniper type of the Great Basin and Colo-
rado Plateau. Unpubl. thesis, Utah St. Univ., Logan. 44
pp.

227. Ives, J. D. 1978. Remarks on the stability of tim-
berline. Pages 313-317 in C. Troll and W. Lauer, eds.,
Geoecological relations between the Southern Temper-
ate Zone and the tropical mountains. Franz Steiner Ver-
lag GMBH, Wiesbaden.

228. Ives, R. L. 1941a. Forest replacement rates in
the Colorado headwaters area. Bull. Torrey Bot. Club
68:407-408.

229. Ives, R. L. 1941b. Vegetative indicators of soli-
fluction. J. Geomorph. 4:128-132.

230. Ives, R. L. 1941c. Rapid identification of the
montane-subalpine zone boundary. Bull. Torr. Bot.
Club. 68:195-197.

231. Ives, R. L. 1942a. Atypical subalpine environ-
ments. Ecology 23:89-96.

232. Ives, R. L. 1942b. The beaver-meadow complex.
J. Geomorph. 5:191-203.

233. Ives, R. L. 1946. Botanical indicators of air drifts.
Ecology 27:364-369.

234. Jack, J. G. 1900. Pikes Peak, Plum Creek, and
South Platte reserves. U.S. Geol. Survey 20th Ann. Rep.,
Part V:39-115.

235. Jackson, A. S. 1965. Wildfires in the Great Plains
grasslands. Tall Timbers Fire Ecol. Conf. 4:241-259.

236. Jackson, J. R. 1972a. Vegetation of the flood
plain of the South Platte River in the proposed Narrows
Reservoir site. Unpubl. thesis, Univ. of N. Colo.,
Greeley. 83 pp.

237. Jackson, J. R. 1972b. The vegetation of the site of
the proposed Narrows Reservoir. J. Colo.-Wyom. Acad.
Sci. 7(2):24 (Abstract).

238. Jackson, J. R., and I. E. Lindauer. 1978. Vegeta-
tion of the flood plain of the South Platte River in the
proposed Narrows Reservoir site. Trans. Missouri Acad.
Sci. 12:37-46.

239. James, J. W., and J. W. Marr. 1966. Route 40
mountain environmental transect: Colorado and Utah.
Report prepared for INSTAAR, Univ. of Colo., Boulder.
218 pp + appendices.

240. James, S. L. 1930. The herbaceous vegetation of
two mesas near Boulder, Colorado. Unpubl. thesis, Univ.
of Colo., Boulder. 41 pp.

241. Jameson, D. A. (Coordinator). 1969. General de-
scription of the Pawnee site. U.S. IBP Grassland Biome
Tech. Rep. No. 1. Colo. St. Univ., Fort Collins. 22 pp.

242. Jefferies, N. W. 1965. Herbage production on a
gambel oak range in southwestern Colorado. J. Range
Mgmt. 18:212-213.

243. Jennings, R. H. 1978. Vegetation. Pages 1-103 in
H. A. Ferchau, ed., Buttes Gas and Oil Iron Hill-
Powderhorn Environmental Study 1977, Western St.
Coll., Gunnison, Colo. Vol. II, Ch. 8.

244. Jennings, R. H. 1979a. Supplement to Chapter
8— Vegetation. In H. A. Ferchau, ed., Pwderhorn Il-Iron
Hill-Powderhorn Environmental Study, Western St.
Coll., Gunnison, Colo.

245. Jennings, R. H. 1979b. Vegetation of the Iron
Hill complex, Powderhorn, Colorado: an environmental
baseline inventory and analysis. Unpubl. thesis. Western
State College, Gunnison, Colo. 211 pp.

246. Johnson, A. W. 1956. Ecology of subalpine forest
communities in the Silver Lake Valley of the Front
Range in Colorado. Unpubl. dissertation, Univ. of Colo.,
Boulder. 121 pp.

247. Johnson, D. L. 1973. Floodplain communities of
the upper Roaring Fork River, near Aspen, Colorado.
Unpubl. report, Dept. of Envir., Pop., and Organismic
Biol., Univ. of Colorado, Boulder.

248. Johnson, D. L., and C. Mutel. 1973. Preliminary
description and inventory of the ecosystems of Walton
Creek canyon and adjacent areas. Steamboat Springs,
Colorado. Unpubl. report prepared for Applied Science
and Resource Planning, Inc., Sacramento, Calif. 29 pp.

249. Johnson, K. L. 1970. Alpine vegetation and soils
of Mesa Seco Plateau, San Juan Mountains, Colorado.
Unpubl. dissertation, Univ. of Illinois, Urbana. 217 pp.

250. Johnson, K. R. 1932a. Ecology of a glacial lake in
central Colorado. Unpubl. thesis, Univ. of Colo., Boul-
der. 30 pp.

251. Johnson, K. R. 1932b. Plant ecology of a glacial
lake. J. Colo.-Wyom. Acad. Sci. 1(4): 13 (Abstract).

252. Johnson, K. R. 1936. Ecology of a glacial lake in
central Colorado. Univ. of Colo. Studies 23:2,35-243.

253. Johnson, K. R. 1939. Plant ecology of north-
western Colorado lakes and surrounding areas. Unpubl.
dissertation, Univ. of Colo., Boulder. 138 pp.

254. Johnson, K. R. 1941. Vegetation of some moun-
tain lakes and shores in northwestern Colorado. Ecology
22:306-316.

255. Johnson, W. M. 1945. Natural revegetation of
abandoned crop land in the ponderosa pine zone of the
Pike's Peak region in Colorado. Ecology 26:363-374.

256. Johnson, W. M. 1953. Effect of grazing intensity
upon vegetation and cattle gains on ponderosa pine-
bunchgrass ranges of the Front Range of Colorado.
USDA Circ. No. 929. 36 pp.

257. Johnson, W. M. 1956. The effect of grazing in-
tensity on plant composition, vigor, and growth of pine-
bunchgrass ranges in central Colorado. Ecology
37:790-798.

258. Johnson, W. M., and G. E. Klipple. 1946. The
natural revegetation of abandoned cropland in the pon-
derosa pine zone of the Pike's Peak region. J. Colo.-
Wyom. Acad. Sci. 3(3):39-40 (Abstract).

259. Johnson, W. M., and C. H. Niederhof. 1941.
Some relationships of plant cover to run-off, erosion, and
infiltration on granitic soils. J. For. 39:854-858.

260. Johnson, W. M., and E. H. Reid. 1958. Herbage
utilization on pine-bunchgrass ranges of Colorado. J.
For. 56:647-651.

261. Johnson, W. M., and E. H. Reid. 1964. Range
condition classification of bunchgrass range at the Man-
itou Experimental Forest in Colorado. J. Range Mgmt.
17:137-141.

Great Basin Naturalist

Vol. 43, No. 1

262. Johnston, B. C. 1978. Vegetation map of Colo-
rado. Appendix I-Part 1, p. 1-5, + map in D. W. Crum-
packer, ed., A nature reserve system for Colorado. Re-
port to the Natural Areas Program of the Department of
Natural Resources, State of Colorado.

263. Judd, B. I. 1974. Plant succession of old fields in
the dust bowl. S.W. Natur. 19:227-239.

264. Kahler, L. J. 1973. Correlation of slope exposure
with differences in the composition of the vegetation
community at 7000 feet in Clear Creek Canyon west of
Golden, Colorado. Unpubl. thesis, Univ. of Colo., Boul-
der. 105 pp.

265. Keammerer, D. B. 1976. Niwot Ridge vegetation
map. 1:24000. Unpubl. manuscript on file at Inst, of
Arct. and Alp. Res., Univ. of Colo., Boulder (reproduced
in Komarkova and Webber 1978. Arct. and Alp. Res.
10:7).

266. Keammerer, W. R. 1974a. Vegetation of the
Grand Valley area. Pages 73-117 in Ecological inventory
of the Grand Valley area. Unpubl. report prepared for
the Colony Development Operation, Atlantic Richfield
Co., Operator, Denver, Colo.

267. Keammerer, W. R. 1974b. Vegetation of Para-
chute Creek Valley. Pages 4-91 in Environmental inven-
tory and impact analysis of a proposed utilities corridor
in Parachute Creek Valley, Colorado. Unpubl. report
prepared for Colony Development Operation, Atlantic
Richfield Co., Operator, Denver, Colo.

268. Keammerer, W. R. 1977. Final report: vegeta-
tion baseline studies, oil shale tract C-b. Unpubl. report,
Stoecker-Keammerer & Assoc, Ecological Consultants,
Boulder, Colo. 183 pp.

269. Keammerer, W. R., and D. B. Keammerer. 1978.
DOW property vegetation studies: endangered species
survey and vegetation mapping 1978. Unpubl. report
prepared for Colony Development Operation, Atlantic
Richfield Co., Operator, Denver, Colo., by Stoecker-
Keammerer & Assoc, Ecological Consultants, Boulder,
Colo. 20 pp..

270. Keammerer, W. R., and D. B. Keammerer. 1980.
Preliminary vegetation studies on the Naval Oil Shale
Reserve. Unpubl. report prepared for TOSCO Corpo-
ration, Denver, Colo., by Stoecker-Keammerer & Assoc,
Ecological Consultants, Boulder, Colo. 14 pp.

271a. Keammerer, W. R., and S. J. Peterson. 1981.
Vegetation studies on the Naval Oil Shale Reserve. Un-
publ. report prepared for TRW Energy Systems Group,
McLean, Virginia, by Stoecker-Keammerer & Assoc,
Ecological Consultants, Boulder, Colo. 77 pp.

271b. Keammerer, W. R., and R. E. Stoecker. 1975.
Vegetation and wildlife studies along proposed corridors
for oil shale tract C-b. Unpubl. report prepared for Shell
Oil Co., Operator, Denver, Colo., by Stoecker-
Keammerer & Assoc, Ecological Consultants, Boulder,
Colo. 86 pp.

272. Keammerer, W. R., and R. E. Stoecker. 1980.
Vegetation and wildlife studies for the Mount Emmons
project. Unpubl. report prepared for AMAX Environ-
mental Services, Inc. by Stoecker-Keammerer & Assoc,
Ecological Consultants, Boulder, Colo. 3 vols.

273. Keith, J. R. 1969. The vegetation of South Table
Mountain, Jefferson County, Colorado. J. Colo.-Wyom.
Acad. Sci. 6(2): 14 (Abstract).

274. Kellogg, R. S. 1905. Forest belts of western Kan-
sas and Nebraska. USDA For. Serv. Bull. 66. 44 pp.

275. Kiener, W. 1935. Structural analysis of alpine
vegetation on Longs Peak, Colorado. J. Colo.-Wyom.
Acad. Sci. 2(1):50-51 (Abstract).

276. Kiener, W. 1937. On the history of the vegeta-
tion of Rocky Mountain National Park. The Colo. For-
ester 1937: 24-25.

277. Kiener, W. 1939. Sociological studies of the al-
pine vegetation on Long's Peak. Unpubl. dissertation,
Univ. of Nebraska, Lincoln.

278. Kiener, W. 1967. Sociological studies of the al-
pine vegetation on Longs Peak. Univ. Nebr. Stud. n.s.
No. 34. 75 pp.

279. Kinraide, T. B. 1977. ChoUa cactus {Opuntia im-
bricata (Haw.) DC.) in El Paso County, Colorado: au-
tecology and preliminary phytosociology. Bull. Ecol.
Soc Am. 58(2):40 (Abstract).

280. Kinraide, T. B. 1978. The ecological distribution
of cholla cactus (Opuntia imbricata (Haw.) DC.) in El
Paso County, Colorado. S.W. Natur. 23:117-134.

281. Klemmedson, J. O. 1953. Interrelations of vege-
tation, soils, and range conditions induced by grazing.
Unpubl. thesis, Colo. St. Univ., Fort Collins. 178 pp.

282. Klemmedson, J. O. 1956. Interrelations of vege-
tation, soils, and range conditions induced by grazing. J.
Range Mgmt. 9:134-138.

283. Klipple, G. E. 1964. Early- and late-season graz-
ing versus season-long grazing of short-grass vegetation
on the central Great Plains. USDA For. Serv. Res. Paper
RM-11, Rocky Mt. For. & Range Exp. Stn., Fort Collins,
Colo. 16 pp.

284. Klipple, G. E., and D. F. Costello. 1960. Vegeta-
tion and cattle responses to different intensities of graz-
ing on short-grass ranges on the central Great Plains.
USDA Tech. Bull. No. 1216. 82 pp.

285. Klish, M. W. 1977. The vegetation of Summit
County, Colorado. Unpubl. thesis, Colo. St. Univ., Fort
Collins. 103 pp.

286. Knowles, P., and M. C. Grant (in press). Age and
size structure analyses of Engelmann spruce, ponderosa
pine, lodgepole pine, and limber pine in Colorado.
Ecology.

287. Komdrkova, V. 1976. Alpine vegetation of the In-
dian Peaks Area, Front Range, Colorado Rocky Moun-
tains. Unpubl. dissertation, Univ. of Colo., Boulder. 655
pp.

288. Komdrkovd, V. 1979. Alpine vegetation of the In-
dian Peaks area. Front Range, Colorado Rocky Moun-
tains. Flora et vegetatio mundi 7, R. Tuxen, ed., 2 vols.
Cramer, Vaduz. 591 pp.

289. Komdrkovd, V. 1980. Classification and ordina-
tion in the Indian Peaks area, Colorado Rocky Moun-
tains. Vegetatio 42:149-163.

290. Komdrkovd, V., and P. J. Webber. 1978. An al-
pine vegetation map of Niwot Ridge, Colorado. Arct.
and Alp. Res. 10:1-29.

291. Krebs, P. H. 1972. Dendrochronology and the
distribution of bristlecone pine (Pinus aristata Engelm.)
in Colorado. Unpubl. dissertation, Univ. of Colorado,
Boulder. 211 pp.

292. Krebs, P. V. 1973. Vegetation. Pages 63-79 +
map in R. F. Madole, ed.. Environmental inventory and
land use recommendations for Boulder county, Colo-
rado. Univ. of Colo., Inst, of Arct. and Alp. Research
Occas. Paper No. 8.

January 1983

Baker: Colorado Bibliography

293. Krebs, P. V., D. P. Groeneveld, and J. P. Spen-
cer. 1976. Ecological overview. Part 1. Vegetation inven-
tory. Pages 81-86 in H. W. Steinhoff and J. D. Ives, eds.,
Ecological impacts of snowpack augmentation in the
San Juan Mountains, Colorado. Final Report, San Juan
Ecol. Proj., Colo. St. Univ., Fort Collins, to the USDI
Bur. Reel, Div. of Atmos. Wat. Res. Mgmt., Denver,
Colo.

294. Kuchler, A. W., and J. McCormick. 1965. Vege-
tation maps of North America. Univ. of Kansas Publ., Li-
brary Series No. 21 [Colorado: p. 162-173].

295. Lambom, C. R. 1962. Theoretical cross-section
transect of the Blue Mesa Reservoir basin in Colorado.
Pages 120-134 in A. M. Woodbury, ed.. Ecological stud-
ies of the flora and fauna of the Curecanti Reservoir ba-
sins, western Colorado. Univ. of Utah, Dept. of Anthro-
pol., Anthropol. Papers No. 59 (Upper Colo. Ser. No. 8).

296. Langenheim, J. H. 1949. Physiography and plant
ecology on a subalpine earthflow, Gunnison County,
Colorado. Unpubl. thesis, Univ. of Minnesota,
Minneapolis.

297. Langenheim, J. H. 1953. Description of the plant
communities and environments of the Crested Butte
area, Gunnison County, Colorado. Unpubl. dissertation,
Univ. of Minn., Minneapolis. 223 pp.

298. Langenheim, J. H. 1955. Flora of the Crested
Butte quadrangle, Colorado. Madrono 13:64-78.

299. Langenheim, J. H. 1956. Plant succession on a
subalpine earthflow in Colorado. Ecology 37:301-317.

300. Langenheim, J. H. 1957. A serial system of classi-
fication applied to the vegetation of the Crested Butte
area, Colorado. Bull. Ecol. Soc. Am. 38:82 (Abstract).

301. Langenheim, J. H. 1962. Vegetation and environ-
mental patterns in the Crested Butte area, Gunnison
County, Colorado. Ecol. Monogr. 32:249-285.

302. Lanier, J. E. 1978. Population dynamics of Rocky
Mountain Douglas-fir (Pseudotsuga menziesii var.
glauca) along an elevational gradient in the Front Range
of Colorado. Unpubl. thesis, Univ. of Colo., Boulder. 97
pp.

303. Lanier, J. E., D. G. Steward, D. A. Boyce, and J.
W. Marr. 1977. A description of the ecosystems of the
Homestake Creek area. Eagle County, Colorado. J.
Colo.-Wyom. Acad. Sci. 9:36 (Abstract).

304. Larson, L. L. 1974. Discriminant and hier-
archical analyses in vegetation classification. Unpubl.
thesis, Colo. St. Univ., Fort Collins. 103 pp.

305. Larson, L. L. 1978. Vegetative variability along a
gradient from lodgepole pine-spruce fir. Unpubl. dis-
sertation, Colo. St. Univ., Fort Collins. 101 pp.

306. Laven, R. D., P. N. Omi, J. G. Wyant, and A. S.
Pinkerton. 1980. Interpretation of fire scar data from a
ponderosa pine ecosystem in the central Rocky Moun-
tains, Colorado. Pages 46-49 in M. A. Stokes and J. Die-
terich (Tech. Coord.) Proceedings of the fire history
workshop. 20-24 Oct. 1980, Tucson, Ariz. USDA For.
Serv. Gen. Tech. Rept. RM-81, Rocky Mountain For.
and Range Exp. Stn., Fort Collins, Colo.

307. Legg, T. E., and R. G. Baker. 1980. Palynology
of Pinedale sediments, Devlin's Park, Boulder County,
Colorado. Arct. and Alp. Res. 12:319-333.

308. Leonard, H. A. 1916. Vegetation of the aspen
groves at Tolland, Colorado. Unpubl. thesis, Univ. of
Colo., Boulder. 17 pp.

309. Lindauer, I. E. 1967. Ecology of phreatophytes
on the Arkansas River in southeastern Colorado. J.
Colo.-Wyom. Acad. Sci. 5(8):65 (Abstract).

310. Lindauer, I. E. 1968. A survey of the woody
phreatophytes in the lower Arkansas River Valley of
Colorado. J. Colo.-Wyom. Acad. Sci. 6(l):51-52
(Abstract).

311. Lindauer, I. E. 1970. The vegetation of the flood
plain of the Arkansas River in southeastern Colorado.
Unpubl. dissertation, Colo. St. Univ., Fort Collins. 92
pp.

312. Lindauer, I. E. 1978. A comparison of the vege-
tative communities of the South Platte and Arkansas
River drainages in eastern Colorado. Pages 56-72 in W.
D. Graul and S. J. Bissel, eds.. Lowland river and stream
habitat in Colorado: a symposium. (Greeley, Colo. 4-5
Oct. 1978). Colo. Chapt. Wildl. Soc. and Colo. Audubon
Council.

313. Lindauer, I. E., and S. J. Christy. 1972. An analy-
sis of the woody vegetation on the South Platte River
flood plain in northeastern Colorado. Unpubl. report to
the U.S. Bureau of Reclamation, Denver, Colo., by the
Dept. of Biol. Sci., Univ. of N. Colo., Greeley.

314. Lindauer, I. E., and J. P. Fitzgerald. 1974. Eco-
logical survey and analysis of terrestrial communities at
the Weld County (Hardin) proposed reservoir site. Un-
publ. report to the U.S. Bureau of Reclamation, Denver,
Colo., by the Dept. of Biol. Sci., Univ. of N. Colo.,
Greeley. 45 pp.

315. Lindauer, I. E., J. P. Fitzgerald, and L. L. Lin-
dauer. 1973. Ecological analyses of flood plain commu-
nities, Narrows Reservoir Site, Colorado. Unpubl. report
to the U.S. Bureau of Reclamation, Denver, Colo, by the
Dept. of Biol. Sci., Univ. of N. Colo., Greeley. 108 pp.

316. Lindauer, I. E., and R. T. Ward. 1968. A survey
of the woody phreatophytes in the lower Arkansas River
Valley of Colorado. Unpubl. report to the U.S. Bureau of
Reclamation, Denver, Colo., by the Dept. of Botany and
Plant Pathology, Colo. St. Univ., Fort Collins, Colo. 46
pp.

317. Livingston, R. B. 1941. The effect of altitude on
the composition of grasslands on the plains of central
Colorado. Unpubl. thesis, Duke Univ., Durham, N.C.

318. Livingston, R. B. 1947. An ecological study of
the Black Forest region and adjacent plains. Unpubl.
dissertation, Duke Univ., Durham, N.C. 134 pp.

319. Livingston, R. B. 1949. An ecological study of
the Black Forest, Colorado. Ecol. Monogr. 19:123-144.

320. Livingston, R. B. 1952. Relict true prairie com-
munities in central Colorado. Ecology 33:72-86.

321. Loder, C. W. 1964. Alpine tundra of Cotton-
wood Pass, Colorado. Unpubl. thesis. Western State Col-
lege, Gunnison, Colo. 139 pp.

322. Loveless, C. M. 1963. Ecological characteristics
of a selected mule deer winter range. Unpubl. dis-
sertation, Colo. St. Univ., Fort Collins. 318 pp.

323. Loveless, CM. 1967. Ecological characteristics
of a mule deer winter range. Colo. Dept. Game, Fish
and Parks, Tech. Publ. No. 20. 125 pp.

324. Lusby, G. C, G. T. Turner, J. R. Thompson, and
V. H. Reid. 1963. Hydrologic and biotic characteristics
of grazed and ungrazed watersheds of the Badger Wash
Basin in western Colorado, 1953-1958. U.S. Geol. Sur-
vey Water-Supply Paper 1532-B.

Great Basin Naturalist

Vol. 43, No. 1

325. Luti, R. 1953. Ecological features of the vegeta-
tion of a ridge in the montane forest of Boulder County,
Colorado. Unpubl. thesis, Univ. of Colo., Boulder. 100
pp.

326. Lutz, F. E. 1922. Altitude in Colorado and geo-
graphical distribution. Bull. Am. Mas. Nat. Hist.
46:335-366.

327. McCuUough, H. A. 1948. Plant succession on
fallen logs in a virgin spruce-fir forest. Ecol. 29:508-513.

328. McCullough, H. A. 1952. The Gothic Natural
Area. Living Wilderness 17(41): 13-20.

329. McCullough, H. A. 1956. Survey of the Gothic
Natural Area. The Scientific Monthly 82:25-32.

330. McEachem, K. 1979. Population biology of Pop-
ulus sargentii in northeastern Colorado. Unpubl. under-
grad. thesis (honor). Univ. of N. Colo., Greeley. 41 pp.

331. McGinnies, W. J., L. W. Osborn, and W. A.
Berg. 1976. Plant-soil-microsite relationships on a salt-
grass meadow. J. Range Mgmt. 29:395-400.

332. McHenry, D. E. 1929. The vegetation of Gregory
Canyon, Colorado. Unpubl. thesis, Univ. of Colo., Boul-
der. 35 pp.

333. McHenry, D. E. 1930. The vegetation of Gregory
Canyon, Colorado. J. Colo.-Wyom. Acad. Sci. 1(2): 14
(Abstract).

334. Mcintosh, A. C. 1923. Vegetation at different
elevations in Boulder Canyon. Unpubl. thesis, Univ. of
Colorado, Boulder. 35 pp.

335. McNeal, D. W., Jr. 1976. Annotated check list of
the alpine vascular plants of Specimen Mountain, Rocky
Mountain National Park, Colorado. S.W. Natur.
20:423-435.

336. Maher, L. J., Jr. 1961. Pollen analysis and post-
glacial vegetation history in the Animas Valley region,
southern San Juan Mountains, Colorado. Unpubl. dis-
sertation, Univ. of Minn., St. Paul. 85 pp.

337. Maher, L. J., Jr. 1963. Pollen analyses of surface
materials from the southern San Juan Mountains, Colo-
rado. Geol. Soc. Am. Bull. 74:1485-1504.

338. Maher, L. J., Jr. 1972. Absolute pollen diagram
of Bedrock Lake, Boulder County, Colorado. Quat. Res.
2:531-553.

339. Mallory, G. F. 1981. A vegetation inventory and
analysis of the Homestake Pitch uranium project, Sa-
guache County, Colorado. Unpubl. thesis. Western State
College, Gunnison, Colo. 129 pp.

340. Markgraf, V. 1980. Lower timberline paleoenvi-
ronmental changes during the last 15,000 years in cen-
tral Colorado. Bull. Ecol. Soc. Am. 61(2):107-108
(Abstract).

341. Marr, J. W. 1958. Lee slope stands in the upper
part of the forest-tundra ecotone on Niwot Ridge, Boul-
der County, Colorado. J. Colo. -Wyoming Acad. Sci.
4(10):41 (Abstract).

342. Marr, J. W. 1959. Forms of tree islands in alpine
tundra. J. Colo.-Wyoming Acad. Sci. 4(11):34 (Abstract).

343. Marr, J. W. 1964a. Utilization of the Front
Range tundra, Colorado. Pages 109-118 in D. J. Crisp,
ed.. Grazing in terrestrial and marine environments.
Blackwell Sci. Publ., Oxford, Eng.

344. Marr, J. W. 1964b. The vegetation of the Boul-
der area. Univ. of Colo. Museum Leaflet 13:34-42.

345. Marr, J. W. 1965. Forest ecology of the Front
Range. Pages 16-20 in C. B. Schulz and H. T. Smith,
eds.. Guidebook for one-day field conferences— Boulder

area, Colorado. Vllth Int. Assoc, for Quat. Res. Con-
gress, Nebraska Acad. Sci., Lincoln.

346. Marr, J. W. 1967. Ecosystems of the east slope of
the Front Range in Colorado. Univ. of Colo. Studies,
Series in Biology No. 8. 134 pp.

347. Marr, J. W. 1977a. The development and move-
ment of tree islands near the upper limit of tree growth
in the southern Rocky Mountains. Ecology
58:1159-1164.

.348. Marr, J. W. 1977b. A classification of tree island
forms in the high mountains of Colorado. J. Colo.-
Wyom. Acad. Sci. 9(1):35 (Abstract).

349. Marr, J. W., D. M. Armstrong, H. Chronic, J.
Chronic, R. W. Pennak, W. A. Weber, R. E. Marr, D.
Steward, and J. C. Meyer. 1980. Natural landmarks of
the southern Rocky Mountain region. Unpubl. report
prepared for the U.S. Dept. Int., Heritage Conserv. and
Recreation Service, Denver, Colo., by Thorne Ecol.
Inst., Boulder, Colo. 736 pp.

.350. Marr, J. W., D. A. Boyce, and J. W. Todd. 1973.
Preliminary report on the Redcliff project. Eagle Coun-
ty, Colorado. Report prepared for the David E. Fleming
Co., Denver, Colo., and the Colo. River Water Conserv.
District, Glenwood Springs, Colo., by the Dept. of EPO
Biol., Univ. of Colo., Boulder. 9 pp.

351. Marr, J. W., and W. S. Boyd. 1979. Vegetation in
the greater Denver area. Front Range Urban Corridor,
Colorado. USGS Misc. Investig. Series, Map I-856-I.

352. Marr, J. W., and D. L. Buckner. 1974a. Revege-
tation of ecosystems modified by pipeline construction
in western Colorado and southeastern Utah. J. Colo.-
Wyo. Acad. Sci. 7(5):30-31 (Abstract).

353. Marr, J. W., and D. L. Buckner. 1974b. Colorado
to Wyoming pipeline corridor study. Unpubl. report
prepared for Colony Development Operation, Atlantic
Richfield Co., Denver by Thorne Ecol. Inst., Boulder,
Colo. 79 pp. -f- appendices.

354. Marr, J. W., D. L. Buckner, and D. L. Johnson.
1974. Ecological modification of alpine tundra by pipe-
line construction. Pages 10-17 in W. A. Berg, J. A.
Brown, and R. L. Cuany, eds., Proceedings of a work-
shop on revegetation of high-altitude disturbed lands.
Colo. St. Univ., Environ. Res. Center Inf. Series No. 10.

355. Marr, J. W., D. L. Buckner, and C. Mutel. 1973.
Ecological analysis of potential oil products corridors in
Colorado and Utah. Unpubl. report prepared for Colony
Development Operation, Atlantic Richfield Co., Den-
ver, Colo. 96 pp. & App. A, B, C.

356. Marr, J. W., D. L. Buckner, and C. Mutel. 1974.
Reaction of vegetation and soils to impact from con-
struction and operation of pipelines in western Colorado
and eastern Utah. Unpubl. report prepared for Colony
Development Operation, Atlantic Richfield Co., Den-
ver, Colo. 138 pp.

357. Marr, J. W., J. Bunin, F. Moore, T. Giese, and C.
Mutel. 1974. Terrestrial ecology: Yampa River Project,
reconnaissance study of ecosystems and environmental
impact. Unpubl. report, dept. of EPO Biology, Univ. of
Colo., Boulder. 100 pp.

358. Marr, J. W., R. Fritz, J. Meyer, and P. Murphy.
1979. Final report— terrestrial plant ecology— stand eco-
system data tables, Juniper/Cross Mountain Project.
Colorado River Water Conservation District, Glenwood
Springs, Colo., by the Dept. of EPO Biology, Univ. of
Colo., Boulder.

January 1983

Baker: Colorado Bibliography

359. Marr, J. W., R. J. Fritz, and P. Murphy. 1979.
The vegetation of selected U.S. Forest Service areas in
the vicinity of the Chmax Mine in Colorado. Unpubl. re-
port prepared for the Climax Molybdenum Co., AMAX,
Inc.

360. Marr, J. W., D. L. Johnson, and D. L. Buckner.
1974. Recovery of alpine tundra ecosystems of Fremont
Pass, Colorado, following pipeline construction. J. Colo.-
Wyom. Acad. Sci. 7(5):31 (Abstract).

361. Marr, J. W., and D. G. Stevk'ard. 1979. Vegeta-
tion in the Colorado Springs-Castle Rock area, Front
Range Urban Corridor, Colorado. USGS Misc. Investig.
Series, Map I-857-C.

362. Marr, J. W., and B. E. Willard. 1970. Persisting
vegetation in an alpine recreation area in the southern
Rocky Mountains, Colorado. Biol. Conserv. 2:97-104.

363. Mase, J. R. 1982. Vegetation survey: Anvil Points
oil shale facility, Garfield County, Colorado. Final re-
port to U.S. Dept. of Energy, Laramie Energy Tech-
nology Center, Laramie, Wyoming, by Radian Corpo-
ration, 8501 Mo-Pac Blvd., Austin, Texas. 57 pp.

364. Masek, J. 1979. Vegetation community descrip-
tions for the South Platte River in Colorado and Nebras-
ka. Water and Power Resources Service, Denver, Colo.
23 pp.

365. Mason, D. T. 1915. The life history of lodgepole
pine in the Rocky Mountains. USDA Bull. No. 154. 35
pp.

366. Maxwell, E. L. 1975. Multispectral analysis of
rangeland conditions. Unpubl. dissertation, Colo. St.
Univ., Fort Collins. 198 pp.

367. May, D. E. 1973. Models for predicting composi-
tion and production of alpine tundra vegetation from
Niwot Ridge, Colorado. Unpubl. thesis, Univ. of Colo.,
Boulder. 99 pp.

368. May, D. E. 1974. Phenology and ecotypic varia-
tion in alpine tundra vegetation, Niwot Ridge, Colorado.
J. Colo.-Wyom. Acad. Sci. 7(5):29-30 (Abstract).

369. May, D. E. 1976. The response of alpine tundra
vegetation in Colorado to environmental variation. Un-
publ. dissertation, Univ. of Colo., Boulder. 164 pp.

370. Mayo, J. E. 1971. A comparative study of the for-
est floor produced by three age-class stands of Pinus
ponderosa. Unpubl. thesis, Univ. of Colo., Boulder. 44
pp.

371. Meyer, J. C. 1978. The terrestrial ecology of Sky-
line Ranch. Unpubl. report on file at The Nature Con-
servancy, Denver, Colo. 72 pp.

372. Michener, M. J. 1964. The high altitude vegeta-
tion of the Needle Mountains of southwestern Colorado.
Unpubl. thesis, Univ. of Colo., Boulder. 77 pp.

373. Miller, P. C. 1964a. Factors influencing the veg-
etation pattern on the White River Plateau in north-
western Colorado. Unpubl. dissertation, Univ. of Colo.,
Boulder. 208 pp.

374. Miller, P. C. 1964b. Factors influencing the veg-
etation pattern of the White River Plateau, north-
western Colorado. Bull. Ecol. Soc. Am. 45(3): 108-109
(Abstract).

375. Miller, P. C. 1970. Age distributions of spruce
and fir in beetle-killed forests on the White River
Plateau, Colorado. Am. Midi. Nat. 83:206-212.

376. Miller, R. L., and G. A. Choate. 1964. The forest
resource of Colorado. USDA For. Serv. Resource Bull.
INT-3., Intermt. For. & Range Expt. Stn., Ogden, Utah.
55 pp.

377. Mills, R. C. 1962. A study of near climax cond-
tions in a spruce-fir ecosystem. Unpubl. thesis, Univ. of
Colo., Boulder. 66 pp.

378. Mitchell, G. C. 1971. Spatial distribution and
successional state of grassland vegetation related to graz-
ing intensity treatments. U.S. Int. Biol. Prog., Grassland
Biome Tech. Rep. No. 101. 70 pp.

379. Moinat, A. D. 1956. Comparative yields of herb-
age from oak scrub and interspersed grassland in Colo-
rado. Ecology 37:852-854.

380. Moir, W. H. 1969a. The lodgepole pine zone in
Colorado. Am. Midi. Nat. 81:87-98.

381. Moir, W. H. 1969b. Steppe communities in the
foothills of the Colorado Front Range and their relative
productivities. Am. Midi. Nat. 81:331-340.

382. Moir, W. H. 1972. Tall grass prairie in Colorado
and its aesthetic value. Pages 40-46 in J. H. Zimmer-
man, ed.. The Second Midwest Prairie Conference,
Univ. of Wise. Arboretum, Madison.

383. Moir, W. H., and M. J. Trlica. 1976. Plant com-
munities and vegetation pattern as affected by various
treatments in shortgrass prairies of northeastern Colo-
rado. S.W. Nat. 21:359-371.

384. Moran, P. 1981. Patterns of plant life at Pawnee
National Grassland: a manual for the interpreter. Un-
publ. report on file at Pawnee National Grasslands,
Greeley, Colo. 88 pp.

385. Morgan, M. D. 1969. Ecology of aspen in Gun-
nison County, Colorado. Am. Midi. Nat. 82:204-228.

386. Morris, M. S. 1932. The structure of the chap-
arral vegetation in northern Colorado. J. Colo.-Wyom.
Acad. Sci. 1(4): 15 (Abstract).

387. Morris, M. S. 1937. Native vegetation of Colo-
rado. J. Colo. -Wyoming Acad. Sci. 2(2):37 (Abstract).

388. Morris, M. S., and R. L. Dix. 1971. Natural vege-
tation of Colorado (Map). Colo. St. Univ., Fort Collins.

389. Moulton, M. P., J. R. Choate, S. J. Bissell, and R.
A. Nicholson. 1981. Associations of small mammals on
the central high plains of eastern Colorado. S.W. Natur.
26:53-57.

390. Moulton, M. P., P. J. Moulton, and J. Sundine.
1980. Site guide to Bonny Reservoir. Colo. Field Or-
nithologists Journ. 14(2):53-55.

391. Mutel, C. F. 1973. An ecological study of the
plant communities of certain montane meadows in the
Front Range of Colorado. Unpubl. thesis, Univ. of Colo.,
Boulder. 77 pp.

392. Mutel, C. F. 1976. From grassland to glacier: an
ecology of Boulder County, Colorado. Johnson Publ.
Co., Boulder. 169 pp.

393. Mutel, C. F. and J. W. Marr. 1973. A vegetative
study of three montane herbaceous basins. J. Colo.-
Wyom. Acad. Sci. 7(4):28 (Abstract).

394. Nelson, J. V. 1973. The environment of the vege-
tation of the Upper Sonoran and Transition zones in the
Uinta Basin. Unpubl. thesis, Univ. of Utah, Salt Lake
City.

395. Nelson, R. A. 1934. Further observations of re-
vegetation on the Twinsisters bum in Rocky Mountain
National Park. J. Colo.-Wyom. Acad. Sci. 1(6):51
(Abstract).

396. Nelson, R. A. 1954. The Twinsisters bum after
twenty years. J. Colo.-Wyom. Acad. Sci. 4(6):45
(Abstract).

Great Basin Naturalist

Vol. 43, No. 1

397. Nelson, R. A. 1970. Plants of Rocky Mountain
National Park. Rocky Mt. Nature Assoc, and U.S. Natl.
Park Service. 3d ed. [Vegetation: p. 17-21].

398. Nelson, R. A., and B. E. Willard. 1962. A survey
of the vegetation of Paradise Park. J. Colo.-Wyom.
Acad. Sci. 5(3):33-34 (Abstract).

399. Nicholson, R. A., and G. K. Hulett. 1969. Rem-
nant grassland vegetation in the central Great Plains of
North America. J. Ecol. 57:599-612.

400. Olgeirson, E. R. 1972. Comparative phys-
iognomic trends in vegetation and soil on a bald near
tree-line in Colorado. Unpubl. dissertation, Univ. of Col-
orado, Boulder. 114 pp.

401. Olgeirson, E. R. 1974a. Parallel conditions and
trends in vegetation and soil on a bald near treeline,
Boreas Pass, Colorado. Arct. & Alp. Res. 6:185-203.

402. Olgeirson, E. R. 1974b. Vascular plants on a
spruce bald in Colorado. Madrono 22:360-363.

403. Olmsted, C. E., Ill and W. R. Taylor. 1977a.
Ecological resource inventory of the Bunker Hill proper-
ty, Boulder County, Colorado. Unpubl. report on file at
The Nature Conservancy, Denver, Colo. 35 pp.

404. Olmsted, C. E., Ill and W. R. Taylor. 1977b.
Ecological resource inventory of the Chambers proper-
ty, Jefferson County, Colorado. Unpubl. report on file at
The Nature Conservancy, Denver, Colo. 41 pp.

405. Osborn, L. W. 1974. Soil-plant relationships in a
saltgrass meadow. Unpubl. thesis, Colo. St. Univ., Fort
Collins. 77 pp.

406. Osbum, W. S., Jr. 1957. Alpine tundra ecosys-
tems of the Front Range in Colorado. Bull. Ecol. Soc.
Am. 38(3):70 (Abstract).

407. Osbum, W. S., Jr. 1958a. Ecology of winter
snow-free areas of the alpine tundra of Niwot Ridge,
Boulder County, Colorado. Unpubl. dissertation, Univ.
of Colo., Boulder. 77 pp.

408. Osbum, W. S., Jr. 1958b. Characteristics of the
Kobresia bellardi meadow stand ecosystem in the Front
Range, Colorado. J. Colo.-Wyom. Acad. Sci. 4(10):38-39
(Abstract).

409. Osbum, W. S., Jr. 1963. The dynamics of fallout
distribution in a Colorado alpine tundra snow accumula-
tion ecosystem. Pages 51-71 in V. Schultz and A. W.
Klement, Jr., eds., Radioecology. Reinhold Publ. Co.,
N.Y. and Am. Inst. Biol. Sci., Washington.

410. Pammel, L. H. 1924. Ecological notes in the
Arapahoe Mountains near Eraser, Colorado. Proc. Iowa
Acad. Sci. 31:133 (Abstract).

411. Pammel, L. H., and R. I. Cratty. 1920. Notes on
some plants of the Arapahoe National Forest and Rocky
Mountain National Park. Proc. Iowa Acad. Sci.
27:51-73.

412. Paulsen, H. A., Jr. 1960. Plant cover and forage
use of alpine sheep ranges in the central Rocky Moun-
tains. Iowa St. J. Sci. 34:731-748.

413. Paulsen, H. A., Jr. 1969. Forage values on a
mountain grassland-aspen range in western Colorado. J.
Range Mgmt. 22:102-107.

414. Paulsen, H. A., Jr. 1970a. The ecological re-
sponse of species in a Thurber fescue community to ma-
nipulative treatments. Unpubl. dissertation, Colo. St.
Univ., Fort Collins.

415. Paulsen, H. A., Jr. 1970b. Competition and suc-
cessional patterns on Thurber fescue grasslands of west-
ern USA. XI Int. Grassl. Congr. [Aug. 1970] Proc.
11:662-665.

416. Paulsen, H. A., Jr. 1975. Range management in
the central and southern Rocky Mountains: a summary
of the status of our knowledge by range ecosystems.
USDA For. Serv. Res. Paper RM-154, Rocky Mt. For.
and Range Expt. Stn., Fort Collins, Colo. 34 pp.

417. Peet, R. K. 1975. Forest vegetation of the east
slope of the northern Colorado Front Range. Unpubl.
dissertation, Cornell Univ., Ithaca, N.Y.

418. Peet, R. K. 1976. Successional patterns and tree
population structure in Rocky Mountain forests. Bull.
Ecol. Soc. Am. 57(1):60 (Abstract).

419. Peet, R. K. 1977. Latitudinal variation in Rocky
Mountain forests. Bull. Ecol. Soc. Am. 58(2):5 (Abstract).

420. Peet, R. K. 1978a. Forest vegetation of the Colo-
rado Front Range: patterns of species diversity. Vegeta-
tio 37:65-78.

421. Peet, R. K. 1978b. Latitudinal variation in south-
em Rocky Mountain forests. J. Biogeog. 5:275-289.

422. Peet, R. K. 1981. Forest vegetation of the Colo-
rado Front Range; composition and dynamics. Vegetatio
45:3-75.

423. Deleted.

424. Penland, C. W. T. 1941. The alpine vegetation
of the southern Rockies and the Ecuadorean Andes.
Colo. Coll. Publ., General Series No. 230, Studies Series
32:1-30.

425. Pennak, R. W. 1963. Ecological and radiocarbon
correlations in some Colorado mountain lake and bog
deposits. Ecology 44:1-15.

426. Petersen, K. L. 1975. Exploratory palynology of a
subalpine meadow, La Plata Mountains, southwestern
Colorado. Unpubl. thesis. Wash. St. Univ., Pullman. 39
pp.

427. Petersen, K. L. 1981. 10,000 years of climatic
change reconstructed from fossil pollen. La Plata Moun-
tains, Southwestern Colorado. Unpubl. dissertation.
Wash. St. Univ., Pullman. 211 pp.

428. Petersen, K. L., and P. J. Mehringer, Jr. 1976.
Postglacial timberline fluctuations. La Plata Mountains,
southwestern Colorado. Arct. and Alp. Res. 8:275-288.

429. Phillips, C. M. 1977. Willow carrs of the upper
Laramie River Valley, Colorado. Unpubl. thesis, Colo.
St. Univ., Fort Collins. 71 pp.

430. Phillips, V. 1981. Response by alpine plants and
soils to microtopography within sorted polygons. J.
Colo.-Wyom. Acad. Sci. 13(1):21 (Abstract).

431. Deleted.

432. Potts, M. K. 1935. Special report on range condi-
tions in Rocky Mountain National Park. Unpubl. report
on file at Rocky Mountain National Park Headquarters,
Estes Park, Colo.

433. Ramaley, F. 1901. Remarks on the distribution of
plants in Colorado east of the divide. Postelsia 1:21-53.

434. Ramaley, F. 1902. Mesa vegetation. Science, n.s.
15:455 (Abstract).

435. Ramaley, F. 1906. Plants of the Florissant region
in Colorado. Univ. Colo. Studies 3:177-185.

436. Ramaley, F. 1907. Plant zones in the Rocky
Mountains of Colorado. Science, n.s. 26:642-643.

437. Ramaley, F. 1908. Botany of northeastern Lari-
mer County, Colorado. Univ. Colo. Studies 5:119-131 +
6 plates.

438. Ramaley, F. 1909a. The silva of Colorado. IV.
Forest formations and forest trees. Univ. Colo. Studies
6:249-281.

January 1983

Baker: Colorado Bibliography

439. Ramaley, F. 1909b. Wild flowers and trees of
Colorado. A. A. Greenman, Univ. Bookstore, Boulder,
Colo. 78 pp.

440. Ramaley, F. 1910. Remarks on some northern
Colorado plant communities with special reference to
Boulder Park (Tolland, Colorado). Univ. Colo. Studies
7:223-236.

441. Ramaley, F. 1914. The amount of bare ground in
some mountain grasslands. Bot. Gaz. 57:526-528.

442. Ramaley, F. 1915. The relative importance of
different species in a mountain grassland. Bot. Gaz.
60:154-157.

443. Ramaley, F. 1916a. Dry grassland of a high
mountain park in northern Colorado. Plant World
19:249-270.

444. Ramaley, F. 1916b. Quadrat studies in a moun-
tain grassland. Bot. Gaz. 62:70-74.

445. Ramaley, F. 1919a. The role of sedges in some
Colorado plant communities. Am. J. Bot. 6:120-130.

446. Ramaley, F. 1919b. Some mountain plant com-
mimities of sandy soil. Plant World 22:313-328.

447. Ramaley, F. 1919c. Xerophytic grasslands at dif-
ferent altitudes in Colorado. Bull. Torr. Bot. Club
46:37-52.

448. Ramaley, F. 1920. Subalpine lake-shore vegeta-
tion in north-central Colorado. Am. J. Bot. 7:57-74.

449. Ramaley, F. 1927. Colorado Plant Life. Univ. of
Colo. Semicentennial Series 1877-1927. Vol. II p.
viii-299.

450. Ramaley, F. 1929. Botany of the San Luis Valley
in Colorado, I. Univ. Colo. Studies 17:27-44.

451. Ramaley, F. 1930. Zonation at the San Luis
Lakes, near Alamosa, Colorado. J. Colo.-Wyom. Acad.
Sci. 1(2): 19-20 (Abstract).

452. Ramaley, F. 1931. Vegetation of chaparral-cov-
ered foothills southwest of Denver, Colorado. Univ.
Colo. Studies 18(4):231-237.

453. Ramaley, F. 1937. Sand-hill vegetation in north-
west-central Colorado. J. Colo.-Wyom. Acad. Sci.
2(2):32-33.

454. Ramaley, F. 1938. Geographic ranges of sand-
hills plants of northeastern Colorado. J. Colo.-Wyom.
Acad. Sci. 2(4) :26 (Abstract).

455. Ramaley, F. 1939a. Sand-hill vegetation of north-
eastern Colorado. Ecol. Monogr. 9:1-51.

456. Ramaley, F. 1939b. Life forms of plants in Colo-
rado sandhills. J. Colo.-Wyom. Acad. Sci. 2(5):33
(Abstract).

457. Ramaley, F. 1940. Vegetation of the San Luis
Valley in Colorado. J. Colo.-Wyom. Acad. Sci.
2(6):35-36 (Abstract).

458. Ramaley, F. 1942. Vegetation of the San Luis
Valley in southern Colorado. Univ. Colo. Studies, Series
D, 1:231-277.

459. Ramaley, F., and L. Kelso. 1931. Autumn vegeta-
tion of the foothills near Boulder, Colorado. Univ. Colo.
Studies 18(4):239-255.

460. Ramaley, F., and L. A. Mitchell. 1911. Ecologi-
cal cross-section of Boulder Park (Tolland, Colorado).
Univ. Colo. Studies 8:277-287.

461. Ramaley, F., and W. W. Robbins. 1908a. Ecolog-
ical notes from north-central Colorado. Univ. Colo.
Studies 5:111-117.

462. Ramaley, F., and W. W. Robbins. 1908b. Plant
zones of the mountain lakes in northern Colorado. Sci-
ence n.s. 27:208 (Abstract).

463. Ramaley, F., and W. W. Robbins. 1908c. Rock-
ridge vegetation of northern Colorado. Science n.s.
27:208 (Abstract).

464. Ramaley, F., and W. W. Robbins. 1909. Studies
in lake and streamside vegetation. I. Redrock Lake near
Ward, Colorado. Univ. Colo. Studies 6:13.3-168 -I- 4
plates.

465. Ratcliff, H. M. 1941. Winter range conditions in
Rocky Mountain National Park. Trans. N. Amer. Wild-
life Conf. 6:132-139.

466. Raven, K. A. 1948. Ecology of a timberline area
in the Colorado Front Range. Unpubl. thesis, Univ. of
Colo., Boulder. 83 pp.

467. Rector, C. D. 1979. Lower Gunnison River Basin
wetland inventory and evaluation. Unpubl. thesis, Univ.
of Colo., Boulder. 71 pp.

468. Rector, C. D., E. W. Mustard, and J. T. Windell.
1978. Preliminary report. Lower Gunnison River Basin
wetland inventory and evaluation. USDA Soil Conserv.
Serv., Denver, Colo. 65 pp.

469. Reed, E. L. 1917. Meadow vegetation in the
montane region of northern Colorado. Bull. Torr. Bot.
Club. 44:97-109.

470. Regelin, W. L., J. G. Nagy, and O. C. Wallmo.
1978. Effects of snowdrifts on mountain shrub commu-
nities. Pages 414-419 in Xlllth Int. Congr. of Game
Biol. [Mar. 11-15, 1977-Atlanta, Ga.]. Wildl. Soc, Wildl.
Manage. Inst., Washington, D.C.

471. Reichhardt, K. L. 1976. An ecological assessment
of the Pawnee Buttes area. Unpubl. report, Dept. of
Biol. Sci., Univ. of N. Colo., Greeley. 58 pp.

472. Reichhardt, K. L. 1978. Succession of old fields
on the Pawnee National Grasslands. Unpubl. thesis,
Univ. of N. Colo., Greeley. 89 pp.

473. Reid, E. H., and L. D. Love. 1951. Range-water-
shed conditions and recommendations for management,
Elk Ridge and Lower Elk Ridge cattle allotments,
Roosevelt National Forest, Colorado. Unpubl. report,
USDA For. Serv. 123 pp. On file at Colo. St. Univ. Li-
brary, A13.2/W31r.

474. Reid, W. H. 1974. Analysis of plant ecological
systems through simulation of individual organisms. Un-
publ. dissertation, Univ. of Colo., Boulder. 265 pp.

475. Rice, R. R. 1974. Productivity of Festuca thurberi
Missionary Ridge, Colorado, 1972. J. Colo.-Wyom. Acad.
Sci. 7(5):41 (Abstract).

476. Richards, J. D., and R. L. Dix. 1979. An environ-
mental model of the subalpine forest of the San Juan
Mountains, Colorado. Bull. Ecol. Soc. Am. 60(2): 103
(Abstract).

477. Risser, P. G. 1971. Plant community structure.
Pages 41-57 in N. R. French, ed.. Preliminary analysis of
structure and fimction in grasslands. Range Sci. Dept.,
Sci. Series No. 10, Colo. St. Univ., Fort Collins.

478. Roach, A. W. 1948. The ecology of vegetational
change in the foothills ecotone near Boulder, Colorado.
Unpubl. thesis, Univ. of Colo., Boulder. 81 pp.

479. Robbins, W. W. 1908. Studies of mesa and foot-
hill vegetation. 4. Distribution of the deciduous trees
and shrubs on the mesas. Univ. Colo. Studies 6:36-49.

480. Robbins, W. W. 1910a. A botanical trip in north-
western Colorado. Univ. Colo. Studies 7:115-125.

481. Robbins, W. W. 1910b. Climatology and vegeta-
tion in Colorado. Bot. Gaz. 49:256-280.

Great Basin Naturalist

Vol. 43, No. 1

482. Robbins, W. W. 1917. Native vegetation and cli-
mate of Colorado in their relation to agricultnre. Colo.
Agr. Exp. Sta Bull. 224. 56 pp.

48.3. Bobbins, W. W. 1918. Successions of vegetation
in Boulder Park, Colorado. Bot. Gaz. 65:493-525.

484. Bobbins, W. W., and G. S. Dodds. 1908. Studies
of mesa and foothill vegetation. 3. Distribution of con-
ifers on the mesa. Univ. Colo. Studies 6:31-36.

485. Bobertson, D. B., J. L. Nielsen, and N. H. Bare.
1966. Vegetation and soils of alkali sagebrush and adja-
cent big sagebrush ranges in North Park, Colorado. J.
Range Mgmt. 19:17-20.

486. Bogers, C. M. 1950. The vegetation of the Mesa
de Maya region of Colorado, New Mexico, and Okla-
homa. Unpubl. dissertation, Univ. of Michigan, Ann Ar-
bor. 125 pp.

487. Bogers, C. M. 1953. The vegetation of the Mesa
de Maya region of Colorado, New Mexico, and Okla-
homa. Lloydia 16:257-291.

488. Boughton, B. D. 1966. Age structure of browse
populations. Unpubl. thesis, Colo. St. Univ., Fort Col-
lins. 154 pp.

489. Boughton, B. D. 1972. Shrub age structures on a
mule deer winter range in Colorado. Ecology
S3:615-625.

490. Bowdabaugh, K. M. 1978. The role of fire in the
ponderosa pine-mixed conifer ecosystems. Unpubl.
thesis, Colo. St. Univ., Fort Collins. 121 pp.

491. Buangpanit, N. 1977. Primary productivity of
sandhills grassland. Unpubl. dissertation, Colo. St. Univ.,
Fort Collins. 165 pp.

492. Bydberg, P. A. 1913. Phytogeographical notes on
the Rocky Mountain region. I. Alpine region. Bull. Torr.
Bot. Club 40:677-686.

493. Bydberg, P. A. 1914a. Phytogeographical notes
on the Bocky Mountain region. II. Origin of the alpine
flora. Bull. Torr. Bot. Club 41:89-103.

494. Bydberg, P. A. 1914b. Phytogeographical notes
on the Rocky Mountain region. III. Formations in the al-
pine zone. Bull. Torr. Bot. Club 41:459-474.

495. Bydberg, P. A. 1915a. Phytogeographical notes
on the Bocky Mountain region. IV. Forests of the sub-
alpine and montane zones. Bull. Torr. Bot. Club
42:11-2.5.

496. Bydberg, P. A. 1915b. Phytogeographical notes
on the Bocky Mountain region. V. Grasslands of the sub-
alpine and montane zones. Bull. Torr. Bot. Club
42:629-642.

497. Bydberg, P. A. 1916a. Phytogeographical notes
on the Bocky Mountain region. VI. Distribution of the
subalpine plants. Bull. Torr. Bot. Club 43:343-364.

498. Bydberg, P. A. 1916b. Vegetative life zones of
the Bocky Mountain region. Mem. N.Y. Bot. Garden
6:477-499.

499. Bydberg, P. A. 1917. Phytogeographical notes on
the Rocky Mountain region. VII. Formations in the sub-
alpine zone. Bull. Torr. Bot. Club 44:431-454.

500. Bydberg, P. A. 1919. Phytogeographical notes on
the Rocky Mountain region. VIII. Distribution of the
montane plants. Bull. Torr. Bot. Club 46:295-327.

501. Bydberg, P. A. 1920. Phytogeographical notes on
the Rocky Mountain region. IX. Wooded formations of
the montane zone of the southern Rockies. Bull. Torr.
Bot. Club 47:441-454.

502. Bydberg, P. A. 1921. Phytogeographical notes on
the Bocky Mountain region. X. Grasslands and other
open formations of the montane zone of the southern
Bockies. Bull. Torr. Bot. Club 48:315-326.

503. Savage, D. A. 1937. Drought survival of native
grass species in the central and southern Great Plains,
19,35. USDA Tech. Bull. No. 549.

504. Savage, D. A., and H. E. Bunyon. 1937. Natural
revegetation of abandoned farm land in the central and
southern Great Plains. Pages 178-182 in Bep. 4th Int.
Grassland Congress, Aberystwyth, Gt. Brit., Sec. 1.
Grassland Ecology.

505. Schaum, J. L. 1979. An assessment of the South
Platte lowland riparian habitat for natural area designa-
tion. Unpubl. thesis, Univ. of N. Colo., Greeley. 129 pp.

506. Scheideman, L. L. 1971. The effects of 2,4-D on
submontane vegetation, southcentral Colorado. Unpubl.
thesis. Western State College, Gunnison, Colo. 77 pp.

507. Schmid, J. M., and T. E. Hinds. 1974. Devel-
opment of spruce-fir stands following spruce beetle out-
breaks. USDA For. Serv. Bes. Paper RM-131, Rocky Mt.
For. and Range Expt. Stn., Fort Collins, Colo. 16 pp.

508. Schmoll, H. M. 19.35. Vegetation of the Chimney
Rock area, Pagosa-Piedra region, Colorado. Priv. Edi-
tion, Distr. by Univ. of Chicago Libraries, Chicago, 111.
58 pp.

509. Schneider, E. C. 1909. The distribution of woody
plants in the Pike's Peak Region. Colo. Coll. Publ. Sci.
Series 12:136-169.

510. Schneider, E. C. 1911. The succession of plant
life on the gravel slides in the vicinity of Pike's Peak.
Colo. Coll. Publ. Sci. Series 12(8):289-.301.

511. Schroeder, R. E. 1977. Species composition
changes on three range types under two grazing systems.
Unpubl. thesis, Colo. St. Univ., Fort Collins. 65 pp.

512. Schwan, H. E., and D. F. Costello. 1951. The
Rocky Mountain alpine type-range conditions, trends
and land use (a preliminary report). Unpubl. report,
USDA For. Serv. Rocky Mtn. Region (R2), Denver, Colo.
18 pp.

513. Scott-Williams, B. W. 1965. The ecology of the
alpine tundra on Trail Bidge. Pages 13-16 in C. B.
Schulz and H. T. Smith, eds., Guidebook for one-day
field conferences-Boulder area, Colorado. Vllth Int. As-
soc, for Quat. Bes. Congress, Nebraska Acad. Sci.,
Lincoln.

514. Shaddle, P. H. 1939. An ecological study of the
vegetation of the Black Forest of Colorado. Unpubl.
thesis, Univ. of Colo., Boulder.

515. Shanks, D. L. 1977. Aerial photo densitometry
for rangeland planning and evaluation. Unpubl. thesis,
Colo. St. Univ., Fort Collins. 66 pp.

516. Shantz, H. L. 1906a. A study of the vegetation of
the Mesa region east of Pike's Peak: the Bouteloua for-
mation: I. Structure of the formation. Bot. Gaz.
42:16-47.

517. Shantz, H. L. 1906b. A study of the vegetation of
the mesa region east of Pike's Peak: the Bouteloua for-
mation: II. Development of the formation. Bot. Gaz.
42:179-207.

518. Shantz, H. L. 1911. Natural vegetation as an in-
dicator of the capabilities of land for crop production in
the Great Plains area. USDA Bur. Plant Ind. Bull. No.
201. 100 pp.

519. Shantz, H. L. 1917. Plant succession on aban-
doned roads in eastern Colorado. J. Ecol. 5:19-42.

January 1983

Baker: Colorado Bibliography

520. Shantz, H. L. 1923. The natural vegetation of
the Great Plains region. Ann. Assoc. Am. Geog.
13:81-107.

521. Shaver, J. C., and H. G. Fisser. 1972. Detailed
vegetation mapping of the IBP Pawnee site. U.S. Int.
Biol. Prog., Grassland Biome Tech. Kept. No. 150. 44
pp.

522. Shepherd, H. R. 1975. Vegetation of two dis-
similar bighorn sheep ranges in Colorado. Colo. Div.
Wildlife Div. Kept. No. 4. 223 pp.

523. Shubert, M. L. 1950. Some problems of aspen
distribution in South Park. J. Colo.-Wyom. Acad. Sci.
4(2):53 (Abstract).

524. Shubert, M. L., W. Binder, and E. L. Todut.
1962a. Ecology of the Black Forest of Colorado re-exam-
ined. J. Colo.-Wyom. Acad. Sci. 5(3):35-36 (Abstract).

525. Shubert, M. L., W. Binder, and E. L. Todut.
1962b. Historical events in an ecological enigma. Bull.
Ecol. Soc. Am. 43(2):52 (Abstract).

526. Simkins, E. C. 1931. The alpine flora of Mt. Ni-
wot, Colorado. Unpubl. thesis, Univ. of Colo., Boulder.
68 pp.

527. Sims, P. L., B. E. Dahl, and A. H. Denham. 1976.
Vegetation and livestock response at three grazing in-
tensities on sandhill rangeland in eastern Colorado.
Colo. Agric. Expt. Stn. Tech. Bull. No. 130. 48 pp.

528. Sims, P. L., J. S. Singh, and W. K. Lauenroth.
1978. The structure and function of ten western North
American grasslands. I. Abiotic and vegetational charac-
teristics. J. Ecol. 66:251-285.

529. Slauson, W. L. and R. T. Ward. 1979. How spe-
cies form communities in alpine tundra. Bull. Ecol. Soc.
Am. 60(2):93 (Abstract).

530. Smith, D. R. 1967. Effects of cattle grazing on a
ponderosa pine-bunchgrass range in Colorado. USDA
For. Serv. Tech. Bull. No. 1371. 60 pp.

531. Smith, E. C. 1942. The willows of Colorado. Am.
Midi. Nat. 27:217-252.

532. Smith, E. L., Jr. 1966. Soil-vegetation relation-
ships of some Artemisia types in North Park, Colorado.
Unpubl. dissertation, Colo. St. Univ., Fort Collins.

533. Somers, P., G. E. Nichols, and R. W. Stransky.
1980. Final report: baseline ecological study of Narra-
guinnep Research Natural Area, San Juan Nat. For., Un-
publ. report. Fort Lewis College, Durango, Colo. 23 pp.

534. Spencer, J. P. 1975. The effects of winter weath-
er modification on alpine plant communities in the San
Juan Mountains, Colorado. Unpubl. thesis, Univ. of
Colo., Boulder. 122 pp.

535. Sperry, O. E. 1936. A study of the growth, tran-
spiration, and distribution of the conifers of the Rocky
Mountain National Park. Bull. Torr. Bot. Club
63:75-103.

536. Springfield, H. W. 1976. Characteristics and
management of southwestern pinyon-juniper ranges: the
status of our knowledge. USDA For. Serv. Res. Paper
RM-160, Rocky Mt. Forest and Range Expt. Sta., Fort
Collins, Colo. 32 pp.

537. Stahelin, R. 1943. Factors influencing the natural
restocking of high altitude bums by coniferous trees in
the central Rocky Mountains. Ecology 24:19-30.

538. Steams-Roger, Inc. 1978a. Rawhide Energy Proj-
ect. Plant site and railroad spur, ecological studies. Vol.
I. Baseline summary, impact assessment and recommen-
dations. Report to Platte River Power Authority. 102 pp.

539. Steams-Roger, Inc. 1978b. Rawhide Energy Proj-
ect. Transmission system, ecological investigations. Vol.
II. Technical Baseline Report. Report to Platte River
Power Authority. 51 pp.

540. Steen, O. A. and R. L. Dix. 1974. A preliminary
classification of Colorado subalpine forests: a working
guide. Unpubl. report on file at Rocky Mt. For. and
Range Expt. Stn., Fort Collins, Colo. 9 pp.

541. Steinhoff, H. W. 1962. Certain aspects of the
ecology of the ponderosa pine community as revealed by
a study of twelve small enclosures in Rocky Mountain
National Park. Unpubl. report on file at Rocky Moun-
tain National Park Hqtrs., Estes Park, Colo.

542. Steinhoff, H. W. 1978. Management of Gambel
oak associations for wildlife and livestock. Unpubl. re-
port on file at Rocky Mountain Regional Office, USDA
Forest Service, Denver, Colo. 119 pp.

543. Stevens, D. R. 1970. Ungulate ecology studies in
Rocky Mountain National Park. U.S. Dept. Int., Nat.
Park Serv. Progress Rep., Unpubl. report on file at the
office of Park Service Biologist, Rocky Mt. Nat. Park
Hqtrs., Estes Park, Colo. 74 pp.

544. Stevens, D. R. 1972. Vegetation map of the east-
em winter range. Unpubl. report on file at Rocky Moun-
tain National Park Hqtrs., Estes Park, Colo.

545. Stevens, D. R. 1976a. Problems of revegetation
of alpine tundra. Proc. 1st Conf. on Sci. Res. in Nat.
Parks 1:241-245. U.S. Dept. Int., Nat. Park Serv., Trans,
and Proc. Series No. 5.

546. Stevens, D. R. 1976b. Ecology of elk on alpine
tundra in Rocky Mountain National Park. Proc. 1st
Conf. on Sci. Res. in Nat. Parks 1:421-426. U.S. Dept.
Int., Nat. Park Serv., Trans, and Proc. Series No. 5.

547. Steward, D. G. 1977. Tree-island communities of
Rocky Mountain National Park. Unpubl. thesis, Univ. of
Colo., Boulder. 140 pp.

548. Stewart, B. K. 1940. Plant ecology and paleo-
ecology of the Creede Valley, Colorado. Unpubl. dis-
sertation, Univ. of Colo., Boulder. 154 pp.

549. Stewart, O. C. 1953. Why the Great Plains are
treeless. Colo. Quart. 2:40-50.

550. Stigall, B. M. 1934. The vegetation of Cuchara
Valley, Colorado. Unpubl. report on file in Univ. of
Colo. Library, 581.9788/St52. 56 pp.

551. Deleted.

552. Strong, L. L. 1980. Estimating phytomass pro-
duction of habitat types on sagebrush steppe. Unpubl.
thesis, Colo. St. Univ., Fort Collins. 133 pp.

553. Strong, N. M. 1978. An ecological study of the
Pierce Preserve, Routt County, Colorado. Unpubl. re-
port on file at The Nature Conservancy, Denver, Colo.
88 pp.

554. Strong, N. M., R. D. Comer, and D. G. Steward.
1978. An ordination and classification of the woody veg-
etation in Boulder County, Colorado. J. Colo.-Wyom.
Acad. Sci. 10(l):35-36 (Abstract).

555. Stroud, D. W., and H. G. Fisser. 1972. Plant pat-
tern, microrelief and Bouteloua gracilis culm density,
studies on the Pawnee site: 1970-71. U.S. Int. Biol.
Prog., Grassland Biome Tech. Rep. No. 186.

556. Sudworth, G. B. 1900a. White River Plateau
Timber Land Reserve. U.S. Geol. Survey 20th Ann.
Rept., Part V: 117-179.

557. Sudworth, G. B. 1900b. Battlement Mesa Forest
Reserve. U.S. Geol. Survey 20th Ann. Rept., Part
V: 181-243.

Great Basin Naturalist

Vol. 43, No. 1

558. Sweeny, J. R. 1975. Effects of snow on oak brush.
Unpubl. dissertation, Colo. St. Univ., Fort Collins, Colo.
101 pp.

559. Sweeny, J. R., and H. W. Steinhoff. 1976. Effects
of snow on oak brush. Pages 343-366 in H. W. Steinhoff
and J. D. Ives, eds.. Ecological impacts of snowpack aug-
mentation in the San Juan Mountains, Colorado. Final
report of the San Juan Ecol. Proj., Colo. St. Univ., Fort
Collins, to the USDI Bur. Reel., Div. of Atmos. Wat.
Res. Mgmt., Denver, Colo.

560. Swift, R. L. 1974. Vegetation-site relations of
ponderosa pine forest in the Front Range of central Col-
orado. Unpubl. thesis, Colo. St. Univ., Fort Collins. 121
pp.

561. Tassel, S., and J. L. Carter. 1980. The vascular
flora of EH Paso County. J. Colo.-Wyom. Acad. Sci.
12(1):29 (Abstract).

562. Terwilliger, C, Jr.. and E. L. Smith. 1978. Range
resource types in North Park, Colorado. Colo. St. Univ.,
Range Sci. Dept. Sci. Series No. 32. 48 pp.

563. Terwilliger, C, Jr., and P. G. Threlkeld. 1974.
Vegetation map, Piceance Basin and Roan Plateau, Col-
orado. In C. W. Cook, ed., Surface rehabilitation of land
disturbances resulting from oil shale development. Env.
Resources Center, Colo. St. Univ., Fort Collins, Tech.
Rept. Series No. 1.

564. Terwilliger, C, Jr., and J. A. Tiedeman. 1978.
Habitat types of the mule deer critical winter range and
adjacent steppe region of Middle Park, Colorado. Un-
publ. report on file at Rocky Mt. For. & Range Expt.
Stn., Fort Collins, Colo. 108 pp.

565. Terwilliger, C, Jr., K. Hess, and C. H. Wasser.
1979. The habitat types of Region II, U.S. Forest Ser-
vice, a preliminary list and description. An initial prog-
ress report for "Habitat Type Classification, Region
Two, United States Forest Service" Cooperative Agree-
ment No. 16-845-CA.

566. Thilenius, J. F. 1975. Alpine range management
in the western United States— principles, practices, and
problems: the status of our knowledge. USDA For. Serv.
Res. Paper RM-157, Rocky Mt. For. & Range Exp. Stn.,
Fort Collins, Cole. 32 pp.

567. Deleted.

568. Thome Ecological Institute. 1973a. Environmen-
tal setting of the Parachute Creek Valley: an ecological
inventory. Unpubl. report prepared for Colony Develop.
Operation, Atlantic Richfield Co., Operator, Denver,
Colo., by Thome Ecol. Inst., Boulder, Colo. [Vegetation:
p. 36-40, map].

569. Thome Ecological Institute. 1973b. Part 1, The
Colony Environmental Study— Parachute Creek, Gar-
field County, Colorado. Unpubl. report prepared for
Colony Development Operation, by Thorne Ecol. Inst.,
Boulder, Colo. [Vegetation: p. 26-30].

570. Tiedeman, J. A. 1978. Phyto-edaphic classifica-
tion of the Piceance Basin. Unpubl. dissertation, Colo.
St. Univ., Fort Collins. 281 pp.

571. Tiedeman, J. A., and C. Terwilliger, Jr. 1978.
Phyto-edaphic classification of the Piceance Basin. Colo.
St. Univ., Range Sci. Dept. Sci. Series No. 31. 265 pp.

572. Tietjen, H. P., C. H. Halvorson, P. L. Hegdal,
and A. M. Johnson. 1967. 2,4-D herbicide, vegetation,
and pocket gopher relationships. Black Mesa, Colorado.
Ecology 48:634-643.

573. Tomanek, G. W., and G. K. Hulett. 1972. A sur-
vey of some remnant prairies of the central Great Plains.
Pages 34-39 in J. H. Zimmerman, ed., The Second Mid-
west Prairie Conference, Univ. of Wise. Arboretum,
Madison.

574. Trammell, J. R., Jr., and J. W. Marr. 1974. Phy-
tosociology of aspen stand-types on the east slope of the
Front Range, Boulder County, Colorado. J. Colo.-
Wyom. Acad. Sci. 7(5):41 (Abstract).

575. Trammell, V. M., and J. R. Trammell, Jr. 1977.
Plant communities of Roxborough Park, Douglas Coun-
ty, Colorado. Unpubl. report, Arapahoe Community
College, Littleton, Colo. 14 pp.

576. Trlica, M. J., and P. Hackney. 1977. Vegetative
and wildlife inventory of Sommerville Table. Unpubl.
report on file at the Bureau of Land Mgmt., Royal
Gorge Resource Area, Canon City, Colo. 20 pp.

577. Turner, G. T. 1964. Vegetation changes on a
mountain grassland in west central Colorado during 19
years. Bull. Ecol. Soc. Am. 45(3): 108 (Abstract).

578. Turner, G. T. 1969. Responses of mountain grass-
land vegetation to gopher control, reduced grazing, and
herbicide. J. Range Mgmt. 22:377-383.

579. Turner, G. T., and E. J. Dortignac. 1954. Infil-
tration, erosion, and herbage production of some moun-
tain grasslands in western Colorado. J. For. 52:858-860.

580. Tumer, G. T. and H. A. Paulsen, Jr. 1976. Man-
agement of mountain grasslands in the central Rockies:
the status of our knowledge. USDA For. Serv. Res. Paper
RM-161, Rocky Mt. For. and Range Expt. Stn., Fort
Collins, Colo. 24 pp.

581. Ueckert, D. N. 1968. Diets of some grasshoppers
common on mountain herbland in northern Colorado.
Unpubl. thesis, Colo. St. Univ., Fort Collins.

582. Ulman, W. J., and H. L. Young, eds., 1970. The
Dodd property natural area study. Unpubl. report,
Dept. of Geogr., Univ. of Colo., Boulder. 68 pp.

583. Ungar, I. A. 1974a. Halophyte communities of
Park County, Colorado. Bull. Torr. Bot. Club
101:145-152.

584. Ungar, I. A. 1974b. The effect of soil factors on
the distribution of halophytic species at South Park, Col-
orado. Bull. Ecol. Soc. Am. 55(2):31-32 (Abstract).

585. Ungar, I. A. 1974c. Inland halophytes of the
United States. Pages 235-305 in R. J. Reimold and W.
H. Queen, eds., Ecology of halophytes. Acad. Press, Inc.,
New York.

586. USDA Soil Conservation Service. 1969. Major
land resource area and generalized land use map— Colo-
rado. SCS, Portland, Oregon.

587. USDA Soil Conservation Service. 1978. Range
site descriptions for Colorado. Tech. Guide, Section II-E
Colo. St. Office, Denver, Colo.

588. USDI Bureau of Land Management. 1979. Pages
2-6 to 2-15 in Grand Junction Resource Area grazing
management— draft environmental statement.

589. USDI Bureau of Reclamation. 1976. Flora and
terrestrial vertebrate studies of the Grand Valley, Colo-
rado. Final Report to U.S. Bur. Reel, from Ecology Con-
sultants, Inc., Fort Collins, Colo. [Vegetation: p. 56-85,
283-354].

590. USDI National Park Service. 1932. Report on
condition of portions of elk and deer winter range in
Rocky Mountain National Park. Unpubl. report on file
at Rocky Mountain National Park Hqtrs., Estes Park,
Colo.

January 1983

Baker: Colorado Bibliography

591. Utah Environmental and Agricultural Consul-
tants. 1973. Environmental setting, impact, mitigation,
and recommendations for a proposed oil products pipe-
line between Lisbon Valley, Utah, and Parachute Creek,
Colorado. Colony Development Operation, Atlantic
Richfield Co., Denver, Colo. [Vegetation: p. 3-38].

592. Vestal, A. G. 1913. Plains vegetation adjoining
the mountains: the region about Boulder in Colorado.
Unpubl. thesis, Univ. of Colo., Boulder. 40 pp.

593. Vestal, A. G. 1914. Prairie vegetation of a moun-
tain-front area in Colorado. Bot. Gaz. 58:377-400.

594. Vestal, A. G. 1915. Phytogeography of the east-
em mountain front in Colorado. Unpubl. dissertation,
Univ. of Chicago, Chicago, 111. 41 pp.

595. Vestal, A. G, 1917. Foothills vegetation in the
Colorado Front Range. Bot. Gaz. 64:353-385.

596. Vestal, A. G. 1919. Phytogeography of the east-
em mountain front in Colorado. I. Physical geography
and distribution of vegetation. Bot. Gaz. 68:153-193.

597. Vories, K. C. 1974. A vegetation inventory and
analysis of the Piceance Basin and adjacent drainages.
Unpubl. thesis. Western State College, Gunnison, Colo.
243 pp.

598. Walker, R. B. 1962. Comparison of a wide south-
facing and a narrow north-facing canyon. Pages 103-119
in A. M. Woodbury, ed.. Ecological studies of the flora
and fauna of the Curecanti Reservoir basins, western
Colorado. Univ. of Utah, Dept. of Anthropol., Anthro-
pol. Papers No. 59 (Upper Colo. Ser. No. 8).

599. Walter, R. G. 1963. The vascular flora of the
north end of South Park. Unpubl. thesis, Univ. of Colo.,
Boulder. 61 pp.

600. Ward, A. L. and J. O. Keith. 1962. Feeding
habits of pocket gophers on mountain grasslands. Black
Mesa, Colorado. Ecology 43:744-749.

601. Ward, R. T. 1961a. An ordination of alpine vege-
tation. Bull. Ecol. Soc. Am. 42:147-148 (Abstract).

602. Ward, R. T. 1961b. Species association in the al-
pine. J. Colo.-Wyom. Acad. Sci. 5(2):38 (Abstract).

603. Ward, R. T. 1966. Ecological studies of alpine
tundra in Rocky Mountain National Park. Bull. Ecol.
Soc. Am. 47(3): 140 (Abstract).

604. Ward, R. T., W. Slauson, and R. L. Dix. 1974a.
The natural vegetation in the landscape of the Colorado
oil shale region. Pages 9-17 in C. W. Cook, ed.. Surface
rehabilitation of land disturbances resulting from oil
shale development. Environ. Resources Center, Colo. St.
Univ., Information Series No. 11« Fort Collins. [Execu-
tive summary of Tech. Rep. Series No. 1:30-66].

605. Ward, R. T., W. Slauson, and R. L. Dix. 1974b.
The natural vegetation in the landscape of the Colorado
oil shale region. Pages 30-66 in C. W. Cook, ed.. Surface
rehabilitation of land disturbances resulting from oil
shale development. Final Rept., Phase I, to the Colo.
Dept. of Nat. Resources. Environ. Resources Center,
Colo. State Univ., Fort Collins, Tech. Rep. Series No. 1.

606. Wardle, P. 1968. Engelmann spmce {Picea engel-
mannii Engel.) at its upper limits on the Front Range,
Colorado. Ecology 49:483-495.

607. Wasser, C. H., and J. L. Retzer. 1965. Ecology
and utility of the central Rocky Mountain alpine zone.
9th Intemat. Grassland Congr. Proc. 9:357-361.

608. Weaver, J. E., and F. W. Albertson. 1944. Na-
ture and degree of recovery of grassland from the great
drought of 1933 to 1940. Ecol. Monogr. 14:393-479.

609. Weaver, J. E., and F. W. Albertson. 1956. Grass-
lands of the Great Plains: their nature and use. Johnsen
Publ. Co., Lincoln, Nebr. 395 pp.

610. Webber, P. J., S. V. Clark, and V. Komdrkovi.
1977. Botanical survey of the environs of the Rocky Flats
(ERDA) Plant: mapping and ordination analyses. Pages
29-34 in D. C. Hunt, ed.. Transactions of meeting on
Rocky Flats buffer zone ecological and environmental
research meeting, 4 March 1977, Rockwell Inter-
national, Rocky Flats Plant, Golden, Colo.

611. Webber, P. J., J. C. Emerick, D. C. E. May, and
V. Komdrkova. 1976. The impact of increased snowfall
on alpine vegetation. Pages 201-264 in H. W. Steinhoff
and J. D. Ives, eds.. Ecological impacts of snowpack aug-
mentation in the San Juan Mountains, Colorado. Final
Report of the San Juan Ecology Project, Colo. St. Univ.,
Fort Collins, to the USDI Bur. Reel., Div. of Atmos.
Wat. Res. Mgmt., Denver, Colo.

612. Webber, P. J. and D. E. May. 1977. The magni-
tude and distribution of belowground plant structures in
the alpine tundra of Niwot Ridge, Colorado. Arct. and
Alp. Res. 9:157-174.

613. Weber, W. A. 1949. Preliminary observations on
plant distribution patterns in Colorado. I. The relict
flora. J. Colo.-Wyom. Acad. Sci. 4(l):45-46 (Abstract).

614. Weber, W. A. 1961. Alpine floristic components
of the southem Rocky Mountains. Bull. Ecol. Soc. Am.
42(4): 164 (Abstract).

615. Weber, W. A. 1965. Plant geography in the
southern Rocky Mountains. Pages 453-468 in H. E.
Wright, Jr. and D. G. Frey, eds.. The Quatemary of the
United States. Princeton Univ. Press, Princeton, N.J.

616. Weber, W. A. 1972. Pages 2, 4-5 in Rocky
Mountain Flora. Colo. Assoc. Univ. Press, Boulder. 4th
ed.

617. Weir, G. H. 1976. Palynology, flora, and vegeta-
tion of Hovenweep National Monument: implications
for aboriginal plant use on Cajon Mesa, Colorado and
Utah. Unpubl. dissertation, Texas A & M Univ., College
Station, Texas. 217 pp.

618. Welden, C. W. 1981. Pattern in alpine tundra.
Unpubl. thesis, Colo. St. Univ., Fort Collins. 144 pp.

619. Wells, P. V. 1965. Scarp woodlands, transported
grassland soils, and concept of grassland climate in the
Great Plains region. Science 148:246-249.

620. Wells, P. V. 1970a. Postglacial vegetational his-
tory of the Great Plains. Science 167:1574-1582.

621. Wells, P. V. 1970b. Historical factors controlling
vegetation patterns and floristic distributions in the cen-
tral plains region of North America. Pages 211-221 in
W. Dort and J. D. Jones, eds.. Pleistocene and recent en-
vironments of the central Great Plains. Univ. of Kansas
Press, Lawrence.

622. Welsh, S. L., J. K. Rigby, and W. K. Hamblin.
1980. A survey of Natural Landmark areas of the north
portion of the Colorado Plateau. Report prepared for
the Heritage Conservation and Recreation Service, U.S.
Dept. Interior. 270 pp.

623. Western Engineers, Inc. 1981. Reconnaissance of
terrestrial vegetation at the Taylor Draw Reservoir site.
Unpubl. report on file at Western Engineers, Inc.,
Grand Junction, Colo. 17 pp.

624. Whipple, S. A. 1973. The species composition
and age-structure of subalpine forests of the Front
Range. Unpubl. thesis, Colo. St. Univ., Fort Collins.

Great Basin Naturalist

Vol. 43, No. 1

625. Whipple, S. A. 1975. The influence of environ-
mental gradients on vegetational structure in the sub-
alpine forest of the southern Rocky Mountains. Unpubl.
dissertation, Colo. St. Univ., Fort Collins.

626. Whipple, S. A. 1976. Successional and popu-
lational dynamics of the major tree species in the Colo-
rado Rocky Mountain subalpine forest. Bull. Ecol. Soc.
Am. 57(1):59 (Abstract).

627. Whipple, S. A. 1977. The importance of buried
seeds to vegetational composition in old-growth Colo-
rado subalpine forest. Bull. Ecol. Soc. Am. 58(2): 15
(Abstract).

628. Whipple, S. A. 1978a. Structural variation in a
Colorado subalpine forest. Bull. Ecol. Soc. Am. 59(2):79
(Abstract).

629. Whipple, S. A. 1978b. The relationship of bur-
ied, germinating seeds to vegetation in an old-growth
Colorado subalpine forest. Can. J. Bot. 56:1505-1509.

630. Whipple, S. A. and R. L. Dix. 1979. Age struc-
ture and successional dynamics of a Colorado subalpine
forest. Am. Midi. Nat. 101:142-158.

6.31. Whitfield, C. J. 1933. The vegetation of the
Pike's Peak region. Ecol. Monogr. 3:75-105.

632. Wilford, B. H. 1967. Conditions and trends of
forest stands within the Park Range weather modifica-
tion study area, Routt National Forest. Unpubl. report
on file at Hahns Peak Dist. Headquarters, Routt N.F.,
Steamboat Springs, Colo. 23 pp.

633. Willard, B. E. 1960a. The ecology and phytoso-
ciology of the Tundra Curves, Trail Ridge, Colorado.
Unpubl. thesis, Univ. of Colo., Boulder. 144 pp.

634. Willard, B. E. 1960b. Application of the methods
of Zurich-Montpellier phytosociology to some alpine
vegetation on Trail Ridge, Rocky Mountain National
Park, Colorado. J. Colo.-Wyom. Acad. Sci. 5(l):36-37
(Abstract).

635. Willard, B. E. 1961. The effects of visitor impact
on Rocky Mountain alpine tundra. Bull. Ecol. Soc. Am.
42:141-142 (Abstract).

636. Willard, B. E. 1963. Phytosociology of the alpine
tundra of Trail Ridge, Rocky Mountain National Park,
Colorado. Unpubl. dissertation, Univ. of Colo., Boulder.

637. Willard, B. E. 1964. Phytosociology of alpine
tundra ecosystems of Trail Ridge, Colorado. Bull. Ecol.
Soc. Am. 45(3):111-112 (Abstract).

638. Willard, B. E. 1967. Effects of visitors on the
ecosystems of Rocky Mountain National Park. Bull.
Ecol. Soc. Am. 48(2):86 (Abstract).

639. Willard, B. E. 1979. Plant sociology of alpine
tundra. Trail Ridge, Rocky Mountain National Park,
Colorado. Colo. Sch. Mines Quart. 74(4):1-119.

640. Willard, B. E., and J. W. Marr. 1970. Effects of
human activities on alpine tundra ecosystems in Rocky
Mountain National Park, Colorado. Biol. Conserv.
2:257-265.

641. Willard, B. E., and J. W. Marr. 1971. Recovery
of alpine tundra under protection after damage by hu-
man activities in the Rocky Mountains of Colorado. Biol.
Conserv. 3:181-190.

642. Willard, B. E., J. W. Marr, and D. Buckner.
1973. Ecological analyses of potential shale oil products
pipeline corridors in Colorado and Utah. Report to the
Colony Development Operation by Thome Ecological
Institute and University of Colo., Lab. of Mt. Ecol. for
Man, Boulder, Colo.

643. Williams, T. E. 1938. Ecology of the Black For-
est. J. Colo.-Wyom. Acad. Sci. 2(4):24 (Abstract).

644. Williams, T. E. 1940. Ecology of the Black For-
est of Colorado. J. Colo.-Wyom. Acad. Sci. 2(6):37
(Abstract).

645. Williams, T. E., and A. E. Holch. 1946. Ecology
of the Black Forest of Colorado. Ecology 27:139-149.

646. Wilson, H. C. 1969. Ecology and successional
patterns of wet meadows. Rocky Mountain National
Park, Colorado. Unpubl. dissertation, Univ. of Utah, Salt
Lake City. 99 pp.

647. Wintemitz, B. L. 1973. Ecological patterns in a
montane breeding bird community. Unpubl. dis-
sertation, Univ. of Colo., Boulder. 109 pp.

648. Woodbury, A. M., S. D. Durrant, and S. Flowers.

1961. A survey of vegetation in the Navajo Reservoir ba-
sin. Univ. of Utah, Dept. of Anthropol., Anthropol. Pa-
pers No. 51, 99 pp.

649. Woodbury, A. M., S. D. Durrant, and S. Flowers.

1962. A survey of vegetation in the Curecanti Reservoir
basins. Univ. of Utah, Dept. of Anthropol., Anthropol.
Papers No. 56. 98 pp.

650. Woodin, H. E. 1953. The pinyon-juniper associ-
ation in the eastern foothills of the Rockies. Unpubl. dis-
sertation, Purdue Univ., Lafayette, Ind.

651. Woodin, H. E., and A. A. Lindsey. 1953. Pinyon-
juniper east of the continental divide, as analyzed by the
line-strip method. Bull. Ecol. Soc. Am. 34(3):84
(Abstract).

652. Woodin, H. E., and A. A. Lindsey. 1954. Juniper-
pinyon east of the continental divide, as analyzed by the
line-strip method. Ecology 35:473-489.

653. Wooten, S. A. 1980. Vegetation recovery follow-
ing disturbance on the Central Plains Experimental
Range. Unpubl. thesis, Colo. St. Univ., Fort Collins.

654. Wright, C. W. 1952. An ecological description of
an isolated pinon pine grove. Unpubl. thesis, Univ. of
Colo., Boulder. 43 pp.

655. Yeager, W. W. 1937. The forest and forestry of
Mesa Verde National Park. Colo. Forester 1937:26-31.

656. Young, R. T. 1907. The forest formations of Boul-
der County, Colorado. Bot. Gaz. 44:321-352.

657. Younkin, W. E., Jr. 1970. A study of the vegeta-
tion of alpine rock outcrops in northern Colorado. Un-
publ. thesis, Colo. St. Univ., Fort Collins. 109 pp.

658. Zarn, M. 1977. Ecological characteristics of pin-
yon-juniper woodlands on the Colorado Plateau— a liter-
ature survey. USDI, Bur. of Land Mgmt., Tech. Note
No. 310. 183 pp.

659. Zimmerman, T. 1978. Skull Creek Study Area
Forestry. Unpubl. report. Bureau of Land Management,
Craig District Office, Craig, Colo. 62 pp.

660. Zuck, R. H. 1974. An ecological versus manage-
rial classification of forest communities. Unpubl. thesis,
Colo. St. Univ., Fort Collins. 110 pp.

'Acknowledgments

I am grateful to the numerous people who
provided access to unpublished reports, and
to J. Scott Peterson, who provided support
and criticism. The author would appreciate
being informed if there are omissions.

EVALUATION OF A PROGRAM TO CONTROL HYDATID DISEASE
IN CENTRAL UTAH^

Ferron L. Andersen^, John R. Crellin^-^, Craig R. Nichols*, and Peter M. Schantz^

Abstract.— A program to control hydatid disease in central Utah was evaluated by: (1) surveillance of infection in
dogs and sheep, (2) questioning adult residents of two Sanpete County communities (Fountain Green and Spring
City) about their knowledge of hydatid disease and their attitudes toward preventive measures, (3) interviewing local
officials to evaluate the proposed implementation of proper community-wide measures, (4) analyzing records of
prophylactic treatment of dogs with praziquantel, and (5) comparing scores of tests given to third and fourth grade
students before and after they colored an educational pamphlet about hydatid disease.

Infection rates of Echinococcus granulosus in dogs brought to volunteer diagnostic clinics dropped from 28.3 per-
cent in 1972 to 1.0 percent in 1979, but increased to 9.8 percent in 1981. This last rise was due mainly to the fact
that some dogs were examined that belonged to range sheepmen who had either not attended a field clinic recently
or had never attended at all. Prevalence of the parasite in slaughtered sheep decreased steadily from 13.2 percent in
1972 to 2.8 percent in 1981. With regard to the questionnaire phase of the project, 87.3 percent and 84.3 percent of
the respondents understood the role of dogs in the life cycle of E. granulosus in Fountain Green and Spring City, re-
spectively. Over 50 percent of the respondents of these two communities had worked directly with sheep sometime
in their life. In general, residents were more willing to practice preventive measures involving sheep than they were
to implement measures involving dogs alone. However, many of the recommended community-wide preventive mea-
sures were not implemented. With the coloring book, students answered an average of 62.5 percent of the questions
correctly before they colored the pamphlet and 83.3 percent afterward. Overall, our results suggest that residents of
Sanpete County are knowledgeable about hydatid disease and its mode of transmission, and that, in general, progress
has been made in control of hydatid disease in central Utah.

Utah has had the most autochthonous cases
of hydatidosis (39) in man reported from the
contiguous United States (Crellin et al.,
1982), with the first reported case diagnosed
in 1944 (Carlquist and Dowell 1951). Studies
on dogs, foxes, and coyotes were undertaken
in the early 1950s to identify the definitive
host, but these were unsuccessful (Butler and
Grundmarm 1951, Grundmann et al. 1953,
Butler and Grundmann 1954). The normal
hosts of Echinococcus granulosus in Utah
(dogs and sheep) were revealed in 1969 as
part of an investigation into the death of a
nine-year old boy in the community of Herri-
man (Kahn et al. 1972), near Salt Lake City.
Concurrently, parasitologists at Brigham
Young University began surveillance of the
parasite in dogs and sheep in central Utah
(Fox et al. 1970, Andersen et al. 1973, Love-
less et al. 1978). As more work was done, it
became apparent that the main foci of in-
fection were in that area of Utah, especially

Sanpete County. As a result, a cooperative
program to study and control hydatid disease
in central Utah was begun in 1971 by per-
sonnel from Brigham Young University (Pro-
vo, Utah), the Utah Department of Health
(Salt Lake City, Utah), and the Centers for
Disease Control (Atlanta, Georgia) (Andersen
et al. 1974).

Materials and Methods

Description of Study Area

Sanpete County is in the center of Utah
and has a total area of 4,136 sq km. A valley
(1700 m elevation) bordered by mountains
(3400 m elevation) runs the length of the
county. There are 14,615 people in the coun-
ty (U.S. Bureau of the Census 1980a), 90 per-
cent of which are Mormons (The Church of
Jesus Christ of Latter-day Saints) (Stinner et
al. 1978). Unlike most farming regions in the

'This project was supported in part by U.S. Public Health Service Grant AI-10588.

'Department of Zoology, Brigham Young University, Provo, Utah 84602.

'Present address: Trinity College, Deerfield, Illinois 60015.

•Bureau of Communicable Disease Control, Utah Department of Health, Salt Lake City, Utah 84103.

'Division of Parasitic Diseases, Center for Infectious Diseases, Centers for Disease Control, Atlanta, Georgia 30333.

Great Basin Naturalist

Vol. 43, No. 1

United States, nearly all residents of Sanpete
County, regardless of occupation, live in one
of 13 communities which vary in population
from 153 to 2807 (U.S. Bureau of the Census
1980a). In 1978 there were approximately
1950 dogs (Loveless et al. 1978) and 90,400
sheep in the county (U.S. Bureau of the Cen-
sus 1980b). Agriculture is the principal in-
dustry, with an emphasis on turkeys and
sheep. Range sheep herds are grazed in the
mountains during summer, on the valley floor
during spring and fall, and on the desert
(100-200 km to the west) during winter. In
addition to the range herds, there are also
semirange herds that are confined during
winter but grazed in the mountains during
summer. There are also many small flocks
confined on a year-round basis, which are
usually made up of ewes obtained from range
sheepmen (Crellin et al., 1982).

Description of Hydatid Disease
Control Program

The main goals of the program have been
reported previously (Andersen et al. 1974)
and are summarized on Table 1. The educa-
tional portion of the program was accom-
plished through press releases, an article in
the National Woolgrower (Andersen and Wal-
lentine 1976), pamphlets distributed county-
wide, talks to civic and church groups, and
displays and personal coimseling done during
community screening clinics. A filmstrip and
coloring book were developed as tools to
educate children. Copies of the filmstrip
were given to the audiovisual departments of

Table 1. Main goals of a program to control hydatid
disease in Sanpete County, Utah.

Community objectives

1. Covering or fencing animal disposal pits at waste dis-
posal sites

2. Eliminating stray or roving dogs

3. Conducting educational programs in local schools

Individual objectives

1. Fostering a widespread understanding of the para-
site's life cycle

2. Advocating proper preventive measures such as not
feeding sheep viscera to dogs, burning or burying
sheep that die on the range, and not allowing dogs to
roam

3. Encouraging individuals to have themselves and their
dogs checked for £. granulosus at yearly screening
clinics

the two school districts in Sanpete County,
and in 1980 coloring books were distributed
to all third and fourth grade students in eight
elementary schools in the county. The overall
goal of this portion of the program was to
educate the residents about the life cycle of
E. granulosus, and the proper measures that
could be taken to prevent the disease.

Local governments were advised as to
proper community-wide control measures,
and it was their choice whether or not such
measures were implemented. Included within
this phase of the program was prophylactic
treatment by a local veterinarian (Dr. M.
John Ramsay) of dogs in the northern half of
Sanpete County with bunamidine hydro-
chloride (1974-1978) and praziquantel
(1979-1981).

Surveillance of dogs, sheep, and human
beings was conducted in several areas of cen-
tral Utah. Starting in 1971, clinics to examine
dogs were held in communities in Sanpete
County, and, in 1980 and 1981, on nearby
mountain range lands (Table 2). In other
areas (Summit and Utah counties), clinics
were usually held at the sheep camp location.
Dogs were dosed with arecoline hydro-
bromide, and the resulting purge was exam-
ined on site. Sheep were checked at slaughter
for the presence of cysts by state meat in-
spectors, and the identification of cysts sus-
pected was confirmed at the parasitology
laboratory at Brigham Young University. For-
ty-nine coyotes (Andersen et al. 1973, Conder
and Loveless 1978) and 74 deer (Jensen et al.,
1982b) were also examined to assess their
possible role as sylvatic reservoirs of in-
fection. Immunodiagnostic clinics for human
beings were conducted usually in conjunction
with clinics for dogs (Klock et al. 1973, Bar-
bour et al. 1978). Clinics were advertised in
local newspapers, by posters placed in com-
mimities, and by letters and phone calls to
sheepmen by individuals from the Utah De-
partment of Health, BYU, and a local veter-
inarian's office.

Evaluation of Control Program

The effectiveness of the control measures
was evaluated by monitoring trends in in-
fection rates in dogs and sheep and the num-
bers of new cases diagnosed in human beings.

January 1983

Andersen et al.: Hydatid Disease

The success of holding screening clinics was
evaluated partially by comparing the list of
all those who brought dogs for examination
with a complete list of all sheepmen in the
region (Crellin et al., 1982). Further eval-
uation was obtained by questioning adult
residents of two Sanpete Coimty commu-
nities (Fountain Green and Spring City) con-
cerning their knowledge of hydatid disease
and attitudes towards proper preventive
measures. The procedures employed in devis-
ing, distributing, and analyzing these ques-
tionnaires were described earlier (Condie et
al., 1981). Implementation of community-
wide measures was evaluated through inter-
views with city officials and visits to the vari-
ous community waste disposal sites.

The effectiveness of the coloring books in
increasing knowledge about hydatid disease
and proper preventive measures was ana-
lyzed by testing the students before they re-
ceived the coloring book and again two
weeks afterward. Pre- and posttest scores
were compared using a paired t-tesi.

Results

Of 15,775 sheep slaughtered in five central
Utah abattoirs since 1971, 1116 (7.1 percent)
were infected with E. granulosus. The range
in yearly prevalence was 13.2 percent in
1972 to 2.8 percent in 1980 and 1981 (Fig. 1).
During the study period, 83 screening clinics

for dogs were held; 109 of 1120 (9.7 percent)
dogs purged were infected with the adult
cestode. Infection rates of the parasite in
dogs declined from 28.3 percent in 1972
(Loveless et al. 1978) to 1.0 percent in 1979,
but rose to 8.9 percent in 1980 and to 9.8
percent in 1981 (Fig. 1) (Jensen et al. 1982a).
This rise, however, was probably due to the
fact that in 1980 7 of the 8 dogs found in-
fected were owned by sheepmen who had
never attended a clinic, and that in 1981 4 of
the 5 dogs infected were owned by sheepmen
who had not attended a clinic in several
years. Sixty of 593 (10.1 percent) owners who
attended a clinic had at least one infected
dog (Fig. 2 and Table 3). Generally, the pro-
portion of owners with at least one infected
dog was higher in communities from the
northern half of the county (Table 3).
Twenty-seven of the 28 (96.4 percent) range
sheepmen presently in the county have taken
some of their dogs to at least one clinic, but
no more than 19 (67.8 percent) have attended
in any one year. Seven clinics were held in
Summit County where 6 of 69 (8.7 percent)
dogs harbored E. granulosus, and one clinic
was held in Utah County where 1 of 21 (4.8
percent) dogs was infected. No echinococ-
cosis infections were found among 49 coyotes
(Andersen et al. 1973, Conder and Loveless
1978) and 74 deer examined (Jensen et al.
1982b).

Table 2. Dogs infected with Echinococcus granulosus in Sanpete County, 1971-1981.

Use of dog

Community

North Sanpete
Fairview
Fountain Green
Mt. Pleasant
Spring City
Other communities

Area total

South Sanpete
Ephraim
Manti
Mayfield
Gunnison

Area total

County total

Tending sheep

House

pet

Dogs

purged

infected (%)

purged

infected (%)

14(20.0)

4(4.4)

153

32(21.7)

3(4.5)

5(5.4)

2(6.5)

16(19.8)

161

12(7.5)

4(7.3)

0(0.0)

451

72(16.0)

383

21(5.5)

8(14.3)

0(0.0)

4(9.3)

1(1.0)

0(0.0)

3(27.3)

0(0.0)

112

15(13.4)

174

1(0.6)

563

87(15.4)

557

22(3.9)

Table 3.

Dog owners in

Great Basin Naturalist

Sanpete County who have had dogs infected with Echinococcus grant

Vol. 43, No. 1

ilosus, 1971-1981.

Rang

e sheepmen

Semirange sheepmen

Not

sheepmen

Region

No. of
owners

No. with
infected dogs (%)

No. of
owners

No. with
infected dogs (%)

No. of
owners

No. with
infected dogs (%)

North
South

25
19

16(64.0)

7(36.8)

67
12

15(22.4)
1(8.3)

285
180

21(7.4)
1(0.6)

Total

23(52.3)

16(20.3)

465

22(4.7)

Fourteen autochthonous cases of hydatid
disease in man have been diagnosed in San-
pete County— an average annual incidence of
3.7 per 100,000 (Table 4). Twelve of these 14
cases were from the northern half of the
county. The 7 cases in Fountain Green since
1952 translate to a prevalence of 50 per
100,000. Six of 2747 individuals in Sanpete
County tested serologically for the presence
of hydatid cysts were diagnosed to be asymp-
tomatic carriers, and subsequently 4 of these
6 have had surgery (Barbour et al. 1978).

In the door-to-door surveys, responses
were obtained from 140 of 156 (89.7 percent)
and 147 of 176 (83.5 percent) households in
Fountain Green and Spring City, respective-
ly. In Fountain Green 256 individuals (1980
population of 578), and in Spring City 228
(1980 population of 675) returned completed
questionnaires. Twenty-nine percent of the

respondents from Fountain Green indicated
that they were aware of hydatid disease be-
fore 1971, but only 9.0 percent from Spring
City were aware of this disease prior to that
year. In Fountain Green, 87.3 percent of the
respondents currently understood the role of
dogs in the life cycle of E. granulosus, and
70.0 percent knew the role of sheep. In
Spring City 84.3 percent imderstood the role
of dogs and 69.0 percent knew the role of
sheep. There were 62.0 percent and 54.0 per-
cent of the adult residents of Fountain Green
and Spring City, respectively, who had
worked with sheep some time in their life. In
Fountain Green 83.0 percent of the adult
residents and in Spring City 77.5 percent
were willing to pay to have a proper animal
disposal pit built and maintained. Spring City
residents were asked the reasons why they
did or did not bring their dogs to clinics. The

35 r

DOGS

SHEEP

— •

\ /

1971 1973 1975 1977 1979 1981

YEAR

Fig. 1. Prevalence oi Echinococcus granulosus in dogs and sheep from central Utah, 1971-1981.

January

1983

Andersen et al.

: Hydatid Disease

Table 4

. Fourteen autochthonous

cases of hydatid disease

in human beings from Sanpete County,

Utah. .

Year of

City of

Connection with

Year of

Location

Name

Sex

birth

residence

sheep raising

surgery

of cyst

N.P.

1944

Fountain Green

Direct"

1952

Lung

J.B.

1916

Fountain Green

Direct

1964

Lung & Liver

P.A.

1946

Fountain Green

Direct

1967

Liver

PJ.

1931

Fountain Green

Direct

1971

Liver

L.C.

1949

Fountain Green

Direct

1971

Liver

M.C.

1908

Mt. Pleasant

Direct

1972

Lung

R.L.

1908

Mt. Pleasant

Direct

1973

Lung

D.J.

1960

Fountain Green

Indirect^

1974

Liver

R.C.

1913

Spring City

Direct

1975

Liver

D.A.

1921

Fountain Green

Direct

1975

Liver

B.S.

1967

Manti

Indirect

1976

Lung

D.D.

1955

Gunnison

Direct

None=

Liver

C.W.

1926

Spring City

Direct

None^

Liver

E.G.

1951

Fairview

Indirect

1979

Lung

"Patient or family members are sheepmen.
"Patient lives near sheepmen,
tase diagnosed in 1976.

desire to know if their dog was infected (93.9
percent) and their desire to eUminate hydatid
disease (90.3 percent) were the reasons most
commonly given for attending. Being un-
aware of the cHnic (55.9 percent) and a lack
of personal time (35.2 percent) were the rea-
sons most frequently listed for not attending.

Data on sources of information (Table 5), at-
titudes and practices of dog and sheep own-
ers (Table 6), and dog and sheep populations
(Table 7) are also presented herein.

Implementation of community-wide pre-
ventive measures and participation in the
praziquantel treatment program were gener-

• RANGE SHEEPMEN

A SEMI-RANGE SHEEPMEN

□ NOT SHEEPMEN

1971

Fig. 2. Percent of individuals in Sanpete County, Utah, having dogs infected with Echinococcus granubsus,
1971-1981.

Great Basin Naturalist

Vol. 43, No. 1

ally poor. Only one community (Spring City)
of the six that had an animal disposal pit, had
a pit cover. Two commimities (Moroni and
Manti) did not allow dead animals to be left
at their waste disposal sites. Laws making it
illegal to leave dead animals on the range
were passed recently in Sanpete and five ad-
joining counties, but, at this time, enforce-
ment is nonexistent. All communities have
leash laws, but only one (Manti) has made an
effort to eliminate stray dogs. Fairview,
Fountain Green, Moroni, and Mt. Pleasant
have enacted laws requiring treatment of
dogs with praziquantel as a stipulation for
obtaining or renewing a dog license; how-
ever, few sheepdogs in Sanpete County are
licensed and compliance is largely voluntary.
In the praziquantel treatment program, only
7 of 63 (11.1 percent) owners participating
had ever had dogs known to be infected.
Only 22 of the 63 (34.9 percent) owners had
ever brought their dogs to a screening clinic.

Discussion

Among the human population at risk in
Sanpete County, cystic hydatid disease has
been diagnosed at a rate (3.7 cases per
100,000 population per year) comparable to
those in other regions regarded as highly en-
demic for this infection. Data for comparison
include rates from the Australian state of
Tasmania prior to the initiation of a state-
wide control program (15 per 100,000 per
year; McConnell and Green 1979); from
Cyprus (12.9 per 100,000 per year; Anony-
mous 1981); from Chile (7.8 per 100,000 per

Table 5. Helpfulness of various sources of informa-
tion about hydatid disease in Fountain Green and Spring
City, Utah.

year; Anonymous 1981); and from Yugoslavia
(3.7 per 100,000 per year; Anonymous 1981).
Also, infection rates in dogs and sheep at the
beginning of the study period herein re-
viewed were similar to those in other endem-
ic regions (Anonymous 1981).

Most successful programs to control E.
granulosus have had both a formal organiza-
tional structure and a mandatory in-
volvement of the local population (Gemmell
1979). In contrast, the control effort in cen-
tral Utah has been a cooperative project in-
volving university, state, and federal
agencies, and participation of the community
has been completely voluntary. The volun-
tary aspect of the program has imposed cer-
tain limitations on the rate of progress of sug-
gested preventive and control measures.
Although the majority of dog owners and
sheep ranchers have responded positively to
recommended measures, and also availed
themselves of the diagnostic services pro-
vided, our questionnaire survey indicated
that some individuals have not. The data in-
dicate that the health educational aspects of
the program have succeeded in making most
adult residents of the two commimities thor-
oughly aware of hydatid disease, but this may
not be adequate to sustain interest and to
motivate necessary changes. We had pre-
viously observed that most dog owners in
Sanpete County were able to respond cor-
rectly to questions about the life cycle of

Table 6. Attitudes and practices of dog and/or sheep
owners in Fountain Green and Spring City, Utah.

Attitude or practice

Percentage of respondents

Fountain Green Spring City

(%) (%)

Attitude— willing to:
take dog to clinic"

83.5

78.2

Percentage of respondents who
perceived source as helpful"

pay tor propnyiacnc
treatment of do^
control do^

86.7
48.8

95.7
65.0

Source of information

Fountain Green

(%)

Spring City

(%)

use animal pit

Practice;

allow dog to have
access to area

88.0

87.8

Screening clinics
Friends

70.8
60.2

56.3
47.8

Pamphlets
Veterinarian

56.5
51.6

50.7

32.7

where sheep are
killed^

21.6

11.0

Newspapers

Filmstrip

Physician

23.5
16.0
12.9

36.7
14.4
13.5

butcher sheep at
home or on
range*"

63.3

90.6

'Individuals who marked a "4" or "5" on a 1 (no help) to 5 (very helpful)
scale.

*Dog owners: Fountain Green, 64, and Spring City, (
"Sheep owners: Fountain Green, 34, and Spring City, 27.

January 1983

Andersen et al.: Hydatid Disease

hydatid disease and how to prevent it; how-
ever, many continue to manage their dogs in
ways that permit them ready access to
sources of infection (Schantz and Andersen
1980).

Another problem associated with the vol-
untary nature of our program involves the in-
terpretation of data collected at diagnostic
field clinics for dogs. Generally, rates of in-
fection measured annually suggested a down-
ward trend, although rates in most recent
years have increased somewhat (Fig. 1). This
apparent increase most likely reflects differ-
ences in the yearly samples of the canine
population, rather than actual increases in
rates of transmission. Some effort was made
in the later years to encourage dog owners
who had not previously attended to bring
their animals to the voluntary clinics. Also,
more diagnostic clinics were held in the sum-
mer sheep range land, rather than in local
communities as was done previously. Thus,
higher rates of infection were found in dogs
not previously examined and in those sheep
dogs that were from high-risk canine popu-
lations associated with herds of sheep.

In contrast to these fluctuating changes
noted in the infection rates of dogs, a more
steady decline was noted for the slaughtered
sheep, and yet no changes in sheep marketing
practices were known to occur during the
study period.

It now seems likely to conclude that, after
10 years, E. granulosus has largely been elim-
inated from "house" dogs (those dogs not
working with sheep), but is still found in
small numbers in dogs owned by the county's
sheepmen. This partial control has probably
been achieved through educational efforts
that resulted in a reduction in availability of
sheep viscera to dogs. However, recom-
mended preventive measures such as control

Table 7. Dog and sheep populations in Fountain
Green and Spring City, Utah.

Fountain Green

Spring City

Households with dogs

64/140

66/147

Number of dogs

109

101

Dogs per dog-owning

household

1.7

1.5

Households with sheep

34/140

27/147

Number of sheep

24,317

3,655

Sheep per sheep-

owning household

714

135

of dogs and proper disposal of dead animals
have not been effectively implemented. The
number of sheep in Sanpete County has
dropped 38.5 percent since 1969 (U.S. Bu-
reau of Census 1980b), and this could be a
factor in the reduction in prevalence of E.
granulosus. A drop in total sheep numbers
was considered a factor in the elimination of
hydatid disease from Iceland (Beard 1973).

In spite of the progress noted in our con-
trol program to date, there is still a continued
potential for transmission of E. granulosus
between dogs and human beings in Sanpete
County. Basically, this is due to: (1) the lack
of a county-wide control program for dogs,
(2) the lack of adequately maintained animal
disposal pits and covers in most of the com-
munities, (3) the persistence of home-slaugh-
tering of sheep, and (4) concentration of most
county residents within communities. Our re-
sults, and those of an earlier study (Condie et
al. 1981), suggest that individuals in central
Utah are generally unwilling to control their
dogs, but would be supportive of community
efforts to build and maintain proper disposal
pits for dead sheep.

In the future we plan to continue the dis-
tribution of all educational aids available,
and also to encourage the community offi-
cials in those areas where hydatid disease has
occurred to build and maintain proper ani-
mal disposal pits and covers. In addition,
most clinics will be held closer to summer
grazing areas, and sheepmen with a poor at-
tendance record at clinics will be encouraged
to have their dogs checked on a regular basis.

Table 8. Efficacy of coloring books as educational
aids for third and fourth grade students in Sanpete
County, Utah.

Average

pretest

posttest

No.

score

School

students

(%)

Fairview
Fountain Green
Moroni
Mt. Pleasant/
Spring City
Ephraim
Gunnison
Manti

57.3

79.6°

51.7

83.3°

66.6

80.8°

107

60.7

78.4°

64.4

88.1°

66.8

87.5°

59.5

86.6°

Total

413

62.5

83.3°

'Posttest score significantly greater than pretest score at a = 0.05 using
paired t-test.

Great Basin Naturalist

Vol. 43, No. 1

Finally, since praziquantel (Droncit®) is now
an approved drug in the United States, and
since dog owners within the two commu-
nities surveyed indicated that they are will-
ing to pay for treatment, we will encourage
continued prophylactic treatment of dogs
with this drug. Finally, surveillance of hyda-
tid disease will be maintained by monitoring
infection rates in slaughtered sheep and by
registering diagnosed human cases.

Literature Cited

Andersen, F. L., P. D. Wright, and C. Mortenson.
1973. Prevalence of Echinococcus granulosus in-
fection in dogs and sheep in central Utah. J. Am.
Vet. Med. Assoc. 163:1168-1171.

Andersen, F. L., J. R. Everett, A. G. Barbour, and F.
J. Schoenfeld. 1974. Current studies on hydatid
disease in Utah. Proc. 78th Ann. Meeting U.S.
Animal Hlth. Assoc. 370-384.

Andersen, F. L., and M. V. Wallentine. 1976. Hydatid
disease. Nat. Woolgrower 66:16-18.

Anonymous. 1981. FAO/UNEP/WHO guidelines on
echinococcosis/hydatidosis surveillance, pre-
vention and control. Wld. Hlth. Org., V.P.H.
80:23, Geneva.

Barbour, A. G., J. R. Everett, F. L. Andersen, C. R.
Nichols, T. Fukushima, and I. G. Kagan. 1978.
Hydatid disease screening: Sanpete County,
Utah, 1971-1976. Am. J. Trop. Med. Hyg.
27:94-100.

Beard, T. C. 1973. The elimination of echinococcosis
from Iceland. Bull. Wld. Hlth. Org. 48:653-660.

Butler, J. M., and A. W. Grundmann. 1951. Fecal ex-
amination of 200 dogs from Salt Lake City, Utah,
for intestinal helminths. J. Am. Vet. Med. Assoc.
118:396-398.

Butler, J. M., and A. W. Grundmann. 1954. The in-
testinal helminths of the coyote Canis latrans
Say, in Utah. J. Parasitol. 40:440-443.

Carlquist, J. H., AND R. J. DowELL. 1951. Echino-
coccus disease. Report of four cases contracted in
the United States. Rocky Mtn. Med. J.
48:773-776.

CoNDER, G. A., AND R. M. LovELESs. 1978. Parasites of
the coyote {Canis latrans) in central Utah. J.
Wildl. Dis. 14:247-249.

CoNDiE, S. J., J. R. Crellin, F. L. Andersen, and P. M.
Schantz. 1981. Participation in a community
program to prevent hydatid disease. Publ. Hlth.
Lond. 95:28-35.

Crellin, J. R., F. L. Andersen, P. M. Schantz, and S.
J. CoNDiE. 1982. Possible factors influencing dis-
tribution and prevalence of Echinococcus gran-
ulosus in Utah. Am. J. Epidemiol. 116:463-474.

Fox, J. C, F. L. Andersen, and K. H. Hoopes. 1970. A
survey of the helminth parasites of cattle and
sheep in Utah valley. Great Basin Nat. 30:
131-145.

Gemmell, M. a. 1979. Hydatidosis control— a global
view. Aust. Vet. J. 55:118-125.

Grundmann, A. W., J. M. Butler, and M. J. Selin.
1953. Examination of commercial silver fox as a
possible host for Echinococcus granulosus (Batsch
1786) Rudolphi 1805 in Salt Lake County, Utah.
J. Am. Vet. Med. Assoc. 123:235-236.

Jensen, L. A., F. L. Andersen, and P. M. Schantz.
1982a. The prevalence of Echinococcus gran-
ulosus and other taeniid cestodes in sheep dogs of
central Utah. Great Basin Nat. 42:65-66.

Jensen, L. A., J. A. Short, and F. L. Andersen. 1982b.
The internal parasites of Odocoileus hemionus of
central Utah. Proc. Helminthol. Soc. Washington
49:317-319.

Kahn, J. B., S. Spruance, J. Harbottle, P. Cannon,
and M. G. Schultz. 1972. Echinococcosis in
Utah. Am. J. Trop. Med. Hyg. 21:185-188.

Klock, L. E., S. L. Spruance, F. L. Andersen, D. D.
JuRANEK, AND I. G. Kagan. 1973. Detection of
asymptomatic hydatid disease by a community
screening program. Am. J. Epidemiol. 97:16-21.

Loveless, R. M., F. L. Andersen, M. J. Ramsay, and R.
K. Hedelius. 1978. Echinococcus granulosus in
dogs and sheep in central Utah, 1971-76. Am. J.
Vet. Res. 39:499-502.

McCoNNELL, J. D., AND R. J. Green. 1979. The control
of hydatid disease in Tasmania. Aust. Vet. J.
55:140-145.

Schantz, P. M., and F. L. Andersen. 1980. Dogs, own-
ers and hydatid disease in Sanpete County. Gr.
Bas. Nat. 40:216-220.

Stinner, W. F., M. B. Toney, and S. Kan. 1978. Popu-
lation changes in rural Utah. Utah Sci.
39:107-109.

U.S. Bureau of the Census. 1980a. 7980 Census. Utah
preliminary report. U.S. Bureau of the Census,
Washington, D.C.

1980b. 1978 Census of Agriculture. Utah prelim-
inary report. U.S. Bureau of the Census, Wash-
ington, D.C.

INFLUENCE OF CRYPTOGAMIC CRUSTS ON MOISTURE RELATIONSHIPS
OF SOILS IN NAVAJO NATIONAL MONUMENT, ARIZONA

Jack D. Brotherson' and Samuel R. Rushforth'

Abstract.— Cryptogamic soil crusts of Betatakin Canyon in Navajo National Monument were investigated to un-
derstand the influence of such crusts on soil moisture relationships and potential sediment production. Crusts sam-
pled were part of the pinyon-juniper community and were studied in paired units. The presence of crusts on soils
significantly increased the depth of water penetration and decreased runoff. Soils showed reduced infiltration of wa-
ter where lichen and algal crusts were present and enhanced infiltration rates where mosses were present. Crusts ap-
pear to cause surface sealing and therefore likely reduce surface evaporation rates as well.

Cryptogamic crusts are nonvascular plant
communities that grow on or immediately
beneath the soil surface. Such communities
are components of most desert ecosystems.
They have been described in several ecosys-
tems in western North America (Anderson
and Rushforth 1976, Anderson et al. 1982a) as
well as in the deserts of the Middle East
(Evenari et al. 1971). Until recently scant at-
tention had been given them and little was
known concerning their role in native ecosys-
tems. Studies of the past decade indicate that
they exert a significant impact on reducing
soil erosion (Evenari et al. 1971, Loope and
Gifford 1972, Kleiner and Harper 1972, Klei-
ner and Harper 1977, Anderson et al. 1982a,
Anderson et al. 1982b). Fletcher and Martin
(1948) found that fungal and algal crusts in-
crease the tensile strength of soil. The algae
appear to be the most effective in binding
the surface soil particles (Durrell and Shields
1961) because of the thick gelatinous sheaths
that enclose the trichomes of several algal
species (Anderson and Rushforth 1976). Such
gelatinous sheaths add strength and aggregat-
ing qualities to the I or 2 mm of surface soil
upon which they grow (Anantani and Ma-
rathe 1974).

Research on the biology of cryptogamic
crusts has also been done in several other
areas. These studies include taxonomy (Ali
and Sandhu 1972, Anderson and Rushforth
1976); nitrogen fixation (MacGregor and
Johnson 1971, Reddy and Gibbons 1975);
land reclamation (Singh 1950); soil fertility

(Shields and Durrell 1964); reproduction,
growth and habitat relations (Evenari et al.
1971, Anderson et al. 1982b); and moisture
(Booth 1941, Loope and Gifford 1972).

The objective of this study was to in-
vestigate the influence of cryptogamic crusts
in the pinyon-juniper woodlands of north-
eastern Arizona on depth of water pene-
tration, infiltration, runoff, and potential
sediment production.

Study Area

Navajo National Monument is located in
northeastern Arizona (Fig. 1) and is the site
of three large Anasazi Indian cliff dwellings.
Betatakin Canyon, the site of the present
study, is a side canyon of the larger Tsegi
Canyon complex and has been described by
Hack (1945). The major geological formation
comprising the canyon is Navajo Sandstone,
which forms sheer towering cliffs 200 m or
more in height. The canyon floor consists of
deep alluvial deposits of sandy Quaternary
fill. Kayenta sandstone outcrops in the lower
reaches of the canyon.

The annual temperatures recorded at the
park headquarters weather station at Betata-
kin canyon ranges from -23 to 38 C with a
mean of 10 C. The number of frost-free days
in the area varies from 107 to 213, with an
average of 155 days. Total annual precipi-
tation ranges from 17 to 48 cm with a yearly
mean of 29 cm. There is a single wet season
lasting from late summer through fall.

'Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602.

Great Basin Naturalist

Vol. 43, No. 1

G A R F I ELD

SAN JUAN

KANE

■\\^i «

-.^^F

, UTAH

POWELL ^^ i;ARIZONA

I \

KEET SIEl tUIN

COCONINO

HOUSE I /BiTATAKIN RUIN^,.-— ^^,3 y

/, Shontj,'

N A V A J 0

APACHE

NAVAJO

INDIAN

RESE RVATI

LEGEND

= STATE LINE

I COUNTY LINE

I ' MAIN HIGHWAY

O N DIRT aOAD

I /-'^ RIVEt

I O TOWN

I ■ *UINS SITE

o
(J

UJ
LU

Fig. 1. Reference map of study area.

Methods

Cryptogamic crusts were sampled in the
pinyon-juniper {Pinus edulis-Juniperus os-
teosperma) community that borders Betatakin
Canyon in Navajo National Monument, Ari-
zona. Cryptogam crusts were studied in
paired units so that varying conditions in
habitat (slope, exposure, soil texture, etc.)
could be kept to a minimum. Pairs consisted
of five sites where crusts were intact and un-
disturbed and five adjacent sites where the
crusts had been heavily disturbed or de-
stroyed. A total of 10 sites were considered

for each measurement. Pairs were always lo-
cated within 2 m of each other.

Water infiltration rates were measured by
using a thin-walled aluminum cylinder 12 cm
tall and 65 mm in diameter. The cylinder was
gently turned into the crust or soil to a depth
of 2 cm and then 50 ml of water was ponded
above the core inside of the cylinder. Infil-
tration into the core was measured as the
number of seconds needed for the ponded
water to disappear into the core.

Depth of water penetration and runoff
were assessed by raining 1.5 liters of water

January 1983

Brotherson, Rushforth: Cryptogamic Crusts

onto the crust or adjacent soil surface
through a perforated 80 mm diameter disk.
The perforations were evenly spaced on a 0.5
cm grid. The disk was placed at a distance of
1.2 m above the ground surface. Total deliv-
ery time for the water to be dispensed onto
the crust or soil surface was 60 seconds.
These rates were designed to approximate or
exceed precipitation at cloudburst propor-
tions (i.e., 10 cm/hr). High intensities of pre-
cipitation, such as those exceeding infiltration
capacities of the soil, are significant because
of their effects on runoff and erosion. Once
the water had disappeared into the crust or
soil surface, depth of penetration was mea-
sured immediately. Five depth measurements
were taken for each watering at each of the
10 areas and then averaged to give a single
value for each site.

Runoff was measured by recording the
across slope and downslope spread of water
rained onto study sites. The area of spread
was computed from these measurements us-
ing the formula for the area of an ellipse.

Soil movement was assessed by estimating
the amount of soil moved during a measured
rain. The following index was used: 1 = no
appreciable movement; 2 = moderate move-
ment—up to 10 percent of soil being dis-
placed; and 3 = heavy movement— between
10 and 20 percent of soil being displaced.

All runoff and soil movement measure-
ments were taken during the third week of
August 1980. Sampling intensity was deter-
mined following the estimation procedures
described by Avery 1975. Significant differ-
ences in the paired measurements were as-
sessed through the use of Students-t statistic.

Results and Discussion

The influence of cryptogamic crusts on six
soil moisture characteristics was assessed. Av-
erage values for all measurements taken dur-
ing this study are given in Table I. All but
one of the measured characteristics showed
significant differences between crusted and
uncrusted soils.

Infiltration measurements on the paired
study sites indicated that well-developed
cryptogamic crusts (Fig. 2) significantly in-
creased the depth of water penetration. This
was also found by Loope and Gifford (1972).
Downslope movement of water was signifi-
cantly greater on the sites that exhibited no
crust development. Likewise, the differences
in total area of surface spread was signifi-
cantly greater on uncrusted soils. These dif-
ferences are probably best explained by the
micro-topographic changes that develop at
the soil surface under the influence of crypto-
gamic crust growth. Well-developed crusts
form pedestals so that the ground surface
looks something like a convoluted brain coral
(Fig. 2-4). Hills and valleys a few centimeters
in relief develop across broad crusted areas.
The small valleys run in all directions and
cause pooling of the water as it hits the soil
surface (Fig. 5). This pooling holds the water
in place for extended periods, thus increasing
the time for infiltration to occur and simulta-
neously decreasing runoff and movement
across the soil surface. With reduced surface
movement, deeper penetration of water oc-
curs. The net effect is to slow the movement
of surface-flowing water, providing longer
periods for infiltration, less opportunity for

Table 1. Relationships of cryptogamic crusts growing on the soil in Navajo National Monument to measured
moisture parameters. Figures represent means and standard deviations (sd).

Characteristic

Crust

Noncrust

Significance

measured

Mean

level

Water penetration depth (cm)

5.46

1.35

3.23

0.69

.05

Downslope spread (cm)

67.62

13.74

95.50

4.24

.001

Across slope spread (cm)

47.24

10.87

45.72

6.48

Area of spread (sq cm)

10434.11

3041.18

13738.50

2185.26

.001

Soil movement"

1.00

0.00

2.60

0.89

.01

Infiltration (seconds)

Moss cover

15.40

3.90

238.00

87.90

.001

Lichen and algae cover

48.00

14.50

31.00

8.10

.001

'Soil movement was assessed as follows: 1 = no movement, 2
between 10-20 percent of soil being displaced.

moderate movement— up to 10 percent of soil being displaced, 3 = heavy movement—

Great Basin Naturalist

Vol. 43, No. 1

Figs. 2-5. Cryptogamic crusts. Left top, moving clockwise: 2. Crusts beneath Utah juniper trees. 3. Well-devel-
oped cryptogamic soil crusts. 4. Close-up of cryptogamic crusts showing typical pinnacle development. 5. Close-up
of cryptogamic crust after experimental rain showing water ponding.

concentration in rills, and decreased power
to cause erosion. In other words, cryptogam
crusting fosters more infiltration and less run-
off of surface water.

Well-developed crust areas also showed
significantly less soil movement (Table 1).
These data support the findings of several
other studies (Fletcher and Martin 1948,
Loope and Gifford 1972, Kleiner and Harper
1977, Anderson et al. 1982b). Cryptogamic
crusts appear to have a protective influence
on the soil in four major ways. First, they
bind the soil surface particles with the inter-
twining growth of algal and fungal filaments
(Durrell and Shields 1961). Second, the moss

and lichen constituents of cryptogam crusts
aid in stabilizing the soil by covering the sur-
face with thalli and penetrating the soil sur-
face with rhizoids (Anderson et al. 1982b).
Third, the irregularities of a well-developed
cryptogamic crust surface tend to break up
microwind patterns and thus reduce wind-
born soil movement (Brady 1974). And
fourth, with less water movement there is
also significantly less soil movement.

Well-developed crusts also influenced wa-
ter movement into the soil. Where moss cov-
er was high, infiltration rates were greatly
enhanced over areas where moss cover had
been removed. The enhancement of infil-

January 1983

Brotherson, Rushforth: Cryptogamic Crusts

tration appeared to be due to the moss thalli
acting as a sponge. On the other hand, where
they had been removed, a .05 to 1 cm thick
layer of silt beneath them acted to retard in-
filtration. Infiltration rates were significantly
reduced or impeded by lichen and algal crust
cover. The highest infiltration rates (most
rapid penetration by water) occurred on soils
with no cryptogamic cover (Table 1), In gen-
eral, where cryptogamic cover was high, in-
creased resistance to infiltration occurred.
Loope and Gifford (1972) noted this pattern
and also found that, when crusts were wetted
previous to infiltration trials, infiltration rates
on crusted soils were retarded by a factor of
two. Fritsch (1922) first suggested that the
highly mucilaginous sheaths of blue-green al-
gae, which are the major components of
cryptogamic crusts in arid environments,
might form a layer at the soil surface that
would both impede water infiltration into the
soil and impede evaporation of soil moisture
caught beneath the algal layer. This would
provide more water to the plants growing in
such areas. Booth (1941) later tested this hy-
pothesis and showed that more moisture was
to be found in the upper layers of soil (i.e.,
the upper 2.5 cm) where cryptogamic crusts
were prominent than in adjacent soils with
no crusts (i.e., 8.9 percent vs. 1.3 percent,
respectively).

Data from several studies indicate that
high cryptogamic crust cover is associated
with high silt in the soil surface (Evenari et
al. 1971, Loope and Gifford 1972, Kleiner
and Harper 1977, Anderson et al. 1982b).
Textural observations on our sites showed
similar patterns. Kleiner and Harper (1977)
also argue that once established the crusts
tend to trap silt at the soil surface. Evenari et
al. (1971) and Blackburn and Skou (1974)
present data that indicate that soils high in
silt often have low permeability rates and
high runoff. They suggested that soils with
high levels of silt in the upper layers often
show high initial infiltration rates, but, as
more wetting occurs, the percolation rates
decrease rapidly and eventually an almost
impenetrable layer can be formed. Beneath
such a sealed surface, air caught in the voids
of the lower layers may have a difficult time
escaping and may therefore further retard in-
filtration (Evenari et al. 1971).

It appears then that at least three factors
tend to reduce water infiltration rates in soils
with cryptogam crusts: (1) the effect of high
levels of silt in the soil and its resultant swell-
ing and sealing action when mixed with wa-
ter (Evenari et al. 1971); (2) the wetting ac-
tion of the water on the gelatinous sheaths of
the algal filaments, causing the filament to
swell and tightly bind the surface soil par-
ticles (Anantani and Marathe 1974, Durrell
and Shields 1961, Fritsch 1922); and (3) air
trapped beneath the sealed surface to further
impede water penetration.

Evenari et al. (1971) also indicated from
their research on micro-watershed irrigation
projects that, as the farm areas receive runoff
water ladened with silt from the watersheds
and as the silt is deposited on the soil surface,
evaporation from the irrigated fields was re-
duced to as little as 7.4 mm over a seven
month period. This kind of reduction in
evaporation in a desert with annual evapo-
ration values from 1700 to 2700 mm would
be highly important relative to moisture re-
tention in the subsurface layers of the soil.

Since cryptogamic crusts tend to seal the
soil surface and since crusts also increase the
depth of water penetration, the effects they
have on reducing moisture stress in desert
ecosystems could prove to be extremely valu-
able. Furthermore, since crust communities
tend to grow in association with high silt lev-
els at the soil surface, these elevated silt lev-
els undoubtedly further reduce water losses
by evaporation. This being the case, crypto-
gamic crusts may be as important in their
role in water conservation in desert systems
as they are in preventing soil erosion.

Literature Cited

Ali, S., and G. R. Sandhu. 1972. Blue-green algae of the
saline soils of the Punjab. Oikos 22:268-272.

Anantani, Y. S., and D. V. Marathe. 1974. Soil aggre-
gating effects of some algae occurring in the soils
of Kutch and Rajasthan. Univ. of Bombay
41:94-100.

Anderson, D. C, K. T. Harper, and S. R. Rushforth.
1982a. Recovery of cryptogamic crusts from graz-
ing on Utah winter ranges. J. Range Manage.
35:355-359.

Anderson, D. C, K. T. Harper, and R. C. Holmgren.
1982b. Factors influencing the development of
cryptogamic soil crusts in Utah deserts. J. Range
Manage. 35:180-185.

Great Basin Naturalist

Vol. 43, No. 1

Anderson, D. C, and S. R. Rushforth. 1976. The
cryptogamic flora of desert soil crusts in southern
Utah. Nova Hedwigia 29:691-729.

Avery, T. E. 1975. Natural resources measurements. 2d
Ed. McGraw-Hill Book Company, New York. 339
pp.

Blackburn, W. H., and C. M. Skou. 1974. Infiltration
rates and sediment production of selected plant
communities in Nevada. J. Range Manage.
27:476-479.

Booth, W. E. 1941. Algae as pioneers in plant succes-
sion and their importance in erosion control.
Ecology 22:38-46.

Brady, N. C. 1974. The nature and properties of soils.
8th ed. Macmillan Publishing Co., Inc., New
York. 639 pp.

DuRRELL, L. W., AND L. M. SHIELDS. 1961. Character-
istics of soil algae relating to crust formation.
Trans. Amer. Micro. Soc. 80:73-79.

Evenari, M., L. Shanan, and N. Tadmor. 1971. The
Negev— the challenge of a desert. Harvard Univ.
Press, Cambridge, Massachusetts. 345 pp.

Fletcher, J. E., and W. P. Martin. 1948. Some effects
of algae and molds in the rain-crust of desert
soils. Ecology 29:95-100.

Fritsch, F. E. 1922. The terrestrial algae. J. Ecology
10:220-236.

Hack, J. T. 1945. Recent geology of the Tsegi Canyon.
Appendix I in R. L. Reals, G. W. Brainerd, and
W. Smith, eds. Archaeological studies in North-
east Arizona. Univ. of California Press, Berkeley
and Los Angeles.

Kleiner, E. F., and K. T. Harper. 1972. Environment
and community organization in grasslands of
Canyonlands National Park. Ecology 53:229-309.

1977. Soil properties in relation to cryptogamic

ground cover in Canyonlands National Park. J.
Range Manage. 30:202-205.

LooPE, W. L., AND G. F. GiFFORD. 1972. Influence of a
soil microfloral crust on select properties of soils
under pinyon-juniper in southeastern Utah. J.
Soil and Water Conser. 27:164-167.

MacGregor, A. N., AND D. E. Johnson. 1971. Capacity
of desert algal crusts to fix atmospheric nitrogen.
Soil Sci. Soc. of Amer. Proc. 35:843-844.

Reddy, G. B., and J. Gibbons. 1975. Nitrogen fixation by
algae on fescuegrass soil crusts. Soil Sci. of Sco.
Amer. Proc. 39:654-656.

Shields, L. M., and L. W. Durrell. 1964. Algae in rela-
tion to soil fertility. Bot. Rev. 30:92-128.

Singh, R. N. 1950. Reclamation of "Usar" lands in India
through bluegreen algae. Nature 165:325-326.

A VASCULAR FLORA OF THE SAN RAFAEL SWELL, UTAH'

James G. Harris^

Abstract.— The vegetation of the San Rafael Swell in southeastern Utah is examined based on personal field col-
lections and previously collected herbarium specimens in the Brigham Young University Herbarium (BRY). An anno-
tated checklist includes information on frequency of occurrence and habitat preference for each entity. Treated are
491 vascular plant taxa from 59 families.

The San Rafael Swell is the eroded rem-
nant of a massive domal anticline, oval in
shape, stretching along a northeasterly axis
from Capitol Reef National Park in northern
Wayne County to the foot of the Tavaputs
Plateau in Carbon County. Folding of the
earth's crust, which formed the domal anti-
cline, and the subsequent erosion of the cen-
tral dome left a huge area of concentric stone
rings, each progressively younger from the
innermost to the outermost. One of these
rings, the Ferron Sandstone Member of the
Mancos Shale Formation, encloses approx-
imately L4 million acres of land and forms
the boundary of this study. In some cases,
where the Ferron Sandstone has been com-
pletely eroded from the structural confines of
the swell, artificial boundaries have been fol-
lowed (Fig. 1).

Throughout the field seasons of 1979 and
1980, I attempted to collect representatives
of all vascular plant species growing in the
San Rafael Swell. A voucher specimen from
each collection has been placed in the Brig-
ham Young University Herbarium (BRY). In-
cluded in the checklist are species not found
by me, but collected by others and vouched
for by specimens in the Brigham Young Uni-
versity Herbarium. The checklist is not defin-
itive; additional taxa will undoubtedly be dis-
covered in the swell.

The following sources were used for identi-
fication and nomenclature: lower vascular
plants and gymnosperms, Cronquist et al.
(1972); dicotyledons, Neese and Welsh

(1981), Welsh (1978, 1980a, 1980b), Welsh
and Atwood (1981), Welsh and Moore (1973),
Welsh and Reveal (1977), Welsh et al. (1981);
monocotyledons, Cronquist et al. (1977).

The checklist includes 478 vascular plant
taxa found in the study area and an addition-
al 13 taxa known to occur on lands immedi-
ately adjacent to the study area and to be ex-
pected in the San Rafael Swell. These 13 taxa
are listed below:

Allionia incarnata L.

Cordylanthus wrightii Gray

Descurainia pinnata (Walter) Britt. var. filipes (Gray)
Peck

Eriogonum smithii Reveal

Euphorbia micromera Boiss.

Haplopappus drummondii (T. & G.) Blake

Helianthus petiolaris Nutt.

Hymenopappus filifolius Hook. var. lugens (Greene)
Jeps.

Lupinus pusillus Pursh var. rubens (Rydb.) Welsh

Phacelia demissa Gray var. heterotricha Howell

Physocarpus alternans (Jones) Howell

Sphaeralcea munroana (Dougl.) Spach

Sporobolus giganteus Nash

The flora of the San Rafael Swell is not
particularly rich in numbers of species. This
is due, at least in part, to the relatively nar-
row elevational range in the area. Although
not rich in numbers of species, the flora is
rather rich in numbers of unique and geo-
graphically restricted species. The following
eight species are strictly endemic to the San
Rafael Swell:

Astragalus rafaelensis Jones

Cryptantha johnstonii Higgins

Cryptantha pnesiana (Payson) Payson

Erigeron maguirei Cronquist

'Part of a master's thesis, originally submitted to the Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602.
'Department of Botany and Range Science, Brigham Young University, Prove, Utah 84602. Present address: Department of Botany, University of Alberta,
Edmonton, Alberta T6G 2E9.

Great Basin Naturalist

Vol. 43, No. 1

Emery
County

Fig. 1. Map of the San Rafael Swell study area.

Pediocactus despainii Welsh & Goodrich
Sclerocactus wrightii Benson
Sphaeralcea psoraloides Welsh
Thelypodiopsis harnebyi Welsh & Atwood
An additional seven species are endemic to
the San Rafael Swell and the adjacent lands
in close proximity to the swell. They are list-
ed as follows:

Astragalus pardalinus (Rydb.) Barneby
Astragalus woodruffii Jones
Hoffmanseggia repens (Eastw.) Cockerell
Hymenoxys depressa (T. & G.) Welsh & Reveal
Lomatium junceum Barneby & Holmgren
Psorothamnus polyadenius (Torr.) Rydb. var. jonesii
Barneby

Townsendia apnea Welsh & Reveal

Vegetative Zones

The vegetation of the San Rafael Swell can
be divided into five major zones: the Sand
Desert Shrub Zone, the Salt Desert Shrub
Zone, the Mixed Desert Shrub Zone, the Pin-
yon-Juniper Zone, and the Ponderosa Pine-
Mountain Brush Zone. These zones are con-
trolled principally by elevation, precipi-
tation, and local edaphic factors. Although in
some areas the zones are clear and well de-
fined, in most areas they interfuse consid-
erably and it becomes difficult to draw a
sharp boundary line between them.

January 1983

Harris: San Rafael Swell Flora

Sand Desert Shrub Zone.— The San Ra-
fael Desert, in the southeastern region of the
San Rafael Swell, ranges from an elevation of
about 4200 ft (1280 m) at the lowest point to
about 5000 ft (1524 m) at the highest point.
It is a sandy desert with moving dunes in
some areas. Common shrubs in this zone in-
clude: Artemisia filifolia. Ephedra torreyana.
Yucca hatrimaniae, Poliomintha incana, Ti-
quilia latior, and Coleogyne ramosissima.
Abronia elliptica, Oenothera pallida, Hoff-
manseggia repens, and Oryzopsis hymenoides
are characteristic herbaceous plants of this
zone.

Salt Desert Shrub Zone.— In the San
Rafael Swell the Salt Desert Shrub Zone is
widespread, occurring mainly on alkaline
clay soils between 4500 and 5800 ft
(1372-1766 m) in elevation. Communities in
this zone are characteristically dominated by
Atriplex confertifolia interspersed with any of
the following subdominant species: Hilaria
jamesii, Oryzopsis hymenoides, Enceliopsis
nudicaulis, Aristida purpurea, Eriogonum in-
flatum, Salsola iherica or Ceratoides lanata.

Highly alkaline flood plains in this zone
are often covered by nearly pure stands of
Sarcobatus vermiculatus, which may at times
be accompanied by thick clumps of Tamarix
ramosissima.

Heavy clay soils of the Mancos Shale For-
mation are often dominated by Atriplex cu-
neata and Atriplex corrugata. During wet
springs these areas may be blanketed by an-
nual species such as Cleome lutea, Lupinus
pusillus, Malcolmia africana, Camissonia sca-
poidea, and Camissonia eastwoodiae.

Mixed Desert Shrub Zone.— Deep, well-
drained soils between 4500 and 5800 ft
(1372-1766 m) in elevation are often domi-
nated by one or several of the following
shrubby species: Artemisia tridentata, Arte-
misia bigelovii, Artemisia spinescens, Xantho-
cephalum sarothrae or Chrysothamnus vis-
cidiflorus. These may be intermingled with
an understory of herbaceous plants including:
Hilaria jamesii, Stipa comata, Boutehua gra-
cilis. Astragalus spp., Sphaeralcea spp., and
Leucelene ericoides.

Pinyon-Juniper Zone.— Pinus edulis and
Juniperus osteosperma are conspicuously
dominant at elevations between 5800 and

7800 ft (1766-2377 m). Within this altitudin-
al range Juniperus osteosperma is dominant at
the lower elevations and Pinus edulis is domi-
nant at the higher elevations. Intermediate
elevations yield a heterogeneous mixture of
both species.

Dense stands have almost no understory,
but more open or scattered stands will in-
clude almost all the species found in the
Mixed Desert Shrub Zone.

PoNDEROSA Pine-Mountain Brush
Zone.— The highest point in the San Rafael
Swell is the San Rafael Knob at 7921 ft (2414
m) in elevation. The area surrounding the
knob and the summit of Cedar Mountain rep-
resent the highest vegetative zone in the
swell. Pinus edulis continues to remain domi-
nant even at the highest elevations, but above
7500 ft (2286 m) plants characteristic of the
Ponderosa Pine-Mountain Brush Zone be-
come increasingly common. Some common
species in this zone are: Pinus ponderosa, Ar-
temisia nova, Cowania mexicana, Mahonia
fremontii, Populus tremuloides, Amelanchier
utahensis, Cercocarpus montanus, Ribes lep-
tanthum, Juniperus scopulorum, Hymenoxys
richardsonii, Symphoricarpos longiflorus and
Philadelphus microphyllus.

In addition to these five major vegetative
zones, smaller communities dependent on
unique local habitats are scattered through-
out the area. These include plants distributed
along seeps and streams, along washes and
roadsides where runoff collects, in hanging
gardens, on rocky outcrops, and on disturbed
sites.

Annotated Checklist

Following each species entry in the check-
list (with the exception of the 13 taxa from
lands adjacent to the study area) a designa-
tion is given concerning the frequency of oc-
currence and habitat preference of the taxon.
Plants of broad distribution that are not lim-
ited to a specific zone or community type are
listed as widespread. The following abbrevia-
tions are used:

C— common

O— occasional

U— uncommon

R— rare

SnD— Sand Desert Shrub Zone

SID— Salt Desert Shrub Zone

MD— Mixed Desert Shrub Zone

PJ— Pinyon-Juniper Zone

Great Basin Naturalist

Vol. 43, No. 1

PM— Ponderosa Pine-Mountain Brush Zone

SS— Seeps and streams

WR— Washes and roadsides

HG— Hanging gardens

RO— Rocky outcrops

DS— Disturbed sites

W— Widespread

ACERACEAE

Acer glabrum Torr. (R;PM)
Anacardiaceae

Rhus trilobata Nutt. var. simplicifolia (Greene) Barkl.
(C;WR)

Rhus trilobata Nutt. var. trilobata (C;WR)

Toxicodendron rydbergii (Small) Greene (C;SS,HG)

Apiaceae

Cymopteris acaulis (Pursh) Raf. (0;SnD,SlD)

Cyrnopteris bulbosus Nels. (C;W)

Cymopteris fendleri Gray (C;W)

Cymopteris purpurascens (Gray) Jones (0;S1D)

Cymopteris purpureus Wats. (0;W)

Lomatium junceum Barneby & Holmgren
(R;S1D,MD,PJ)

Stum suave Walt. (R;WR)
Apocynaceae

Amsonia eastwoodiana Rydb. (C;SnD)

Ainsonia jonesii Woodson (0;MD,PJ)

Apocynum cannabintim L. (C;SS,HG)

Cycladenia humilis Benth. var. jonesii (Eastw.) Welsh
& Atwood (R;MD)

ASCLEPIADACEAE

Asclepias cryptoceras Wats. (C;W)
Asclepias labriformis Jones (C;WR,SD,MD)
Asclepias macrosperma Eastw. (C;SnD,MD)
Asclepias ruthiae Maguire & Woodson (0;W)
Asclepias speciosa Torr. (C;WR)

ASTERACEAE

Ambrosia acanthicarpa Hook. (0;S1D,MD,PJ)

Ambrosia artemisiifolia L. (C;S1D,MD,PJ)

Artemisia biennis Willd. (U;WR)

Artemisia bigelovii Gray (C;MD,PJ)

Artemisia campestris L. (U;SnD,MD)

Artemisia dracunculus L. (C;WR)

Artemisia filifolia Torr. (C;SnD)

Artemisia frigida Willd. (R; WR)

Artemisia ludoviciana Nutt. (0;WR)

Artemisia nova Nels. (C;PJ,PM)

Artemisia pygmaea Gray (0;S1D,MD,PJ)

Artemisia spinescens Eaton (0;S1D, MD)

Artemisia tridentata Nutt. (C;MD,PJ,WR)

Aster glaucodes Blake (U;PM)

Baccharis emoryi Gray (U;WR)

Brickellia longifolia Wats. (U;WR)

Brickellia microphylla (Nutt.) Gray (U;WR)

Brickellia oblongifolia var. linifolia (Eaton) Robins

(0;PJ,WR)

Brickellia scabra (Gray) Nels. (R;HG,WR)
Chaenactis stevioides Hook. & Arn. (C;S1D,MD)
Chamaechaenactis scaposa (Eastw.) Rydb. (0;PJ)
Chrysothamnus greenei (Gray) Greene (0;S1D)
Chrysothamnus linifolius Greene (0;WR)
Chrysothamnus nauseosus (Pall.) Britt. var. consimilis

(Greene) Hall (C;WR)

Chrysothamnus nauseosus (Pall.) Britt. var. gnapha-

loides (Greene) Hall (U;WR)

Chrysothamnus nauseosus (Pall.) Britt. var. junceus

(Greene) Hall (U;WR)

Chrysothamnus nauseosus (Pall.) Britt. var. leio-

spermus (Gray) Hall (R;WR)

Chrysothamnus pulchellus (Gray) Greene (U;WR)
Chrysothamnus viscidiflorus (Hook.) Nutt. var. steno-

phyllus (Gray) Hall (U;WR)

Chrysothamnus viscidiflorus (Hook.) Nutt. var. vis-
cidiflorus (C;WR)

Cirsium pulchellum (Greene) Woot. & Standi. (U;WR)

Cirsium undulatum (Nutt.) Spreng. (C;WR)

Dicoria canescens Gray (U;SnD)

Encelia frutescens Gray (U;WR)

Enceliopsis nudicaulis (Gray) Nels. (C;S1D, MD)

Enceliopsis nutans (Eastw.) Nels. (C;S1D,MD)

Erigeron argentatus Greene (0;S1D,MD,PJ)

Erigeron compactus Blake var. consimilis Cronquist

(0;MD,PJ)

Erigeron divergens T. & G. (U;PJ)

Erigeron eatonii Gray (U;PJ,PM)

Erigeron maguirei Cronquist (R;WR)

Erigeron pulcherrimus Heller var. pulcherrimus

(C;S1D,MD,PJ)

Erigeron pumilus Nutt. ssp. concinnoides Cronq. var.

condensatus (Eaton) Cronq. (C;S1D,MD,PJ)
Erigeron utahensis Gray (C;WR,RO)
Gaillardia pinnatifida Torr. (C;S1D,MD)
Gaillardia spathulata Gray (C;S1D,MD,PJ)
Grindelia squarrosa (Pursh) Dunal. (0;WR)
Haplopappus armerioides (Nutt.) Gray (C;MD,PJ,RO)
Haplopappus drummondii (T. & G.) Blake
Helianthella microcephala (Gray) Gray (C;PJ,PM)
Helianthus annutts L. (C;WR)
Helianthus anomalus Blake (C;SnD)
Helianthus petiolaris Nutt.

Heterotheca villosa (Pursh) Shinners (C;S1D,MD,PJ)
Hymenopappus filifolius Hook. var. lugens (Greene)

Jeps.

Hymenopappus filifolius Hook. var. megacephalus

Turner (C;S1D,MD)

Hymenopappus filifolius Hook. var. pauciflorus

(Johnst.) Turner (0;S1D)

Hymenoxys acaulis (Pursh) Parker var. arizonica

(Greene) Parker (C;MD,PJ)

Hymenoxys depressa (T. & G.) Welsh & Reveal

(0;R0)

Hymenoxys richardsonii (Hook.) Cockerell (C;PJ,PM)
Iva axillaris Pursh (0;SS,WR)
Iva xanthifolia Nutt. (0;WR)
Leucelene ericoides (Torr.) Greene (C;W)
Lygodesmia arizonica Tomb (C;S1D,MD,WR)
Lygodesmia grandiflora (Nutt.) T. & G. (C;S1D)
Lygodesmia juncea (Pursh) D. Don (R;SnD)
Machaeranthera canescens (Pursh) Gray (R;SnD)
Machaeranthera grindelioides (Nutt.) Shinners (C;W)
Machaeranthera linearis Greene (U;WR)
Machaeranthera tanacetifolia (H.B.K.) Nees (C;WR)
Malacothrix sonchoides (Nutt.) T. & G. (C;S1D,MD)
Oxytena acerosa Nutt. (C;SS,WR)
Parthenium ligulatum (Jones) Barneby (U;S1D,MD,PJ)
Petradoria pumila (Nutt.) Greene (C;PJ,WR,RO)
Platyschkuhria integrifolia (Gray) Rydb. var. oblongi-
folia (Gray) Ellison (C;WR)

Prenanthella exigua (Gray) Rydb. (C;WR,RO)
Senecio multicapitatus Greenm. (U;PJ,PM,WR)

January 1983

Harris: San Rafael Swell Flora

Senecio multilobatus T. & G. (0;PJ,WR)

Stephanomeria exigua Nutt. (0;WR)

Stephanomeria runcinata Nutt. (R;WR)

Stephanomeria spinosa (Nutt.) Tomb (U;WR)

Stephanomeria tenuifolia (Torr.) Hall (C;WR,RO)

Taraxacum officinale Weber (C;SS)

Tetradymia canescens DC. (0;PJ,PM)

Tetradymia glabrata Gray (C;MD,PJ)

Tetradymia spinosa Hook. & Am. (0;WR)

Thelesperma subnudum Gray (C;W)

Townsendia annua Beaman (U;MD)

Townsendia aprica Welsh & Reveal (U;S1D,MD,PJ)

Townsendia incana Nutt. (C;S1D,MD,PJ)

Tragopogon dubius Scop. (C;W)

Vanclevea stylosa (Eastw.) Greene (C;SnD)

Wyethia scabra Hook. (C;WR)

Xanthium strumarium L. (0;SS)

Xanthocephalum microcephalum (DC.) Shinners (R;PJ)

Xanthocephalum sarothrae (Pursh) Shinners

(C;S1D,MD,PJ)

Xylorhiza venusta (Jones) Heller (C;RO)

Berberidaceae

Mahonia fremontii (Torr.) Fedde (C;MD,PJ)

Betulaceae
Betula occidentalis Hook. (R;SS)

Boraginaceae

Cryptantha cinerea Greene (R;PJ,PM)
Cryptantha confertifolia (Greene) Payson (R;SnD)
Cryptantha crassisepala (T. & G.) Greene var. ela-

chantha Johnst. (C;W)

Cryptantha flava (Nels.) Payson (C;W)
Cryptantha flavoculata (Nels.) Payson (C;PJ)
Cryptantha fulvocanescens (Wats.) Payson var. ech-

inoides (Jones) Higgins (0;S1D)

Cryptantha fulvocanescens (Wats.) Payson var. ful-
vocanescens (C;MD,PJ)

Cryptantha gracilis Osterh. (U;PJ)

Cryptantha humilis (Gray) Payson var. nana (Eastw.)

Higgins (0;PJ)

Cryptantha jamesii (Torr.) Payson var. disticha

(Eastw.) Payson (U;MD,WR)

Cryptantha jamesii (Torr.) Payson var. setosa (Jones)

Johnst. (C;PJ)

Cryptantha johnstonii Higgins (U;S1D)
Cryptantha jonesiana (Payson) Payson (U;S1D)
Cryptantha mensana (Jones) Payson (0;S1D)
Cryptantha paradoxa (Nels.) Payson (0;PJ)
Cryptantha recurvata Coville (0;MD,PJ,WR)
Cryptantha rollinsii Johnst. (0;MD,PJ,WR)
Cryptantha tenuis (Eastw.) Payson (0;WR)
Cryptantha wetherillii (Eastw.) Payson (0;S1D)
Heliotropium convulvulaceum (Nutt.) Gray (U;SnD)
Lappula occidentalis (Wats.) Greene var. cupulata

(Gray) Higgins (R;MD,PJ)

Lappula occidentalis (Wats.) Greene var. occidentalis

(C;W)

Lithospermum incisum Lehm. (C;PJ)

Tiquilia latior (Johnst.) Richardson (0;S1D,MD)

Brassicaceae

Arabis demissa Greene var. lanugida Rollins

(R;PJ,PM)
Arabis perennans Wats. (0;PJ)
Arabis pulchra Jones var. pallens Jones (0;PJ)
Arabis selbyi Rydb. (C;PJ)
Caulanthus crassicaulis (Torr.) Wats. (C;PJ)

Chorispora tenella (Pall.) DC. (U;DS)

Descurainia pinnata (Walt.) Britt. var. filipes (Gray)

Peck

Descurainia pinnata (Walt.) Britt. var. intermedia

(Rydb.) Hitchc. (U;MD)

Descurainia pinnata (Walt.) Britt. var. osmarium

(Cockerell) Shinners (C;W)

Descurainia sophia (L.) Webb. (0;WR)
Dithyrea wislizenii Engelm. in Wisliz. (0;SnD)
Lepidium densiflorum Schrad. var. pubicarpum (Nels.)

Thell. (R;PJ)

Lepidium densiflorum Schrad. var. ramosum (Nels.)

Thell. (0;MD,PJ)

Lepidium montanum Nutt. var. jonesii (Rydb.) Hitchc.

(C;W)

Lesquerella alpina (Nutt.) Wats. var. alpina

(0;PJ,PM)

Lesquerella intermedia (Wats.) Heller (0;PJ,PM)
Lesquerella kingii Wats. (R;PJ)
Lesquerella rectipes Woot. & Standi. (C;W)
Lesquerella subumbellata Rollins (U;PJ,PM)
Malcolmia africana (L.) R. Br. in Ait. (C;S1D,DS)
Physaria acutifolia Rydb. (C;PJ)
Physaria chambersii Rollins (U;PJ)
Schoencrambe linifolia (Nutt.) Greene (0;PJ,PM)
Sisymbrium altissimum L. (C;PJ,PM)
Stanleya pinnata (Pursh) Britt. (C;S1D)
Stanleya viridiflora Nutt. in T. & G. (U;MD,PJ)
Streptanthella longirostris (Wats.) Rydb. (C;W)
Streptanthus cordatus Nutt. ex T. & G. (C;PJ)
Thelypodiopsis barnebyi Welsh & Atwood (R;MD)
Thelypodiopsis divaricata (Rollins) Welsh & Reveal

(C;S1D,MD,PJ)

Cactaceae

Coryphantha vivipara (Nutt.) Britt. & Rose (0;PJ)
Echinocereus triglochidiatus Engelm. var. melana-

canthus (Engelm.) Benson (C;S1D,MD,PJ)

Opuntia basilaris Engelm. & Bigel. (0;SnD,MD,PJ)
Opuntia erinacea Engelm. (0;SnD)
Opuntia polyacantha Haw. (C;W)
Pediocactus despainii Welsh & Goodrich (R;PJ)
Pediocactus simpsonii (Engelm.) Britt. & Rose (R;PJ)
Sclerocactus parviflorus Clover & Jotter var. inter^

medius (Peebles) Woodruff & Benson (C;W)
Sclerocactus wrightiae Benson (U;S1D,MD,PJ)

Capparidaceae

Cleome lutea Hook. (C;S1D)
Cleome serrulata Pursh (0;PJ)
Cleomella palmerana Jones (0;PJ)

Caprifoliaceae

Sambucus coerulea Raf. (U;PJ,PM)
Symphoricarpos longiflorus Gray (0;PJ,PM)

Caryophyllaceae

Arenaria eastwoodiae Rydb. (C;PJ)

Arenaria fendleri Gray (C;S1D)

Arenaria hookeri Nutt. var. desertorum Maguire (R;PJ)

Paronychia sessiliflora Nutt. (0;PJ,WR)

Celastraceae

Forsellesia meionandra (Koehne) Heller (0;PJ,RO)

Chenopodiaceae

Allenrolfea occidentalis (Wats.) Kuntze (0;S1D)
Atriplex argentea Nutt. (U;S1D)
Atriplex canescens (Pursh) Nutt. (C;S1D,MD)
Atriplex confertifolia (Torr. & Frem.) Wats. (C;S1D)
Atriplex corrugata Wats. (C;S1D)

Great Basin Naturalist

Vol. 43, No. 1

Atriplex cuneata Nels. (C;S1D)

Atriplex graciliflora Jones (U;S1D)

Atriplex powellii Wats. {C;S1D)

Atriplex saccaria Wats. (U;MD)

Bassia hysopifolia Pallas (U;WR)

Ceratoides lanata (Pursh) Howell (C;S1D)

Chenopodium fremontii Wats. (C;W)

Chenopodium glauciim L. (0;S1D)

Chenopodium pratericola Rydb. (C;SS)

Grayia brandegei Gray (U;MD)

Halogeton glomeratus (Bieb.) Meyer (C;DS)

Kochia iranica Bomm. (C;WR)

Kochia americana Wats. (R;S1D,WR)

Monolepis nuttalliana (Schult.) Greene (0;WR)

Sahola iberica Sennen & Pan (C;W)

Sarcobatus vermiculatus (Hook.) Torr. (C;S1D,SS,WR)

Suaeda torreyana Wats. (C;SS,WR)

CONVOLVULACEAE

Convoltmlus arvensis L. (C;WR)
Cuscuta cuspidata Engelm. (C;MD)

CuPRESSACEAE

Juniperus osteosperma (Torr.) Little (C;PJ,PM)

Juniperus scopulorum Sarg. (U;PM)
Cyperaceae

Carex filifolia Nutt. (C;SS)

Carex parryana Dewey (R;SS)

Eleocharis palustris (L.) Roemer & Schult. (0;SS)

Eleocharis rostellata (Torr.) Torr. (U;SS)

Scirpus maritimus L. (0;SS)

Scirpus pungens Vahl. (C;SS)

Scirpus validus Vahl. (0;SS)
Elaeagnaceae

Shepherdia rotundifolia Parry (U;PJ)

Shepherdia argentea (Pursh) Nutt. (U;WR)
Ephedraceae

Ephedra torreyana Wats. (C;W)

Ephedra viridis Coville (C;W)
Equisetaceae

Equisetum laevigatum A. Br. (C;SS)
Euphorbiaceae

Euphorbia albomarginata T. & G. (U;PJ)

Euphorbia fendleri T. & G. (C;S1D,MD,PJ)

Euphorbia micromera Boiss.

Euphorbia nephradenia Bameby (R;S1D,WR)

Euphorbia palmeri Engelm. (R;S1D)

Euphorbia parryi Engelm. (G;SnD)

Euphorbia robusta (Engelm.) Small (0;PJ,PM)

Euphorbia serphyllifolia Pers. (C;PJ,PM)
Fabaceae

Astragalus amphioxys Gray var. amphioxys
(C;S1D,MD,PJ)

Astragalus amphioxys Gray var. vespertinus (Sheld.)
Jones (C;W)

Astragalus argophyllus Nutt. var. martinii Jones
(U;PJ,PM)

Astragalus asclepiadoides Jones (C;S1D)

Astragalus brandegei Porter (U;S1D,MD,PJ)

Astragalus calycosus Torr. (0;S1D,MD,PJ)

Astragalus ceramicus Sheldon (U;WR)

Astragalus coltonii Jones var. coltonii (C;S1D,MD,PJ)

Astragalus convallarius Greene var. convallarius
(U;PJ,PM)

Astragalus cymboides Jones (C;S1D,MD,PJ)

Astragalus desperatus Jones var. desperatus
(C;SnD,MD,PJ)

Astragalus desperatus Jones var. petrophilus Jones
(C;MD,SnD,PJ)

Astragalus episcopus Wats. (C;MD,PJ)

Astragalus flavus Nutt. var. argillosus (Jones) Bameby
(C;S1D)

Astragalus flavus Nutt. var. flavus (C;S1D,MD,PJ)

Astragalus flexuosus (Hook.) Don. var. diehlii (Jones).
Barneby (0;S1D,MD,PJ)

Astragalus gey eri Gray (0;MD)

Astragalus kentrophyta Gray var. cohradoensis Jones
(U;WR,RO)

Astragalus kentrophyta Gray var. elatus Wats. (R;S1D)

Astragalus lentiginosus Dougl. var. araneosus (Sheld.)
Barneby (G;MD,PJ)

Astragalus lentiginosus Dougl. var. palans (Jones)
Jones (C;MD,PJ)

Astragalus lonchocarpus Torr. (C;S1D,MD,PJ)

Astragalus moencoppensis Jones (G;S1D,MD,PJ)

Astragalus mollissimus Torr. var. thompsonae (Wats.)
Barneby (C;W)

Astragalus musiniensis Jones (C;S1D,MD,PJ)

Astragalus pardalinus (Rydb.) Barneby

(0;SnD,MD,PJ)

Astragalus praelongus Sheld. var. ellisiae (Rydb.)
Barneby (C;S1D)

Astragalus praelongus Sheld. var. praelongus (U;S1D)

Astragalus preussii Gray var. preussii (C;S1D,MD,PJ)

Astragalus rafaelensis Jones (0;S1D)

Astragalus sabulonum Gray (0;PJ,PM)

Astragalus spatulatus Sheld. (0;PJ,PM)

Astragalus subcinereus Gray var. basalticus Welsh

(R;PJ)

Astragalus tenellus Pursh (0;MD,PJ)
Astragalus wingatanus Wats. (R;S1D,MD,PJ)
Astragalus woodruffii Jones (U;SnD,MD)
Dalea flavescens (Wats.) Welsh (0;SnD,MD,PJ)
Dalea oligophylla (Torr.) Shinners (0;WR,RO)
Glycyrrhiza lepidota Pursh (C;WR,SS)
Hedysarum boreale Nutt. (C;MD,PJ,PM)
Hoffmanseggia repens (Eastw.) Cockerell (C;SnD)
Lathyrus brachycalyx Rydb. var. eucosmus (Butters &

St. John) Welsh (U;WR)

Lupinus argenteus Pursh var. argenteus (R;WR)

Lupinus argenteus Pursh var. tenellus Dougl. (U;MD)

Lupinus brevicaulis Wats. (U;S1D,MD)

Lupinus pusillus Pursh var. pusillus (C;S1D,MD,PJ)

Lupinus pusillus Pursh var. rubens (Rydb.) Welsh

Medicago sativa L. (C;WR)

Melilotus alba Medicus (0;SS,WR)

Melilotus officinalis (L.) Lam. (0;SS,WR)

Oxytropis lambertii Pursh var. bigelovii Gray

(R;PJ,PM)

Oxytropis sericea Nutt. (0;MD,PJ)

Psoralea lanceolata Pursh var. lanceolata (U;MD,PJ)

Psoralea lanceolata Pursh var. stenophylla (Rydb.)

Toft & Welsh (0;WR)

Psorothamnus polyadenius (Torr.) Rydb. var. jonesii

Barneby (R;S1D,MD)

Psorothamnus thompsonae (Vail) Welsh & Atwood

(R;MD)

Robinia pseudoacacia L. (R;WR)
Sophora stenophylla Gray (0;MD)

Fumariaceae

Corydalis aurea Willd. (R;PJ)

January 1983

Harris: San Rafael Swell Flora

GEI>rrtANACEAE

Swertia albomarginata (Wats.) Kuntze (R;PM)
Swertih utahensis (Jones) St. John (C;SnD,MD)
Hydrophyllaceae
Phacelia constancei Atwood (R;S1D)
Phacelia corrugata Nels. (C;W)
Phacelia demissa Gray var. demissa (0;S1D,PJ)
Phacelia demissa Gray var. heterotricha Howell
Phacelia indecora Howell (R;S1D)
Phacelia ivesiana Torr. (C;S1D,MD,PJ)
Phacelia rafaelensis Atwood (C;MD,PJ)

JUNCACEAE

Juncus arcticus Willd. (C;SS)
Juncus balticus Willd. (C;SS)
Juncus nodosus L. (U;SS)
Juncus torreyi Coville (0;SS)

JUNCAGINACEAE

Triglochin maritima L. (U;SS)
Lamiaceae

Poliomintha incana (Torr.) Gray (C;SnD)
Liliaceae

Allium geyeri Wats. (C;PJ,PM)

Allium macropetalum Rydb. (U;S1D)

Allium textile Nels. & Macbr. (C;SnD)

Androstephium breviflorum Wats. (C;S1D,MD,PJ)

Asparagus officinalis L. (0;WR,SS)

Cahchortus nuttallii T. & G. (C;S1D,MD)

Eremocrinum albomarginatum (Jones) Jones (C;SnD)

Smilacina stellata (L.) Desf. (C;HG)

Yucca harrimaniae Trel. (C;SnD,MD,PJ)

LiNACEAE

Linum aristatum Engelm. (U;WR)
Linum perenne L. (C;WR)
Linum puberulum (Engelm.) Heller (0;S1D)
Linum subteres Winkler (0;WR,PJ)

LOASACEAE

Mentzelia albicaulis Dougl. (C;S1D,MD,PJ)
Mentzelia humilis (Gray) DarL (U;WR)
Mentzelia multiflora (Nutt.) Gray (C;SnD)
Mentzelia pterosperma Eastw. (U;WR)
Mentzelia pumila (Nutt.) T. & G. (0;WR)

Malvaceae

Sphaeralcea coccinea (Nutt.) Rydb. (C;W)
Sphaeralcea grossulariifolia (Hook. & Arn.) Rydb.

(0;SnD,MD)

Sphaeralcea leptophylla (Gray) Rydb. (R;WR)
Sphaeralcea munroana (Dougl.) Spach
Sphaeralcea parvifolia Nels. (C;W)
Sphaeralcea psoraloides Welsh (R;S1D,MD)

Nyctaginaceae
Abronia elliptica Nels. (C;SnD,MD,WR)
Abronia nana Wats. (U;S1D,PJ)
Allionia incarnata L.

Oxybaphus linearis (Pursh) Robins. (C;W)
Tripterocalyx micranthus (Torr.) Hook. (0;SnD)

Oleaceae

Fraxinus anomala Torr. in Wats. (C;WR)

Onagraceae

C^lylOphus lavandulaefolia (T. & G.) Raven

(0;PT,RO)

Cafnissonia eastwoodiae (Munz) Raven (C;SnD,SlD)
CdviU^hhia scapoidea (T. & G.) Raven (C;SnD,SlD)
Cafnissonia walkeri (Nels.) Raven (U;S1D,MD)
Oenothera brachycarpa (Gray) Britt. (U;MD,PJ,WR)
Oenothera caespitosa Nutt. (C;SnD,MD,WR)
Oenothera pallida Lindl. ssp. pallida (C;SnD,MD,WR)

Oenothera pallida Lindl. ssp. trichocalyx (Nutt.) Munz
& Klein (C;S1D,MD)
Orchidaceae

Epipactis gigantea Dougl. ex Hook. (C;HG,SS)
Orobanchaceae

Orobanche fasciculata Nutt. (0;W)
Papaveraceae

Argemone corymbosa Greene (C;SnD)

Argemone munita Dur. & Hilg. (C;SnD)

PiNACEAE

Pinus edulis Engelm. (C;PJ,PM)
Pinus ponderosa Lawson (C;PM)
Pseudotsuga menziesii (Mirb.) Franc. (C;PM)
Plantaginaceae

Plantago patagonica Jacq. (C;S1D,MD)

POACEAE

Agropyron caninum (L.) Beauv. (U;MD,RO)

Agropyron cristatum (L.) Gaertn. (C;MD,PJ)

Agropyron spicatum (Pursh) Scribn. & Smith (U;PJ)

Agropyron trachycaulum (Link) Malte (U;WR)

Andropogon hallii Hackel (R;WR)

Aristida purpurea Nutt. (C;W)

Bouteloua curtipendula (Michx.) Gray (R;MD)

Bouteloua gracilis (H.B.K.) Lag. ex Steud.
(C;S1D,MD,PJ)

Bromus japonicus Thunb. (R;PJ)

Bromus tectorum L. (C;WR,DS)

Calamagrostis scopuhrum Jones (U;WR)

Dichanthelium lanuginosum (Elliott) Gould (U;SS)

Distichlis spicata (L.) Greene var. striata (Torr.)
Scribn. (C;S1D,WR)

Elymus canadensis L. (U;WR)

Elymus cinereus Scribn. & Merr. (0;MD,PJ)

Elymus junceus Fischer (R;S1D,MD)

Elymus salina Jones (C;PJ)

Erioneuron pilosum (Buckley) Nash (C;MD,RO,WR)

Erioneuron pulchellum (H.B.K.) Tateoka (R;PJ)

Festuca pratensis Huds. (U;SS)

Hilaria jamesii (Torr.) Benth. (C;W)

Hordeum jubatum L. (C;WR)

Muhlenbergia asperifolia (Nees & Mey.) Parodi
(0;WR)

Muhlenbergia pungens Thurber (C;SnD)

Munroa squarrosa (Nutt.) Torr. (0;PJ)

Oryzopsis hymenoides (Roem. & Schult.) Ricker
(C;W)

Oryzopsis micrantha (Trin. & Rupr.) Thurber (U;PJ)

Panicum virgatum L. (0;SS,WR)

Phragmites australis (Car.) Trin. ex Steud. (C;SS)

Poa fendleriana (Steud.) Vasey (0;PJ,PM)

Poa pratensis L. (U;SS,WR)

Poa sandbergii Vasey (U;PJ)

Polypogon monspeliensis (L.) Desf. (U;SS,WR)

Schedonnardus paniculatus (Nutt.) Trel. (R;S1D)

Schizachyrium scoparium (Michx.) Nash (0;PJ,WR)

Sitanion hystrix (Nutt.) Smith (C;W)

Spartina gracilis Trin. (U;SS)

Sporobolus airoides (Torr.) Torr. (C;S1D,MD)

Sporobolus contractus Hitchc. (0;MD,PJ,WR)

Sporobolus cryptandrus (Torr.) Gray (G;S1D,MD,PJ)

Sporobolus flexuosus (Thurb.) Rydb. (U;SnD,WR)

Sporobolus giganteus Nash

Stipa arida Jones (U;WR)

Stipa comata Trin. & Rupr. (C;MD,PJ)

Stipa Columbiana Macoun. (U;PJ)

Great Basin Naturalist

Vol. 43, No. 1

SHpa speciosa Trin. & Rupr. (U;PJ)
' Vulpia octoflora (Walt.) Rydb. (0;PJ,WR)

. t'oLEMONIACEAE

Gilia aggregata (Pursh) Spreng. (C;W)

Gilia congesta Hook. (0;S1D)

Gilia gunnisonii T. & G. (C;MD,WR)

Gilia inconspicua (Smith) Sweet (C;PJ)

Gilia latifolia Wats. (R;RO,WR)

Gilia leptamena Gray (C;SnD,MD,PJ)

Gilia polycladon Torr. in Emory (0;S1D)

Gilia pumila Nutt. (0;S1D)

Gilia roseata Rydb. (C;WR,RO)

Gilia stenothyrsa Gray (0;PJ)

Leptodactylon caespitosum Nutt. (U;WR)

Leptodactylon pungens (Torr.) Nutt. (0;MD,WR,RO)

Leptodactylon watsonii (Gray) Rydb. (U;WR,RO)

Phlox austromontana Gov. (0;PJ)

Phlox hoodii Rich. (C;MD,PJ)

Phlox hngifolia Nutt. (0;S1D,MD)

POLYGALACEAE

Polygala acanthoclada Gray (R;MD)
Polygala subspinosa Wats. (G;SlD,PJ,RO)

POLYGONACEAE

Eriogonum alatum Torr. in Sitgr. (C;PJ)

Eriogonum hatemanii Jones (C;PJ)

Eriogonum. bicolor Jones (C;MD,PJ)

Eriogonum cernuum Nutt. (C;S1D)

Eriogonum corymbosum Benth. var. corymbosum
(G;S1D,MD)

Eriogonum fasciculatum Benth. var. polifolium
(Benth. in DG.) T. & G. (R;MD)

Eriogonum flexum Jones (R;S1D)

Eriogonum gordonii Benth. (U;S1D,MD)

Eriogonum hookeri Wats. (0;MD,PJ)

Eriogonum inflatum Torr. & Frem. var. fusiforme
(Small) Reveal (0;S1D,MD,PJ)

Eriogonum inflatum Torr. & Frem. var. inflatum
(G;S1D,MD)

Eriogonum jamesii Benth. var. flavescens Wats.
(G;MD,PJ,WR)

Eriogonum leptocladon T. & G. var. leptocladon
(C;S1D)

Eriogonum microthecum Nutt. var. foliosum (T. & G.)
Reveal (0;S1D)

Eriogonum ovalifolium Nutt. var. ovalifolium
(C;W,RO)

Eriogonum salsuginosum (Nutt.) Hook. (0;MD,PJ)

Eriogonum shockleyi Wats. var. longilobum (Jones)
Reveal (C;S1D,MD)

Eriogonum smithii Reveal

Eriogonum tumulosum (Bameby) Reveal (U;PJ)

Eriogonum wetherillii Eastw. (0;W)

Polygonum aviculare L. (0;WR,DS)

POLYPODIACEAE

Adiantum capillus-veneris L. (U;HG)
Cheilanthes feei Moore (0;HG)
Pellaea glabella Mett. ex Kuhn (R;HG)

PORTULACEAE

Portulaca oleracea L. (R;PJ,PM)
Talinum validulum Greene (0;PJ)
Ranunculaceae
Aquilegia micrantha Eastw. (C;HG)
Clematis ligusticifolia Nutt. (G;WR)
Delphinium nuttallianum Pritz. ex Walp. (0;W)
Delphinium scaposum Greene (0;PJ)

Ranunculus cymbalaria Pursh (U;SS)

Ranunculus testiculatus Grantz (0;DS)
Rosaceae

Amelanchier utahensis Koehne (C;WR,SS)

Cercocarpus intricatus Wats. (G;PJ,RO)

Cercocarpus montanus Raf. (G;PJ,PM,WR)

Coleogyne ramosissima Torr. (G;SnD)

Cowania mexicana D. Don (C;PJ)

Fallugia paradoxa (D. Don) Endl. (0;WR)

Physocarpus alternans (Jones) Howell

Physocarpus monogynus (Torr.) Gou. (U;PJ)

Purshia tridentata (Pursh) DC. (0;WR)

Rosa woodsii Lindl. (R;PM)
Rubiaceae

Galium multiflorum Kellogg (0;WR)
Salicaceae

Populus x acuminata Rydb. (U;WR,SS)

Popidus angustifolia James ex Torr. (0;WR)

Populus fremontii Wats. (G;WR,SS)

Populus tremuloides Michx. (0;PM)

Salix exigua Nutt. (G;SS)
Santalaceae

Comandra umbellata (L.) Nutt. var. pallida (DG.)
Jones (G;W,WR)
Saxifragaceae

Philadelphus microphyllus Gray (G;PJ,PM)

Ribes cereum Dougl. (R;PJ,PM)

Ribes leptanthum Gray (0;PM)

Scrophulariaceae

Castilleja chromosa Nels. (0;W)

Castilleja linariaefolia Benth. (0;SS)

Castilleja scabrida Eastw. (G;W)

Cordylanthus kingii Wats. (0;PJ)

Cordylanthus wrightii Gray

Mimulus rubellus Gray (R;PM)

Penstemon carnosus Pennell (G;PJ)

Penstemon cyanocaulis Payson (U;WR)

Penstemon eatonii Gray (0;PJ,PM)

Penstemon lentus Pennell (U;WR)

Penstemon utahensis Eastw. (G;PJ)
Selaginellaceae

Selaginella mutica Eaton ex Underw. (U;HG)
Tamaricaceae

Tamarix ramosissima Ledeb. (G;SS,WR)
Typhaceae

Typha latifolia L. (G;SS)
Ulmaceae

Celtis reticulata Torr. (U;WR)

Ulmus pumila L. (R;WR)

Literature Cited

Cronquist, a., a. Holmgren, N. Holmgren, and J.
Reveal. 1972. Intermountain flora: vascular
plants of the Intermountain West. vol. I. Hafner
Publishing Gompany, Inc., New York.

Gronquist, a., a. Holmgren, N. Holmgren, J. Reveal,
AND P. Holmgren. 1977. Intermountain flora:
vascular plants of the Intermountain West. vol.
VI. Golumbia University Press, New York.

Neese, E., and S. L. Welsh. 1981. Astragalus section
Desperati (Leguminosae) and a new species from
the Uinta Basin of Utah. Rhodora 83:455-460.

Welsh, S. L. 1978. Utah flora: Fabaceae (Leguminosae).
Great Basin Nat. 38:225-367.

January 1983

Harris: San Rafael Swell Flora

1980a. Utah flora: Malvaceae. Great Basin Nat.

40:27-37.
1980b. Utah flora: miscellaneous families. Great

Basin Nat. 40:38-58.
1981. New taxa of western plants— In tribute.

Brittonia 33:294-303.
Welsh, S. L., N. D. Atwood, S. Goodrich, E. Neese,

K. H. Thorne, and B. Albee. 1981. Preliminary

index of Utah vascular plant names. Great Basin

Nat. 41:1-108.
Welsh, S. L., and G. Moore. 1973. Utah plants:

Tracheophyta. 3d ed. Brigham Young University

Press, Provo, Utah.
Welsh, S. L., and J. L. Reveal. 1977. Utah flora: Brassi-

caceae (Cruciferae). Great Basin Nat. 37:297-364.

PRONGHORN RESPONSES TO HUNTING COYOTES

Timothy D. Reynolds'

Abstract.— Six accounts of pronghorn antelope {Antilocapra americana) chasing or attacking coyotes (Canis lat-
rans) are described: three chases by individual pronghorn does, two by herds of antelope, and one joint effort by a
pronghorn doe and a Short -eared Owl {Asio flammeus). Modifications of Berger's (1979) ungulate antipredatory de-
fense model are proposed.

Coyotes {Canis latrans) in the western
United States feed on pronghorn antelope
{Antilocapra americana). Pubhshed accounts
indicate that coyote predation on pronghorn
is not a particularly rare event (Thompson
1949, Arrington and Edwards 1951, Udy
1953, Beale and Smith 1973). In fact.
Springer and Smith (1981) recorded prong-
horn remains in more than 50 percent of the
summer coyote scats they examined. Con-
trariwise, until recently, published accounts
of responses of pronghorn to predators were
uncommon, and records of pronghorn chas-
ing cr attacking coyotes were lacking. Berger
(1979) described a "previously unknown de-
fense strategy in pronghorn" in which a
group of antelope chased a coyote. From this
observation he developed a schematic repre-
sentation of antipredatory defenses in un-
gulates, and concluded that predator harass-
ment is beneficial to the prey by (1) giving
naive individuals the opportunity to recog-
nize predators in a low risk situation, (2) al-
lowing the prey to safely monitor the pred-
ator's position, and (3) making the predator
reluctant to attack in the future. Lipetz and
Bekoff (1980) analyzed 25 antelope-coyote
chases and concluded only that such encoun-
ters appear to have direct survival value for
proiighom fawns.

Pescribed here are six observations of
pronghorn, either singly or in groups, chasing
coyotes. One event, detailed below, included
a joint effort between a Short-eared Owl
{Asio flammeus) and a pronghorn doe. A re-
finement of Berger's (1979) antipredatory de-
fense model is proposed for pronghorn. All

observations were recorded in the sagebrush
{Artemisia tridentata) dominated habitat at
the National Environmental Research Park
on the U.S. Department of Energy's Idaho
National Engineering Laboratory (INEL) Site
in southeastern Idaho.

Observations

Group Response

On 14 July 1978 and 18 November 1979, I
witnessed groups of pronghorn chase coyotes.
The first occasion was similar to Berger's
(1979) report. A coyote was observed stalking
a small band of antelope (4 does, 1 fawn) that
was loafing and feeding about 300 m from a
larger group (5 does, 2 bucks, 2 fawns). One
feeding doe from the smaller band apparent-
ly sighted the coyote at a distance of nearly
100 m, stared toward the coyote for a few
seconds, and sounded an alarm call. The re-
maining antelope of both groups were then
alert and directed their attention toward the
vicinity of the coyote. When the stalking
coyote approached within 40 m, the group of
5 quickly joined the larger group. The coyote
followed, maintaining a distance of 40-50 m
from the antelope, and sat down as the
groups merged. One doe (thought to be a
yearling) took a few steps toward the coyote,
then returned to the main group. She re-
peated this investigative sequence twice. On
the fourth foray she was accompanied by the
13 other pronghorn. All antelope stopped
momentarily about 30 m from the coyote,
then burst into a full run toward the coyote.

'Department of Biology, Idaho State University, Pocatello, Idaho 83209; and U.S. Department of Energy, Radiological and Environmental Science Labo-
ratory, 550 2nd Street, Idaho Falls, Idaho 83401. Present address: Biology Department, Boise State University, Boise, Idaho 83725.

January 1983

Reynolds: Pronghorn Responses

The coyote fled and was pursued for 300-400
m before disappearing from view.

The November 1979 encounter differed
from the previous one in that 2 coyotes were
observed moving near a large group of about
120 pronghorn. The pronghorn sighted the
coyotes at a distance of about 200 m. A large
buck left the group and walked directly to-
ward the coyotes. He was followed by about
20 animals (both does and bucks). This group
began running toward the coyotes when the
distance had closed to less than 100 m. The
remaining 100 or so pronghorn simply ob-
served the chase. The coyotes immediately
took flight. The pronghorn stopped chasing
after running about 150 m. The coyotes con-
tinued their retreat, but at a slow run or trot,
while repeatedly looking back at the prong-
horn. The coyotes vanished from view at
about 600 m.

My first example above, and Berger's
(1979) report, suggest that pronghorn groups
must contain sufficient numbers of animals
before a chase will be initiated. This lower
limit or threshold concept may be valid un-
der certain conditions, but as evidenced by
the following accounts it is by no means a
universal trend in antelope antipredatory
behavior.

Doe with Fawns

On 20 June 1978, a female pronghorn was
observed nursing two fawns, seemingly una-
ware of a coyote furtively approaching her
from the rear. While licking one fawn, the
doe apparently noticed the coyote 30 m
away and gave an alarm call. The fawns
promptly dropped into the immobility re-
sponse (Autenrieth and Fichter 1975) and the
doe stared intently at the advancing coyote.
The coyote approached to within 20 m, and
then made a dash toward the antelope. The
doe responded by charging the coyote, caus-
ing it to veer away from the fawns. The doe
pursued the coyote for nearly 50 m, and then
returned to a position about halfway between
the fawns and the predator. Twice more the
coyote ran toward the fawns, and each time
was thwarted by the charging doe, who again
positioned herself between her young and the
coyote. The coyote slowly moved away from
the antelope while the doe intently watched

its progress. When the coyote had withdrawn
to a distance of 80 m, it abruptly changed its
direction, putting itself on a course that
would bring it within 20-30 m of the fawns.
The doe again charged the coyote and pur-
sued it for nearly 400 m before both dis-
appeared from view. The doe returned to the
area 85 minutes later, called her fawns from
seclusion, and resumed nursing them.

Another postparturient doe and coyote in-
teraction was observed on 30 June 1978.
Other than the fact that this doe had only
one fawn, this encounter closely followed the
sequence described above: the coyote ap-
proached to within 30 m before charging,
only to be charged by the doe. Second and
third attacks followed; each time the coyote
was chased a short distance away by the doe.
On the fourth attempt, as the coyote veered
from its course, the doe actually butted it in
the side, rolling it over. The coyote regained
its footing without losing momentum, and
was vigorously pursued by the doe for about
150 m. The doe stopped, watched the coyote
run away, then intermittently fed, or sham-
fed (Autenrieth and Fichter 1975), for nearly
30 minutes before returning to the vicinity of
the fawn.

A third antelope doe was observed defend-
ing two neonates, approximately two weeks
old, from a pair of coyotes on 8 June 1979.
When observations began (0925 hours MST)
the doe was feeding and the fawns were ca-
vorting nearby. At 0932 hours the doe
spotted coyote No. 1 about 50 m to the north
and gave an alarm call. The fawns immedi-
ately lay down, separated from each other by
a distance of 8-10 m. As the doe focused her
attention on the now stationary coyote, coy-
ote No. 2 appeared behind coyote No. 1 and
began moving in an arc toward the east. Coy-
ote No. 2 had approached to within 20 m of
the fawns when the doe charged it, causing it
to move further eastward from the fawns. Al-
most simultaneously, coyote No. 1 dashed to-
ward the fawns and was within 10 m of them
before the doe whirled and charged, forcing
it to the west of the secluded young. Coyote
No. 2 then advanced and was driven off,
again to the east. Coyote No. 1 again at-
tacked, this time advancing within 1-2 m of
one of the fawns before being repulsed by the
doe. Bleating, the fawn burst from its bed,

Great Basin Naturalist

Vol. 43, No. 1

and ran in a southerly direction accompanied
by the doe. At the sound of the bleat, the sec-
ond fawn immediately took flight, but was
quickly brought down from behind by coyote
No. 2. The doe and the surviving fawn ran at
full speed for nearly 200 m and abruptly
stopped. The fawn lay down and the doe
moved in a seemingly leisure manner,
roughly in a southwestward direction, fre-
quently looking back toward the feeding
coyotes.

Joint Interspecific Response

At 0545 hours on 13 May 1977, an extraor-
dinary predator-prey encounter was observed
and recorded. A single pronghorn doe was
observed feeding about 400 m west of my po-
sition and about 80 m west of a Short-eared
Owl nest known to contain two yoimg. A
Short-eared Owl was noticed flying oddly
about 100 m north of the doe. The owl was
flying in a southerly direction and repeatedly
"dive bombing" from a height of 10-15 m to
the top of the sage. The owl continued this
undulating flight toward the now alert
pronghorn. As the owl closely approached,
the doe ran through the sagebrush in the
same direction as the owl's flight, alternating
a head up and head down posture. The latter
was coordinated with short bursts of speed.
The animals continued this pattern for about
100 m. As they emerged from the sagebrush
into a crested wheatgrass (Agropyron crista-
tum) planting, a coyote was seen running
ahead of the antelope and below the owl. To-
gether, they pursued the coyote for nearly
300 m before the coyote reentered the sage-
brush. Both the pronghorn and the owl then
abandoned the chase. The doe looked in the
direction of the coyote for nearly five min-
utes, then resumed feeding. The owl circled
to a height of about 50 m and began hunting
activities. Carrying a prey item, it visited the
nest 15 minutes later. Further investigation
indicated that the owl's mate had been on or
near the nest throughout the joint anti-
predatory defense. It is doubtful that the an-
telope participating in the chase was pro-
tecting a fawn. The earliest record of
pronghoms fawning on the Idaho National
Engineering Laboratory Site is 23 May
(1980), with the peak of fawning normally

occurring the last week of May and the first
week of June each year.

Conclusions

Figure 1 is an adaptation of Berger's (1979)
antipredatory defense model for ungulates,
and represents my proposed spectrum of
pronghorn responses to hunting coyotes. The
wide solid arrows indicate the responses most
likely to occur in pronghorn coyote encoun-
ters. Narrow solid lines represent docu-
mented responses that occur less often, and
the wavy arrows account for the rare obser-
vation of concurrent, interspecific chasing.
The dashed arrows indicate some possible re-
actions of pronghorn to hunting coyotes that
were not recorded in my observations.

The actions taken by pronghorn when con-
fronted by coyotes appear to be generally re-
lated to the size and composition of the
pronghorn group. Individuals unaccompanied
by fawns, or small groups of pronghorn, tend
to retreat from coyote predators, often join-
ing other bands of pronghorn. Larger groups
of pronghorn exhibit a continuum of respon-
ses ranging from mild interest, or curiosity, to
actual attack that in the broadest context
represents mobbing behavior (Harvey and
Greenwood 1978). My observations indicate
the postparturient does, with fawns nearby,
invariably attack or chase coyotes advancing
toward them. The intensity of these attacks,
and the context in which they occur, closely
resembles the antipredatory response of
"snarling" described by Curio (1975). The
constancy of this behavior supports the thesis
that, in certain situations (i.e., does with
fawns nearby), predator harassment has di-
rect survival value for pronghorn fawns
(Lipetz and Bekoff 1980). The significance of
the joint (cooperative?) chase by a pronghorn
doe and a Short-eared Owl is unknown.

There are two plausible interpretations of
this event. First, as several instances of
pronghorn chasing Short-eared Owls in an
antipredatory context have been observed
(Fichter, pers. comm., Copeland, in litt.), it is
possible that the doe was responding to both
the coyote and the owl as potential pred-
ators. However, if the doe was not protecting
a fawn (the date of this encounter suggests
she was not), the adaptive advantage, or evo

January 1983

Reynolds: Pronghorn Responses

Individual

or small group

Interspecific
cooperation

DOE with fawn(s)

Attack

Fig. 1. A proposed model of the responses of pronghorn antelope to coyotes. See text for the explanation of
arrows.

lutionary significance of her actions is not ob-
vious. On the other hand, as this paper and
that by Lipetz and Bekoff (1980) suggests,
pronghorn chasing coyotes is not an excep-
tionally rare event. Although published re-
cords are few. Short-eared Owls have not in-
frequently been observed mobbing predatory
species, including coyotes (pers. obs., Trost,
pers. comm., Clark 1975). It is likely that the
antelope-owl-coyote interaction described

here represents a mutual, albeit fortuitous, ef-
fort by the antelope and owl to hustle the
coyote. The proximity of the chase to the owl
nest makes the reasons behind the owl's in-
volvement obvious. The factors precipitating
the antelope's behavior are less certain. Fich-
ter (unpublished data) witnessed a buck band
of over a dozen pronghorn chase a coyote in
mid- June 1965. These animals pursued the
predator for 1-1.5 km, passing in front of and

Great Basin Naturalist

Vol. 43, No. 1

circling the running coyote at least twice, a
sequence frequently associated with moving
vehicles in pronghom country. It is possible
that in certain low-risk situations, such as
when a coyote is already fleeing from harass-
ment, antelope may participate in the chase
as a playlike exercise. This might represent a
learning experience for the prey and/ or
predator, lending support to any or all of
Berger's (1979) explanations for ungulates at-
tacking predators. However, the rarity of
birds and mammals jointly mobbing a poten-
tial predator precludes a convenient analysis
of the role of thjs interspecific behavior in
the relationship of predators and prey. More
data are required before the evolutionary sig-
nificance of this and similar observations can
be properly assessed.

Acknowledgments

This is a contribution from the INEL Site
Ecological Studies Program, supported by
the Office of Health and Environmental Re-
search, U.S. Department of Energy. I thank
R. E. Autenrieth for his suggestions and M.
W. Barrett, P. T. Bromley, E. Fichter, O. D.
Markham, J. M. Peek, F. L. Rose, and G. E.
Svendsen for improving the drafts of this
manuscript.

Literature Cited

Arbington, O. N., and A. E. Edwards. 1951. Predator
control as a factor in antelope management.
Trans. N. Am. Wildl. Conf. 16:179-195.

Autenrieth, R. E., and E. Fichter. 1975. On the be-
havior and socialization of pronghom fawns.
Wildl. Monogr. No. 42. Ill pp.

Beale, D. R., and a. D. Smith. 1973. Mortality of
pronghorn antelope fawns in western Utah. J.
Wildl. Manage. 37:343-352.

Berger, J. 1979. "Predator harassment" as a defensive
strategy in ungulates. Am. Midi. Nat.
102:197-199.

Clark, R. J. 1975. A field study of the Short-eared Owl
Asio flammeus (Pontoppidan) in North America.
Wildl. Monogr. No. 47. 67 pp.

Curio, E. 1975. The functional organization of anti-
predator behavior in the Pied Flycatcher, a study
of avian visual perception. Anim. Behav.
23:1-115.

Harvey, P. H., and P. J. Greenwood. 1978. Anti-
predator defense strategies: some evolutionary
problems. Pages 129-151 in J. R. Krebs and N. B.
Davies, eds.. Behavioral ecology: an evolutionary
approach. Blackwell Scientific Pubs., Oxford. 494
pp.

Lipetz, V. E., and M. Bekoff. 1980. Possible functions
of predator harassment in pronghom antelope. J.
Mammal. 61:741-743.

Thompson, W. K. 1949. Predation on antelope. J. Wildl.
Manage. 13:313-314.

Udy, J. R. 1953. Effects of predator control on antelope
populations. Utah State Dept. Fish and Game
Publ. No. 5. 48 pp.

FLORISTICS OF THE UPPER WALKER RIVER, CALIFORNIA AND NEVADA

Matt Lavin'

Abstract.— A checklist of the vascular flora of the upper Walker River is presented. Listed are 1078 taxa from
this 4000 km^ area. The upper Walker River encompasses a portion of the boundary between the Intermountain and
Sierra Nevada floristic regions, and hence displays much floristic diversity within a relatively small area. Due to its
location along the east slope of the Sierra Nevada, the Walker River drainage is unique in that it contains much
elevational variation eastward into the Intermountain Region. This elevational extension is due to the presence of
large mountain ranges including the Sweetwater Mountains, the Bodie Hills, and the Wassuk Range. As a result of
this elevational variation, there is much overlapping of the two floristic regions. Additionally, the easternmost expo-
sures of the Sierran granodiorites occur within the Walker River basin and may enhance the eastward migration of
Sierran plants. The 90 percent floristic similarity (Sorenson's) between the Sweetwater Mountains, lying to the east
of the Sierra, and the east slope of the Sierra Nevada (within the Walker River drainage) indicates the Sweetwaters
to be more affiliated with the Sierran flora instead of the Intermountain flora.

The upper Walker River drainage includes
an area of approximately 4000 km 2, located
at the north end of Mono Coimty, California,
the southern ends of Douglas and Lyon coun-
ties, Nevada, and the western edge of Miner-
al County, Nevada. The western boundary of
this drainage generally follows the crest of
the Sierra Nevada from the Conway Summit-
Virginia Lakes area, northward to the Topaz
Lake-Monitor Pass area. The eastern bound-
ary is delimited by the Bodie, Masonic, and
Pine Grove hills and the southwest slopes of
the Pine Nut Range. The Sweetwater Moim-
tains and the Wellington hills lie between
these hills and the Sierra Nevada.

East slope Sierran vegetation present here
has been generally described by Billings
(1951) and Rundel et al. (1977). The area is
dominated by mixed conifer forests composed
of Pinus jeffreyi and Abies concolor at the
low elevations, and Abies magnifica, Tsuga
mertensiana, Pinus monticola, and P. albi-
caulis at the higher elevations. The alpine
vegetation of the Sierra is uniquely adapted
for extreme svmimer drought (Chabot and
Billings 1972).

To the east of the Sierra, Intermountain
vegetation (high elevation sagebrush steppe
situated above woodlands composed of Pinus
monophylla and Juniperus osteosperma) pre-

dominates. This vegetation has also been
briefly described by Billings (1951). The
Sweetwater Mountains, situated between the
Intermountain and Sierran floristic regions,
display characteristics of both.

The vascular flora of the upper Walker
River is modestly represented in literature.
Works such as Hinton (1975), Reveal (1968),
Cox (1972), Reveal and Ertter (1980), Hard-
ham and True (1972), Strother (1974), Munz
(1968), Bameby (1964), Dempster and Ehren-
dorfer (1965), Halse (1981), and others cite
specific collections made within this area.
Davis (1979) compiled a plant list and keys to
the plants occurring on the east central
Sierra (Owens Lake to Lake Tahoe) and asso-
ciated desert ranges. This work was based
solely on distributions given in existing floras.

Sharsmith (1940) includes the Walker Riv-
er portion of the Sierra Nevada as the north-
ernmost boundary of the Sierra alpine floris-
tic region. This area includes Leavitt Peak
south to Dunderberg Peak. Thome (1982) de-
fines the upper Walker River basin as the
northern boundary of the transmontane Cali-
fornian floristic region.

Major and Taylor (1977) conducted a vege-
tation study of the alpine zone of the Sweet-
water Mountains. They list 43 species from
the area. Taylor (1977) indicates a 40-50

'Department of Biology, University of Nevada, Reno, Nevada 89557. Present address: P.O. Box 13494, Reno, Nevada 89507.

Great Basin Naturalist

Vol. 43, No. 1

percent floristic similarity (Sorenson's) be-
tween the Carson Pass area of the Sierra Ne-
vada and the Sweetwater Mountains. Since a
50 percent turnover rate in plant species oc-
curs every 650 km along the Cascade-Sierran
axis (Taylor 1977), the Walker River portion
of the Sierra must have a very high degree of
similarity with Carson Pass, 60 km to the
north. An interpretation of this would mean
the Walker River portion of the Sierra has
approximately a 50 percent floristic sim-
ilarity to the Sweetwaters.

Bell (1980) described the alpine flora of the
Wassuk Range, located in the lower Walker
River drainage just west of Walker Lake (50
km east of the Sweetwater Mountains). She
lists 70 vascular plants from the alpine flora
of this region. Bell has also recently com-
pleted a study of the alpine flora of the
Sweetwater Mountains, but this information
is unavailable.

Messick (1982) completed a flora of the
Bodie Hills, and the results of this work are
incorporated in Table 1 .

The upper Walker River drainage is the
region of several type collections. The Sweet-
water Mountains are the type locale for
Cordylanthus ramosus ssp. setosus, Senecio
pattersonensis and Draba lemmonii var. in-
crassata. In the Bodie Hills, we have the type
locale for Draba quadricostata, Pinus mon-
ophylla, Arabis bodiensis (see Rollins 1982),
Streptanthus oliganthus, and Phacelia mon-
oensis. A. bodiensis, S. pattersonensis, D. lem-
monii var. incrassata, and D. quadricostata
are endemic to the Walker River drainage.

The Wellington Hills are the type locale
for Astragalus oophorus var. lavinii, and So-
nora Pass for Raillardella argentea, Cymop-
terus cinerarius, and Wyethia mollis. "A dry
rocky mountain near Sonora Pass" is the type
location for Astragalus platytropis, A. lentigi-
nosus var. ineptus, and A. whitneyi. Bameby
suggests that the type locale for these three
Astragali may be the Sweetwater Moujitains.
However, during the course of this in-
vestigation, all three, including A. platy-
tropis, were found on Leavitt and Emma
Peak of the Sierra Nevada. Therefore, the
type locale given by Gray could have possi-
bly referred to the Sierra instead of the
Sweetwaters.

Methods

The checklist (Table 1) was developed
from collections made during the course of
this study: July 1979 to August 1982. Addi-
tional collections were recorded from the
herbaria at the University of Nevada, Reno
(RENO), The New York Botanical Garden
(NY), and the California Academy of Sciences
(CAS). Collections from the upper Walker
River made by Arnold Tiehm and Margaret
Williams, Reno, Nevada; Steve Wharff,
Tonopah, Nevada; Frank Smith, Smithfield,
Utah; Dennis Breedlove, CAS; Joe Robertson,
E. F. Kleiner, Tom Lugaski, Pat and Ham
Vreeland, H. N. Mozingo, and Fred Ryser, all
from the University of Nevada, Reno, were
also recorded in this checklist. Approximately
3850 numbers were recorded. Distributional
information contained within the checklist
was continually refined while in the field.
Various documentary works, mentioned in
the introduction, were used to determine
those plants that have a probable distribution
within the upper Walker River drainage, but
were not observed during this study.

Almost all of the taxa listed can be found
on deposit at the University of Nevada, Reno,
herbarium and the Toiyabe National Forest
Supervisor's Office, Reno, Nevada.

The purposes of the checklist (Table 1) are
to both document the flora present in the up-
per Walker River and to document the geo-
graphical and altitudinal distribution of each
taxon within this area. Nomenclature gener-
ally follows Kartesz and Kartesz (1980).

Results and Discussion

Table 1 lists 1078 taxa of vascular plants
from the upper Walker River. The upper
Walker River drainage is unique in that it ex-
tends much elevational variation of the east
slope of the Sierra Nevada eastward into the
Intermountain Region, due to the presence
and close proximity of such large mountain
ranges as the Sweetwater Mountains, the
Bodie Hills, and the Wassuk Range. Along
any floristic boundary there is bound to be
some overlap of unique plant species or char-
acteristic vegetation of one flora into anoth-
er. However, many plants having their center
of distribution in the Intermountain Region

January 1983

Lavin: Walker River Floristics

also have outlying populations in the mon-
tane environments of the Sierra Nevada. This
phenomenon is discussed by Taylor (1976) for
the Carson Pass area of the Sierra. He attri-
butes the occurrence of many Intermountain
plants on the east slope of the Sierra to Xero-
thermic invasion. A list of these plants at the
headwaters of the Walker River follow:

Agoseris glauca var. monticola

Allium biceptrum

Allium parvum

Arabis bodiensis

A. pulchra var. pulchra

Artemisia arbuscula

Aster ascendens

Astragalus platytropis

A. iodanthus

Amelanchier utahensis

Antennaria dimorpha

Balsamorhiza sagittata

Calyptridium roseum

Cercocarpus ledifolius

Cirsium utahense

Cheilanthes gracillima

Chenopodium overi

Chorizanthe brevicomu var. spathulata

Chrysothamnus nauseosus ssp. albicaulis

C. viscidiflorus

Crepis acuminata

C. modocensis ssp. subacaulis

Castilleja linariifolia

Cordylanthus ramosus ssp. setosus

Cryptantha circumscissa

Cryptantha echinella

Erigeron aphanactis

E. nevadincola

E. breweri var. porphyreticus

E. eatonii ssp. plantagineus

Eriogonum elatum

E. microthecum var. ambiguum

E. ovalifolium var. nevadense

E. ivrightii var. subscaposum

Galium multiflorum

Gilia leptantha ssp. salticola

Grayia spinosa

Haplopappus acaulis

Heuchera duranii

Hydrophyllum capitatum var. alpinum

Leptodactylon pungens

Linanthus nuttallii

Lupinus caudatus ssp. caudatus

L. nevadensis

Lomatium nevadense var. nevadense

Lomatium nevadense var. parishii

Melica striata

Mentzelia congesta

Mimulus densus

M. rubellus

Navarretia breweri

Opuntia polyacantha var. rufispina

Poa nevadensis var. juncifolia

P. nevadensis var. nevadensis

Paeonia brownii

Penstemon bridgesii

Phoenicaulis cheiranthoides

Phacelia humilis

Phlox covillei

Plagiobothrys hispidus

P. kingii var. harknessii

Prunus andersonii

Purshia tridentata

Pinus monophylla

Ribes velutinum

Rosa woodsii var. ultramontana

Scrophularia desertorum

Senecio canus

S. pattersonensis

S. spartioides

Sisyrinchium halophilum

Sphaeromeria cana

Stephanomeria spinosa

Streptanthus oliganthus

Tetradymia canescens

Thelypodium crispum

Zigadenus paniculatus

These 77 plant taxa represent approx-
imately 10 percent of the flora present on the
east slope of the Sierra within the Walker
River drainage. This is below Taylor's (1976)
estimate of 20 percent for the east slope
within the Carson River drainage just to the
north. However, when distribution records
become more complete, it would not be sur-
prising to find 20 percent of the Sierran flora
within the Walker River basin being com-
posed of Intermountain elements.

The Xerothermic climate may also have
been responsible for the northern migration
of Amelancheir pallida var. covillei, Cea-
nothus greggii var. vestitus, Cercocarpus ledi-
folius var. intricatus, Ivesia purpurascens ssp.
congdonis, Cryptantha confertiflora, Phacelia
peirsoniana, and Plagiobothrys jonesii. The
Walker River drainage may be the north-
ernmost location for these plants. {Cryp-
tantha confertiflora has been fovmd just to
the north in the Carson River drainage.)

It is documented that the eastward migra-
tion of Sierran plant species into the Inter-
mountain Region is small relative to the
westward migration of Rocky Mountain
plants into this region (Harper et al. 1978).
However, the Walker River drainage has
many Sierra or Pacific cordilleran plant spe-
cies occurring well into the Intermountain
Region, some of which occur as far east as
Masonic Mountain or the Wassuk Range. A
list of these plants follows:

Agropyron pringlei
Allium campanulatum

Great Basin Naturalist

Vol. 43, No. 1

Allophyllum gilioides

A. violaceum

Amelanchier pallida

Anelsonia eurycarpa

Angelica lineariloba

Arabis inyoensis

A. platysperma var. howellii

Arnica nevadensis

Astragalus kentrophyta var. danaus

A. lentiginosus var. ineptus

A. purshii var. lectulus

Calochortus leichtlinii

Carex tahoensis

Chaenactis nevadensis

Chrysothamnus parryi ssp. monocephalus

Claytonia nevadensis

Cryptantha glomeriflora

Cryptantha nubigena

Cymopteris cinerarius

Draba lemmonii (var. incrassata)

D. oligosperma var. subsessilis

D. stenoloba var. ramosa
Erigeron petiolaris

E. pygmaeus
Eriogonum rosense
Gentianopsis hohpetala
Gentiana newberryi
Geum canescens

Galium hypotrichium ssp. hypotrichium

Haplopappus apargioides

Hieracium hcrridum

Ivesia lycopodioides

I. purpurascens ssp. congdonis

Juniperus occidentalis ssp. australis

Kalmia microphylla

Ledum glandulosum var. californica

Leucophysalis nana

Lupinus andersonii

L. caudatus ssp. montigenus

L. confertus

L. hypolasius

L. meionanthus

L. caudatus ssp. montigenus

L. sellulus var. lobbii

L. tegeticulatus (breweri bryoides)

Luzula divaricate

Mimulus coccineus

Penstemon davidsonii var. davidsonii

Pinus jeffreyi

P. monticola

Polygonum douglasii var. latifolia

Raillardella argentea

Rhamnus rubra ssp. rubra

Scirpus dementis

Senecio fremontii var. occidentalis

S. scorzonella

Sisyrinchium idahoense var. occidentale

Symphoricarpos parishii

Trifolium andersonii ssp. andersonii

Additionally, work done by Goodrich
(1981) in central Nevada has shown that sev-
eral other Pacific cordilleran plants extend

their ranges far into the Intermountain Re-
gion. These plants, also found extending east-
ward within the Walker River basin, include
Artemisia rothrockii, Carex helleri, Silene sar-
gentii, Astragalus whitneyi (see Reveal 1979)
and Astragalus purshii var. tinctus (see Bam-
eby 1964). Silene sargentii has been listed as
endemic to the Sierra Nevada. However, the
type collection, as given by Hitchcock and
Maguire (1947), is from Table Mountain of
the Monitor Range in central Nevada.

The eastward distribution of these Sierran
plants may be enhanced by the very eastern
exposures of the Sierran granodiorites. These
granites can be found as far east as the Was-
suk Range (Bateman 1967). However, the
most likely factor enhancing the migration of
these Sierran plants is the abundance of mon-
tane and alpine habitats found eastward into
the Intermountain Region. The floristic com-
ponents of the Sweetwater Moujitains point
to this eastward migration.

The Sweetwaters, situated between the
Sierra and Intermountain floristic regions, are
regarded as belonging to the Intermountain
flora (Cronquist et al. 1972). Sierra conifer
forests are extensive on the Sweetwaters, es-
pecially toward the southern end. The tim-
berline vegetation is dominated solely by
Pinus albicaulis, a Sierran characteristic.

Using the information provided in Table 1,
Sorenson's index of similarity (Billings 1978)
between the Sierra and Sweetwaters can be
determined with regard to floristic elements
in both the alpine and the montane conifer
forests. These are calculated to be 90 and 93
percent, respectively. This indicates the
Sweetwater Mountains to be more affiliated,
floristically, with the Sierra than previously
thought.

However, it might be considered that the
east slope of the Sierra, within the Walker
River drainage, being influenced by Inter-
mountain vegetation, is bound to yield a high
index of similarity to the Sweetwaters. In
other words, the whole of the upper Walker
could be considered "Intermountain" with
regard to vegetational composition. This is
reinforced by the fact that the Walker River
portion of the Sierra does lack typical "un-
derstory" brush vegetation that is found just
to the north in the Carson River drainage.
Arctostaphylos patula, Ceanothus cordulatus

January 1983

Lavin: Walker River Floristics

and C. prostratus have not been found here
(or are at least not abundant) and A. neva-
densis was foujid only twice in very small
populations. Instead, the dominant brush
cover consists of Purshia tridentata, Cea-
nothus velutinus, Artemisia tridentata ssp.
vaseyana, and Symphoricarpos oreophilus.

In comparing all the upper Walker River
flora with the flora of central Nevada (Good-
rich 1981), both floras being just about equal
in number of taxa, approximately a 50 per-
cent similarity can be determined. The
Sweetwaters could, therefore, be easily con-
sidered as part of the east slope Sierran flora.
Aside from the high indices of similarity, the
Sweetwaters are dominated by many Sierra
or Pacific cordillera plant species; Lupinus
hypolasius, Chrysothamnus parryi ssp. mon-
ocephalus, and Draba oligosperma var. sub-
sessilis at the highest elevations, and Pinus
jeffreyi and P. contorta var. murrayana at the
lower elevations.

Acknowledgments

This study was funded by the U.S. Forest
Service, Toiyabe National Forest. I am very
grateful to them for this and for their cooper-
ation. Thanks to Sue Sullivan, U.S. Forest
Service, the checklist (Table 1) and its trans-
fer to Brigham Young University Press was
made possible. For their help with identi-
fication of plant specimens, I am indebted to
Warren Wagner (Botrychium), Lincoln Con-
stance (Apiaceae), Ted Barkley (Senecio), Al-
mut Jones (Aster), Guy Nesom (Erigeron),
Gerald Ownbey (Cirsium and Argemone),
Reed Rollins (Brassicaceae), Arthur Cronquist
(miscellaneous), Alfred Schuyler (aquatic
plants), Gary Wallace (Pyrola), Rupert Barn-
eby (Fabaceae), Richard Halse (Phacelia),
Duane Atwood (Phacelia), Douglass Hender-
son (Sisyrinchium), Paul Fryxell (Malvaceae),
David Boufford (Circaea), Alva Day (Polemo-
niaceae), Lauramae Dempster (Galium), Har-
lan Lewis (Gayophytum), Lawrence Heckard
(Castilleja, Cordylanthus, Orthocarpus), Fred-
rick Meyer (Valeriana), John T. Howell
(Carex and miscellaneous), Margaret Wil-
liams (miscellaneous), Arnold Tiehm (mis-
cellaneous), and Ken Genz (miscellaneous). I
am especially grateful to John Thomas How-
ell, Margaret Williams and Arnold Tiehm for

sharing with me their intimate knowledge of
the Intermountain and Sierran floras, and to
the curators at NY and CAS for allowing ac-
cess to their herbaria.

Literature Cited

Barneby, R. C. 1964. Atlas of North American Astra-
galus. New York Botanical Gardens. 2 Vols.

Bateman, p. C. 1967. The Sierra Nevada Batholith. Sci-
ence 158: 1407-1417.

Bell, K., and R. Johnson. 1980. Alpine flora of the
Wassuk Range, Mineral County, Nevada.
Madroiio 27: 25-35.

Billings, W. D. 1951. Vegetation zonation in the Great
Basin of western North America. Pages 101-122
in Les Bases e regeneration do la des zones
arides. International Colloquium. Intemat. Union
of Bio. Sci. Ser. B.9.

1978. Alpine phytogeography across the Great

Basin. Great Basin Nat. Mem. 2: 105-117.

Chabot, B. F., and W. D. Billings. 1972. Origins and
ecology of the Sierran alpine flora and vegeta-
tion. Ecol. Monogr. 42: 163-199.

Cox, B. J. 1972. Biosystematics of Lupinus lepidus-L.
caespitosus complex. Unpublished dissertation.
Univ. of Missouri, Columbia. 443 pp.

Cronquist, A., A. Holmgren, N. Holmgren, and J.
Reveal. 1972. Intermountain flora. Vol. 1. Haf-
ner. New York. 584 pp.

Davis, B. 1979. Draft: Plant list and keys. East Side Cen-
tral Sierra and associated desert Ranges. U.S.
Forest Service, Toiyabe National Forest, Reno,
Nevada. 124 pp.

Dempster, L. T., and F. Ehrendorfer. 1965. Evolution
of the Galium multiflorum complex in western
North America. II. Critical taxonomic revision.
Brittonia 17: 289-334.

Goodrich, S. 1981. A floristic study of central Nevada.
Unpublished thesis, Brigham Young Univ. 400
pp.

Halse, R. R. 1981. Taxonomy of Phacelia Sect. Miltitzia
(Hydrophyllaceae) Madrono 28: 121-132.

Hardham, C. B., and G. H. True. 1972. Malacothrix tor-
reyi (Compositae) in California. Madrono 21: 535.

Harper, K. T., D. C. Freeman, W. K. Ostler, and L.
C. Klikoff. 1978. The flora of the Great Basin
mountain ranges: diversity, sources and dispersal
ecology. Great Basin Nat. Mem. 2: 81-103.

HiNTON, W. F. 1975. Systematics of the Calyptridium
umbellatum complex (Portulacaceae). Brittonia
27: 197-208.

Hitchcock, C. L., and B. Maguire. 1947. A revision of
the North American species of Silene. Univ. of
Washington Pub. Biol. 13: 1-73.

Kartesz, J., AND R. Kartesz. 1980. A synonymized
checklist of the vascular flora of the United
States, Canada and Greenland. Univ. of North
Carolina Press. Chapel Hill. 500 pp.

Lavin, M. 1981. Floristics of the headwaters of the
Walker River, California and Nevada. Unpub-
lished thesis. Univ. of Nevada, Reno. 141 pp.

Major, J., and D. W. Taylor. 1977. Alpine. Pages
601-675 in M. G. Barbour and J. Major, eds..

Great Basin Naturalist

Vol. 43, No. 1

Terrestrial vegetation of California. John Wiley
Interscience, New York.

Messick, T. 1982. The flora and phytogeography of the
Bodie Hills of Mono County, CA and Mineral
County, NV. Unpublished thesis, Humboldt State
Univ.

MuNZ, P. A., AND D. Keck. 1968. A California flora.
Univ. of California Press. Berkeley. 1681 pp. and
supplement.

Reveal, J. L. 1968. Notes of Eriogonum— IV. A revision
of the Eriogonum deflexum complex. Brittonia
20: 13-33.

1979. Biogeography of the Intermountain Re-
gion. Mentzelia 4: 1-92.

Reveal, J. L., and B. Ertter. 1980. Noteworthy collec-
tions. Madrono 27: 142.

Rollins, R. C. 1982. Studies on Arahis (Cruciferae) of
Western North America II. Contr. Gray Her-
barium of Harvard. No. 212:111-114.

RuNDEL, P. W., D. J. Parsons, and D. T. Gordon. 1977.
Montane and subalpine vegetation of the Sierra
Nevada and Cascade ranges. Pages 559-599 in M.
G. Barbour and J. Major, eds., Terrestrial vegeta-
tion of California. John Wiley Interscience, New
York.

Sharsmith, C. W. 1940. A contribution to the history of
the alpine flora of the Sierra Nevada. Unpub-
lished dissertation. Univ. of California, Berkeley.
274 pp.

Strother, J. L. 1974. Taxonomy of Tetradymia (Com-
positae: Senecioeae). Brittonia 26: 177-202.

Taylor, D. W. 1976. Disjunction of Great Basin plants
in the northern Sierra Nevada. Madrono 23:
301-310.

1977. Floristic relationships along the Cascade-

Sierran axis. Amer. Midi. Natur. 97: 333-349.

Thorne, R. F. 1982. The desert and other transmontane
plant communities of southern California. Aliso
10:219-257.

Table 1. Checklist of the vascular flora of the upper Walker River. For the columns under the heading of
"AREA":N = Sierra Nevada; S= Sweetwater Mountains; W = Wellington Hills; M = Masonic Hills, Bodie Hills, Pine
Grove Hills, and the southwest slopes of the Pine Nut Range. For the columns under the heading of "HABITAT":
1 = riparian; 2= low elevation sagebrush-grass zone; 3=pinyon-juniper woodland; 4 = high elevation sagebrush -grass
zone; 5 = Jeffrey pine-white fir forests; 6 = red fir forests; 7 = lodgepole pine forests; 8 = whitebark pine forests; 9= al-
pine zone (see Billings 1951, Lavin 1981, for a general description of these). An X indicates the plant has been ob-
served in the field or by herbarium specimen; an O indicates the plant has a probable distribution within the area
due to information obtained in the literature.

AREAS

HABITATS

N S W M

123456789

Adiantaceae - pteridophyta
Aspidotis

densa (Brack.) Lellinger
Cheilanthes

gracillima D.C. Eat.
Cryptogramma

crispa (L.) R. Br. ex Hook.

ssp. acrostichoides (R. Br.) Hulten
Pellaea

breweri D.C. Eat.

bridgesii Hook.

Aspleniaceae
Athyrium

distentifolium Tausch ex Opiz

var. americanum (Butters) Boivin
Cystopteris

fragilis (L.) Bemh.
Woodsia

oregana D.C. Eat.

scopulina D.C. Eat.

Dennstaedtiaceae
Pteridium

aquilinum (L.) Kuhn

var. pubescens Underwood

Equisetaceae
Equisetum

arvense L.

hyemale L.

var. affine (Engelm.) A. A. Eat.

laevigatum A. Braun

X X

X X
X X

X X

X X X X X

X
X X

X XXX

X XX

X X X o

X XX

XXXX X XXX

January 1983

Lavin: Walker River Floristics

Table 1 continued.

AREAS

HABITATS

N S W M

12 3 4 5

7 8 9

ISOETACEAE

Isoetes

bolanderi Engelm.
var. bolanderi

Marsileaceae
Marsilea

vestita Hook. & Grev.

Ophioglossaceae
Botrychium

lunaria (L.) Sw.

var. minganese (Victorin) Dole

simplex E. Hitchcock

Selaginellaceae
Selaginella

watsonii Underwood

CUPRESSACEAE - PINOPHYTA

Juniperus
communis L.
occidentalis Hook.

ssp. australis Vasek
osteosperma (Torr.) Little

Ephedraceae
Ephedra

nevadensis S. Wats.

viridis Coville

Pinaceae
Abies

concoloT (Gord. & Glend.) Hildebr.

magnifica A. Murr.
Pinus

albicaulis Engelm.

contorta Dougl. ex Loud.

var. murrayana (Grev. & Balf.) Engelm.

flexilis James

jeffreyi Grev. & Balf.

monophylla Torr. & Frem.

monticola Dougl. ex D. Don
Tsuga

mertensiana (Bong.) Carr.

AcERACEAE - ANTHOPHYTA

Acer

glabrum Torr.

var. torreyi (Greene) Smiley

Alismataceae
Sagittaria
cuneata Sheldon

Amaranthaceae
Amaranthus

alhus L.

blitoides S. Wats.

retroflexus L.

Apiaceae
Angelica

breweri Gray

lineariloba Gray

X X

XX o

X X X X

X X

X X
X X X X
X X

XX X

XXX

X X

XXX

X O X X X X X
O XX

XXX

X XX

XXX

o o o

X X X X

X
X X

X X

100

Great Basin Naturalist

Vol. 43, No. 1

Table 1 continued.

AREAS

N S W M

HABITATS

123456789

Berula

erecta (Huds.) Coville
Cicuta

douglasii (DC.) Coult. & Rose
Conium

maculatum L.
Cymopterus
cinerarius Gray
globosus (S. Wats.) S. Wats.
panamintensis Coult. & Rose
var. panamintensis
Heracleum

lanatum Michx.
Ligusticum

grayii Coult. & Rose
Lonuitiiim

dissectum (Nutt.) M. & C.

var. multifidum (Nutt.) M. & C.
foeniculaceum (Nutt.) Coult. & Rose

ssp. macdougalii (Coult. & Rose) Theobald
nevadense (S. Wats.) Coult. & Rose
var. nevadense

var. parishii (Coult. & Rose) Jepson
plummerae (Coult. & Rose) Coult. & Rose
var. sonnei (Coult. & Rose) Jepson
Osmorhiza

chilensis Hook. & Am.
occidentalis (Torr. & Gray) Torr.
Perideridia

bolanderi (Gray) A. Nels. & J.F. Macbride

ssp. bolanderi
lemmonii (Coult. & Rose) Chuang & Const.
parishii (Coult. & Rose) A. Nels.

ssp. latifolia (Gray) Chuang & Const.
Podistera

nevadensis (Gray) S. Wats.
Pteryxia

terebinthina (Hook.) Coult. & Rose

var. californica (Coult. & Rose) Mathias
Sphenosciadium
capitellatum Gray

Apocynaceae
Apocynum

androsaemifolium L.

ssp. pumilum (Gray) Boivin

X medium Greene

ASCLEPIADACEAE

Asclepias

cryptoceras S. Wats.

ssp. cryptoceras

ssp. davisii (Woods.) Woods.
fascicularis Dene.
speciosa Torr.

ASTERACEAE

Achillea

millefolium L.

var. alpicola (Rydb.) Garrett
var. lanulosa (Nutt.) Piper

X
X X

X X

X
X

X X

X o

X X

X X X X X

X X X X
X X

X X X X
X X X X

X X

XXXX XXX X
XXXX X xxxxxx

XXX
O X

XXX

X X

XXXX

XXX

XX X
XXXX

X X
XXX

XXX

January 1983

Lavin: Walker River Floristics

101

Table 1 continued.

AREAS

HABITATS

N S

X X

Agoseris

aurantiaca (Hook.) Greene
elata (Nutt.) Greene
glauca (Pursh) Raf.

var. nwnticola (Greene) Cronquist
var. laciniata (D.C. Eat.) Smiley
Ambrosia

acanthicarpa Hook.
Anisocoma

acaulis Torr. & Gray
Antennaria

alpina (L.) Gaertn.

var. media (Greene) Jepson
dimorpha (Nutt.) Torr. & Gray
micTophylla Rydb.
umbrineUa Rydb.
Arnica

chamissonis Less.

ssp. foliosa (Nutt.) Maguire

var. andina (Nutt.) Ediger & Barkley
ssp. foliosa (Nutt.) Maguire
var. incana (Gray) Hulten
cordifolia Hook.

var. cordifolia
diversifolia Greene
longifolia D.C. Eat.
mollis Hook.
nevadensis Gray
parry i Gray

var. sonnei (Greene) Cronquist
sororia Greene
Artemisia

arbuscula Nutt.
cana Pursh
dougtasiana Bess.
dracunculus L.
ludoviciana Nutt.

ssp. incompta (Nutt.) Keck
ssp. ludoviciana
norvegica Fries

var. saxitalis (Bess.) Hook.
nova A. Nels.
rothrockii Gray
spinescens D.C. Eat.
tridentata Nutt.

ssp. vaseyana (Rydb.) Beetle
ssp. wyomingensis Beetle & Young
ssp. tridentata
Aster

ascendens Lindl. in Hook.
alpigenus (Torr. & Gray) Gray
ssp. andersonii (Gray) Onno
campestris Nutt.

var. bloomeri Gray
eatonii (Gray) T.J. Howell
integrifolius Nutt.
occidentalis (Nutt.) Torr. & Gray

var. occidentalis
scopuhrum Gray

X X

102

Great Basin Naturalist

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Table 1 continued.

AREAS

habitats

N S W M

12 3 4 5 6 7

Bahamorhiza

hookeri (Hook.) Nutt.
sagittata (Pursh) Nutt.
Bidens

cemua L.
Brickellia
grandiflora (Hook.) Nutt.
var. petiolaris Gray
microphylla (Nutt.) Gray
oblongifolia Nutt.

var. linifolia (D.C. Eat.) B.L. Robins.
Chaenactis

alpigerm C.W. Sharsmith
douglasii (Hook.) Hook. & Am.

var. rubricaulis (Rydb.) Ferris
nevadensis (Kellogg) Gray
xantiana Gray
Chrysothamnus

nanseosus (Pall.) Britton

ssp. consimilis (Greene) Hall & Clem,
ssp. albicaulis (Nutt.) Hall & Clem,
ssp. hololeucus (Gray) Hall & Clem.
parryi (Gray) Greene

ssp. monocephalus (Nels. & Kenn.) Hall & Clem,
ssp. nevadensis (Gray) Hall & Clem.
viscidiflorus (Hook.) Nutt.

ssp. puberulus (D.C. Eat.) Hall & Clem,
ssp. viscidiflorus
Cichorium

intybus L.
Cirsium

andersonii (Gray) Petrak
congdonii Moore & Frankton
eatonii (Gray) B.L. Robins.
pastoris J.T. Howell
tioganum (Congd.) Petrak
utahense Petrak
vulgare (Savi) Tenore
Conyza

canadensis (L.) Cronquist

var. glabrata (Gray) Cronquist
Crepis

acuminata Nutt.
ssp. acuminata
intermedia Gray
modocensis Greene

ssp. subacaulis (Kell.) Babcock & Stebbins
nana Richards.

ssp. ramosa Babcock & Stebbins
ssp. nana
occidentalis Nutt.

ssp. pumila (Rydb.) Babcock & Stebbins
ssp. occidentalis

ssp. conjuncta (Jepson) Babcock & Stebbins
runcinata (James) Torr. & Gray
ssp. hallii Babcock & Stebbins
Dugaldia

hoopesii (Gray) Rydb.

XXX
X X X X

X X
XXX

XX X

X
X X

X X

X
X

X X

January 1983

Lavin: Walker River Floristics

103

Table 1 continued.

AREAS

HABITATS

NSWM 123456789

Eatonella

nivea (D.C. Eat.) Gray
Erigeron

aphanactis (Gray) Greene

var. aphanactis
barbellulatus Greene
bloomeri Gray

var. bhameri
breweri Gray
var. breweri

var. porphyretictis (M.E. Jones) Cronquist
clokeyi Cronquist
compositus Pursh

var. glabratus Macoun
coulteri Porter
divergens Torr. & Gray
eatonii Gray

ssp. plantagineus (Greene) Cronquist
lonchophyllus Hook.
nevadincola Blake
peregrinus (Pursh) Greene

ssp. callianthemus (Greene) Cronquist
var. angustifolius (Gray) Cronquist
ssp. callianthemus (Greene) Cronquist
var. hirsutus Cronquist
petiolaris Greene
pygmaeus (Gray) Greene
tener (Gray) Gray
vagus Payson
Eriophyllum

lanatum (Pursh) Forbes

var. integrifolium (Hook.) Smiley
Eupatorium

occidentale Hook.
Glyptopleura

marginata D.C. Eat.
Gnaphalium
palustre Nutt.
microcephalum Nutt.

var. thermale (E. Nels.) Cronquist
Grindelia

squarrosa (Pursh) Ehinal
var. squarrosa
Gutierrezia

sarothrae (Pursh) Britt. & Rusby
Haplopappus

acaulis (Nutt.) Gray
apargioides Gray
bloomeri (Hook.) Gray
lanceolatus (Hook.) Torr. & Gray

var. lanceolatus
macronema Gray
racemosus (Nutt.) Torr.

ssp. glomeratus (Nutt.) Hall
suffruticosus (Nutt.) Gray
uniflorus (Hook.) Torr. & Gray
ssp. uniflorus

X X

X
X

X X

X
X

104

Great Basin Naturalist

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Table 1 continued.

AREAS

habitats

NSW

X X

XXX

X
X

X X

X X
X X

X
X

Helianthus

annuus L.
Heterotheca

breweri AGray) Shinners
Hieracium

albiflorum Hook.

gracile Hook,
var. gracile

horridum Fries
Hulsea

algida Gray

heterochroma Gray
Hymenopappus

filifolius Hook.

var. nanus (Rydb.) B.L. Turner
Hymenoxys

coaperi (Gray) Cockerell

var. canescens (D.C. Eat.) Parker
Iva

axillaris Pursh
Lactuca

serriola L.

tartarica (L.) C.A. May

ssp. pulchella (Pursh) Stebbins
Layia

glandulosa (Hook.) Hook. & Am.
ssp. glandulosa
Leucanthemum

vulgare Lam.
Machaeranthera

canescens (Pursh) Gray

shastensis Gray

var. montana (Greene) Cronquist & Keck
var. gossophylla (Piper) Cronquist & Keck
Madia

ghmerata Hook.

gracilis (Sm.) Keck
Malacothrix

sonchoides (Nutt.) Torr. & Gray
var. torreyi (Gray) E. WiUiams
Microseris

lindleyi (DC.) Gray
Nothocalais

alpestris (Gray) Chambers
Psihcarphus

brevissimus Nutt.
var. brevissimus
Raillardella

argentea (Gray) Gray

scaposa (Gray) Gray
Senecio

canus Hook.

cymbalarioides Beuk

fremontii Torr. & Gray
var. occidentalis Gray

hydrophilus Nutt.

integerrimus Nutt.

var. exaltatus (Nutt.) Cronquist

multilobatus Torr. & Gray ex Gray

X X

o o o

January 1983

Lavin: Walker River Floristics

105

Table 1 continued.

AREAS

HABITATS

X
X

o
o

X
X

pattersonensis Hoover

scorzonelUi Greene

serra Hook,
var. serra

spartioides Torr. & Gray

streptanthifolius Greene

triangularis Hook.

wemeriifolius (Gray) Gray
Solidago

canadensis L.

var. salebrosa (Piper) M.E. Jones

multiradiata Ait.

occidentalis (Nutt.) Torr. & Gray

spectahilis (D.C. Eat.) Gray
Sonchus

asper (L.) Hill
Sphaeromeria

cana (D.C. Eat.) Heller

potentilloides (Gray) Heller
var. potentilloides
Stephanomeria

exigua Nutt.

ssp. coronaria (Greene) Gottlieb
ssp. exigua

spinosa (Nutt.) S. Tomb
Taraxacum

officinale Weber
Tetradymia

axillaris A. Nels.

var. longispina ( M.E. Jones) Strother

canescens DC.

glahrata Torr. & Gray

spinosa Hook. & Am.

tetrameres (Blake) Strother
Townsendia

condensata Parry ex Gray

scapigera D.C. Eat.
Tragopogon

dubius Scop.
Wyethia

mollis Gray

Betulaceae
Alnus

incana (L.) Moench

ssp. tenuifolia (Nutt.) Breittung

BORAGINACEAE

Amsinckia

tessellata Gray
Cryptantha

affinis (Gray) Greene

circumscissa (Hook. & Am.) Johnston

var. hispida (J.F. Macbr.) Johnston

var. circumscissa
confertiflora (Greene) Payson
echinella Greene
flavoculata (A. Nels.) Payson
glomeriflora Greene
humilis (Gray) Payson

var. humilis

o o

X X

106

Great Basin Naturalist

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Table 1 continued.

AREAS

habitats

N S W M

123456789

jamesii (Torr.) Payson

var. abortiva (Greene) Payson
nuhigena (Greene) Payson
pterocarya (Torr.) Greene

var. pterocarya

torreyana (Gray) Greene

var. torreyana

var. pumila (Heller) Johnston
watsonii (Gray) Greene
sp. nov.
Hackelia
floribunda (Lehm.) Johnston
micrantha (Eastw.) J.L. Gentry
Lappula

redowskii (Homem.) Greene
var. redowskii
Mertensia

oblongifolia (Nutt.) G. Don

var. nevadensis (A. Nels.) L.O. Williams
Pectocarya

setosa Gray
Plagiobothrys
hispidus Gray
jonesii Gray
kingii (S. Wats.) Gray

var. harknessii (Greene) Jepson
var. kingii
scouleri (Hook. & Am.) Johnston
var. scouleri
Tiquilia

nuttallii (Benth. ex Hook.) Richards.

Brassicaceae
Anelsonia

eurycarpa (Gray) Macbr. & Payson
Arabis

bodiensis Rollins
cobrensis M.E. Jones
davidsonii Greene
divaricarpa A. Nels.
drummondii Gray
fernaldiana Rollins

var. stylosa (S. Wats.) Rollins
glabra (L.) Bemh.
hirsuta (L.) Scop.

var. glabrata Torr. & Gray
holboellii Homem.

var. penduhcarpa (A. Nels.) Rollins

var. pinetorum (Tidestrom) Rollins

var. retrofracta (Grahm.) Rydb.

var. holboellii
inyoensis Rollins
lemmonii S. Wats.

var. lemmonii

var. depauperata (A. Nels. & Kenn.) Rollins
lyallii S. Wats.

var. lyallii
platysperma Gray

var. platysperma

var. howellii (S. Wats.) Jepson

X
X

X X

X X
X

X X

XXX

X X

XXX

X X

X
X

X X

January 1983

Table 1 continued.

Lavin: Walker River Floristics

107

puberula Nutt.

pulchra M.E. Jones ex S. Wats,
var. pulchra
var. gracilis M.E. Jones
sparsiflora Nutt.

var. subvillosa (S. Wats.) Rollins
var. sparsiflora
Barbarea

orthoceras Ledeb.

var. dolichocarpa Fern,
var. orthoceras
Capsella

bursa-pastoris (L.) Medic.
Cardamine
breweri S. Wats,
var. breweri
Cardaria

pubescens (C.A. Mey) Jarmolenko
Caulanthus

pilosus S. Wats.
Descurainia

califomica (Gray) O.K. Schulz
pinnata (Walt.) Britt.

ssp. filipes (Gray) Detling
ssp. halictorum (Cockerell) Detling
ssp. menziesii (DC.) Detling
richardsonii (Sweet) O.E. Schulz
ssp. incisa (Engelm.) Detling
ssp. viscosa (Rydb.) Detling
sophia (L.) Webb ex Prantl
Draba

albertina Greene
breweri S. Wats.
densifolia Nutt.
douglasii Gray

var. crockeri (Lemmon) C.L. Hitchc.
lemmonii S. Wats,
var. lemmonii
var. incrassata Rollins
oligosperma Hook.

var. subsessilis (S. Wats.) O.E. Schulz
var. oligosperma
paysonii J.F. Macbride
quadricostata Rollins
stenoloba Ledeb.

var. ramosa C.L. Hitchcock
Erysimum

argillosum (Greene) Rydb.
capitatum (Dougl.) Greene
perenne (S.Wats, ex Coville) Abrams
repandum L.
Hymenolobus

procumbens (L.) Nutt. ex Torr. & Gray
Lepidium

lasiocarpum Nutt.
perfoliatum L.
virginicum L.

var. pubescens (Greene) C.L. Hitchc.

AREAS

]

HABITATS

)

N S W M

1 2

7 8 9

X X X X

X X

X X X X

XXXX X X X X
XXXX X X X XXX

XXXX

XXX

XXXX X xxxxxxx
o o

x x

XXXX X XXXX

XXXX XX X X

X X
XXXX X X

o o

XX O O O X X

XXXX xxxxxxxx

X X O X XXX

X X O XX

XXXX XXX XXX

X XX

X X
X X

XXX
X X

X
X

X X

XXX

XXXX XXX X
XXX X XXX

XX XX

X XX
XXXX XXXX

XXXX XXXX

108

Great Basin Naturalist

Vol. 43, No. 1

Table 1 continued.

AREAS

habitats

N S W M

123456789

Lesquerella

kingii S. Wats,
var. kingii
Nasturtium

officinale R. Br.
Phoenicaulis

cheiranthoides Nutt.
Polyctenium

fremontii (S. Wats.) Greene
Rorippa

curvisiliqua (Hook.) Bess, ex Britton

teres (Michx.) R. Stuckey
Sisymbrium

altissimum L.
Stanleya

pinnata (Pursh) Britton
var. pinnata
Streptanthus

cordatus Nutt. ex Torr. & Gray

oliganthus Rollins

tortuosus Kellogg

var. orbiculatus (Greene) Hall
Thelypodium

crispum Greene ex Payson

integrifolium (Nutt.) Endl.
ssp. complanatum Al-Shehbaz

laciniatum (Hook.) Endl.

Cactaceae
Opuntia

polyacantha Haw.

var. rufispina (Engelm. & Bigelow) Benson

pulchella Engelm.

Callitrichaceae
CalUtriche

heterophylla Pursh emend. Darby
var. bolanderi (Hegelm.) Fassett.

verna L. emend. Kuetz.

Campanulaceae
Nemacladus

rigidus Curran
Porterella

carnosula (Hook. & Am.) Torr.

Capparidaceae
Cleomella

hillmanii A. Nels.

parviflora Gray

Caprifoliaceae
Lonicera

involucrata (Rich.) Banks ex Spreng.
Sambucus

caerulea Raf.

racemosa L.

ssp. pubens (Michx.) House

var. microbotrys (Rydb.) Kearney & Peebles
Symphoricarpos

longifloTus Gray

X X

X
X

X X

X
X

X X X X
X X

X X X X XX

0
X

X
X

January 1983

Lavin: Walker River Floristics

109

Table 1 continued.

oreophilus Gray

var. oreophilus
parishii Rydb.

Caryophyllaceae
Arenaria

aculeate S. Wats.
Cerastium
alpinum L.
vulgatum L.
Minuartia

nuttallii (Pax) Briq.

ssp. fragilis (Maguire & Holmgren) McNeill
ssp. gracilis (B.L. Robins.) McNeill
obtusiloba (Rydb.) House
rossii (R. Br.) Graebn.
rubella (Wahlenb.) Hiem
Pseudostellaria

jamesiana (Torr.) Weber & Hartman
Sagina

saginoides (L.) Karst.
Saponaria

officinalis L.
Silene

bemardina S. Wats,
ssp. maguirei Bocquet

var. maguirei
ssp. bemardina
menziesii Hook.

ssp. dorrii (Kellogg) C.L. Hitchc.& Maguire
nuda (S. Wats.) C.L. Hitchc. & Maguire

ssp. insectivora (Henders.) C.L. Hitchc. & Maguire
sargentii S. Wats.
Stellaria
crispa Cham. & Schlecht.
longipes Goldie
umbellata Turcz. ex Kar. & Kir.

Chenopodiaceae
Atriplex

argentea Nutt.
ssp. argentea
canescens (Pursh) Nutt.

ssp. canescens
confertifolia (Torr. & Frem.) S. Wats.
heterosperma Bunge
patula L.

ssp. hastata (L.) H. & S.
rosea L.
Bassia

hyssopifolia (Pallas) Kuntze
Ceratoides

lanata (Pursh) J.T. Howell
var. lanata
Chenopodium
album L.
atrovirens Rydb.
botrys L.
dessiccatum A. Nels.

var. leptophylloides (J. Murr) H.A. Wahl.
var. dessiccatum

AREAS

HABITATS

NSW

4 5

XXX

X X

XXX

X X

XXX

O X

X X

X ox

X X

X x

X X

X X
X X
X

X
X
X

X
X

X
X
X

X X

X X X X

X X

XXX
X O X X
X X

X X

X o o o

110

Great Basin Naturalist

Vol. 43, No. 1

Table 1 continued.

AREAS

habitats

fremontii S. Wats.

leptophyllum (Moq.) Nutt. ex S. Wats.

overi Aellen
Grayia

spinosa (Hook.) Moq.
Halogeton

glomeratus (Stephan ex Bieb) C.A. Mey.
Kochia

scoparia (L.) Schrad.
Monolepis

nuttalliana (Roem. & Schult.) Greene

spathulata Gray
Sakola

iberica Sennen & Pau
Sarcobatus

vermiculatus (Hook.) Torr.
var. vermiculatus
Sueda

occidentalis S. Wats.

Clusiaceae
Hypericum
formosum H.B.K.

ssp. scouleri (Hook.) C.L. Hitchc.

CONVOLVULACEAE

Calystegia

polymorpha (Greene) Munz
Convolvulus

arvensis L.

CORNACEAE

Cornus
sericea L.
ssp. sericea

Crassulaceae
Sedum

integrifolium Coult. & A. Nels.

lanceolatum (Nutt.) Britton & Rose

obtusatum Gray

Crossosomataceae
Forsellesia

nevadensis (Gray) Greene

CUSCUTACEAE

Cuscuta

suksdorfii Yunker

var. subpedicellata Yunker

Cyperaceae
Carex

abrupta Mackenzie

aquatilis Wahlenb.

athrostachya Olney

aurea Nutt.

brevipes W. Boott.

canescens L.

capitata L.

congdonii Bailey

disperma Dewey

douglasii Boott

1 2

8 9

X
X

X
X
X

X
X

X X X X

XXX
X

X X

XXX

XX o

X X
X

X X

XXX
X

XXX

X X

O X X
X XXX

X X

X o

X X

o o

X X

O X

X X

January 1983

Lavin: Walker River Floristics

111

Table 1 continued.

exserta Mackenzie
festivelhi Mackenzie
fissuricola Mackenzie
haydeniana Olney
helleri Mackenzie
heteroneura W. Boott

var. epapillosa (Mackenzie) F.J. Herm.

var. heteroneura
hoodii Boott
jepsonii J.T. Howell
jonesii Bailey
lanuginosa Michx.
leporinella Mackenzie
luzulifolia W. Boott
microptera Mackenzie
nebraskensis Dewey
nervina Bailey
nigricans C.A. Mey.
pachystachya Cham, ex Steudel.
phaeocephala Piper
praegracilis W. Boott
rossii Boott ex Hook.
rostrata Stokes ex With.
scopulorum T. H. Holm.

var. bracteosa (Bailey) F.J. Herm.

var. scopulorum
simulate Mackenzie
specifica Bailey
spectabilis Dewey
straminiformis Bailey
subnigricans Stacey
tahoensis Smiley
vallicola Dewey
vemacula Bailey
vesicaria L.
Eleocharis

engelmannii Steud.
palustris (L.) Roemer & Schultes
pauciflora (Lightf.) Link
Eriophorum

crinigerum (Gray) Beetle
Scirpus
acutus Muhl. ex Bigelow
americanus Pers.
dementis M.E. Jones
microcarpus Presl.
nevadensis S. Wats.
pungens Vahl.

Elaeagnaceae
Elaeagnus

angustifolia L.
Shepherdia

argentea (Pursh) Nutt.

Elatinaceae
Elatine

rubella Rydb.

AREAS

HABITATS

N S

1 2

X X

o o

X X

0 o

X X

X X
X

X
X

X X

o o

112

Table 1 continued.

Great Basin Naturalist

Vol. 43, No. 1

AREAS

habitats

NSWM 12 3 456789

Ericaceae
Arctostaphylos

tievadensis Gray
Cassiape

mertensiana (Bong.) D. Don
Kalmia

microphylla (Hook.) Heller
var. microphylla
Ledum

glandulosum Nutt.

var. californicum (Kellogg) C.L. Hitchc.
Orthilia

secunda (L.) House
ssp. secunda
Phyllodoce

breweri (Gray) Heller
Pterospora

andromedea Nutt.
Pyrola

californica Krisa

dentata Sm.

var. dentata

minor L.
Sarcodes

sanguinea Torr.
Vaccinium

caespitosum Michx.

var. paludicola (Camp) Hulten

uliginosum L.

ssp. occidentale (Gray) Hulten

EUPHORBIACEAE

Chamaesyce

serpyllifolia (Pers.) Small

Fabaceae
Astragalus

andersonii Gray

bolanderi Gray

calycosus Torr. ex S. Wats.

var. calycosus
canadensis L.

var. brevidens (Gand.) Bameby
casei Gray
curvicarpus (Heller) J.F. Macbr.

var. curvicarpus
gibbsii Kellogg
iodanthus S. Wats.

var. iodanthus
johnnis-howellii Bameby
kentrophyta Gray

var. danaus Bameby
lentiginosus Dougl. ex Hook.

var. ineptus (Gray) M.E. Jones
malacus Gray
obscurus S. Wats.
oophorus S. Wats.

var. lavinii Bameby in ed.
platytropis Gray

X
X

X X X X

X X

XX X

XXX
X

X X

O O X X X

X X

XXX

X X

X X
XXX

X o

XXX
X

X o

XXX

XX X

X o

X X

X X X X X X X
X X
X

X X

January 1983

Lavin: Walker River Floristics

113

Table 1 continued.

AREAS

HABITATS

N S W M

123456789

purshii Dougl. ex Hook.

var. lectulus (S. Wats.) M.E. Jones
var. tinctus M.E. Jones
whitneyi Gray
var. whitneyi
Dalea

omata (Dougl. ex Hook.) Eat. & Wright
Lupinus

andersonii S. Wats.
arbustus Dougl. ex Lindl.

ssp. calcaratus (Kellogg) Dunn
brevicaulis S. Wats.
caudatus Kellogg
ssp. caudatus

ssp. montigenus (Heller) Hess & Dunn
confertus Kellogg
hypolasius Greene
lyallii Gray

var. danatis (Gray) S. Wats,
var. lyallii
meionanthus Gray
nevadensis Heller
polyphyllus Lindl.

ssp. superbus (Heller) Munz
sellulus Kellogg

var. lobbii (S. Wats.) Cox
var. sellulus
tegeticulatus Eastw
X inyoensis Heller
Medicago
lupulina L.
sativa L.
Melilotus
albus Medic.
officinalis (L.) Pallas
Oxytropis

parryi Gray
Robinia

pseudoacacia L.
Trifolium
andersonii Gray

ssp. beatleyae Gillett
ssp. andersonii
cyathiferum Lindl.
hybridum L.
longipes Nutt.
ssp. longipes
monanthum Gray

var. monanthum
pratense L.
productum Greene
repens L.

wormskjoldii Lehm.
Vicia
americana Muhl. ex Willd.
ssp. americana

Fagaceae
Castanopsis

sempervirens (Kellogg) Dudley

X X
X X

X X

X X
X

X X

X
X X

X X

XXX
XXX

XXX

X X X X X X
XX XX

X X

XXX

X X

XXX

X X

X X X X

114

Great Basin Naturalist

Vol. 43, No. 1

Table 1 continued.

AREAS

HABITATS

N S W M

123456789

Gentianaceae
Frasera

speciosa Dougl. ex Griseb.
Gentiana

calycosa Griseb.

newberryi Gray
Gentianopsis

holopetala (Gray) litis

Geraniaceae
Erodium

cicutarium (L.) L'Her.
Geranium

richardsonii Fisch. & Trautv.

Haloragidaceae
Hippuris
vulgaris L.

Hydrocharitaceae
Elodea

canadensis Michx.

Hydrophyllaceae
Hesperochiron

califomicus (Benth.) S. Wats.
Hydrophyllum

capitatum Dougl. ex Benth.
var. alpinum S. Wats.
Noma

aretioides (Hook. & Am.) Brand

densum Lemmon

rothrockii Gray
Nemophila

spatulata Coville
Phacelia

bicolor Torr. ex S. Wats.

curvipes Torr. ex S. Wats.

frigida Greene

hastata Dougl. ex Lehm.

ssp. compacta (Brand) Heckard

heterophylla Pursh

ssp. virgata (Greene) Heckard

humilis Torr. & Gray
var. humilis

hydrophylloides Torr. ex Gray

linearis (Pursh) Holz.

monoensis Halse

peirsoniana J.T. Howell

ramosissima Dougl. ex Lehm.
var. ramosissima

tetramera J.T. Howell
Tricardia

watsonii Torr. ex S. Wats.

Iridaceae
Iris

missouriensis Nutt.
Sisyrinchium

fialophilum Greene

idahoense Bickn.

var. occidentale (Bickn.) Henderson

X X

o o

X X

XXX

O X

X X

X X X X

X X

XXX

O X

XX X

X X X X

X X

XXX
XX o

XX X

XXX

X X X X X

XXX

O X

X X

XXX

X X

XXX

X X

X
X

X X

X
X

X XX

XXXX XXXXX XX

X X X X X X X

XXXX X X X XX

January 1983

Lavin: Walker River Floristics

115

Table 1 continued.

AREAS

HABITATS

N S W M

123456789

JUNCACEAE

Juncus

balticus Willd.

bryoides F.J. Herm.

bufonius L.

var. occidentalis F.J. Herm.
var. bufonius

capillaris F.J. Herm.

chlorocephalus Engelm.

drumniondii E. Mey.

ensifolius Wikstr.

var. montanus (Engelm.) C.L. Hitchcock

hemiendytus F.J. Herm.

kello^ii Engelm.

longistylis Torr.

macrandrus Coville

megaspermus F.J. Herm.

mertensianus Bong.

mexicanus Willd.

nevadensis S. Wats.

orthophyllus Coville

parry i Engelm.

saximontanus A. Nels.
Luzula

ditxiricata S. Wats.

multiflora (Retz.) Lej.

ssp. comosa (E. Mey.) Hulten

spicata (L.) DC.

subcongesta (S. Wats.) Jepson

JUNCAGINACEAE

Triglochin
concinna Burtt-Davy

var. debilis (M.E. Jones) J.T. Howell
maritima L.

Lamiaceae
Agastache

urticifolia (Benth.) Kuntze
Marrubium

vulgare L.
Mentha

arvensis L.

ssp. haplocalyx Briq.
Monardella

odoratissima Benth.

ssp. glauca (Greene) Epling
Salvia

dorrii (Kellogg) Abrams
var. dorrii
Trichostema

austromontana Lewis

Lemnaceae
Lemna

gibba L.

minuta H.B.K.

trisulca L.

Lentibulariaceae
Utricularia
vulgaris L.

X X

X X X X

O X X X

XX X

X X

XXX

o o o

X O X X

X X XX
X X X X

XX XXX

X X

116

Great Basin Naturalist

Vol. 43, No. 1

Table 1 continued.

AREAS

N S W M

HABITATS

123456789

LiLIACEAE

Allium

amplectens Torr.

anceps Kellogg

atroTubens S. Wats,
var. atrorubens

bisceptrum S. Wats.

campanulatum S. Wats.

lemmonii S. Wats.

parvum Kellogg

validum S. Wats.
Cahchortus

bruneaunis A. Nels. & J.F. Macbride

leichtlinii Hook. f.
Camassia

leichtlinii (Baker) S. Wats.
Fritillaria

atropurpurea Nutt.

pinetorum A. Davids.
Lilium

parvum Kellogg
Muilla

transmontana Greene
Smilacina

stellate (L.) Desf.
var. stellata
Triteleia

gracilis (S. Wats.) Greene

ixioides (Ait. f.) Greene
ssp. analina (Greene) Lenz
Veratrum

califomicum Durand
var. californicum
Zigadenus

paniculatus (Nutt.) S. Wats.

venetiosus S. Wats,
var. venenosus

LiMNANTHACEAE

Floerkea

proserpinacoides Willd.

LiNACEAE

Linum

lewisii Pursh
var. lewisii

LOASACEAE

Mentzelia

albicaulis (Hook.) Torr. & Gray
congesta (Nutt.) Torr. & Gray
dispersa S. Wats.
laevicaulis (Hook.) Torr. & Gray
montana (A. Davids.) A. Davids.
torreyi Gray

var. torreyi
veatchiana Kellogg

LORANTHACEAE

Arceuthobium

divaricatum Engelm.
Phoradendron

juniperinum Engelm. ex Gray
ssp. juniperinum

X
X

X X

O X O X

X X O X X X

X
X

January 1983

Lavin: Walker River Floristics

117

Table 1 continued.

AREAS

HABITATS

NSWM 123456789

Malvaceae
Malva

neglecta Wallr.
Sidalcea

glaucescens Greene

multifida Greene

oregana (Torr. & Gray) Gray
ssp. spicata (Kegel) C.L. Hitchc.
Sphaeralcea

atnbigua Gray

ssp. monticola Kearney

grossulariifolia (Hook. & Am.) Rydb.

MORACEAE

Humulus
lupulus L.

var. lupuhides E. Small

Nyctag inaceae
Abronia

turbinata Torr. ex S. Wats.
Mirahilis

bigelovii Gray

var. retroTsa (Heller) Munz

Nymphaeaceae
Nuphar

luteum (L.) Sibthorp. & Sm.

ssp. polysepalum (Engelm.) E.O. Real

Oleaceae
Menodora

spinescens Gray

Onagraceae
Boisduvalia

densiflora (Lindl.) S. Wats.
Camissonia

claviformis (Torr. & Frem.) Raven
ssp. integrior (Raven) Raven

nevadensis (Kellogg) Raven

pubens (S. Wats.) Raven

pusilla Raven

subacaulis (Pursh) Raven

tanacetifolia (Torr. & Gray) Raven
ssp. tanacetifolia
Circaea

alpina L.

ssp. pacifica (Aschers. & Magnus) Raven
Epilobium

anagallidifolium Lam.

angustifolium L.

brachycarpum Presl.

ciliatum Raf.
ssp. ciliatum

glaberrimum Barbey

lactiflorum Hausskn.

latifolium L.

obcordatum Gray
ssp. obcordatum

oregonese Hausskn.

X X X X

X X
XX X

X X
X X

X X

X X
X X

X X

o o o o

X X

X o o

X X

118

Great Basin Naturalist

Vol. 43, No. 1

Table 1 continued.

AREAS

habitats

Gayophytum

decipiens Lewis & Szweykowski
diffusum Ton. & Gray

ssp. parviflorum Lewis & Szweykowski
heterozygum Lewis & Szweykowski
ramosissimum Torr. & Gray
Oenothera
caespitosa Nutt.

ssp. marginata (Nutt.) Munz
hookeri Ton. & Gray

ssp. angustifolia (R.R. Gates) Munz

ORCmOACEAE

Corallorhiza

maculata Raf.
Epipactis

gigantea Dougl. ex Hook.
Listera

convallarioides (Sw.) Nutt.
Platanthera

dilatata (Pursh) Lindl. ex Beck

var. leucostachys (Lindl.) Luer

sparsiflora (S. Wats.) Schlechter
Spiranthes

romanzoffiana Cham.

Orobanchaceae
Orobanche

corymbosa (Rydb.) Ferris

var. corymbosa
fasciculata Nutt.

var. lutea (Parry) Ashey
var. fasciculata

Paeoniaceae
Paeonia

brownii Dougl. ex Hook.

Papaveraceae
Argemone

munita Dur. & Hilg.

ssp. rotundata (Rydb.) G. Ownbey
Corydalis

aurea Willd.
Dicentra

uniflora Kellogg
Eschscholzia

calif omica Cham.

Plantag in aceae
Plantago

lanceolata L.

POACEAE

Agropyron

dasystachum (Hook.) Scribn. & Sm.

var. dasystachum
desertorum (Link.) Schultes
pringlei (Scribn. & Sm.) A.S. Hitchc.
scribneri Vasey
trachycaulum (Link.) Malte ex H. Lewis

var. latiglume (Scribn. & Sm.) Beetle

var. trachycaulum

X X

X X X X

X X

X X X X

X X

XXX
X X
X

XX X

X X

o o

XX X XX

XX X

XXX

XX X

O X X
X X

X X X X

X X
X X

X X

January 1983

Lavin: Walker River Floristics

119

Table 1 continued.

AREAS

HABITATS

N S

X X

X
X

X X

Agrostis

capillaris L.

exarata Trin.

filiculmis M.E. Jones

lepida A.S. Hitchc.

scabra Willd.
Alopecurus

aequalis Sobol.

geniculatus L.
Beckmannia

syzigachne (Stendl.) Feam.
Bromus

anomalus Rupr. ex Foum.

carinatus Hook. 6f Am.

catharticus Vahl

commutatus Schrad.

inermis Leyes.

rubens L.

suksdorfii Vasey

tectorum L.
CakiTnagrostis

breweri Thurb.

canadensis (Michx.) Beauv.

purpurascens R. Br.
Dactylis

glomerata L.
Danthonia

intermedia Vasey
Deschampsia

cespitosa (L.) Beauv.
ssp. cespitosa

danthonioides (Trin.) Munro ex Benth.

elongata (Hook.) Munro ex Benth.
Distichlis

spicata (L.) Greene

var. stricta (Torr.) Beetle
Echinochloa

crusgalli (L.) Beauv.
Elymus

cinereus Scribn. & Merr.

glaucus Buckl.
ssp. glaucus
ssp. virescens (Piper) Gould

triticoides Buckl.
var. triticoides
var. pubescens A.S. Hitchc.
Elysitanion

X hansenii (Scribn.) Bowden
Eragrostis

orcuttiana Vasey
Festuca

brachyphylla Schultes

pratensis Huds.

rubra L.
Glyceria

elata (Nash) M.E. Jones
Hilaria

jamesii (Torr.) Benth.
Hordeum

brachyantherum Nevski

jubatum L.

X X

X
X

X X

120

Great Basin Naturalist

Vol. 43, No. 1

Table 1 continued.

AREAS

habitats

N S

5 6

X X

X
X

X X

X
X

X X

Koeleria

nitida Nutt.
Leucopoa

kingii (S. Wats.) W.A. Weber
Melica

bulbosa Geyer ex Porter & Coult.

stricta Boland.
Muhlenbergia

asperifolia (Nees & Meyem) Parodi

filiformis (Thurb.) Rydb.

richardsonis (Ttin.) Rydb.
Oryzopsis

hymenoides (Roemer & Schultes) Ricker
Panicum capillare L.
Phleum

alpinum L.

pratense L.
Phragmites

australis (Cav.) Trin. ex Steud.
Poa

ampla Merr.

annua L.

bolanderi Vasey

bulbosa L.

compressa L.

cusickii Vasey

epilis Scribn.

fendleriana (Steud.) Vasey

gracillima Vasey

incurva Scribn. & Williams

interior Rydb.

leibergii Scribn.

leptocoma Trin.

nervosa (Hook.) Vasey

nevadensis Vasey ex Scribn.

var. juncifolia (Scribn.) Beetle
var. nevadensis

palustris L.

pratensis L.

rupicola Nash, ex Rydb.

scabrella (Thurb.) Benth. ex Vasey

secunda Presl.

suksdorfii (Beal) Vasey ex Piper
Polypogon

monspeliensis (L.) Desf.
Puccinellia

distans (Jacq.) Pari.

lemmonii (Vasey) Scribn.

nuttalliana (Schultes) A.S. Hitchc.
Secale

cereale L.
Sitanion

hystrix (Nutt.) J.G. Smith

var. brevifolium (J.G. Smith) C.L. Hitchc.

jubatum J.G. Smith
Sphenopholis

obtusata (Michx.) Scribn
var. obtusata
Sporobolus

cryptandrus (Torr.) Gray

X X

X X X X
X X

X X

XXX
X

O 0

January 1983

Lavin: Walker River Floristics

121

Table 1 continued.

AREAS

HABITATS

N S

X X

X
X
X

X X

X
X

X X

Stipa

californica Merr. & Burtt-Davy
Columbiana Macoun.
comata Trin. & Rupr.

var. comata
nevadensis B.L. Johnson
occidentalis Thurb. ex S. Wats.
pinetorum M.E. Jones
speciosa Trin. & Rupr.
thurberiana Piper
webberi (Thurb.) B.L. Johnson
Trisetum
spicatum (L.) Richt.

ssp. spicatum
triflorum (Bigelow) Love & Love
ssp. molle (Michx.) Love & Love
Vulpia
megalura (Nutt.) Rydb.
octoflora (Walt.) Rydb.

var. hirtella (Piper) Henr.
var. octoflora

POLEMONIACEAE

Allophyllum

gilioides (Benth.) A. & V. Grant
integrifolium (Brand) A. & V. Grant
violaceum (Heller) A. & V. Grant
Collomia

grandiflora Dougl. ex Lindl.
linearis Nutt.
tinctoria Kellogg
Eriastrum

sparsiflorum (Eastw.) Mason
wilcoxii (A. Nels.) Mason
Gilia

brecciarum M.E. Jones

ssp. brecciarum
capillaris Kellogg
inconspicua (Sm.) Sweet
interior (Mason & A. Grant) A. Grant
leptalea (Gray) Greene

ssp. leptalea
leptantha Parish

ssp. salticola (Eastw.) A. & V. Grant
leptomeria Gray
malior Day & V. Grant
modocensis Eastw.
ophthalmoides Brand
sinuata Dougl. ex Benth.
Gymnosteris

parvula (Rydb.) Heller
Ipomopsis

aggregata (Pursh) V. Grant

ssp. ag^regata

ssp. attenuata (Gray) V. & A. Grant
congesta (Hook.) V. Grant

ssp. montana (A. Nels. & Kennedy) V. Grant

ssp. palmifrons (Brand) Day

122

Great Basin Naturalist

Vol. 43, No. 1

Table 1 continued.

AREAS

habitats

N S W M

123456789

Leptodactylon

pungens (Torr.) Nutt. ex Rydb.
ssp. pulchriflonim (Brand) Mason
Linanthus

ciliatus (Benth.) Greene

var. neglectus (Greene) Jepson
nuttallii (Gray) Greene ex Milliken

ssp. nuttallii
septentrionalis Mason
MicTosteris

gracilis (Hook.) Greene
ssp. gracilis

ssp. humilfs (Greene) V. Grant
Navarretia

breweri (Gray) Greene
divaricata (Torr. ex Gray) Greene
var. divaricata
Phlox

caespitosa Nutt.

ssp. pulvinata Wherry
covillei E. Nels.
diffusa Benth.

ssp. subcarinata Wherry
ssp. diffusa
hoodii Richards

ssp. canescens (Torr. & Gray) Wherry
longifolia Nutt.
Polemonium
caeruleum L.

ssp. amygdalinum (Wherry) Munz
califomicum Eastw.
chartaceum Mason
eximium Greene
pulcherrimum Hook.

POLYGALACEAE

Polygala

intermontana Wendt
subspinosa S. Wats.

POLYGONACEAE

Chorizanthe

brevicomu Torr.

var. spathulata (Rydb.) C.L. Hitchc.
watsonii Torr. & Gray
Eriogonum

baileyi S. Wats.

var. baileyi
beatlleyae Reveal
brachyantherum Coville
caespitosum Nutt.

var. cemuum
elatum Dougl. ex Benth.

var. elatum
esmeraldense S. Wats.
heermannii Dur. & Hilg.

var. humilius (Stokes) Reveal
hookeri S. Wats.
incanum Torr. & Gray

X X X X

X X X X XXX

X X

X
X

X X

January 1983

Lavin: Walker River Floristics

123

Table 1 continued.

\REAS

HABITATS

5 6

7 8

lobbii Torr. & Gray

var. lobbii

X X

maculatum Heller

marifolium Torr. & Gray

X X

micTothecum Nutt.

var. laxiflorum Hook.

X X

var. ambiguum (M.E. Jones) Reveal

nidularium Coville

nudum Dougl. ex Benth.

var. deductum (Greene) Jepson

X X

nutans Torr. & Gray

var. nutans

ochrocephalum S. Wats.

var. alexandrae Reveal in ed

ovalifolium Nutt.

var. ovalifolium

X X

var. nivale (Canby) M.E. Jones

var. nevadense Gandoger

pusillum Torr. & Gray

rosense A. Nels. & Kennedy

X X

spergulinum Gray

var. reddingianum (M.E. Jones) J.T. Howell

X X

strictum Benth.

ssp. proliferum (Torr. & Gray) Stokes

umbellatum Torr.

var. nevadense Gandoger

var. umbellatum

X X

vimineum Dougl. ex Benth.

wrightii Torr. ex Benth.

var. subscaposum S. Wats.

Oxyria

digyna (L.) Hill

Oxytheca

dendroidea Nutt.

ssp. dendroidea

Polygonum

amphibium L.

var. stipulaceum Coleman

aviculare L.

bistortoides Pursh

X X

douglasii Greene

var. douglasii

var. johnstonii Munz

O 0

var. latifolium (Engelm.) Greene

kelloggii Greene

X X

minimum S. Wats.

X X

persicaria L.

shastense Brewer ex Gray

X X

Rumex

acetosella L.

calif omicus Rech. f.

o o

crispus L.

lacustris Greene

occidentalis S. Wats.

paucifolius Nutt. ex S. Wats.

ssp. paucifolius

ssp. gracilescens (Rech. f.) Rech. f.

salicifolius Weijnm.

triangulivalvis (Danser) Rech. f.

124

Great Basin Naturalist

Vol. 43, No. 1

Table 1 continued.

AREAS

habitats

N S W M

123456789

PORTULACACEAE

Calyptridium

roseum S. Wats.

umbellatum (Torr.) Greene
Claytonia

nevadense S. Wats.

rubra (T.J. Howell) Tidestrom

umbellata S. Wats.
Leivisia

nevadensis (Gray) B.L. Robins.

pygmaea (Gray) B.L. Robins,
ssp. pygmaea

rediviva Pursh
ssp. rediviva
ssp. minor (Rydb.) A. Holmgren

sierrae Ferris

triphylla (S. Wats.) B.L. Robins.
Montia

chamissoi (Ledeb. ex Spreng.) Greene

POTAMOGETONACEAE

Potamogeton
gramineus L.
richardsonii (Benn.) Rydb.

Primulaceae
Androsace

septentrionalis L.

var. subumbellata A. Nels.
Dodecatheon

alpinum (Gray) Greene
ssp. alpinum

ssp. majtis H.J. Thompson
jeffreyi Van Houtte
Primula

suffrutescens Gray

Ranunculaceae
Aconitum

columbianum Nutt.
Actaea

rubra (Ait.) WiUd.

ssp. arguta (Nutt.) Hulten
Anemone

drummondii S. Wats.
Aquilegia

formosa Fisch.

var. paucifhra (Greene) Boothman
var. formosa
pubescens Goville
Caltha

leptosepala DC.

ssp. howellii (Huth) P.G. Smith
Clematis

ligusticifolius Nutt.

var. brevifolia Nutt.
Delphinium

andersonii Gray
ssp. andersonii
glaucum S. Wats.

X X

O X

X X

X X X X
X X

X X

XXX

X X X X

X X X X X

O O 0 o

o o

ox X

X X

o o

X X

X X X X

X X

X X X X X

X X O X X

January 1983
Table 1 continued.

Lavin: Walker River Floristics

125

AREAS

HABITATS

NSWM 123456789

parishii Gray
ssp. parishii
polycladon Eastw.
Myosurus
aristatus Benth. ex Hook.
minimus L.

ssp. montanus Campbell
Ranunculus

alismifolius Geyer ex Benth.

var. alismellus Gray
andersonii Gray
aquatilis L.

var. capillaceus (Thuill.) DC.
cymbalaria Pursh

var. saximontanus Fern.
eschscholtzii Schlecht.

var. eschscholtzii

var. oxynotis (Gray) Jepson
glaberrimus Hook.

var. ellipticus Hook.
occidentalis Torr. & Gray

var. ultramx)ntantis Greene
testiculatus Crantz
Thalictrum
fendleri Gray

var. fendleri
sparsiflorum Turcz. ex Fisch. & Mey.

var. saximontanum Boivin

Rhamnaceae
Ceanothus
greggii Gray

var. vestitus (Greene) McMinn
velutinus Dougl. ex Hook.
Rham.nus

rubra Greene
ssp. rubra

ROSACEAE

Amelanchier
pallida Greene
pumila Torr. & Gray
utahensis Koehne

var. covillei (Standi.) Clokey
var. utahensis
Cercocarpus

ledifolius Nutt. ex Torr. & Gray
var. ledifolius

var. intricatus (S. Wats.) M.E. Jones
Chamaebatiaria

millefolium (Torr.) Maxim.
Fragaria

virginiana Duchesne

ssp. platypetala (Rydb.) Staudt
Geum

canescens (Greene) Munz
macrophyllum Willd.

X X

X X
XX X

X X

XXX

X X

X X X X X

X X

O O 0 o

X X

XXX

X X X X

XXX XX

126

Great Basin Naturalist

Vol. 43, No. 1

Table 1 continued.

AREAS

habitats

N S W M

123456789

Holodiscus

dumosa (Nutt.) Heller

var. glabrescens (Greenm.) C.L. Hitchc.
Horkelia
fusca Lindl.

ssp. pseudocapitata (Rydb.) Keck
Ivesia

gordonii (Hook.) Torr. & Gray
lycopodioides (Hook.) Torr. & Gray
ssp. lycopodioides
ssp. megalopetala (Rydb.) Keck
muirii Graj^

purpurascens (S. Wats.) Keck
ssp. congdonis (Rydb.) Keck
shockleyi S. Wats.
Potentillc
anserina L.

ssp. anserina
biennis Greene
breweri S. Wats.
diversifolia Lehm.
drummondii Lehm.

ssp. bruceae (Rydb.) Keck
ssp. drummondii
fhbellifolia Hook, ex Torr. & Gray

var. fhbellifolia
fruticosa L.

ssp. floribunda (Pursh) Elkington
glandulosa Lindl.

ssp. nevadensis (S. Wats.) Keck
gracilis Dougl. ex Hook.

var. glabrata (Lehm.) C.L. Hitchc.
var. flabelliformis (Lehm.) Torr. & Gray
newberryi Gray
pectinisecta Rydb.
pseudosericea Rydb.
Prunus

andersonii Gray

emarginata (Dougl. ex Hook.) Walp.

virginiana L.

var. demissa (Nutt.) Sarg.
var. melanocarpa (A. Nels.) Sarg.
Purshia

tridentata (Pursh) DC.
Rosa

woodsii Lindl.

var. ultrarrumtana (S. Wats.) Jepson
Rubus

parviflorus Nutt.

var. bifarius Fern.
Sibbaldia

procumbens L.
Sorbus

califomica Greene
Spiraea
densiflora Nutt. ex Torr. & Gray
ssp. densiflora

X X X X

XXX XX

X X

O X

X X

XXXX X xxxxxx

X XX

X X O X XX

X O O X X

X XXX

January 1983

Lavin: Walker River Floristics

127

Table 1 continued.

AREAS

HABITATS

N S W M

123456789

RUBIACEAE

Galium
aparine L.
bifolium S. Wats.
hypotrichium Gray

ssp. ebbettsense Dempster & Ehrend.

ssp. hypotrichium
multiflorum Kellogg
trifidum L.

ssp. pacificum Wieg.

ssp. subbiflorum (Wieg.) Puff
trifhrum Michx.
Kelloggia

galioides Torr.

Salicaceae
Populus

balsamifera L.

ssp. trichocarpa (Torr. & Gray) Brayshaw
fremontii S. Wats.
tremuloides Michx.
Salix
arctica Pallas

drummondiana Barratt ex Hook.
eastwoodiae Heller
exigtia Nutt.
geyeriana Anderss.

var. geyeriana

var. argentea (Bebb) Schneid.
lasiandra Benth.

var. lasiandra

var. caudata (Nutt.) Sudworth
lasiolepis Benth.
lemmonii Bebb

ligulifolia (Ball) Ball ex Schneid.
lutea Nutt.
melanopsis Nutt.
myrtillifolia Anderss.

var. myrtillifolia
orestera Schneid.
planifolia Pursh

var. monica (Bebb) Schneid.
reticulata L.

ssp. nivalis (Hook.) Love, Love & Kapoor
scouleriana Barratt ex Hook.

Saxifragaceae
Heuchera

duranii Bacig.

rubescens Torr.

var. alpicola Jepson
Lithophraffna

glabrum Nutt.
Mitella

breweri Gray

pentandra Hook.
Pamassia

palustris L.

var. califomica Gray

X X
X X

X
X

X X

X X
X X

X
X

X X

X
X

X X

128

Great Basin Naturalist

Vol. 43, No. 1

Table 1 continued.

AREAS

habitats

N S W M

123456789

Ribes

aureutn Pursh

var. aureum
cereum Dougl.
inebrians Lindl.
inerme Rydb.

var. inerme
montigenum McClatchie
velutinum Greene

var. velutinum

var. glanduliferum (Heller) Jepson
viscosissimum, Pursh

var. viscosissim,um.
Saxifraga

aprica Greene
bryophora Gray
nidifica Greene
odontoUmui Piper
oregana T.J. Howell

var. sierrae Coville
tolmiei Torr. & Gray

var. ledifolia (Greene) Engl. & Irmsch.

SCROPHULARIACEAE

Antirrhinum

kingii S. Wats.
Castilleja

applegatei Fern.

var. fragilis (Zeile) N. Homgren
var. pallida (Eastwood) N. Holmgren

chromosa A. Nels.

exilis A. Nels.

lemmonii Gray

linariifolia Benth. ex DC.

miniata Dougl. ex Benth.

nana Eastwood

peirsonii Eastwood

pilosa (S. Wats.) Rydb.
Collinsia

parviflora Dougl. ex Lindl.
Cordylanthus

helleri (Ferris) J.F. Macbride

ramosus Nutt. ex Benth.
ssp. setosus Pennell
Keckiella

breviflora (Lindl.) Straw

ssp. glabrisepala (Keck) Straw
Limosella

aquatica L.
Mimetanthe

pilosa (Benth.) Greene
Mimulus

breweri (Greene) Coville

coccineus Congd.

densus A.L. Grant

floribundus Dougl. ex Lindl.

guttatus Fisch. ex DC.

lewisii Pursh

mephiticus Greene

X X

X
X

o
o

o o

January 1983
Table 1 continued.

Lavin: Walker River Floristics

129

AREAS

HABITATS

NSWM 1234567

moschatus Dougl.
primuloides Benth.

var. primuloides
rubellus Gray
suksdorfii Gray
tilingii Kegel
Orthocarpus

copelandii Eastwood

var. cryptanthus (Piper) Keck
hispidus Benth.
luteus Nutt.
Pedicularis
attollens Gray
ssp. attollens
groenlandica Retz.
ssp. groenlandica
semibarbata Gray
var. semibarbata
Penstemon
bridgesii Gray
dovidsonii Greene

var. davidsonii
deustus Dougl. ex Lindl.

ssp. deustus
heterodoxus Gray

var. heterodoxus
humilis Nutt. ex Gray
newberryi Gray
ssp. newberryi
procerus Dougl. ex Grahm.

ssp. formosus (A. Nels.) Keck
rubicundis Keck
rydbergii A. Nels.

var. varians (A. Nels.) Cronq.
speciosus Dougl. ex Lindl.
Scrophularia

desertorum (Munz) R.J. Shaw
Verbascum

thapsus L.
Veronica
americana (Raf.) Schwein. ex Benth.
anagallis-aquatica L.
beccabunga L.
peregrina L.

ssp. xalapensis (H.B.K.) Pennell
serpyllifolia L.

ssp. humifusa (Dickson) Syme
ssp. serpyllifolia
wormskjoldii Roemer & Schultes
ssp. alterniflora (Fern.) Pennell

SOLANACEAE

Leucophysalis

nana (Gray) Averett
Nicotiana

attenuata Torr. ex S. Wats.
Solanum

triflorum Nutt.

X X

X X O X X X X

XX X X X X

XXXX XXXX XXX

XXXX X X X XXX

XXX XXX

XX O X O X X

o o o o

X X

XXX

X X

XX X X XXX X
XXXX XXX XXX

XXXX XXX

XXXX XXXXX X

XX X X X X

X o o

o o o

X X O X O X X

XX X X XX XX

XXX X X X

X X XX

XX XXXX

XXXX XX X

o o o

130

Great Basin Naturalist

Vol. 43, No. 1

Table 1 continued.

AREAS

habitats

NSWM 123456789

Typhaceae
Typha
latifolia L.

Urticaceae
Urtica
dioica L.

var. holosericea (Nutt.) C.L. Hitchc.

Valebianaceae
Plectritis

macrocera Torr. & Gray
ssp. grayit (Suksdorf) Morey
Valeriana

californica Heller

Verbenaceae
Verbena
bracteata Lag. & Rodr.

ViOLACEAE

Viola
adunca Sm.

var. adunca
bakeri Greene

ssp. bakeri
beckwithii Torr. & Gray

ssp. beckwithii
macloskeyi Lloyd
ssp. macloskeyi
nephrophylla Greene

var. nephrophylla
purpurea Kellogg

ssp. atriplicifolia (Greene) Baker 6f Clausen
ssp. aurea (Kellogg) Clausen

Zygophyllaceae
Tribulus
terrestris L.

X X

X X X X

X X
XX o

X X

ox XXX

X X

X X
X

X
X

X X

AGROPYRON ARIZONICUM (GRAMINEAE: TRITICEAE)
AND A NATURAL HYBRID FROM ARIZONA

Grant L. Pyrah'

Abstract.— The new hybrid X Agrositanion pinalenoensis (Gramineae: Tribe Triticeae) is found in disturbed, for-
ested areas of higher elevations in southern Arizona. In the Pinaleno Mountains where logging has been heavy, nu-
merous disturbed habitats have permitted frequent hybrid populations to persist. Intermediate phenotype, chromo-
some behavior, lack of seed set, and pollen sterility were used to interpret the status of this hybrid derivative.
Introgression and/or segregation are not apparent.

Of the many natural and experimental in-
tergeneric hybrids reported in the Triticeae,
none have been reported between Agropyron
arizonicum Scribner & Smith and Sitanion
hystrix var. brevifolium (J. G. Smith) C. L.
Hitchcock. The present paper describes ex-
tensive hybridizations between these two
taxa.

Agropyron arizonicum has flat leaves, dis-
tinct flexuous spikes, one spikelet per node,
and more or less ascending awns (although
they are somewhat divergent at maturity). Si-
tanion hystrix has long been recognized as an
extremely variable species; however, erect
spikes, very long awns on the glumes and
lemmas, and 2 spikelets per rachis node are

typical. The variation in .Arizona has been
treated by Wilson (1963) as S. longifolium.

Although Sitanion hystrix has very exten-
sive distribution in western North America, it
is limited to higher elevations of isolated
mountain ranges in the southwestern United
States and northern Mexico. On these iso-
lated mountain ranges, the range of this spe-
cies frequently overlaps that of Agropyron
arizonicum, a species of high elevations re-
stricted to west Texas, southwestern New
Mexico, and southeastern Arizona (Fig. 1). In
the Pinaleno Mountains, Graham Co., Ari-
zona, and the Santa Catalina Mountains,
Pima Co., Arizona (Fig. 2), where extensive
areas have been distiubed by logging (Fig. 3),
recreation, summer home development, and
road building, numerous hybrids between
Agropyron arizonicum and Sitanion hystrix
var. brevifolium are formed. Eleven hybrid

Fig. 1. Cross-hatched area represents sympatric range
of Sitanion hystrix var. brevifolium and Agropyron

anzonicum.

Fig. 2. Cross-hatched areas represent the distribution
of X Agrositanion pinalenoensis Pyrah in Arizona.
(Larger area is the Pinaleno Mountains, smaller area is
the Santa Catalina Mountains).

'Southwest Missouri State University, Springfield, Missouri 65802.

131

132

Great Basin Naturalist

Vol. 43, No. 1

Fig. 3. Typical disturbed forest with dense stands oi Agropyron arizonicum and Sitanion hystrix var. brevifolium.

populations were found and studied in the
Pinaleno Mountains at elevations generally
between 7000 and 10,000 ft.

Materials and Methods

Eleven hybrid populations in the Pinaleno
Mountains and two hybrid populations from
the Santa Catalina Mountains were studied.
Prepared herbarium specimens from each are
deposited in the Southwest Missouri State
University herbarium. Five additional mature
inflorescences were obtained from each hy-
brid and four parent specimens from the
High Peak population. These were put in en-
velopes and used for comparative measure-
ments of length of glume, lemma, awn, and
rachis joint, and also determination of the
number of spikelets per rachis node.

A few late-flowering inflorescences were
fixed in a solution of one part glacial acetic
acid to three parts absolute ethyl alcohol.
These were stored in 70 percent ethyl alco-
hol and used for cytological studies.

Pollen grains were obtained from mature
spikes and viewed with the scanning electron

microscope to determine viability. Soil pH
was determined by sampling five sites each
for parental species and the hybrid. The soil
sample was carefully obtained by taking soil
from the entire soil profile of 0 to 5 inches
and mixed. A soil-water slurry was prepared
and the pH determined by a standard pH
meter.

In an attempt to assess the pollen parent
and the seed parent and hybrid success, a
large population near High Peak was studied
in the following way. A circular area 6 m in
diameter was marked around each of 25
hybrids. The number of specimens of each
parental species as well as other hybrids
within this circle were recorded.

Results and Discussion

Pure stands of Sitanion typically grow in
rather open, unshaded, shallow soil, with top-
soil and litter depths from 1 to 3 inches and
clay with scattered rock constituting the re-
maining root zone. Soil pH range is 5.4 to
5.8. Agropyron arizonicum grows in richer
soils, with the topsoil and litter occupying

January 1983

Pyrah: Agropyron Hybrid

133

the upper 10 inches and only a limited
amount of clay and rock toward the bottom
of the root zone. Soil pH range is 5.9 to 6.4.
Characteristically, this species is more vigor-
ous in partially shaded areas, but it also
grows in open sun. Nearly all combinations
of the above soil conditions and other habitat
requirements have been created by logging
and road building, resulting in numerous dis-
turbed habitats, as well as habitats for each
parental species in very close proximity. In
many of these situations, plants of both spe-
cies either touch each other or are within
only a few inches. Since flowering occurs
over the same time period, this allows for
showers of pollen to accomplish hybridiza-
tion. In nearly all these situations hybrids are
found.

The frequency of hybridization between
these species is difficult to assess, since a ma-
ture hybrid plant is the only indication that
hybrid pollination occurred. A circular area
(6 m in diameter) around each of 25 hybrids
near High Peak was examined and the num-
ber of associated parental and hybrid plants
was counted (Table 1). It is suggestive from
the columns opposite hybrids 8, 9, 10, 11, and

Table 1. Comparison of the number of parental and
hybrid plants within an area (6 meters in diameter)
around 25 hybrids in a population near High Peak (see

text).

Hybrid No.

Sitanion

Agropyron

Hybrids

1 .

13 that hybrid success is dependent upon the
density and proximity of both parents. Al-
though this may appear to be obvious, there
are situations in some dicots in which hy-
bridization is abundant but one parent is rare
(Stutz 1964, Pyrah 1965). A series of artificial
hybridizations would be necessary to deter-
mine whether or not Agropyron or Sitanion is
more important as the seed or pollen parent
of the natural hybrids.

Hybrid plants are readily recognized be-
cause of their robust size and slightly nodding
spikes (Fig. 4), and most are growing in dis-
turbed habitats (Fig. 3). Soil pH ranges from
5.7 to 6.5. Of the 11 populations studied,
nearly every hybrid specimen fell within an
intermediate range in morphological features
and fragility of the rachis. A diagramatic il-
lustration (Fig. 5) of Sitanion hystrix var. bre-
vifolium and Agropyron arizonicum with
their putative hybrids shows average lengths
of the glume, lemma, awn, and rachis joint

Fig. 4. Spikes of Sitanion hystrix var. brevifolium
(left), hybrid (center), and Agropyron arizonicum (right).

134

Great Basin Naturalist

Vol. 43, No. 1

\^
\/
\/
\/

\/
\/

N//r

x=io.o

•X=66.5

X=11.0
=80. 1

\/
\/

X=10.7

X=15.3

-X=36.4

\//\T::;:::Yfi

X=26.6

■K=13.7
►-X=14.2

Sitanion

Hybrid

Agropyron

Fig. 5. Schematic representation of some spike and spikelet characters of S. hystrix var. brevifolium, A. arizoni-
cum, and their natural hybrid. Measurements used in this figure are means (X) from one population. A = Spike; cen-
ter line between oblique lines (spikelets) represent the rachis joint. B = Lemma (triangle) with attached awn. C =
Glume.

and the number of spikelet pairs per spike.
Measurements were derived from a popu-
lation near High Peak consisting of 25 hy-
brids and 4 parental specimens (Table 2).
Field observation and examination of numer-

ous herbarium specimens reveal clearly that
the 2 parents are rather uniform with regard
to the characters used and that where varia-
tion exists the range does not overlap that of
the hybrid.

January 1983

Pyrah: Agropyron Hybrid

135

Table 2. Summary of measurements of spike characters used from one population to distinguish Agropyron ari-
zonicum, Sitanion hystrix var. brevifolium, and their natural hybrid.

No. nodes

No. single

No. double

Glume

Lemma awn

Lemma

Rachis joint

per spike

spikelets

length

Agropyron X

11.900

11.600

0.300

14.156

26.588

13.739

15.349

40.000

109.000

119.000

109.000

2.329

2.318

0.648

2.290

4.758

9.122

3.059

XSE

0.368

0.367

0.103

0.219

0.436

0.836

0.293

Sitanion X

10.900

1.900

9.000

80.091

66.505

11.037

10.020

40.000

99.000

109.000

99.000

1.236

0.709

1.013

11.639

8.759

0.849

2.162

XSE

0.195

0.112

0.160

1.170

0.839

0.081

0.217

Hybrid X

13.360

8.730

4.630

28.071

36.372

11.609

10.737

400.000

1236.000

1336.000

1236.000

2.380

2.888

2.733

4.741

7.441

1.220

1.969

XSE

0.119

0.144

0.137

0.135

0.204

0.033

0.056

The hybrid did not set seed and no viable
pollen was produced because the pollen
grains were collapsed and empty (compare
Figs. 6, 7, and 8). Preliminary cytological ex-
amination of one hybrid showed 14 bivalents
at metaphase; however, some bivalents show
irregular pairing of chromosome segments
that could cause cryptic structural hybridity
(Fig. 9; arrows indicate conspicuous asynap-
tic and synaptic pairing within two biva-
lents). Morphology of spike characters was
generally intermediate. These evidences
strongly suggest that the plants studied are
first generation hybrids and that little or no
introgression occurs. These hybrid plants
conform to the generic description of X
Agrositanion as reported by Bowden (1967).

Descriptions of the Hybrids

X Agrositanion pinalenoensis Pyrah, Hyb.
nov. (Agropyron arizonicum Scribner &
Smith X Sitanion hystrix var. brevifolium (J.
G. Smith) C. L. Hitchcock).

Hybrida sterilis, inter Sitanion hystrix var.
brevifolium et Agropyron arizonicum proba-
biliter sed differt ab utroque spica moderate
nutanti et inferioribus sex nodis spicae cum
binatis vel binatis singulisque spiculis et supe-
rioribus nodis spicae ciun singulus spiculus;
differt a prima articulis rhachis et lemma-
tibus longioribus et glumis et aristis lemma-
tum brevioribus; differt a secunda articulis
rhachis et lemmatibus brevioribus et glumis
et aristis lemmatum longioribus.

Figs. 6, 7, and 8. Electron micrographs (SEM) of pollen from Sitanion hystrix var. brevifolium (Fig. 6), hybrid (Fig.
7), and Agropyron arizonicum (Fig. 8). ca 600X.

136

Great Basin Naturalist

Vol. 43, No. 1

Fig. 9. Meiotic metaphase I of hybrid. Arrows in-
dicate two pairs of chromosomes with synaptic and asy-
naptic regions.

Sterile hybrids differing from both parents
by having moderately nodding spikes and
usually having the lower 6 to 8 spike nodes
with paired spikelets or a combination of
paired and single spikelets and only single
spikelets at the upper nodes; differing from
Sitanion by having longer rachis joints and
lemmas but shorter glumes and lemma awns;
differing from Agropyron by having shorter
rachis joints and lemmas but longer glumes
and lemma awns.

Type.— Open grassy meadows of disturbed
forests along State Rt. 366 near mile marker
141 on Mt. Graham, Pinaleno Mountains,
Graham Co., Arizona, 13 Aug. 1975, Grant
L. Pyrah 3051 (SMS).

Discussion

The classification of genera, species, and
hybrids in the Triticeae is still open to ques-
tion. Many taxonomists question the validity
of recognizing all the genera now published,
although Gould (1947) and Church (1967) are
two of only a few who have initiated some
consolidation. Widespread hybridization,
similar chromosome behavior, and several

variable spike characteristics were the pri-
mary bases for consolidating Sitanion, Agro-
pyron, and Hystrix with Elymus. Hitchcock
et al. (1969) defend the retention of estab-
lished separate generic names for this agro-
nomically important tribe, primarily on the
basis of practicality. Agropyron arizonicum is
probably closely allied to A. spicatum and
there are still conflicts about the variation
found in Sitanion hystrix. Wilson (1963)
treats the plants of this region as Sitanion
longifolium, but Hitchcock et al. (1969) pre-
fer S. hystrix var. brevifolium.

Confusion is now arising in the literature
for numerous named hybrids because of the
lack of agreement as to the generic status of
each of the parents involved. If a consoli-
dation of genera were accepted, hybrids
would bear the appropriate specific designa-
tion within the genus Elymus. However,
Bowden (1967) recognized the classical treat-
ment of this tribe and made appropriate no-
menclatural designations for intergeneric
hybrids in conformity with the International
Code of Botanical Nomenclature. Hybrid
naming in this case had resulted in such ge-
neric combinations as Agroelymus, Agrohor-
deum, Agrositanion, Elyhordeum, and Sitor-
deum. I have chosen to use the classification
of Bowden and to create the binomial X
Agrositanion pinalenoensis Pyrah.

Literature Cited

Bowden, W. 1967. Taxonomy of intergeneric hybrids of
the tribe Triticeae from North America. Cana-
dian J. Bot. 45:720.

Church, G. L. 1967. Taxonomic and genetic relation-
ships of eastern North American species of
Elymus. Rhodora 69:135.

1967. Pine Hills Elymus. Rhodora 69:330-351.

Gould, F. W. 1947. Nomenclatorial changes in Elymus
with a key to the California species. Madrono
9:120-128.

1974. Nomenclatural changes in the Poaceae.

Brittonia 26:60.

Hitchcock, C. L., A. Cronquist, M. Owenby, and J.
W. Thompson. 1969. Page 701 in Vascular plants
of the Pacific Northwest. Part I.

Pyrah, G. 1964. Cytogenetic studies of Cercocarpus in
Utah. Unpublished thesis. Brigham Young Univ.,
Provo, Utah.

Stutz, H. C, and L. K. Thomas. 1963. Hybridization
and introgression in Cowania and Purshia. Evolu-
tion 18:183-195.

Wilson, F. D. 1963. Revision of Sitanion (Triticeae,
Gramineae). Brittonia. 15:303-323.

SPECIES COMPOSITION, DISTRIBUTION, AND PHYTOSOCIOLOGY OF
KALSOW PRAIRIE, A MESIC TALL-GRASS PRAIRIE IN IOWA

Jack D. Brotherson'

Abstract.— Species composition, distribution, and phytosociology of an 8 hectare area of tall grass prairie was in-
tensively studied. Elevation and soils data were correlated with species distribution patterns. All species showed a
response. Nine general patterns of distribution were observed in relation to elevation and soil types. Ordination and
interspecific association analyses were used to identify clusters or groups of species having similar ecological ampli-
tudes. Sporobolis heterolepis is the dominant plant of the upland prairie. The vegetation of the prairie is best de-
scribed and represented by the continuum concepts of phytosociology.

A government survey started in March
1832, when Iowa was still a territory, and
completed in August 1859 first documented
the original extent of Iowa's prairie. The sur-
vey indicated that in the 1850s grassland cov-
ered about 85 percent of Iowa (U.S. Govern-
ment 1868, Hayden 1945, Hewes 1950, and
Dick-Peddie 1955). Today there are only a
few tracts of this once vast Iowa prairie
remaining.

Provision for state-owned prairies was
made in 1933 when the Iowa State Con-
servation Commission prepared a report
known as the Iowa Twenty-five Year Con-
servation Plan. This plan led to the purchase
of several prairies that are now owned by
state agencies. The prairies were purchased
and set aside as natural areas with the intent
that the various typical landscapes, wild
flowers, and wild life of the native tall-grass
prairie region be preserved for posterity. It
was also intended that these areas would be
useful as '^ame and wild life sanctuaries; as
examples of the native prairie soil types,
where comparisons could be made with culti-
vated soils of the same soil association; and as
reserves of prairie where scientific in-
vestigations could be made on problems con-
cerning the native vegetations, floras, and
faunas of the various topographic, climatic,
and prairie districts throughout Iowa. There-
fore, they were meant to serve as a reference
point by which future generations could
compare the influences of man on Iowa since

settlement (Hayden 1946, Moyer 1953, Aii-
man 1959, Landers 1966).

Kalsow Prairie, 64.8 ha (160 acres) of un-
plowed grassland in Pocahontas County,
Iowa, is one such area. Criteria for its pur-
chase dictated that this area satisfy the re-
quirements of a game preserve, contain one
or more soil types of an association, and in-
clude several regional vegetation types (Hay-
den 1946). Since its purchase in 1949 it has
been the object of several studies on the na-
ture and description of its vegetation, soils,
management, insects, response to fire, mam-
mals, and nematodes (Moyer 1953, Eh-
renreich 1957, Esau 1968, Richards 1969,
Brennan 1969, Norton and Ponchillia 1968,
Schmitt 1969).

The characteristics of Iowa prairie in terms
of vegetation types, structure, and general
ecology of the dominant species was the sub-
ject of several papers during the 1930s and
1940s (Steiger 1930, Rydberg 1931, Weaver
and Fitzpatrick 1934, Hayden 1943). These
authors recognized the existence of six major
types of grassland or vegetative communities
and generally concluded that water relations,
as affected by climate, soil, and topography,
are responsible for local variations in the
structure and distribution of Iowa prairie
vegetation. Weaver and Fitzpatrick (1934)
state:

In varying the water relations of soil and air they merely
bring about changes in the groupings of the dominant
grasses and accompanying segregations and rearrange-
ments of the forbs.

'Department of Botany and Range Science, Brigham Young University, Prove, Utah 84602.

137

138

Great Basin Naturalist

Vol. 43, No. 1

The major grassland types, as alluded to in
the above studies, were labeled "Con-
sociations" after Weaver and Clements
(1938) and were designated as follows:

1. Big Bluestem type {Andropogon gerardi)—
found on the lower moist slopes and well-
aerated lowlands.

2. Slough Grass type {Spartina pectinata)—
found on poorly aerated and wet soils of
sloughs and natural drainage systems.

3. Tall Panic Grass- Wildrye type {Panicum
virgatum and Elymus canadensis)— found
to occur on soils intermediate between
Slough Grass and Big Bluestem types.

4. Little Bluestem type {Schizachyrium sco-
parius)— most important upland type
(well-drained soils).

5. Needle Grass type {Stipa spartea)— found
on the uplands, often occurring as a nar-
row zone following the shoulders of the
ridges.

6. Prairie Dropseed type {Sporobolus hetero-
lepis)—iound locally on the dryest upland
sites.

Moyer (1953), Aikman and Thome (1956),
Ehrenreich (1957), and Kennedy (1969) in re-
cent studies present ecological and taxonomic
descriptions of four state-owned native
prairie tracts. All accounts contain extensive
reviews of prairie literature. The vegetation
complex as treated in these studies is limited
basically to upland prairie. The studies also
include information on soils, microclimate,
topography, and management. Aikman
(1959) reviewed in some detail the state of
prairie research in Iowa.

Investigations involving the distribution of
individual species within the prairie associ-
ation began with the work of Shimek (1911,
1915, 1925). Weaver (1930) and Weaver and
Fitzpatrick (1932) discuss the role of the ma-
jor grasses and forbs within the commimity.
Steiger (1930) and Cain and Evans (1952)
mapped the spatial distributions of several
species. They conclude that the principal fac-
tors affecting the local distribution patterns
of prairie species are as follows: (1) micro-
climatic conditions, (2) edaphic variations, (3)
the biology of the species concerned, particu-
larly methods of reproduction and dispersal,
(4) the relations of the species and other or-
ganisms, animal as well as plant, occurring in
the community, and (5) the element of

chance in the dispersal and establishment of
new individuals. Local distribution patterns
of species have been of interest to many ecol-
ogists (Curtie 1955, Kenshaw 1964, Sanders
1969).

Species in general show varying degrees of
aggregation or association due to exhibited
preferences for or tolerances of certain envi-
ronmental conditions. The distributional pat-
terns and interactions of the component spe-
cies of a community express its
phytosociological structure. Studies of grass-
land phytosociology have been concerned
with either classification or ordination of ba-
sic species groups (Crawford and Wishart
1968).

This investigation was undertaken to pro-
vide information on the phytosociology of an
8 ha tract of Kalsow prairie in relation to
edaphic and topographic variation. It in-
cludes information on species composition
and distribution, factors affecting the distri-
butional patterns of these species, community
types, and interrelationships within and be-
tween these communities.

Materials and Methods

Study Site

Kalsow Prairie is one of several state-
owned Iowa prairies. It is 5 miles northwest
of Manson, Iowa, and comprises the NE V4 of
Section 36, Belleville Township, T 90N, R
32W, Pocahontas County. It occurs in a part
of northcentral Iowa that was glaciated dur-
ing the most recent advances of the Wiscon-
sin Glacier and within the Clarion-Nicollet-
Webster soil association area (Ruhe 1969).
The area was chosen for study on the basis of
its vegetational composition (i.e., floristic
richness and the presence of several plant
community types).

The Vegetation

Taxonomy

Voucher specimens from the prairie were
collected in duplicate throughout the grow-
ing seasons. All specimens were identified,
and identical sets have been deposited in the
herbaria of Iowa State University, Ames,
Iowa, and Brigham Young University, Provo,

January 1983

Brotherson: Kalsow Prairie

139

Utah. Nomenclature follows Pohl (1966) for
the grasses, Gilly (1946) for the sedges, and
Gleason (1952) for the forbs.

Community Types

The 8 ha tract within Kalsow Prairie is ad-
jacent to the south boundary of Kalsow
Prairie and contains within its borders two
vegetation types or subcommunities. These
vegetation units were identified and delimit-
ed as follows (Fig. 1):

1. Upland prairie— includes a major por-
tion of the study area on the ridges and
adjacent lower slopes.

2. Potholes and drainage— found in the
swales and lowlands of the study site.

The vegetation of these community types
was analyzed using two separate approaches.
The first involved the identification and list-
ing of all species foimd within their bound-
aries. The second utilized random plots to de-
termine percent cover, composition, and

-^WEST

MAP OF THE KALSOW PRAIRIE

A AREAS AFFECTED BY SOIL DRIFT FROM ADJACENT FIELDS

■ 20 ACRES OF. PRAIRIE INVOLVED IN SOIL AND PLANT

DISTRIBUTION STUDIES
d©^ POTHOLES AND DRAINAGE

Fig. 1. Map of Kalsow Prairie.

140

Great Basin Naturalist

Vol. 43, No. 1

interspecific relationships of species within
these subcommunities.

Quadrat Analysis

The vegetation of each area was sampled
by using 20 X 50 cm (1000 cm 2) quadrats.
The quadrats were located on a restricted
basis to reduce bias and to keep adjacent
quadrats equal distances apart. Sampling was
done between August 1 and September 15,
when most species had reached their max-
imimi growth. Cover estimates were made
for each quadrat through use of Dauben-
mire's (1959) method.

Coverage was determined separately for
all species overlapping the plot regardless of
where the individuals were rooted. Coverage
was projected to include the perimeter of
overlap of each species regardless of super-
imposed canopies of other species. The ca-
nopies of different species are commonly in-
terlaced or superimposed over the same area;
therefore coverage percents often total great-
er than 100 percent.

Community Analysis

Plant distribution and topographic studies.
An 8 ha (20 acres) plot of prairie (Figure 1)
containing a large segment of potholes and
drainage was selected and staked off in a 9 X
9 m grid. Each 27 square-meter block was
then surveyed and a presence list compiled
for all plant species found within the area. A
total of 968 blocks was thus surveyed, and
distribution data were tabulated for 160 spe-
cies. Topographic readings were taken at 968
points and recorded in tenths of feet on the
same 8 ha grid. Points were located at the
comers of the 27 square meter plots.

Soil mapping. Soils were mapped on the 8
ha intensive study area. Mapping was done in
cooperation with the Iowa State University
Soils Survey under the supervision of Dr.
Thomas E. Fenton, with Mr. J. Herbert Hud-
dleston doing the actual mapping in the field.
The mapping criteria for decisions on soil
series delineation were as follows:

A Clarion— typical well-drained soil on
convex ridges. Surface color lOYR
2/2-3/2; subsoil color lOYR 4'3-5'4.

A" Clarion-Nicollet— an intergrade with
respect to drainage as interpreted
from the color profile. Surface is still

lOYR 2/2-3/3, but the subsoil colors
are duller, not exceeding /3 chroma.
Profile is not mottled as in Nicollet.

N Nicollet-typical Nicollet, lOYR
21^-2/2 surface color, /2 chromas in
the subsoil, which is mottled. Depth
to carbonates generally greater than
30 inches.

Na Calcareous Nicollet— as above, but
calcareous at some depth less than
30 inches.

W Webster— typical Webster with
black (N2/-10YR 2/1) surface colors
and gray (lOYR 4/1-4/2-5/2) sub-
soil colors. Depth to gray subsoil
ranges from 23/41 inches, but is
commonly 30-35 inches. Carbonates
occur at some depth below 22 inch-
es, but the usual range in depth to
carbonates is 22-36 inches. Some
soils identified as Webster are non-
calcareous in the entire probe depth
(42 inches).

Wh Heavy Webster— typical colors of
Webster but heavier textures, strong-
er development in the B and a lack
of carbonates in 42 inches. In many
places spots of Wh are included in
the regular Webster mapping unit.
On the other hand, some areas iden-
tified as regular Webster but non-
calcareous to 36 inches or more
might better have been called heavy
Webster. The Webster soils, as
mapped, include a rather broad
range of texture and depths to carbo-
nates, which could be more precisely
subdivided only with further
investigations.

N- Webster-Nicollet— an intergrade
whose surface color and friability is
like Nicollet but whose subsoil is
darker or grayer than true Nicollet.
The soil is drier than Webster.

Na- Calcareous Webster-Nicollet— as
above but calcareous somewhere
above 22 inches.

H Harps— typical Harps, a loamy,
weakly developed soil that effer-
vesces strongly to violently from the
surface downward. Calcium carbo-
nate equivalent probably in the
range 20-40 percent.

January 1983

Brotherson: Kalsow Prairie

141

C Canisteo— this is essentially cal-
careous Webster. As mapped, it may
be noncalcareous in the surface, but
carbonates must be detected some-
where in the 0-15 inch layer. It has
lower calcium carbonate equivalent,
heavier textures, and stronger devel-
opment than Harps.

H" Harps-Canisteo— an intergrade that
has either Harps-like characteristics
in the surface and becomes more Ca-
nisteo-like with depth or Canisteo-
like surface characteristics and a
Harps-like subsoil.

Wa Webster-Canisteo— an intergrade in
which carbonates are first detected
in the 15-22 inch layer. All other
characteristics of Wa, W, and C are
essentially the same.

C" Inverted Canisteo-Heavy Webster—
this represents a rather peculiar con-
dition that tends to occur as a nar-
row band around the potholes. The
surface is moderately to strongly cal-
careous, but carbonates decrease
with depth to a noncalcareous,
heavy, well-developed subsoil like
that of heavy Webster.

G Glenco— a poorly drained soil that
occupies small potholes, the outer
portions of large potholes, or con-
necting drainage-ways. It has a
black, highly organic surface but a
gray, mineral, heavy, well-developed
subsoil. In many respects it is similar
to heavy Webster except for the or-
ganic surface and lack of grit and
pebbl-.s. Depth to carbonate is gen-
erally greater than 42 inches, but
may be up to 36 inches.

Ga Calcareous Glenco— Glenco that be-
comes calcareous above 36 inches. It
usually lacks the heavy textures and
good development of regular Glenco
as well.

O Okoboji— a black, mucky silt loam,
very weakly developed soil occur-
ring in the deepest areas of the
potholes.

GO Glenco-Okoboji— an intergrade that
may have the heavy textures of
Glenco, but is darker, more organic,
less well developed, and wetter than
Glenco.

Seventeen soil series were recognized and
mapped in the field, utilizing soil samples ob-
tained with a 42-inch hand probe.

Data Analysis

General descriptive data. Data collected
from quadrat studies, mapping studies, soil
studies, and topographic studies were used to
describe generally the vegetation. Frequency
values and average cover values were deter-
mined for all species in every stand.

Ordination analysis. An ordination tech-
nique proposed by Orloci (1966) was em-
ployed to ordinate vegetation units within
the different subcommunities listed above.
Through this technique the entities to be or-
dinated (i.e., plant species or stands of vege-
tation) are projected as points into n-dimen-
sional space. Such points are positioned by
attribute scores through the application of
the R and Q techniques of factor analysis.
Once established, this multidimensional array
of points is then reduced to a three dimen-
sional system. This is accomplished by select-
ing the two most different stands or species
and placing one at zero and the other at
some distance along the abscissa. All other
stands or species under consideration are
then positioned linearly in relationship to
these two extremes. This action thus estab-
lishes the X-axis. The above process is repeat-
ed until all points have been established in
three dimensional space (i.e., Y and Z axes
have been added). Coordinate values for the
X, Y, and Z axes are given as output from the
computer.

Interspecific association analysis. Expres-
sions of interspecific association were at-
tempted utilizing Cole's Index (1949). Step
one in the computation of the index involves
the accumulation of 2 X 2 contingency ta-
bles. Actual calculation of the index involves
the following three sets of formulas:

when ad = be:

C7± e =

ad - be

(a + c)(c + d)

(a + b)(b + d) n(a + b)(b + c)
when be > ad and d = a:

C7± c =

ad - be

(b + d)(c + d)

(a + b)(a + c) n(a + b)(a + c)

142

Great Basin Naturalist

Vol. 43, No. 1

when be > ad and a > d:

f^ , _ ad - be (a + b)(a + c)

(b + d)(e + d) n(b + d)(e + d)

where C7 = Cole's Index of Interspecific
Association

c = standard deviation Cole's
Index

n = total number of samples
and a, b, c, and d represent the four cells of
the 2x2 contingency table.

Tests of statistical significance were per-
formed by means of the Chi-square test. The
chi-squares were computed by the formula:

X2 =

(ad - bc)2n

(a + b)(a + c)(c + d)(b + d)

where X^ = Chi-square value
n = number of samples
and a, b, c, and d represent the different cells
of the 2x2 contingency table.

In all cases a single degree of freedom was
used. Chi-square values greater than 3.84
were considered to be significant at the 5
percent level, and values greater than 6.63
were considered to be significant at the 1
percent level.

Data representation. Graphic representa-
tion of data obtained from topographic stud-
ies and from ordination analysis was drawn
by the computer. Such representation was
accomplished through the use of a plotting
technique developed and programmed by
Mr. Howard Jesperson, Agricultural Experi-
mental Station, Iowa State University.

Results and Discussion

Species composition

Information on species sampled in the up-
land regions of Kalsow Prairie is presented in
Table 1. Cover, composition (i.e., based on
cover), and frequency values of Sporobolus
heterolepis, Andropogon gerardi, Poa pra-
tensis, and Panicum leibergii indicate these
are the dominant grasses of the upland sites.
Important or subdominant forbs include Sol-
idago canadensis, Solidago rigida, Helianthus
grosseserratus, Amorpha canescens. Aster eri-
coides, Desmodium canadense, Zizia aurea,
Helianthus laetiflorus. Aster laevis, Ratibida
pinnata, Ceanothus americanus, and Rosa
suffulta.

Average cover values (Table 1) ranged
from a high of 25.4 for Sporobolus heterolepis
to a low of 0.01 for several species. Percent-
age frequency values, on the other hand,
ranged from 73.1 for Andropogon gerardi to
0.1 for many species. No tests of correlation
were made between average cover values
and percentage frequency, but those species
showing the highest cover values generally
showed correspondingly higher percentage
frequency values.

Since Sporobolus heterolepis is the domi-
nant plant of the upland sites, Kalsow Prairie
is placed within the "Consociation" desig-
nated by Weaver and Fitzpatrick (1934) as
the Prairie Dropseed type {Sporobolus hetero-
lepis). Weaver and Fitzpatrick (1934) de-
scribed this particular consociation as being
the least extensive and least important tall-
grass subcommunity. It was found to occupy
drier upland sites and included the two sub-
cominants Stipa spartea and Schizachyrium
scoparius. Although these two species were
present (Table 1), they were not foimd in suf-
ficient quantity to be labeled subdominants.
The important grass species found with
Sporobolus heterolepis in this study (i.e., An-
dropogon gerardi, Poa pratensis, and Panicum
leibergii) suggest that the present-day upland
regions of Kalsow Prairie are vegetatively
distinct from the Prairie Dropseed Con-
sociation of similar areas described earlier by
Weaver. Both the species and their charac-
teristics suggest that this difference is due ei-
ther to change in the original vegetation, to
differences in community characteristics, or
to variations in the more recently glaciated
land. Poa pratensis, for example, is an in-
troduced species whose characteristics are
such that it is able to compete well within
the environment of prairie protected from
fire and, under conditions of grazing, mow-
ing, and other disturbance, is known to in-
crease in importance (Weaver 1954). An-
dropogon gerardi, on the other hand, is a
native grass described by Weaver and Fitz-
patrick (1934) as the dominant of the most
extensive tall-grass consociation that occu-
pied the lowlands and lower moist slopes of
the tall-grass prairie region.

Historical information, as well as evidence
obtained in this study, indicates that much of
the Kalsow Prairie has been subjected to

January 1983

Brotherson: Kalsow Prairie

143

Table 1. Cover, composition, and frequency percentages for species sampled on upland prairie sites.

Cover

Composition

Frequency

Frequency*

Species

(%)

Sporobolus heterolepis

25.42

27.81

66.7

85.0

Andropogon gerardi

15.93

17.43

73.1

87.5

Poa pratensis

12.36

13.52

54.5

90.0

Solidago canadensis

4.12

4.51

38.9

15.0

Solidago rigida

2.57

2.81

19.4

5.0

Panicum leibergii

2.56

2.80

34.0

12.5

Helianthus grosseserratus

2.30

2.52

31.6

10.0

Amorpha canescens

2.08

2.28

15.0

27.5

Aster ericoides

1.99

2.19

40.7

37.5

Desmodium canadense

1.66

1.82

23.0

5.0

Zizia aurea

1.62

1.77

36.1

67.5

Helianthus laetiflorus

1.49

1.63

11.4

7.5

Aster laevis

1.30

1.42

16.9

7.5

Ratibida pinnata

1.26

1.39

20.6

10.0

Ceanothus americanus

1.15

1.26

2.5

Rosa suffulta

1.07

1.17

18.4

45.0

Lysimachia hybrida

.84

.92

5.0

Convolvulus sepium

.76

.83

1.2

2.5

Silphium laciniatum

.72

.79

12.4

15.0

Achillea lanulosa

.68

.74

11.7

45.0

Galium obtusum

.65

.71

29.1

5.0

Spartina pectinate

.60

.66

10.3

17.5

Artemisia ludoviciana

.59

.65

4.4

2.5

Comandra umbellata

.59

.65

9.1

45.0

Schizachyrium scoparius

.52

.57

1.7

85.0

Fragaria virginiana

.52

.57

20.7

7.5

Physalis heterophylla

.47

.51

2.1

Elymus canadensis

.45

.49

19.4

12.5

Stipa spartea

.45

.49

6.6

20.0

Aster simplex

.43

.47

13.1

Muhlenbergia racemosa

.37

.40

11.2

Panicum virgatum

.36

.39

12.1

75.0

Senecio pauperculus

.36

.39

9.3

Lithospermum canescens

.35

.38

13.0

17.5

Heliopsis helianthoides

.31

.34

5.8

17.5

Psoralea argophylla

.29

.32

3.8

5.0

Solidago missouriensis

.28

.31

2.7

5.0

Apocynum sibiric^ .i

.27

.30

3.3

Asclepias tuberosa

.26

.28

3.5

2.5

Setaria viridis

.23

.25

Cirsium altissimum

.23

.25

4.6

17.5

Sorghastrum nutans

.21

.23

5.4

42.5

Liatris pycnostachya

.21

.23

9.8

27.5

Petalostemum purpureum

.20

.22

29.5

32.5

Pycnanthemum virginianum

.20

.22

7.0

5.0

Lythrum alatum

.20

.22

Phlox pilosa

.16

.18

6.3

10.0

Physalis virginiana

.16

.18

3.1

7.5

Viola pedatifida

.14

.15

5.5

5.0

Setaria lutescens

.13

.14

Viscia americana

.12

.13

4.7

7.5

Lathyrus venosus

.12

.13

2.5

5.0

Equisetum kansanum

.11

.12

4.7

42.5

Eryngium yuccifolium

.11

.12

1.0

2.5

Petalostemum candidum

.11

.12

3.9

7.5

^Figures taken from Moyer (1953) for comparison purposes.

144

Great Basin Naturalist

Vol. 43, No. 1

Table 1 continued.

Species

Cover

(%)

Composition

(%)

Frequency Frequency*

Baptisia leucophaea
Asclepias syriaca
Ambrosia artemisifolia
Baptisia leucantha
Carex gravida
Oxalis stricta
Teucrium canadense
Viola sp.

Gentiana andrewsii
Potentilla arguta
Scutellaria leonardii
Thalictrum dasycarpum
Lespedeza capitata
Solidago riddellii
Anemone cylindrica
Helenium autumnale
Pedicularis canadensis
Bouteloua curtipendula
Chenopodium album
Lathyrus palustris
Liatris aspera
Lycopus americanus
Lysimachia chiliata
Mentha arvensis
Solidago gymnospermoides
Vemonia fasciculata
Taraxacum officinale
Echinacea pallida
Agropyron repens
Agropyron smithii
Anemone canadensis
Arabis hirsuta
Asclepias sullivantii
Asclepias verticillata
Astragalus canadensis
Cicuta maculata
Helianthus maximiliani
Juncus tenuis
Lactuca scariola
Lysimachia quadrifhra
Panicum capillare
Phleum pratense
Rudbeckia hirta
Veronicastrum virginicum
Allium sp.
Aster novae-angliae
Cacalia tuberosa
Prenanthes racemosa
Solidago nemoralis
Trifolium pratense

.09

.10

15.0

.07

.08

1.4

.06

.07

1.4

.06

.07

.06

.07

2.0

.06

.07

.06

.07

1.7

.06

.07

3.2

.05

.06

1.1

2.5

.05

.06

.05

.06

2.2

.05

.06

1.6

7.5

.04

1.0

2.5

.04

2.4

.03

35.0

.03

1.2

22.5

.03

7.5

.02

1.1

.02

25.0

.02

1.4

5.0

.02

10.0

.01

1.2

2.5

.01

1.7

.01

5.0

.01

2.5

.01

10.0

.01

85.0

.01

2.6

2.5

.01

12.5

^Figures taken from Moyer (1953) for comparison purposes.

mowing, grazing to some extent, and abun-
dant pocket gopher activity. Both Mima
moimds and pocket gopher {Geomys bur-
sanins) activity are widely scattered across
the prairie. The Mima mounds are poorly un-
derstood areas of disturbance. Other disturb-
ance areas are along the south and west
boimdaries of the prairie, where dust from
adjacent plowed fields has been deposited in
depths up to two or three feet.

The distribution of soil types in the 8 ha
intensive study site is here of interest. Our
survey showed that much of the 8 ha is of
lowland soil types. In fact, a large part of the
upland prairie may occupy lowland soil
types. The disturbance will give possible ex-
planation to the high incidence of Poa pra-
tensis found with Sporobolus heterolepis, and
the large tracts of lowland soil types might
well explain the abundance of Andropogon

January 1983

Brotherson: Kalsow Prairie

145

gerardi. Why Sporobolus heterolepis is found
growing in such abundance on the lowland
areas is difficult to explain, but it might be
due to the high amount of calcareous soil
types found within Kalsow Prairie.

Early studies (Weaver and Fitzpatrick
1934, Shimek 1925) suggest the distribution
of Sporobolus heterolepis as restricted to
driest uplands. Because these areas often
show a lack of soil profile development or
outcroppings of parent material often high in
carbonates (Oschwald et al., 1965), it is fea-
sible that Sporobolus heterolepis is adapted to
grow on soils of high carbonate content and
that it might easily be extended to lowland
soils high in carbonate content.

Moyer (1953), in a study of the Kalsow
Prairie vegetation, gave percentage frequen-
cy values for many of the species included in
Table 1 of this paper. His figures are report-
ed in column four of Table 1 for comparison.
These figures suggest that there have been
some changes in the species composition of
the upland prairie since 1953. Some of the
species that show increases in percentage fre-
quency in the past are Solidago canadensis.

Solidago rigida, Panicum leibergii, Helianthus
grosseserratus, Desmodium canadense, Ga-
lium obtusum, and Fragaria virginiana. Spe-
cies that show decreases in percentage fre-
quency over this same period are Phleum
pratense, Poa pratensis, Zizia aurea, Rosa
suffulta, Schizachyrium scoparius, Panicum
virgatum, Sorghastrum nutans, Equisetum
kansanum. Anemone cylindrica, Liatris as-
pera, and Sporobolus heterolepis. Such
changes are not easily explained but might be
related to general fluctuations of the vegeta-
tion over a period of years, to fluctuations in
climatic conditions (i.e., time and duration of
rainfall, drought, etc.), to interspecific com-
petition, to differences in the technique and
intensity of sampling, and to the possible in-
fluence of slight disturbance upon the prairie
due to increased populations of pocket goph-
ers, dust accumulation from adjacent fields,
public visitors, and management practices.

To describe in greater detail the inter-
relationships of species in the upland prairie
a three-dimensional stand and species ordina-
tion treatment was attempted using Orloci's
(1966) method. The results are shown in Fig-
ures 2, 3, and 4. Data used in the ordination

Fig. 2. Three-dimensional ordination of 37 upland prairie sites.

146

Great Basin Naturalist

Vol. 43, No. 1

were from 444 samples taken from 37 sites in
the upland prairie community. These 37 sites
are shown as points in three-dimensional
space in Figure 2 and as dots in two-dimen-
sional space in Figure 3. Initially, attempts
were made to place the individual sites into
specific groups. Groups were designated on
the basis of where the stands fell when
plotted in three-dimensional space (i.e., those
sites which fell close together were consid-
ered to be the most similar and were placed
within the same group). Attempts to under-
stand the meaning of such groupings were
unsuccessful. Fiuther attempts to understand
the ordering pattern led to the conclusion
that discrete grouping within these upland
prairie regions is not feasible. It appears that
the ordering of the stands into three-dimen-
sional space was controlled by the response
of several of the major species to environ-
mental gradients. Of these species, Andropo-
gon gerardi (Fig. 5) and Sporobolus hetero-
lepis (Fig. 6) were plotted against the X and
Y coordinates of the ordination. As can be
seen, both species show continuous distribu-
tion in relationship to the axes. Stands
plotted near the origin and adjacent to the Y-
axis were found to be from drier sites, and
those found away from the origin were found
on wetter, more moist sites. These facts tend
to support the hypothesis that the vegetation
of the upland prairie is a continuum as ear-
lier described by Curtis (1955) and Dix and
Butler (1960). Kennedy (1969), in studying an
upland prairie in Guthrie County, Iowa, also
concluded that prairie vegetation there is
best described through the use of the contin-
uum-index concept.

Attempts at environmental factor correla-
tion with the ordination axes were not made
since only general information on environ-
mental gradients was available. It seems,
however, that these axes represent environ-
mental gradients and that the ordering of
stands or species along these axes is accom-
plished through the response of the different
stands or species to certain factors such as
moisture, texture, soil carbonates, or other
soil factors.

The species ordination is shown in Figure
4. Spartina pectinata and Ceanothus ameri-
canus are the most different entities on the
X-axis, and Andropogon gerardi is the most
distinct entity on the Y-axis. Other species
having distinct distribution patterns are Phys-
alis virginiana, Silphium laciniatum, Oxalis
stricta, Amorpha canescens, Solidago mis-
souriensis, Desmodium canadense, He-
lianthus grosseserratus. Aster ericoides, Vicia
americana, Pycnanthemum virginianum,
Ratibida pinnata. Aster laevis, and He-
lianthus laetiflorus. All other species either
showed no definite distribution patterns or
were too rare to establish a meaningful pat-
tern. The circles A, B, and C in Figure 4 rep-
resent the points where 76 of the 92 species
fell. This ordering of species has not deli-
neated associated groups but has pointed out

Fig. 3. Two-dimensional ordination of 37 upland
prairie sites. Factors responsible for pattern are
unknown.

Fig. 4. Two-dimensional ordination of species found
in upland prairie study sites, factors responsible for pat-
terns unknown: A, B, and C. = Clusters of species not
showing distinct distribution patterns, d. = Helianthus
laetiflorus. e. = Aster laevis. f. = Ratibida pinnata. g.
= Pycnanthemum virginianum. h. = Aster ericoides. i.
= Vicia americana. j. = Helianthus grosseserratus. k. =
Desmodium canadense. 1. = Amorpha canescens. m. =
Oxalis stricta. n. = Silphium laciniatum. o. = Physalis
virginiana. p. = Andropogon gerardi. q. = Spartina
pectinata.

January 1983

Brotherson: Kalsow Prairie

147

those species that exhibit pecuHar distribu-
tion patterns or that show a definite response
to one or several environmental stimuli.
Sanders (1969) found that the R-analysis of
Orloci (1966) did give him some fairly dis-
tinct groups of associated species as well as
groups of species that could not be consid-
ered associated. Collins (1968) used the tech-
nique to identify taxa that were distinct and
different and used them as indicator species
in his interpretation of the ecological rela-
tionships of fossil diatom populations. It is
evident from Figure 4 that the method has
not provided information on groups of associ-
ated species but rather has indicated taxa
that are distinct and therefore may have
some usefulness as indicator species.

Attempts to discover groups of positively
associated species within the upland regions
of Kalsow Prairie were made using Cole's In-
dex (1949). Those species showing positive as-
sociation with other taxa are shown in Table
2. A total of 298 significant associations were
found. Some species, such as Achillea lanu-
losa, Agropyron smithii, Amorpha canescens,
Andropogon gerardi, Asclepias tuberosa,
Carex gravida, Comandra umbellata, Helen-
ium autumnale, Lespedeza capitata, Phleum
pratense, and Solidago gymnospermoides, ex-
hibit positive association with only a limited
number of species. Other species, however,
show positive association with a large num-
ber of species. Some of these species are As-
ter ericoides, Desmodium canadense, Fragaria
virginiana, Galium obtusum, Helianthus
grosseserratus, Poa pratensis, Solidago cana-
densis, Solid go rigida, Sporobolus hetero-
lepis, and Zizia aurea. Many species showed

no significant association or expressed values
of high negative association. Positive values
of Cole's Index indicate that species occur to-
gether more often than would otherwise be
expected due to chance (Hale 1955, Hurlbert
1969). Therefore, through the use of such an
index one can deduce groups of species that
consistently show positive values of associ-
ation with one another. Figures 7, 8, and 9
were constructed from values taken from
Table 2 to illustrate the existence of such
groups within the upland prairie. In all three
cases one species was picked and the corre-
sponding figure was then built up around this
species.

Species Distribution Patterns

Eight hectares of the prairie adjacent to its
southern boundary (Figure 1) were selected
for intensive study of the distribution of plant
species in relation to soils and topography.
The area was chosen because it included
within its boundaries a representation of all
vegetation types occurring on Kalsow
Prairie. The area was staked on a 9 X 9 m
grid that placed 968 points within the 8 ha.
From these points all factors included in this
study were examined.

The presence of all plant species found in
the area was recorded in relation to each 27
square-meter section of the grid. From these
present figures, distribution maps for 160
species were constructed. Examples of these
maps are shown in Figures lOA through
lOHH. These figures illustrate examples of
distribution patterns often shared by several
species. Andropogon gerardi (Fig. lOD) illus-
trates a type of pattern typical of many spe-
cies commonly found in the upland prairie.

Fig. 5. Two-dimensional ordination of upland prairie
with percentage cover values of Sporobolus heterolepis
for each site shown relating directly to the Y-axis.

Fig. 6. Two-dimensional ordination of upland prairie
with percentage cover values of Andropogon gerardi for
each site shown relating directly to the X-axis.

148

Great Basin Naturalist

Vol. 43, No. 1

Table 2. Cole's Index values expressing positive interspecific association on upland prairie.

Species

X2a

Ct"

o^-

Achillea lanulosa

Agropyron repens
Agropyron smithii

Ambrosia artemisifolia

Amorpha canescens

Andropogon gerardi

Schizachyrium scoparius

Anemone canadensis

Apocynum sibiricum

Artemisia ludoviciana

Asclepias sullivantii

Andropogon gerardi
Aster ericoides
Poa pratensis
Solidago rigida
Sporobolus heterolepis
Carex gravida
Convolvulus septum
Physalis heterophylla
Andropogon gerardi
Aster ericoides
Galium obtusum
Helianthus grosseserratus
Muhlenbergia racemosa
Petalostemum purpureum
Pycnanthemum virginianum
Solidago canadensis
Solidago riddellii
Asclepias syriaca
Helianthus grosseserratus
Senecio pauperculus
Setaria lutescens
Setaria viridis
Stipa spartea
Taraxacum officinale
Andropogon gerardi
Panicum leibergii
Solidago rigida
Sporobolus heterolepis
Aster ericoides
Poa pratensis
Solidago canadensis
Sporobolus heterolepis
Comandra umbellata
Lithospermum canescens
Panicum leibergii
Petalostemum purpureum
Phlox pilosa
Solidago rigida
Sporobolus heterolepis
Zizia aurea
Artemisia ludoviciana
Petalostemum candidum
Poa pratensis
Solidago missouriensis
Aster simplex
Calamagrostis canadensis
Carex aquatilis
Carex retrorsa
Carex lasiocarpa
Fragaria virginiana
Heliopsis helianthoides
Polygonum coccimeum
Spartina pectinata
Aster ericoides
Convolvulus sepium
Helianthus laetiflorus
Poa pratensis
Elymus canadensis

18.77

.73

.17

6.84

.22

.08

5.03

.27

.11

13.41

.21

.05

11.06

.51

.15

4.91

.22

.09

7.04

.44

.16

5.31

.22

.09

5.96

.73

.29

8.97

.45

.15

4.33

.34

.16

5.43

.43

.18

24.25

.27

.05

8.26

.20

.06

5.65

.18

.07

13.22

.53

.14

25.22

.24

.04

83.22

.43

.04

5.12

.23

.10

155.77

.77

.06

58.48

.28

.03

165.69

.57

.04

4.00

.24

.12

172.49

.55

.04

15.62

.51

.13

28.29

.24

.04

19.00

.20

.04

41.54

.77

.11

126.73

.18

.01

176.68

.30

.02

85.47

.15

.01

238.41

.46

.02

10.40

.21

.06

19.30

.35

.08

5.62

.41

.17

7.53

.18

.06

11.12

.19

.05

6.01

.25

.10

8.23

.77

.26

5.23

.31

.13

5.53

.19

.08

6.16

.19

.07

5.57

.80

.33

25.23

.16

.03

14.68

.26

.06

102.95

.70

.06

44.71

.35

.05

70.51

.55

.06

66.57

.39

.04

3.83

.38

.19

4.39

.27

.12

15.26

.18

.04

13.73

.18

.05

4.53

.42

.19

51.32

.22

.03

14.98

.28

.07

15.37

.78

.19

5.94

.67

.27

*Chi -square

"Cole's Index

^Standard deviation Cole's Index

January 1983

Table 2 continued.

Brotherson: Kalsow Prairie

149

Species

X2a

<if

Asclepias syriaca

Asclepias tuberosa

Asclepias verticillata
Aster ericoides

Aster laevis

Aster simplex

Bouteloua curtipendula

Calamagrostis canadensis

Carex atherodes

Carex aquatilis

Carex gravida

Aster simplex
Cirsium altissimum
Desmodium canadense
Equisetum kansanum
Lithospermum canescens
Rosa suffulta
Senecio pauperculus
Taraxacum officinale
Thalictrum dasycarpum
Aster ericoides
Desmodium canadense
Elymus canadensis
Sporobolus heterolepis
Zizia aurea

Lithospermum canescens
Helianthus grosseserratus
Poa pratensis
Solidago canadensis
Sporobolus heterolepis
Zizia aurea

Desmodium canadense
Panicum leibergii
Poa pratensis
Sporobolus heterolepis
Calamagrostis canadensis
Carex gravida
Carex retrorsa
Fragaria virginiana
Galium obtusum
Helianthus grosseserratus
Poa pratensis
Senecio pauperculus
Silphium laciniatum
Spartina pectinata
Comandra umbellata
Helianthus laetiflorus
Phlox pilosa
Carex aquatilis
Carex retrorsa
Carex lasiocarpa
Phalaris arundinacea
Polygonum coccineum
Spartina pectinata
Carex retrorsa
Polygonum coccineum
Scirpus fluviatilis
Carex retrorsa
Carex lasiocarpa
Phalaris arundinacea
Polygonum coccineum
Spartina pectinata
Desmodium canadense
Fragaria virginiana
Galium obtusum
Helianthus grosseserratus
Liatris pycnostrachya
Muhlenbergia racemosa
Petalostemum purpureum

4.71

.38

.17

5.77

.21

.08

9.84

.66

.21

6.87

.21

.08

8.15

.31

.10

9.99

.32

.10

47.45

.43

.06

25.23

.20

.04

12.88

.23

.06

4.80

.27

.12

17.17

.42

.10

9.21

.29

.09

17.15

.92

.22

7.13

.30

.11

14.39

1.00

.26

17.76

.17

.03

105.26

.46

.04

49.17

.22

.03

70.97

.49

.05

40.95

.19

.02

23.69

.30

.06

53.78

.36

.04

27.26

.54

.10

9.76

.41

.13

42.41

.21

.03

34.75

.21

.03

30.80

.17

.03

4.89

.19

.08

54.31

.37

.05

95.05

.56

.05

7.48

.58

.21

61.08

.19

.02

32.99

.29

.05

34.20

.32

.05

5.23

.29

.12

7.18

.41

.15

7.47

.31

.11

427.86

.50

.02

580.80

.74

.03

404.31

.45

.02

171.76

.20

.01

64.45

.18

.02

154.17

.30

.02

18.21

.27

.06

370.61

.87

.04

145.67

.34

.02

469.33

.88

.04

406.70

.59

.02

57.45

.16

.01

33.93

.17

.02

66.04

.26

.03

17.26

.49

.11

30.76

.64

.11

21.59

.56

.11

36.90

.71

.11

8.21

.22

.07

4.90

.23

.10

9.82

.23

.07

*Chi-square

"Cole's Index

^Standard deviation Cole's Index

150

Table 2 continued.

Great Basin Naturalist

Vol. 43, No. 1

Species

X2a

07C

Carex lasiocarpa

Carex retrorsa

Cirsium altissimum

Comandra umbellata

Convolvulus sepium
Desmodium canadense

Elymus canadensis

Equisetum kansanum

Eryngium yuccifolium

Fragaria virginiana

Silphium laciniatum
Solidago canadensis
Lathyrus palustris
Lysimachia hybrida
Phalaris arundinacea
Polygonum coccineum
Spartina pectinata
Carex lasiocarpa
Phalaris arundinacea
Polygonum coccineum
Spartina pectinata
Fragaria virginiana
Galium obtustim
Helianthus grosseserratus
Petalostemum candidum
Physalis virginiana
Solidago canadensis
Desmodium canadense
Elymus canadensis
Fragaria virginiana
Panicum leihergii
Petalostemum purpureum
Poa pratensis
Hatibida columnifera
Solidago rigida
Sporobolus heterolepis
Zizia aurea
Solidago nemoralis
Poa pratensis
Elymus canadensis
Fragaria virginiana
Galium obtusum
Helianthus grosseserratus
Muhlenbergia racemosa
Poa pratensis
Solidago rigida
Sporobolus heterolepis
Zizia aurea
Fragaria virginiana
Galium obtusum
Poa pratensis
Sporobolus heterolepis
Heliopsis helianthoides
Lithospermum canescens
Petalostemum candidum
Phlox pilosa
Sporobolus heterolepis
Zizia aurea
Panicum leibergii
Rosa suffulta
Solidago rigida
Sporobolus heterolepis
Galium obtusum
Helianthus grosseserratus
Muhlenbergia racemosa
Poa pratensis
Solidago canadensis
Sporobolus heterolepis
Zizia aurea

19.94

.30

.06

6.94

.57

.21

110.24

1.00

.09

147.33

1.00

.08

92.24

.21

.02

27.12

.17

.03

72.67

.30

.03

465.71

.50

.02

158.23

.20

.01

79.04

.20

.02

117.52

.27

.02

8.78

.21

.07

7.65

.29

.10

8.53

.34

.11

8.80

.15

.05

24.36

.28

.05

5.09

.21

.09

47.92

.50

.07

11.91

.23

.06

15.97

.26

.06

16.18

.24

.05

19.78

.18

.03

10.28

.39

.12

26.47

.26

.05

15.47

.23

.05

15.25

.62

.15

28.32

.43

.08

32.92

.17

.02

5.30

.66

.28

80.81

.27

.02

75.91

.40

.04

81.55

.43

.04

26.91

.24

.04

31.07

.23

.04

38.84

.34

.05

20.28

.21

.04

99.46

.70

.06

66.09

.29

.03

72.47

.27

.03

21.27

.26

.05

44.15

.39

.05

42.70

.49

.07

7.25

.19

.07

20.66

.20

.04

11.42

.21

.06

6.58

.19

.07

20.81

.83

.18

11.12

.31

.09

7.80

.31

.11

10.29

.22

.06

4.57

.24

.11

7.90

.85

.30

69.59

.39

.04

40.39

.31

.04

37.67

.26

.04

29.55

.33

.05

13.28

.33

.08

39.10

.48

.07

4.89

.18

.08

*Chi-square

''Cole's Index

'^Standard deviation Cole's Index

January 1983

Table 2 continued.

Brotherson: Kalsow Prairie

151

Species

X2a

Galium obtusum

Gentiana andrewsii

Helenium autumnale

Helianthus grosseserratus
Helianthus laetiflorus

Helianthus maonmiliana
Heliopsis helianthoides

Lactuca scariola
Lathyrus palustris

Lespedeza capitata

Liatris aspera
Liatris pycnostachya

Lithospermum canescens
Lycopus americanus
Lysimachia hybrida

Lysimachia quadriflora

Helianthus grosseserratus
Muhlenbergia racemosa
Silphium laciniatum
Solidago canadensis
Zizia aurea

Heliopsis helianthoides
Liatris pycnostachya
Lithospermum canescens
Zizia aurea

Helianthus grosseserratus
Lythrum alatum
Muhlenbergia racemosa
Poa pratensis

Pycnanthemum virginianum
Senecio pauperculus
Solidago canadensis
Solidago canadensis
Panicum leibergii
Phlox pilosa
Sporobolus heterolepis
Scutellaria leonardii
Taraxacum officinale
Poa pratensis

Pycnanthemum virginianum
Ratibida pinnata
Solidago canadensis
Pedicularis canadensis
Rosa suffulta
Lysimachia hybrida
Senecio pauperculus
Silphium laciniatum
Spartina pectinata
Viola sp.

Lithospermum canescens
Panicum leibergii
Rosa suffulta
Silphium laciniatum
Physalis virginiana
Poa pratensis
Silphium laciniatum
Solidago canadensis
Sporobolus heterolepis
Zizia aurea
Panicum leibergii
Sporobolus heterolepis
Zizia aurea
Lythrum alatum
Senecio pauperculus
Spartina pectinata
Muhlenbergia racemosa
Polygonum coccineum
Scirpus fluviatilis
Spartina pectinata
Viola sp.

Muhlenbergia racemosa
Pedicularis canadensis
Petalostemum purpureum
Poa pratensis
Senecio pauperculus

91.71

.35

.03

25.27

.21

.04

41.48

.17

.02

13.09

.31

.08

5.17

.18

.07

12.46

.45

.12

4.04

.26

.13

4.52

.40

.18

4.34

.71

.34

15.18

.88

.22

48.25

.19

.02

10.43

.22

.06

11.28

.86

.25

7.30

.25

.09

40.42

.62

.09

5.32

.41

.17

73.49

.21

.02

42.24

.37

.05

25.63

.18

.03

5.53

.36

.15

11.06

.19

.05

11.06

.19

.05

6.77

.52

.19

20.89

.32

.07

6.37

.21

.08

8.97

.41

.13

12.13

.19

.05

7.31

.35

.12

35.58

.24

.04

3.94

.21

.10

10.63

.46

.14

14.78

.39

.10

6.30

.23

.08

12.60

.45

.12

3.80

.38

.19

5.52

.28

.11

5.93

.28

.11

21.69

.40

.08

6.21

.24

.09

17.79

.19

.04

17.40

.28

.06

39.56

.78

.12

26.25

.32

.06

26.20

.25

.04

37.09

.81

.13

23.59

.33

.06

52.06

.21

.02

18.03

.43

.10

19.71

.43

.09

5.77

.60

.25

53.59

1.00

.13

95.59

.83

.08

13.03

.64

.17

9.00

.31

.10

8.08

.22

.07

54.24

.50

.06

4.08

.20

.09

4.41

.63

.30

17.76

.48

.11

■Chi-square

"Cole's Index

(^Standard deviation Cole's Index

152

Great Basin Naturalist

Vol. 43, No. 1

Table 2 continued.

Species

X2a

C^b

Lythrum alatum

Mentha arvensis

Muhlenbergia racemosa
Panicum capillare

Panicum leibergii
Panicum virgatum
Pedicularis canadensis

Petahstemum candidum
Petalostemum purpureum

Phalaris arundinacea

Phleum pratense

Phlox pilosa

Physalis virginiana

Poa pratensis
Polygonum coccineum
Potentilla arguta
Psoralea argophylla
Pycnanthemum virginianum

Ratibida pinnata

Rosa suffulta
Rudbeckia hirta
Senecio pauperculus

Setaria lutescens
Silphium laciniatum

Solidago canadensis
Solidago gymnospermoides

Senecio pauperculus
Spartina pectinata
Phalaris arundinacea
Polygonum coccineum
Zizia aurea

Pycnanthemum virginianum
Senecio pauperculus
Silphium laciniatum
Solidago canadensis
Zizia aurea
Poa pratensis
Sporobolus heterolepis
Poa pratensis
Sporobolus heterolepis
Pycnanthemum virginianum
Senecio pauperculus
Solidago rigida
Zizia aurea
Ratibida pinnata
Rosa suffulta
Poa pratensis
Solidago canadensis
Solidago rigida
Sporobolus heterolepis
Zizia aurea

Polygonum coccineum
Spartina pectinata
Phlox pilosa
Ratibida pinnata
Ratibida pinnata
Sporobolus heterolepis
Rosa suffulta
Solidago rigida
Sporobolus heterolepis
Scirpus fluviatilis
Solidago missouriensis
Stipa spartea
Senecio pauperculus
Silphium laciniatum
Solidago canadensis
Sporobolus heterolepis
Zizia aurea
Sporobolus heterolepis
Zizia aurea
Sporobolus heterolepis
Solidago rigida
Solidago canadensis
Taraxacum officinale
Setaria viridis
Solidago canadensis
Spartina pectinata
Sporobolus heterolepis
Viola sp.
Zizia aurea
Sporobolus heterolepis
Solidago rigida
Sporobolus heterolepis
Zizia aurea

5.70

.27

.11

6.37

.28

.11

6.67

.21

.08

6.64

.30

.11

7.48

.25

.09

20.53

.20

.04

27.20

.24

.04

12.59

.20

.05

6.31

.21

.08

11.91

.28

.08

12.96

.24

.06

56.41

.65

.08

12.96

.24

.06

56.41

.65

.08

37.72

.49

.07

5.78

.22

.09

5.11

.29

.12

20.45

.79

.17

4.02

.21

.10

4.27

.23

.10

11.86

.34

.09

16.33

.28

.06

20.61

.22

.04

30.35

.71

.12

24.91

.33

.06

78.87

.41

.04

25.65

.26

.05

5.50

.34

.14

7.39

.73

.26

8.27

.25

.08

7.60

.64

.23

5.93

.21

.08

4.93

.32

.14

54.66

.31

.04

350.08

.37

.01

5.48

.30

.12

6.50

.18

.07

30.62

.19

.03

25.56

.21

.04

25.73

.31

.06

11.25

.39

.11

7.48

.69

.25

22.69

.48

.10

19.66

.23

.05

13.85

.54

.14

7.17

.75

.28

51.17

.42

.05

100.89

.32

.03

525.50

.89

.03

4.85

.34

.15

15.95

.21

.05

32.83

.52

.09

38.07

.19

.03

43.57

.31

.04

21.62

.28

.06

3.87

.22

.11

8.50

.86

.29

9.77

.47

.15

*Chi-square

"Cole's Index

'^Standard deviation Cole's Index

January 1983

Brotherson: Kalsow Prairie

153

Table 2 continued.

Species

X2a

Solidago rigida
Sorghastrum nutans

Sporobolus heterolepis

Viola sp.

Panicum itnplicatum

Sporobolus heterolepis
Sporobolus heterolepis
Zizia aurea
Zizia aurea
Zizia aurea
Solidago nemoralis

96.94

.84

.08

17.68

.81

.19

5.46

.23

.09

130.17

.19

.01

8.97

.35

.11

8.63

.19

.06

^hi-square

"Cole's Index

'^Standard deviation Cole's Index

This pattern indicates that these species have
wide ecological amplitudes and are limited
basically by conditions peculiar to the drain-
age areas of the prairie. Other species that
showed similar distribution patterns were As-
ter ericoides, Elymus canadensis, Equisetum
kansanum, Lithospermum canescens, Peta-
lostemum purpureum, Poa pratensis, Ratibida
pinnata, Rosa suffulta, Solidago canadensis,
Solidago rigida, Sporobolus heterolepis, and
Zizia aurea.

A pattern closely resembling that of An-
dropogon gerardi but also showing limited
distribution on the higher and drier ridges of
the area is that exemplified by Silphium la-
ciniatum (Fig. lOJ). Species included under
this type of pattern were Desmodium cana-
dense, Fragaria virginiana, Galium obtusum,
Helianthus grosseserratus, Heliopsis helian-
thoides, Liatris pycnostachya, Panicum virga-
tum, and Spartina pectinata.

The pattern showed by Ambrosia artemisi-
folia (Fig. lOB) is limited to the border weed
communities. Other species found limited to

Fig. 7. Solidago rigida and associated species of up-
land prairie as determined by Cole's (1949) Index, the
more lines between species, the greater the association.
So ri = Solidago rigida, Ru hi = Rudbeckia hirta, Pe ca
= Pedicularis canadensis, Ph vi = Physalis virginiana.
An Sc = Schizachyrium scoparius, Er yu = Eryngium
yuccifolium, Pe pu = Petalostemum purpureum. So gy
= Solidago gymnospermoides, Sp he = Sporobolus het-
erolepis, Ac la = Achillea lanulosa. Am ca = Amorpha
canescens, De ca = Desmodium canadense, Co um =
Comandra umbellata.

these areas were Amaranthus tamariscinus.
Ambrosia trifida, Brassica nigra, Chenopo-
dium album, Helianthus annuus. Polygonum
pennsylvanicum. Polygonum persicaria, Se-
taria lutescens, and Setaria viridis.

Figure IOC (Amorpha canescens) illustrates
a pattern common to species limited to
growth on the ridges and lower slopes of the
area. This would correspond to areas com-
posed mainly of Clarion, Nicollet, and Web-
ster soil types (Fig. 11). When compared with
the pattern exhibited by Andropogon gerardi,
this type shows a narrowing ecological am-
plitude and decrease in the ability of species
exhibiting this type of pattern to compete in
lowland areas. Other species showing this
type of pattern were Achillea lanulosa,
Arabis hirsuta, Asclepias syriaca, Asclepias
tuberosa. Aster laevis, and Panicum leibergii.

Several species found limited in distribu-
tion to the mid- and upland slopes of the
prairie exemplify the pattern shown by Soli-
dago nemoralis (Fig. lOE). These species
were Eryngium yuccifolium, Solidago gym-
nospermoides, Solidago riddellii, and Viola
pedatifida. Such species show rather narrow
ecological amplitudes when compared with
the groups discussed earlier.

Fig. 8. Amorpha canescens and associated species of
upland prairie as determined by Cole's (1949) Index, the
more lines between the species, the greater the associ-
ation. Am ca = Amorpha canescens. Pa li = Panicum
leibergii, An ge = Andropogon gerardi, Sp he = Sporo-
bolus heterolepis.

154

Great Basin Naturalist

Vol. 43, No. 1

Another group exhibiting rather narrow
ranges in distribution are characterized by
the patterns shown in Figures ION and 100.
These species, Artemisia ludoviciana, Cea-
noihus americanus. Echinacea pallida, He-
lianthus laetiflorus, Lathyrus venosus, Les-
pedeza capitata, Liatris aspera, Petalostemum
candidum, Potentilla arguta, Psoralea argo-
phylla, Solidago missouriensis, and Stipa
spartea, are found occupying the ridges and
drier sites of the prairie. This would corre-
spond to the Clarion, Clarion-Nicollet, and
Nicollet areas of Figure 11.

A finai group of species limited from
growth in the drainage areas of the prairie
show a pattern characteristic of those found
in Figure lOA (Agropyron smithii) and Figure
lOF (Helenium autumnale). Here again the
ecological amplitudes of these species are
narrow when compared with Andropogon ge-
rardi or Sporobolus heterolepis. As can be
seen, the distribution of these species corre-
sponds closely to the borders of the pothole
and drainage complex; thus these species
mainly occupy soils that are characterized by
being highly calcareous to the surface. Other
species exhibiting this type of distribution are
Agrostis alba. Aster simplex, Lycopus ameri-
canus, Lysimachia quadriflora, Lythrum ala-
tum, Senecio pauperculus and Viola sp.

Species restricted in occurrence to the
potholes and drainage ways of the area were
found to exhibit two types of distributional
patterns. The first, shown by Calamagrostis

Fig. 9. Aster ericoides and associated species of up-
land prairie as determined by Cole's (1949) Index, the
more lines between the species, the greater the associ-
ation. As er = Aster ericoides, Ar lu = Artemisia ludovi-
ciana. He gr = Helianthus grosseserratus, Po pr = Poa
pratensis. As tu = Asclepias tuberosa, Sp he = Sporo-
bolus heterolepis, Zi au = Zizia aurea. An gr = An-
dropogon gerardi. So ca = Solidago canadensis, Ac la =
Achillea lanulosa, Ag sm = Agropyron smithii.

canadensis in Figure lOF, corresponds gener-
ally to the shallower areas of the drainage
system. The pattern shown by Figure lOF
also includes the species Apocynum sibiri-
cum, Asclepias incamata, Carex aquatilis,
Carex lasiocarpa, Carex retrorsa, Phalaris
arundinacea, Teucrium canadense, and Ver-
nonia fasiculata. The areas covered by these
species correspond generally to the Glenco
soils as shown in Figure 11. The second, illus-
trated by Carex atherodes and Scirpus fluvia-
tilis in Figure lOG and lOH, is more restrict-
ed in extent than the above and corresponds
to the deeper areas within the drainage sys-
tem. Species occupying areas equivalent to
those shown in Figures lOG and lOH were
Lysimachia hybrida. Polygonum coccineum,
and Mentha arvensis. These areas correspond
to Glenco-Okoboji and Okoboji soil locations
as shown in Figure 11.

In several cases it was noted that two spe-
cies belonging to the same genus showed op-
posing patterns of distribution. Examples of
this phenomenon are illustrated by the spe-
cies Aster laevis and Aster simplex. Figures
lOK and lOL; Helianthus grosseserratus and
Helianthus laetiflorus. Figures lOS and lOT;
and Liatris aspera and Liatris pycnostachya.
Figures lOU and lOV.

Other species were shown to have patterns
corresponding to the distribution of Mima
mounds found within the area. Such patterns
are shown by Convolvulus sepium (Fig. lOF)
and by Oxalis stricta (Fig. lOY).

Many factors affect the distribution of a
species within the community. It has been
shown that individuals of different taxa sel-
dom have identical spatial arrangements
within an area (Greig-Smith 1964), yet, as
shown above, the distribution patterns of
some species may be similar and often show
overlapping boundaries. Such species may be
closely associated due to preferences for sim-
ilar microenvironments or, as in the case of
Andropogon gerardi, because of wide ecologi-
cal amplitude. Generally these differences in
the local distribution of species have been at-
tributed to local microenvironments (i.e.,
Mima mounds, animal burrows, ridge tops,
and drainage ways), interspecific competition
(i.e., allelopathy, shade tolerance, etc.), spe-
cies biology (i.e., modes of reproduction, seed
dispersal, immigration rates, etc.), or one to

January 1983

Brotherson: Kalsow Prairie

155

,-^#

........ 1

. ..llll ' 'r.'rl'".'.:

'■ ;

• ;i:Pi!rrt:

.;!-=

-" (- •

: ' /^^^ —

^~^c^

^~^'

■-=••/..::■

=-"^

•fee

kC^'-

II:...

-"C^jl

it :r:H:

iiiiiic;

~i::hl-" :

lit..'"

::lnii.v

....

•■■

-^ ..

::•

{ \

\\ •::

' r^^

rC^...

'i ■•:

-J '■

ilj:

^^j

:r^:r-

JZ^H. J •

." • jf^r^

■^

•>i

"^l:"'-'

'■^v

-.1

^iif

jp> -IV

^iiii:;

2ii ^

:::y

iu.j^-";:;ij'

--;^

Fig. 10. Distribution patterns in 8 hectare study area of: A. Agropyron smithii. B. Ambrosia artemisifolia. C.
Amorpha canescens. D. Andropogon gerardi. E. Solidago nemoralis. F. Calamagrostis canadensis. G. Carex atherodes.
H. Scirpus fluviatilis. I. Helenium autumnali. J. Silphium laciniatum. K. Aster iaeuis. L. Aster simplex. M. Apocynum
sibericum. N. Artemisia ludoviciana. O. Ceanothus americanus. P. Conuo/uuZus sepium. Q. Desmodium canadense. R.
Fragaria virginiana. S. Helianthus grosseserratus. T. Helianthus laetiflorus. U. Liatris aspera. V. Liatris pyonostachya.
W. Lisimachia hybrida. X. Lycopus americanus. Y. OxaZis striata. Z. Panicum leibergii. AA. Polygonum coccineum.
BB. Phalris arundinacea. CC. Psoralea aryophylla. DD. Senecio aurens. EE. Spartina pectinata. FF. Sporobolus heter-
olepis. GG. Teucrium canadense. HH. Vemonia fasciculata.

156

Great Basin Naturalist

Vol. 43, No. 1

Fig. 11. Soil series map of 20-acre intensive study area, abbreviations described in Methods section. A = Clarion,
A~ = Clarion-Nicollet, N = Nicollet, Na = calcareous Nicollet, N~ = Nicollet-Webster, Na~ = calcareous Nicol-
let-Webster, W = Webster, Wh = heavy Webster, Wa = calcareous Webster, C = Canisteo, H = Harps, H" =
Harps-Canisteo, C" = inverted Canisteo-heavy Webster, G = Glenco, Ga = calcareous Glenco, GO = Glenco-
Okoboji, O = Okoboji.

several edaphic factors (soil and water re-
gimes, macronutrients, micronutrients, tex-
ture, organic matter, etc.) (Curtis 1959,
Greig-Smith 1964, Kershaw 1964). From this
we can conclude that species showing similar
patterns of distribution may be equally well
adapted in their response to one or more en-
vironmental stimuli and yet differ greatly in
their basic ecological amplitudes. The re-
sponse of individuals to the environmental

complex is measured in a species distribution
pattern as well as in its importance within
the community.

Attempts were made to access the response
of the species included in this study to the
factors of soil and topography. Soil and ele-
vation readings were recorded at all 968
points of the grid. From these readings a soils
map (Figure 11) and contour and elevation
maps (Figs. 12, 13) were constructed for the

Fig. 12. Topographic map of 20-acre intensive study area.

WEST-«-

January 1983

Brotherson: Kalsow Prairie

157

8 ha plot. This made it possible to group all
plant samples according to 0.5 ft changes in
elevation or according to soil series. Once
grouped, average cover values were com-
puted for all participating species and re-
corded in Tables 3 (elevation data) and 4
(soils data). As can be seen from these tables,
all species showed response to these factors.
Several species, Andropogon gerardi,
Amorpha canescens. Aster ericoides, Elymus
canadensis, Panicum leibergii, Poa pratensis,
Solidago canadensis, Sporobolus heterolepis,
and Zizia aurea, showed wide tolerance in
relation to both soil and elevation, but all ex-
hibited peak^ or plateaus of occurrence.
These peaks or plateaus are interpreted to
represent the optimum conditions under
which a particular species can reach its high-
est importance within the community in rela-
tion to the entire species complex.

Other species showed rather narrow ranges
of tolerance. Some of these were Schiz-
achyrium scoparius, Apocynum sibiricum,
Asclepias sullivantii, Calamagrostis cana-
densis, Carex atherodes, Eryngium yuccifo-
lium, Lysimachia hybrida, Physalis hetero-
phylla, Viola pedatifida and Ceanothus
americanus. Those exhibiting narrow ranges
also showed peaks of occurrence. For species
exhibiting narrow tolerances, four basic types
of distribution patterns as related to elevation
(Table 3) are recognizable: (1) pothole and

drainage, (2) lower slopes, (3) mid- and upper
slopes, and (4) ridges.

For species showing response to the soil
factor (Table 4) three basic classes are recog-
nizable: (1) Glenco, Glenco-Okoboji, and
Okoboji, (2) calcareous, and (3) noncalcareous
and ridge. Species indicating preference for
class 1 were Calamagrostis canadensis, Carex
atherodes, Carex aquatilis, Carex lasiocarpa,
Carex retrorsa, Lysimachia hybrida. Poly-
gonum coccineum, and Scirpus fluviatilis.
Species showing preference for the cal-
careous soils (class 2) were Agropyron smithii,
Desmodium canadense, Galium obtusum,
Helenium autumnale, Petalostemum purpu-
reum, Senecio pauperculus, Silphium lacinia-
tum, Solidago canadensis, Solidago nemoralis
and Solidago riddellii. Examples of species
preferring class 3 were Amorpha canescens,
Artemisia ludoviciana, Asclepias tuberosa,
Baptisia leucophaea, Eryngium yuccifolium,
Lathyrus palustris, Panicum leibergii, Poa
pratensis, Solidago missouriensis, Vicia ameri-
cana, and Ceanothus americanus.

These groups of recognizable patterns,
each involving several species, suggest the ex-
istence of subcommunities within the prairie
area. To ascertain the existence of such com-
munities, the data from Tables 3 and 4 were
treated using Orloci's (1966) method of ordi-
nation. When the results from the soils analy-
sis were plotted (Fig. 14), four basic groups

Fig. 13. Topographic map of 20-acre intensive study
area plotted by computer.

IQ y .

9 of

Fig. 14. Two-dimensional ordination of vegetation
found on the different soil types in the 20-acre study
area; cluster A indicates vegetation on Nicollet and Ni-
collet-Webster soil types; B indicates vegetation on
Clarion, Clarion-Nicollet, Webster, heavy Webster, cal-
careous Nicollet, calcareous Nicollet-Webster, cal-
careous Webster, Canisteo, Harps, Harps-Canisteo, and
inverted Canisteo-heavy Webster soil types; C indicates
vegetation on Glenco-Okoboji and Okoboji soil types; D
indicates vegetation on Glenco and calcareous Glenco.

158 Great Basin Naturalist Vol. 43, No. 1

Table 3. Average cover values for species in relation to elevation in 20-acre intensive study area.

12 3 4 5 6

Species .7-1.2 1.3-1.8 1.9-2.4 2.5-3.0 3.1-3.6 3.7-4.2

Achillea lanulosa

Agropyron smithii

Ambrosia artemisifolia

Amorpha canescens

Andropogon gerardi

Schizachyrium scoparius

Anemone canadensis

Anemone cylindrica

Apocynum sibiricum

Arabis hirsuta

Artemisia ludoviciana

Asclepias incamata

Asclepias sullivantii

Asclepias syriaca

Asclepias tuberosa

Aster ericoides

Aster laet>is

Aster simplex

Baptisia leucantha

Baptisia leucophaea

Calamagrostis canadensis

Carex atherodes

Carex aquatilis

Carex gravida

Carex lasiocarpa

Carex retrorsa

Chenopodium album

Cicuta maculata

Cirsium altissimum

Comandra umbellata

Convolvulus sepium

Desmodium canadense

Elymus canadensis

Equisetum kansanum

Eryngium yuccifolium

Fragaria virginiana

Galium obtusum

Gentiana andrewsii

Helenium autumnale

Helianthus grosseserratus

Helianthus laetifhrus

Helianthus maximiliani

Heliopsis helianthoides

Lactuca scariola

Lathyrus palustris

Lathyrus venosus

Lespedeza capitata

Liatris pycnostachya

Lithospermum canescens

Lycopus americanus

Lysimachia chiliata

Lysimachia hybrida

Lysinwchia quadriflora

Lythrum alatum

Mentha arvensis

Muhlenbergia racemosa

Oxalis stricta

Panicum capillare .01

.23

.28

.02

.12

.09

.01

.55

.20

1.49

4.27

.52

5.57

9.06

9.49

10.10

.25

.92

.75
.02

.55
.03

1.36

1.98
.52

1.15

.65
.08

.04
.01
.06

.02

.19
.02

.13

.07

.28

.02

.44

.71

.41

1.87

2.09

1.72

.10

.77

1.84

.56

2.72

2.55

1.66
.13

.82

1.59

33.65

36.13

9.52

1.09

34.55

16.35

4.28

.23

2.05

3.22

1.33

.19
.01

.02

1.63

1.91

.53

.08

4.51

7.24
.02

2.67

.57
.14

.02

.54

.55

.67

.44

.03

.02

.14

.38
.09

.32
.03

.72

2.37

3.49

2.96

.11

.49

1.05

.85

.04

.14

.21
.05

.24
.91

.89

.92

1.91

1.11

.90

1.91

1.69

1.84

.93

.21

.01

.06

.13

.03

.28

.19

.18

.02

2.95

8.89

8.88

7.46

4.59

.01

.43
.01

1.39

.28

.22

.24

.02

.06

.03

.02

.08

.06

.01

.11

.20

.80

.63

.65

.07

.26

.34

.33

.24

.13

.01

.03

21.50

1.42

.07

.04

.03

.24

.20

.12

.28

.08

.52

.78
.18

.25

.22

January 1983 Brotherson: Kalsow Prairie 159

Table 3 continued.

7 8 9 10 11 12 13 14 15

4.3-4.8 4.9-5.4 5.5-6.0 6.1-6.6 6.7-7.2 7.3-7.8 7.9-8.4 8.5-9.0 9.1-9.6

.71 1.00 1.07 .13 8.00

.05

.71

2.03

4.50

1.79

1.71

7.86

.83

7.50

1.25

11.70

14.67

10.95

26.21

14.29

57.92

40.00

20.00

1.42

.33 .08 .71 2.50 4.29 3.00

7.89 2.50 3.50 1.25

7.37 2.50 5.83 1.00

.71

1.46

.25

.71

3.35

3.08

2.02

1.37

2.42

4.76

.09

.28

.33

1.08

.33

.17

.28

.08

.24

.08

.47

.25

.12

.33

.24

.12

2.36

2.08

2.50

.71

.25

.48

.09

.08

.09

.08

.71

.99

.08

.52

.25

.12

3.82

.83

.24

3.16

2.67

2.50

.50 .71

.05 .71

.05

2.14

.79 2.14 2.92 3.00 7.50

.26 2.14

.26 .71 .42 1.25

.36 .50

.13

.13 .71

8.16 5.71

.14 .79

.12 .39

.24 .53 .42

160

Great Basin Naturalist

Vol. 43,

, No. 1

Table 3 continued.

Species

.7-1.2

1.3-1.8

1.9-2.4

2.5-3.0

3.1-3.6

3.7-4.2

Panicum leihergii

.02

.51

1.19

2.31

Panicum virgatum

.26

.66

.86

.60

Pedicularis canadensis

.47

.09

.13

Petabstemum candidum

.02

Petalostemum purptireum

.03

.13

.59

.40

.35

Phalaris arundinacea

7.05

5.49

3.98

.49

Phlox pilosa

.04

.05

.13

Physalis heterophylla

Physalis virginiana

.01

.04

.03

Poa pratensis

1.24

2.82

3.57

4.24

Polygonum coccineum

27.27

16.81

4.02

1.52

.01

Potentilh arguta

.02

Psoralea argophylla

.02

.09

Pycnanthemum virginianum

.37

1.83

.77

.35

Ratibida pinnata

.21

.30

1.65

1.93

1.50

Rosa suffulta

.24

.46

.91

Rudbeckia hirta

.10

.01

.03

Scirpus atrovirens

.52

Scirpus fluviatilis

2.05

6.22

1.41

.08

Scutellaria leonardii

.07

.04

.13

.08

Senecio pauperculus

.42

3.15

3.92

.59

.35

Setaria lutescens

.13

.01

.09

Setaria viridis

.01

.03

.24

Silphium laciniatum

2.09

4.84

2.75

2.10

Solidago canadensis

.66

3.98

6.02

5.68

2.12

Solidago gymnospermoides

.01

.31

.11

Solidago missouriensis

.01

.09

Solidago rigida

1.81

3.20

5.44

Sorghastrum nutans

.09

.17

.42

.08

Spartina pectinata

1.36

4.27

3.74

1.79

.84

.30

Sporobolus heterolepis

.52

7.76

23.83

40.98

49.78

Stipa spartea

.05

Teucrium canadense

.03

1.07

.31

.01

Thalictrum dasycarpum

.04

.44

.33

Vernonia fasiculata

.59

Veronicastrum virginicum

.02

Viola pedatifida

.04

.08

Viola sp.

.07

.12

.08

Vicia americana

.05

Zizia aurea

.21

1.43

3.28

4.18

2.74

Allium sp.

.02

Aster novae-angliae

.23

.02

Cacalia tuberosa

.02

Ceanothus americana

.01

Panicum implicatum

.06

.09

Prenanthes racemosa

.08

Solidago nemoralis

.02

.13

.45

1.69

Solidago riddellii

.03

.13

.69

.18

.03

Taraxacum officinale

.09

Echinacea pallida

.02

were recognizable. These groups are labeled
A, B, C, and D, with group A corresponding
to the noncalcareous and ridge entity de-
scribed previously and made up of plants
showing preference for Nicollet and Nicollet-
Webster soils. Group B includes all but one
of the calcareous soil types plus four non-
calcareous types. The noncalcareous types

are found at the periphery of the group and
include Clarion, Clarion-Nicollet, Webster,
and heavy Webster soil types. Group C in-
cludes the Glenco-Okoboji and Okoboji soils,
and group D includes Glenco and calcareous-
Glenco soils. These last two groups corre-
spond to class 1 for species showing response
to the soil factor described above.

January 1983

Brotherson: Kalsow Prairie

161

Table 3 continued.

4.3^.8

4.9-5.4

5.5-6.0

6.1-6.6

6.7-7.2

7.3-7.8

7.9-8.4

8.5-9.0

9.1-9.6

2.64

2.16

2.62

9.74

2.50

2.92

.50

.24

.67
.17

.13
.13

.09

.58

.12

4.87

2.14

5.00

.09

.12

.13

2.50

7.36

9.25

26.55
.71

22.50
.13

27.86

26.67

38.50

61.25

.05

1.00

.83

.13

.71

2.92

.50

.24

.17

1.46

2.75

2.11

2.03

1.50

2.62

1.18

1.25

.09

.36

.57

2.50

.79

2.41

4.88

.13

.61

2.22

3.50

2.38

4.34
1.58

11.43

2.92
2.50

7.50

5.66

3.17

2.74

.79

.09

.26

.05

42.74

43.08

26.90

17.63

33.21

8.75

8.00

.09

.08

.95

.26
.13

.50

1.25

.42

.09

.05

.09

2.36

.08

.25

1.31

.26

.13

.42
.42

.05
.05

.50

1.97

2.14

2.50

24.50

Ordination of elevation data (Fig. 15)
showed no recognizable groupings. Instead it
separated the different elevation classes
(Table 3) along a curve, point 14 represent-
ing the ridge tops and point 1 representing
the bottom of the potholes. This would tend
to support statements made earlier that the
vegetation of Kalsow Prairie is best repre-

sented by the continuum concept of Curtis
and Mcintosh (1951).

The definable subcommunities or groups
(Fig. 14) as based on soils data represent the
response of the different taxa in the vegeta-
tion to an environmental stimulus (i.e., carbo-
nate soils) that is not distributed along
gradients (i.e., at 9 X 9 m sampling levels)

162 Great Basin Naturalist Vol. 43, No. 1

Table 4. Average cover values for species in relation to soil series in 20-acre intensive study area.

Species A A" N Na N" Na" W

Achillea lanulosa .14 .19 .57 .44 .21 .37

Agropyron smithii .01
Ambrosia artemisifolia

Amorpha canescens 3.47 2.87 4.16 4.06 6.57 2.56

.56

3.47

2.87

4.16

4.06

6.57

39.72

19.25

12.26

21.56

10.04
.08

Andrcypogon gerardi 39.72 19.25 12.26 21.56 10.04 10.63 6.70

Schizachyrium scoparius .08 .80

Anemone canadensis .03

Apocynum sibiricum .09

Arabis hirsuta

Artemisia ludoviciana 3.19 .56 .38 1.88 .12

Asclepias incamata

Asclepias sullivantii

Asclepias syriaca

Asclepias tuberosa

Aster ericoides

Aster laevis

Aster simplex

Baptisia leucophaea

Calamagrostis canadensis

Carex atherodes

Carex aquatilis

Carex gravida

Carex lasiocarpa

Carex retrorsa

Chenopodium album

Cirsium altissimum

Comandra umbellata

Convolvulus sepium

Desmodium canadense

Elymus canadensis

Equisetum kansanum

Eryngium yuccifolium

Fragaria virginiana

Galium obtusum

Gentiana andrewsii

Helenium autumnale

Helianthus grosseserratus

Helianthus laetiflorus

Helianthus maximiliani

Heliopsis helianthoides

Lactuca scariola

Lathyrus palustris

Lathyrus venosus

Lespedeza capitata

Liatris pycnostachya

Lithospermum canescens

Lycopus americanus

Lysimachia chiliata

Lysimachia hybrida

Lysimachia quadriflora

Lythrum alatum

Mentha arvensis

Muhlenbergia racemosa

Oxalis stricta

Panicum capillare

Panicum leibergii

Panicum virgatum

Pedicularis canadensis

Petalostemum candidum . 19 .09

Petalostemum purpureum .65 .12 .21

.28

.01

.65

.61

1.88

.73

.19

3.89

1.94

3.58

2.19

.42

1.50

3.47

4.44

1.56

9.68

1.29
.08

2.50

.46
2.12

.14

.85
.33

1.88

.24
.48

.03

10.48

.75

1.11

.01

.28

2.07

.01

.60

.14

.19

.08

.21

.05

.97

.28

.04

.65

1.88

2.19

2.71

1.64

.28

.19

.24

2.19

.44

1.04

.60

.14

.05
.61

.31

.44
.65

.42

.12

.38

.14

.19

.24

.16

2.15

.14

.65

.42

1.83
.01
.03

1.65

.31

2.58

1.25

9.26

3.19

6.20

3.87
.28

6.87

2.18
.24

4.79

.18

.03
.01
.08

.56

.09
.65

.24

1.88

.21

.14

.31

1.21

.40

.28

.56
.09

.52

.31
1.88

.56
.04

1.04

.21
.21

.17
.11

.03
.01
.12
.23

.01

6.67

7.50

2.64

4.06

.77

3.33

1.20

.14

.44
.04

2.71

.49

January 1983

Brotherson: Kalsow Prairie

163

Table 4 continued.

H-

.42

.40

.07

.02

.09

.65

.10

.09

.22

.17

.09

.73

1.93

.33

.04

.02

12.19

9.09

10.87

14.42

11.57

14.25

.05

.13

.63

.77

.46

1.74
.02

1.31

.02

.73

.17
.03
.71

.09

.02
.18
.64

.25
.02

.16
.07

.81

.74

2.06

3.50

2.63
1.88

2.81

1.22
1.70

2.81
1.18

2.59
.31

2.42
.55

1.76

3.54

1.02
.17

1.36

1.63

1.18

3.98

1.80

2.38

11.15

2.46

1.41

.81

13.61

45.26

56.88

1.50

.63

.25

7.73

8.25

32.38

1.88

.20

.11

.64

1.48

3.90

3.00

1.04

.04

.16

.25

.39

2.48

3.88

.25

6.88

.37

.58

1.23

1.38

8.27

10.63

6.00

.10

.77

.50

.67

.68

.19

.35

.45

.64

.16

.30

.09

.10

.48

.14

1.35

4.66

3.53

2.79

4.15

2.69

.14

.31

1.08

1.10

.56

1.10

.37

.07

.10

.31
.26

.15
.02

.11

.17

.52

2.24

1.62

.96

.98

1.57

.07

.13

2.08

1.42

2.43
.18
.31

1.52
.58

2.16
.30

1.85
.09

.14

.13

13.85

5.17

9.96

7.86

6.31

12.41

3.36

4.63

.10

.20

.02

.04

.34

.31

.18

.34

.65

.06

.02

.10

.06

.05

.09

.02

.10

.23

.63

1.02

.72

.67

.42

.28

.10

.48

.33

.22

.21

.09

.10

.03

.02
.02

.18

.04

.09

.05

.13

.10

.31

.03

.28

.02

.11

.04

.20

.42

1.47

.43

2.70

1.57

.05

.68

.80

1.03

.51

.30

.20

.46

.45

.25

.06

.02

.60

.50

.87

.42

.09

.93

.83

.28

.19

.25

.02

1.31

.05

.75

21.50

17.25

164

Great Basin Naturalist

Vol. 43,

No. 1

Table 4. Average cover values for

species in

relation to soil

series in 20-i

acre intensive study

area.

Species

N-

Na-

Phalaris arundinacea

1.09

Phlox pilosa

.19

.16

.03

Physalis heterophylla

2.31

.28

1.88

Physalis virginiana

.14

.09

.28

.08

.03

Poa pratensis

16.53

31.09

5.99

32.81

5.21

3.22

Polygonum coccineum

.44

Potentilla arguta

.14

.56

.05

Psoralea argophylla

1.81

1.30

.15

.31

.08

.21

Pycnanthemum virginianum

.28

.43

Ratibida pinnata

2.92

3.24

.05

.32

1.46

1.29

Rosa suffulta

.14

3.06

1.56

2.19

1.67

.29

Rudbeckia hirta

.04

Scirpus atrovirens

Scirpus fluviatilis

.01

Scutellaria leonardii

.09

.05

.08

.14

Senecio pauperculus

1.24

Setaria lutescens

.56

Setaria viridis

.56

.31

Silphium laciniatum

.38

1.25

2.15

Solidago canadensis

.14

2.41

3.21

.31

2.86

1.25

3.16

Solidago gymnospermoides

.09

.69

1.88

Solidago missouriensis

1.67

.56

.28

Solidago rigida

.56

3.25

.31

4.27

2.50

3.79

Sorghastrum nutans

.31

.20

.11

Spartina pectinata

.04

.83

Sporobolus heterolepis

12.08

21.76

62.69

37.19

58.95

27.92

37.41

Stipa spartea

.97

.09

.14

.08

.21

.05

Teucrium canadense

.05

.21

.23

Tahlictrum dasycarpum

.05

Vemonia fasciculata

.09

Veronicastrum virginicum

.01

Viola pedatifida

.09

.12

.05

Viola sp.

.03

Vicia americana

.28

.09

.05

.31

.04

.03

Zizia aurea

.14

.93

4.58

4.06

1.73

3.33

1.52

Allium sp.

Aster novae-angliae

.09

Cacalia tuberosa

.01

Cearwthus americana

10.56

.56

.04

Panicum implicatum

.04

.15

Prenanthes racemosa

.09

Solidago nemoralis

.28

.93

.75

Solidago riddellii

.01

Taraxacum officinale

but in mappable units with fairly discrete
boundaries. This would tend to cause vegeta-
tion sensitive to carbonate influence to group
accordingly.

An ordination of species, utilizing the data
from Tables 3 and 4, isolated taxa having dis-
tinct distribution patterns. These species are
Amorpha canescens, Andropogon gerardi. As-
ter ericoides. Aster laevis, Calamagrostis
canadensis, Carex athorodes, Carex aquatilis,
Desmodium canadense, Helianthus grosseser-
ratus, Helianthus laetiflorus, Panicum lei-

bergii, Phalaris arundinacea, Poa pratensis.
Polygonum coccineum, Ratibida pinnata,
Scirpus fluviatilis, Silphium laciniatum, Sol-
idago canadensis, Solidago rigida, Spartina
pectinata, Sporobolus heterolepis, Zizia aurea,
and Ceanothus americanus, all of which show
distinct distribution patterns and in many
cases high preference for certain soil groups
or elevations.

The relationships between elevation and
soil series are shown in Figure 16. The soil
types are positioned along the base line as

January 1983
Table 4 continued.

Brotherson: Kalsow Prairie

165

H-

1.56
.63

1.35

.10
.10
.10

.21
3.65

7.40
2.50

.63

.10

1.88

29.38

.73

.21

.31
.10

.20

.06
2.81

.06

.45

1.73

.68

.11
.48

2.95

4.12

.26

5.34

.17

.28

44.40

.28

.09

3.41

.09

.02
3.86

.99

3.05

.42

.09

.04

2.45

3.84
7.30

3.77

.35

.15

25.42

.15
.20

.31

.02

5.31

.13

.07

.05
3.15

2.17

1.94

.20

.16

.09
3.59

4.69
7.39

.05

1.36

.63

.54

24.08

.38
.38

.09
.02
4.64
.02
.05

.17

3.18
.25

1.99

1.91

.25

.09
2.84

.25
4.79
5.30

.09

.34

.85

32.80

.25
.17

.04

.25

5.13
.04

.09

5.74

.83
.09

.09
9.17

3.61
13.43

1.20
.09
.56

9.44

.09
.09

.09

3.52

.56

6.24

.05

8.74

6.63

.75

40.50

36.75

.35

1.87

.14

.28
.05

7.22
.79

.30

.49

13.63

17.50

.13

.75

.13

3.38

.13

.73
2.92

1.02
.09

.04
.31
.50

.47
.45

.17
.03

.19

they appeared in the field. In all cases where
the noncalcareous soils had adjacent cal-
careous variants the calcareous variants
showed higher average elevations.

Summary and Conclusions

1. Sporobolus heterolepis is the dominant
plant of the upland prairie that places Kal-
sow Prairie within the "Consociation" desig-
nated by Weaver and Fitzpatrick (1934) as
the Prairie-Dropseed type.

2. The vegetation of the upland prairie
communities is best described and represent-
ed by the continuum concept as described by
Curtis (1955).

3. The vegetation of the upland prairie has
changed since Moyer's 1953 study. Species
showing increased importance in my study
are Solidago canadensis, Solidago rigida, Pan-
icum leibergii, Helianthus grosseserratus, and
Fragaria virginiana. Species decreasing in im-
portance were Phleum pratense, Poa pra-
tensis, Zizia aurea, Schizachyrium scoparius.

166

Great Basin Naturalist

Vol. 43, No. 1

Fig. 15. Two-dimensional ordination of vegetation
found at different elevations in the 20-acre study area; 1
= .7-1.2 feet elevation, 2 = 1.3-1.8 feet elevation, 3 =
1.9-2.4 feet elevation, 4 = 2.5-3.0 feet elevation, 5 =
3.1-3.6 feet elevation, 6 = 3.7-4.2 feet elevation, 7 =
4.3-4.8 feet elevation, 8 = 4.9-5.4 feet elevation, 9 =
5.5-6.0 feet elevation, 10 = 6.1-6.6 feet elevation, 11 =
6.7-7.2 feet elevation, 12 = 7.3-7.8 feet elevation, 13 =
7.9-8.4 feet elevation, 14 = 8.5-9.0 feet elevation.

Panicum virgatum, Sorghastrum nutans, and
Sporobolus heterolepis.

4. Soil series, elevations, and species distri-
bution patterns were mapped on an 8 ha in-
tensive study plot. Elevation and soils data
are correlated with species distribution pat-
terns. All species show a response. Nine gen-
eral patterns of distribution are described,
with the following species as examples:

a. Andropogon gerardi—species of wide
distribution, limited only by conditions
peculiar to the drainage areas of the
prairie.

b. Silphium laciniatum—a. pattern closely
resembling that of Andropogon gerardi
but showing limited distribution on the
higher and drier ridges.

c. Ambrosia artemisifolia— species limited
to the border weed communities.

d. Amorpha canescens—a. pattern common
to species limited to the ridges and
lower slopes.

e. Solidago nemoralis— species limited to
mid- and upland slopes of the prairie.

f. Ceanothus americanus—a. pattern limit-
ed to the ridges and drier sites of the
prairie.

g. Helenium awfumna/e— limited to
growth on soils that are highly cal-
careous to the surface.

h. Calamagrostis canadensis— limited to
growth along the shallower areas of the
pothole and drainage system.

i. Scirpus fluviatilis— growth corresponds
to deeper areas within the drainage
system.

5. Species occurring in the intensive study
were ordinated using Orloci's (1966) method.
The technique did not delineate associated
groups of species, yet it pointed out species
exhibiting peculiar distribution patterns.
Such species are useful as indicator species.

6. Indices of interspecific association were
computed for all participating species (Cole
1949) and found to be extremely useful in
identifying clusters or groups of species hav-
ing similar ecological amplitudes.

Literature Cited

AiKMAN, J. M. 1959. Prairie research in Iowa. The Amer-
ican Biology Teacher 21:7-8.

Aikman, J. M., AND R. F. Thorne. 1956. The Cayler
Prairie: an ecologic and taxonomic study of a
northwest Iowa prairie. Iowa Acad. Sci.
63:177-200.

Brennan, K. M. 1969. Vertebrate fauna of Kalsow
Prairie. Unpublished thesis. Iowa State Univ.

G„ GO O

A A- N N„ N- 1^ W V^t, \% C

SOIL SERIES

Fig. 16. Relationships between elevation and soil series as found in 20-acre intensive study area; mean value in-
dicated for each soil by short horizontal line.

January 1983

Brotherson: Kalsow Prairie

167

Cain, S. A., and F. C. Evans. 1952. The distribution
patterns of three plant species in an old-field
community in southeastern Michigan. Contribu-
tions from the Laboratory of Vertebrate Biology
No. 52.

Cole, L. C. 1949. The measurement of interspecific as-
sociation. Ecology 30:411-424.

Collins, G. B. 1968. Implications of diatom succession
in postglacial sediments from two sites in north-
em Iowa. Unpublished dissertation. Iowa State
Univ.

Crawford, R. M. M., and D. Wishart. 1968. A rapid
classification and ordination method and its ap-
plication to vegetation mapping. Ecology
56:385-404.

Curtis, J. T. 1955. A prairie continuum in Wisconsin.
Ecology 36:558-566.

Daubenmire, R. 1959. A canopy-coverage method of
vegetational analysis. Northwest Science
33:43-66.

Dick-Peddie, W. a. 1955. Presettlement forest types in
Iowa. Unpublished thesis. Iowa State Univ.

Due, R. L., AND J. E. Butler. 1960. A phytosociological
study of a small prairie in Wisconsin. Ecology
41:316-327.

Ehrenreich, J. H. 1957. Management practices for
maintenance of native prairies in Iowa. Unpub-
lished dissertation. Iowa State Univ.

Esau, K. L. 1968. Carabidae (Coleoptera) and other ar-
thropods collected in pitfall traps in Iowa corn-
fields, fencerows, and prairies. Unpublished dis-
sertation. Iowa State Univ.

GiLLY, C. L. 1946. The Cyperaceae of Iowa. Iowa State
Coll. J. Sci. 21:55-151.

Gleason, H. a. 1952. The new Britton and Brown illus-
trated flora of the northeastern United States and
adjacent Canada. Lancaster Press, Lancaster,
Pennsylvania. 3 vols.

Greig-Smith, p. 1964. Quantitative plant ecology. But-
terworth. Washington, D.C.

Hale, M. E., Jr. 1955. Phytosociology of corticolous
cryptograms in the upland forests of southern
Wisconsin. Ecology 36:45-62.

Hayden, Ada. 1943. A botanical survey in the Iowa lake
region of Clay and Palo Alto counties. Iowa State
Coll. J. Sci. 17:277-416.

1945. The selection of prairie areas in Iowa

which should be preserved. Proc. Iowa Acad. Sci.
52:127-148.

1946. A progress report on the preservation of

prairie. Proc. Iowa Acad, of Sci. 53:45-82.

Hewes, L. 1950. Some features of early woodland and
prairie settlement in a central Iowa county. An-
nals Assoc. Amer. Geog. 40:40-57.

HuRLBERT, S. H. 1969. A coefficient of interspecific asso-
ciation. Ecology 50:1-9.

Kennedy, R. K. 1969. An analysis of tall-grass prairie
vegetation relative to slope position, Sheeder

Prairie, Iowa. Unpublished thesis. Iowa State
Univ.

Kershaw, K. A. 1964. Quantitative and dynamic ecolo-
gy. Edward Arnold, Ltd., London, England.

Landers, R. Q. 1966. Visit the virgin prairie. Iowa Farm
Science 21:418-419.

MoYER, John F. 1953. Ecology of native prairie in Iowa.
Unpublished dissertation. Iowa State Univ.

Norton, D. C, and P. E. Ponchillia. 1968. Stylet-bear-
ing nematodes associated with plants in Iowa
prairies. J. Iowa Acad. Sci. 75:32-35.

Orloci, L. 1966. Geometric models in ecology. I. The
theory and application of some ordination meth-
ods. J. Ecology 54:193-215.

Oschwald, W. R. et al. 1965. Principal soils of
Iowa. Special Report No. 42, Department of
Agronomy, Iowa State Univ.

Pohl, R. W. 1966. The grasses of Iowa. Iowa. Iowa State
J. Sci. 40:341-566.

Richards, M. S. 1966. Observations on responses of
prairie vegetation to an April fire in central
Iowa. Unpublished thesis. Iowa State Univ.

RuHE, R. V. 1969. Quaternary landscapes in Iowa. Iowa
State Univ. Press, Ames Iowa. 249 pp.

Rydberg, p. A. 1931. A short phytogeography of the
prairies and great plains of central North Ameri-
ca. Brittonia 1:57-66.

Sanders, D. R. 1969. Structure and pattern of the her-
baceous understory of deciduous forests in cen-
tral Iowa. Unpublished dissertation. Iowa State
Univ.

ScHMiTT, D. P. 1969. Plant parasitic nematodes and
nematode populations in the Kalsow Prairie. Un-
published thesis. Iowa State Univ.

Shimek, B. 1911. The prairies. Bull. Lab. Nat. Hist.,
State Univ. Iowa 6:169-240.

Shimek, B. 1915. The plant geography of the Lake Oko-
boji region. Bull. Lab. Nat. Hist., State Univ.
Iowa 7:1-90.

Shimek, B. 1925. The persistence of the prairie. Univ. of
Iowa Studies 11:3-24.

Steiger, T. L. 1930. Structure of prairie vegetation.
Ecology 11:170-217.

U.S. Government. 1868. First survey of the State of
Iowa. Plats deposited in the State House, Des
Moines.

Weaver, J. E. 1930. Underground plant development in
its relation to grazing. Ecology 11:543-557.

Weaver, J. E. 1954. North American prairie. Lincoln,
Nebraska, Johnsen Publishing Company. 348 pp.

Weaver, J. E., and F. E. Clements. 1938. Plant Ecolo-
gy. 2d ed. McGraw-Hill, New York.

Weaver, J. E., and T. J. Fitzpatrick. 1932. Ecology and
relative importance of the dominants of the tall-
grass prairie. Bot. Gaz. 93:113-50.

Weaver, J. E., and T. J. Fitzpatrick. 1934. The prairie.
Ecol. Monogr. 4:109-295.

DEER MOUSE, PEROMYSCUS MANICULATUS, AND ASSOCIATED RODENT FLEAS
(SIPHONAPTERA) IN THE ARCTIC-ALPINE LIFE ZONE
OF ROCKY MOUNTAIN NATIONAL PARK, COLORADO

R. B. Eads' and E. G. Campos'

Abstract.— Peromyscus maniculatus and related small rodents have been examined for ectoparasites in the tun-
dra region of the Rocky Mountain National Park 1974-1979. One hundred and ninety-four P. maniculatus were ex-
amined from two tundra sites. Flea infestation rates were 1.9 fleas per mouse examined and 4.1 fleas per infested
mouse. Species taken in significant numbers were Monopsyllus thambus (51 percent), Peromyscopsylla hesperomys
(34 percent), Malaraeus euphorbi (9 percent), and Catallagia calisheri (4 percent). Peromyscus maniculatus host/flea
relationships in the tundra are compared with those in other life zones in the park.

Personnel of the Vector-Borne Diseases Di-
vision have studied ectoparasite/host rela-
tionships in the Rocky Mountain National
Park (RMNP) for a number of years. Of prin-
cipal interest have been vectors and reser-
voirs of Yersinia pestis and Colorado tick fe-
ver virus. All life zones in the Park have been
investigated. Because of a dearth of published
information concerning fleas and other ecto-
parasites of rodents in the arctic-alpine life
zone of the Rocky Mountains, data obtained
during the period 1974-1979 in the zone are
presented here. Emphasis has been placed on
the deer mouse, Peromyscus maniculatus, as
it is by far the most abundant rodent in the
zone. Perom.yscus maniculatus fleas in the
tundra are compared with those in other life
zones.

Study Area

The RMNP is located in Larimer and Boul-
der counties in north central Colorado. It
covers 1046 km^ of mountainous terrain, with
elevations ranging from 2400 to over 4300 m.

Regional ecosystems of north central Colo-
rado, with approximate elevations as defined
by Marr (1961), include the grassland-lower
montane ecotone region (1707-1829 m).

lower montane forest climax region
(1829-2347 m), lower montane-upper mon-
tane ecotone region (2347-2438 m), upper
montane forest climax region (2438-2743 m),
upper montane-subalpine ecotone region
(2743-2835 m), subalpine forest climax re-
gion (2835-3353 m), subalpine-alpine eco-
tone region (3353-3475 m), and alpine tun-
dra climax region (3475 m-mountain tops).

Collections in the tundra were concen-
trated at sites 1 and 2, located as shown in
Figure 1. Site 1 is 2.5 km west of Rainbow
Curve on Trail Ridge Road at an elevation of
around 3475 m. Site 2 is on Fall River Road,
some 1.5 km below the Alpine Visitors Cen-
ter at an elevation of approximately 3523 m.
Sites 1 and 2 are in areas with limited human
activity. Trapping was confined to an area of
5 or 6 ha in both locations. In the tundra
(Fig. 2) short grasses, sedges, and forbes pre-
dominate in the exposed meadows. Some-
what taller grasses and dwarf shrubs are
found in low-lying areas partially protected
from the violent winds that sweep the
tundra.

Methods

Mammals were captured in the tundra,
primarily in Sherman live traps (7.62 X 7.62

'Veclor-Bome Diseases Division, Center for Infectious Diseases, Centers for Disease Control, Public Health Service, U.S. Department of Health and Hu-
man Services, Post Office Box 2087, Fort Collins, Colorado 80522-2087.

168

January 1983

Eads, Campos: Colorado Fleas

169

V "■■■ f

^////////////////////////////////i//////////////////////////A

Fig. 1. Flea collection sites in the Rocky Mountain National Park, Larimer County, Colorado.

X 22.86 cm), with National live traps (12.7
X 12.7 X 40.64 cm) also used on occasion.
No attempts were made to collect larger
mammals. Traps baited with rolled oats were
set in parallel lines of 20 stations at approx-
imately 20 m intervals. Traps were placed
adjacent to rocky outcroppings, when avail-
able, to provide trapped rodents some pro-
tection from the elements. The number of
lines varied, but there were usually 4 or 5, 25
m apart. Traps were set in the morning and
inspected the following day.

Trapping in the tundra was limited to the
summer months. Trail Ridge and Fall River
roads were only open to vehicular traffic
June-September, with some closures at ir-
regular intervals during this period due to
hazardous driving conditions.

As the Sherman traps were picked up,
those containing rodents were placed in plas-
tic bags to prevent loss of fleas during trans-
port to a field laboratory. Each mammal was
lightly anesthetized witii ether and brushed
in a white enamel pan to remove ectopara-

170

Great Basin Naturalist

Vol. 43, No. 1

Fig. 2. Tundra region, Rocky Mountain National Park.

sites. Certain animals were bled, ear tagged,
and released, and serological tests for plague
and Colorado tick fever were performed.
Others were held for more extensive ecto-
parasite recovery efforts, including exam-
ination of the mice under a dissecting micro-
scope. Fleas collected were held in 2 percent
saline solution for plague testing or in 70 per-
cent alcohol for definitive taxonomic studies.

Results and Discussion

Trapping over a six-year period (1974-79)
has provided considerable information on
deer mouse host/ flea relationships in the Col-
orado tundra region. As shown in Table 1,
156 P. maniculatus were captured in this pe-
riod during the summer months at Site 1. An
average of 8 P. maniculatus were taken per
100 trap nights. About 47 percent were in-
fested with fleas, with a mean number of 2.1
fleas per animal examined and 4.4 per in-
fested animal. Site 2 was trapped less fre-
quently than Site 1. As shown in Table 2, the
flea burden was 1.1 fleas per mouse and 2.9
fleas per infested mouse.

Ectoparasites other than fleas were in-
frequently encountered on deer mice in the
timdra. Fewer than 10 percent were infested,
usually lightly, with the sucking louse, Hopl-
opleura hesperomydis. Larval chigger mites
were somewhat more prevalent. Neo-
trombicula harperi, N. microti, and Eu-
schoengastia guntheri were taken. A single
larval argasid tick was recovered from a P.
maniculatus above the tree line, probably an
Argas sp.

The second most prevalent rodent taken in
the tundra was the heather vole, Phenacomys
intermedius. Twenty-three were examined,
and 65 percent were infested with three spe-
cies of fleas at the rate of 2.4 fleas per animal
and 3.6 per infested animal. Species included
Peromyscopsylla selenis (67 percent), Mega-
bothris abantis (19 percent), and Monopsyllus
thambus (15 percent). Two species of chigger
mites, N. harperi and E. guntheri, were recov-
ered from heather voles. One vole was heavi-
ly infested and several lightly infested with
the dermanyssid mite Hirstionyssus
isabellinus.

January 1983

Eads, Campos: Colorado Fleas

171

Table 1. Peromyscus maniculatus examined for fleas in the tundra region of the Rocky Mountain National Park
(Site 1).

Percent

Mean

traps with

Number

Number with

Total

Mean number

number fleas/

Date

P. manic.

fleas

fleas/animal

infested animal

7-16-74

1.73

3.17

7-24-74

5.17

10.33

8-14-74

1.1

2.75

8-15-74

2.67

5.33

9-26-74

.33

1.5

8-22-75

1.12

1.73

8-18-76

1.0

2.33

6-29-77

5.0

8-17-77

2.2

5.5

9-02-77

1.5

2.0

9-09-77

1.0

4.0

6-27-78

4.33

6.5

7-07-78

.67

2.0

7-25-78

3.0

6.0

8-03-78

1.5

9-07-78

1.29

3.14

7-10-79

2.29

5.33

7-18-79

9.0

18.0

8-24-79

2.57

6.0

8.11

156

325

2.08

4.39

Lesser numbers of the least chipmunk, Eu-
tamias minimus (7); golden-mantled ground
squirrel, Spermophilus lateralis (3); yellow-
bellied marmot, Marmota flaviventris (2); and
pika, Ochotona princeps (3) were captured
and examined.

Eleven fleas, 10 Monopsyllus eumolpi, and
one M. thambus were removed from 4 of the
7 chipmunks. The 2 marmots were infested
with 7 Thrassis stanfordi and 2 Oropsylla ru-

pestris. Three Oropsylla idahoensis were re-
covered from 2 of the 3 golden-mantled
ground squirrels. The 3 pikas examined were
infested with the following fleas: 46 Am-
phalius necopinus, 46 Ctenophyllus terribilis,
and 1 M. thambus. The chigger mite, N. mi-
croti, was also taken from all 3 pikas.

Peromyscus maniculatus host/flea relation-
ships in the tundra have proven to be dis-
tinctive relative to the other life zones in the

Table 2. Peromyscus maniculatus examined for fleas in the tundra region of the Rocky Mountain National Park
(Site 2).

Number of mice

Number with

Number and species

Date

trapped

fleas

of fleas

7-17-74

2 Monopsyllus thambus

7-24-74

3 M. thambus

1 Malaraeus euphorbi

8-17-77

9 M. thambus

7 M. euphorbi

4 Peromyscopsylla hesperomys

1 Megabothris abantis

1 Catallagia calisheri

9-02-77

5 P. hesperomys
2 M. thambus
1 C. calisheri

9-09-77

3 M. thambus
2 P. hesperomys
1 M. euphorbi

9-24-77

1 Peromyscopsylla selenis

Totals

172

Great Basin Naturalist

Vol. 43, No. 1

RMNP. As shown in Table 3, 21 species were
recovered in 1974 and 20 in 1975 in all life
zones. This is well over twice the number of
species found to be parasitizing deer mice in
the tundra. The 2 X ratio remains virtually
unchanged when the recoveries are limited
to normal or true parasites of deer mice, ex-
cluding species that have strayed from envi-
rormiental associates.

Monopsylhis wagneri was the principal
flea below the treeline (68 percent); over 60
percent of the deer mice were infested in
1974-1975. The only other prevalent species
were Malaraeus telchinum (10 percent of to-
tal fleas) and Opisodasys keeni (8 percent of
total fleas). Each of the other 11 species nor-
mally parasitic on deer mice made up 1-5
percent of the total fleas. As shown in Table
4, mean infestation rates for the 2,090 P. ma-
niculatus examined in all life zones in the
RMNP in 1974-76 were 1.2 fleas per mouse
and 2.5 fleas per infested mouse.

Higher flea infestation rates per deer
mouse were obtained in the tundra than in

the other life zones. This may be a reflection
of more intensive examination of the mice in
the tundra. Also, tundra mice were usually
held for several examinations, and in the
other life zones normal procedure was to in-
spect them once, tag and release them. A to-
tal of 194 deer mice were examined from the
two tundra sites, and 368 fleas were recov-
ered from 89 of them. This is a rate of 1.9
fleas per mouse examined and 4.1 fleas per
infested mouse.

In the tundra, fewer species were involved
in parasitizing deer mice than in the other
life zones. Only four were taken in significant
numbers. Monopsyllus wagneri disappears
completely from the mice in the tundra and
is replaced by M. thambus (51 percent of to-
tal fleas at Site 1 and 44 percent at Site 2).
Overlapping of the two species occurred at
Rainbow Curve, elevation approximately
3290 m. This scenic overlook consists of an
exposed rocky slope with large boulders bor-
dered by subalpine forest.

Table 3. Species of fleas from Peromyscus maniculatus in all life zones in Rocky Mountain National Park in
1974-1975.

1974

1975

Number

offleas-1,018

Number

of fleas- 1,357

Percent of

Percent of animals

Percent of

Percent of animals

Species

total fleas

with species

total fleas

with 5

ipecies

Ceratophyllidae

Malaraeus euphorbi
M. telchinum

5.9

1.3

7.3

1.5
14.5

1.3

17.5

Megabothris abantis
Monopsyllus eumolpi
M. tliambus

<
<

4.1

<
<

1
1
3.6

<
<
<

1
1
1

<
<
<

M. wagneri
M. vison

(

84.8

60.8
1

70.3

1
<

Opisocrostis Labis
Opisodasys keeni
Orchopeas leucopus
Oropsylla idahoensis

<
<

9.9

9.5

1.7

<
<

5.8

7.3
1.6

Hystrichopsyllidae

Callistopsyllus deuterus
Catallagia calisheri

—

<
<

1
1

<
<

C. decipiens
C. neweyi
Epitedia wenmanni

4.4
2.4

3.9

4.1

1.8

1
2

3.6

Hystrichopsylla occidentalis
Megarthroglossus sp.
M. divisus

<
<

1
1

<
<

Rhadinopsylla sectilis

Leptopsyllidae

Amphipsylla sibirica
Peromyscopsylla hesperomys
P. selenis

<
<

4.2

<
<

4.5

<
<

1
1

1.9

January 1983

Eads, Campos: Colorado Fleas

173

Table 4. Summary of flea collections from Peromyscus maniculatus from all life zones in Rocky Mountain Nation-
al Park, 1974-1976.

Number of

mice

Number of

Percent

Total

Number of

examined

mice

of mice

number of

fleas/

Year

for fleas

with fleas

infested

fleas

mouse

infested mouse

1974

955

431

45.1

1,018

1.1

2.4

1975

957

507

53.0

1.357

1.4

2.7

1976

178

44.9

183

1.0

2.3

Totals

2,090

1,018

48.7

2,558

1.2

2.5

Peromyscopsylla hesperomys was the sec-
ond most prevalent P. maniculatus flea in the
tundra (35 percent of the total fleas at Site 1
and 26 percent at Site 2). This flea is found
on deer mice in all life zones of the RMNP
but in much smaller numbers at the lower
elevations, making up <5 percent of the to-
tal fleas collected in the park.

The third most common species on deer
mice in the tundra, Malaeraeus euphorbi (7
percent of the total fleas at Site 1 and 21 per-
cent at Site 2) was also more prevalent than
at lower altitudes. Less than 2 percent of to-
tal fleas collected in the park were this
species.

A nest flea, Catallagia calisheri, was a poor
fourth on tundra deer mice. However, the
fact that 4 percent of the fleas at Site 1 and 5
percent at Site 2 were this species is in-
dicative of much greater numbers in the
nests. Catallagia calisheri was not taken be-
low the tree line. At lower elevations in
RMNP, the common deer mouse nest Catal-
lagia was decipiens. Single specimens of the
nest fleas, Callistopsyllus deuterus and Me-
garthroglossus sp. (female) were taken from
deer mice in the timdra. Both genera were
also rarely encountered in other RMNP life
zones. Orchopeas leucopus, a common Per-
omyscus parasite at lower elevations, was
rare in the park. It was taken but once in the
timdra and once in other RMNP life zones.
The only other fleas taken from deer mice in
the tundra were a few Megabothris abantis
and Peromyscopsylla selenis, strays from
heather voles.

Conclusions

Our data concerning the fleas of P. ma-
niculatus support the conclusions of Wenzel

and Tipton (1966) that the altitudinal ranges
of many ectoparasites do not necessarily
coincide with those of the hosts. Peromyscus
maniculatus is prevalent in the RMNP at all
elevations. However, vertical stratification is
evident in the species of fleas parasitizing
them, especially at the upper levels. In the
tundra, around 3475 m, the number of flea
species on P. maniculatus was greatly re-
duced from the lower elevations, but the
mean number per animal was somewhat
higher. Only four species were encountered
in significant numbers on P. maniculatus in
the tvmdra. Monopsyllus thambus and Catal-
lagia calisheri, found only on the tundra, re-
placed M. wagneri and C. decipiens of lower
elevations, respectively. Peromyscopsylla hes-
peromys and Malaraeus euphorbia, found at
all elevations studied in RMNP, were signifi-
cantly more prevalent on P. maniculatus in
the tundra than at lower elevations.

Acknowledgments

Excellent cooperation has been received
from naturalists of the National Park Service,
U.S. Department of Interior, during these
studies. Ray Bailey, Vector-Borne Diseases
Division statistician, was most helpful in sup-
plying computerized data. Gary Maupin took
the tundra photograph. Numerous persons,
presently and formerly with VBDD, contrib-
uted to the field investigations. Included
were Dr. R. G. McLean, Dr. A. M. Barnes,
Ronald Shriner, Karen Pokorny, Gary Mau-
pin, Leon Carter, Edwin Heidig, William Ar-
chibald, and David Pegg.

Taxonomic assistance was kindly provided
by the following authorities: Dr. J. E. Keirans
(ticks). Dr. R. B. Loomis (chigger mites), and
Dr. K. C. Emerson (sucking lice).

174 Great Basin Naturalist Vol. 43, No. 1

Literature Cited Wenzel, R. L., and v. J. Tipton. 1966. Some relation-

ships between mammal hosts and their ectopara-
Marr, J. W. 1961. Ecosystems of the east slope of the sites. Pages 677-723 in Ectoparasites of Panama,

front range in Colorado. Univ. of Colorado Stud- Field Mus. Nat. Hist. Chicago, Illinois,

ies (Series in Biology) No. 8. Univ. of Colorado
Press. Boulder.

FOOD OF LARVAL TUI CHUBS, GILA BICOLOR, IN PYRAMID LAKE, NEVADA'

David L. Galat* and Nancy Vucinich^

Abstract.— Four near-surface locations in Pyramid Lake, Nevada, were sampled for larval tui chubs {Gila bicolor)
during summer and early fall 1979. Numbers of larvae collected were highest in mid-July. Zooplankton was the only
food eaten throughout the survey; the cladoceran Moina hutchinsoni was the major species eaten at all locations. An-
other cladoceran, Diaphanosoma leuchtenbergianum, was also important to the diet of pelagic larvae, and the cope-
pod Cyclops vemalis was eaten in significant quantities by nearshore fish. Changes in diet composition of larval tui
chubs during summer corresponded to seasonal succession of zooplankton species in Pyramid Lake.

The tui chub, Gila bicolor, is a polytypic
cyprinid native to drainage systems in Ore-
gon, California, and Nevada (Moyle 1976). It
is the most abundant fish in Pyramid Lake,
Nevada, where adults constitute over 90 per-
cent of gill net catches (Vigg 1978, 1981). A
major role of this species in the economy of
Pyramid Lake is as forage for the primary
sport fish, the Lahontan cutthroat trout (Sal-
mo clarki henshawi), which commonly attains
trophy weights of over 3 kg (Snyder 1917,
Kucera 1978, Galat et al. 1981).

LaRivers (1962), Langdon (1979), and Vu-
cinich et al. (1981) provided information on
food of adult and juvenile tui chubs in Pyra-
mid Lake; Miller (1951), Kimsey (1954), and
Cooper (1978) presented similar information
for this species in other waters; and Williams
and Williams (1980) described the food of re-
lated Gila species. We present data on abun-
dance, distribution, and food of the pre-
viously uninvestigated larval phase of the tui
chub in Pyramid Lake.

Methods

Two shallow littoral stations in Pyramid
Lake (40° 00' N, 119° 35' W), one northeast
and the other southeast of Sutcliffe, Nevada,
were quantitatively sampled for larval tui
chubs between 18 June and 9 August 1979. A
metered 1 mm-mesh net, 0.5 m in diameter,
was hand-towed along the surface at about
the 1 m depth contour and parallel to shore.

However, only 2 of 10 attempts at collecting
larval tui chubs in the shallow littoral region
were successful and only three larvae were
captured. Because so few fish were recovered
from these locations they are not discussed
further.

Two deeper littoral locations were also
quantitatively sampled for larval fishes: a sur-
face pelagic station, northeast of Sutcliffe,
Nevada, at the 72 m depth contour, and a
surface nearshore station, southeast of Sut-
cliffe at the 5-10 m depth contour, were
sampled every two weeks from 26 June
through 26 October 1979. Larvae were col-
lected with the 0.5 m net described above,
towed 1-3 m below the surface behind a
boat. Fish collected were immediately killed
in MS-222 to minimize regurgitation and
preserved in 10 percent formalin.

After identifying larvae as tui chubs and
measuring their fork lengths (FL), the entire
digestive tract was removed. Contents from a
maximum of 10 nonempty larval tracts were
pooled for each date and station where more
than two fish were collected. All zooplank-
ters recovered from larval fish were identi-
fied to species and enumerated under a com-
pound microscope at 40-200X. Algae and
unidentifiable matter (e.g., detritus and di-
gested material) were never observed in sub-
stantial amounts and were not quantified.

Numbers of organisms recovered from guts
were converted to carbon equivalents based
on the average carbon content of whole

'The Colorado Cooperative Fishery Research Unit is supported by the U.S. Fish and WildUfe Service, Colorado Division of Wildlife, and Colorado State
University.
"Cooperative Fishery Research Unit, Colorado State University, Fort Collins, Colorado 80523.
'Pyramid Lake Indian Tribal Enterprises, Star Route, Sutcliffe, Nevada 89510.

175

176

Great Basin Naturalist

Vol. 43, No. 1

55 ■

'>< 25

« 20 H

^ 15 H

10 -

5 -

• Pebgic station

o- — o Near- shore station

,.--- c-.

Jun Jul Aug Sep Oct

Fig. 1. Numbers of tui chub larvae collected near the surface at two locations in Pyramid Lake, Nevada, 1979.

organisms as measured with a Hewlett-Pack-
ard model 185B CHN analyzer, following
Sharp's (1974) procedures. Because carbon
values were not obtained for Daphnia schod-
leri or copepod nauplii, these organisms were
assigned approximate carbon equivalents rel-
ative to their intact size. We selected this ap-
proach over volumetric or weight techniques
because it more accurately reflected the
bioenergetic significance of a food item to
the fish's diet.

Results and Discussion

The length range of larval tui chubs cap-
tured was 6-15 mm. Following Snyder's
(1976) terminology, only mesolarvae (<12
mm FL) and metalarvae (>12 mm) were
identified from our collections. Yolk sacs
were not apparent in any larvae. The alimen-
tary tract was straight and tubelike in larvae
up to 13 mm long; in progressively larger fish
it began to loop and swell anteriorly. The
transition of fish from metalarva to juvenile
occurred at a length of about 15 mm.

Estimated larval abundance at the pelagic
and nearshore stations peaked in mid- July at
0.06 and 0.03 larvae/m^ respectively (Fig. 1),
when water temperature from the surface to
a depth of 5 m was 21 C. This period coin-
cides with the reported time of peak tui chub

spawning in Pyramid Lake (Kucera 1978). By
September few larvae were collected at the
surface nearshore station, and none at the
surface pelagic site.

Digestive tracts from tui chubs taken on
four pelagic and three nearshore sampling
dates, totaling 25 and 14 larvae, respectively,
were examined. Guts were usually one-half to
three-quarters full and contained only zoo-
plankton (Tables 1 and 2).

Moina hutchinsoni was the dominant zoo-
plankter recovered from all pelagic larvae
sampled and was also of greatest significance
to larvae in two of three nearshore collec-
tions. Diaphanosoma leuchtenbergianum
ranked second in importance among food
items for pelagic larvae but was of minor im-
portance among nearshore larvae. In con-
trast, Cyclops vernalis appeared in all three
nearshore samples but was insignificant in
pelagic larval stomachs. Alona costata was
observed in larvae from two nearshore sam-
ples but was not recovered from pelagic lar-
vae. Food items eaten in small amounts were
Eucypris sp., Branchionus spp., Daphnia
schodleri, and copepod nauplii.

We anticipated finding more rotifers and
copepod nauplii in larval tui chub stomachs,
particularly since these groups were abun-
dant in littoral zooplankton samples (Vucin-
ich et al. 1981). Kimsey (1954) reported that

January 1983

Galat, Vucinich: Food of Tui Chubs

177

Table 1. Pooled stomach contents of pelagic larval tui chubs from Pyramid Lake, Nevada. Carbon values are ap-
proximate and represent reconstructed organisms. Numbers in parentheses are subtotals for the various categories.

June 26

July 17

July 31

August 15

Total C

Percent

Total C

Percent

Total C

Percent

Total C

Percent

Food item

No.

(Mg)

Total C

No.

(Mg)

Total C

No.

(Mg)

Total C

No.

(Mg)

Total C

Cladocera

(8)

(22)

(98.7)

(32)

(88)

(99.7)

(55)

(150)

(98.4)

(50)

(139)

(96.8)

Moina

98.7

75.7

62.5

126

87.7

Diapharwsoma

20.5

35.9

9.1

Alona

Daphnia

2.8

3.2

Copepoda

(2)

(1.0)

(0.7)

(4)

(4.5)

(3.2)

Cyclops

3.0

2.1

Nauplii

1.0

0.7

1.1

Rotatoria

Brachionus

0.3

1.3

0.5

0.6

1.4

0.9

Ostracoda

Eucypris

Grand total

152

144

No. tracts

examined

No. tracts with

food

Mean fish

length (mm)

10.2

10.9

11.9

newly hatched tui chubs in Eagle Lake, Cali-
fornia, fed on rotifers, diatoms, desmids, and
other microscopic material. Perhaps tui chub
larvae smaller than those captured in the
present study fed on these organisms.

Digestive tracts from larvae captured at
the surface nearshore station contained more
C. vernalis, A. costata, and Eucypris sp. than
were recovered from larvae collected at the

surface pelagic station. Conversely, D. leuch-
tenbergianum was more abundant in guts
from pelagic tui chubs. A probable explana-
tion for these differences is that the first
three zooplankton taxa named prefer a ben-
thic habitat and hence would be more avail-
able than D. leuchtenbergianum, a limnetic
species, to nearshore tui chubs (Pennak 1978).
Shifts in larval tui chub diet composition

Table 2. Pooled stomach contents of near-shore larval tui chubs from Pyramid Lake, Nevada. Carbon values are
approximate and represent reconstructed organisms. Numbers in parentheses are subtotals for the various categories.

June 26

July 17

July 31

Total C

Percent

Total C

Percent

Total C

Percent

Food item

No.

^g)

Total C

No.

(Mg)

Total C

No.

Total C

Cladocera

(7)

(18)

(33.2)

(66)

(183)

(80.2)

(31)

(87)

(96.7)

Moina

5.6

10.9

168

73.6

96.7

Diaphanosoma

4.6

Alona

22.3

4.6

2.0

Daphnia

Copepoda

(23)

(35)

(66.8)

(8)

(19)

(8.3)

(1)

(3.0)

(3.3)

Cyclops

66.8

7.9

3.0

3.3

Nauplii

1.0

0.4

Rotatoria

Brachionus

5.7

2.5

Ostracoda

Eucypris

9.0

Grand total

228

No. tracts

examined

No. tracts with

food

Mean fish

length (mm)

10.3

12.8

178

Great Basin Naturalist

Vol. 43, No. 1

from C. vemalis to D. leuchtenbergianum
and M . hutchinsoni as the summer progressed
paralleled seasonal changes in the relative
abundance of these zooplankton species in
Pyramid Lake (Galat et al. 1981), suggesting
that larval tui chubs, like adults (Langdon
1979), are opportunistic feeders.

ACKNOW^LEDGMENTS

We thank the Pyramid Lake Paiute Indian
Tribe for granting us permission to conduct
this study. We also thank E. P. Bergersen, K.
Hamilton-Galat, E. P. Eschmeyer, and D.
Snyder for their critical review of the manu-
script. Mark Coleman assisted with field col-
lections and zooplankton identification.
Study funding was provided by the Bureau of
Indian Affairs under Contract 14-16-0008-
974 to the U.S. Fish and Wildlife Service and
Colorado State University.

Literature Cited

Cooper, J. J. 1978. Contributions to the life history of
the Lahontan tui chub, Gila hicolor obesa (Gi-
rard), in Walker Lake, Nevada. Unpublished
thesis. Univ. of Nevada, Reno. 98 pp.

Galat, D. L., E. L. Lider, S. Vigg, and S. R.
Robertson. 1981. Limnology of a large, deep.
North American terminal lake, Pyramid Lake,
Nevada, USA. Hydrobiologia 82:281-317.

KiMSEV, J. B. 1954. The life history of the tui chub, Siph-
ateles bicolor (Girard), from Eagle Lake, Califor-
nia. California Fish Game 40:395-410.

Kucera, p. a. 1978. Reproductive biology of the tui
chub, Gila bicolor, in Pyramid Lake, Nevada.
Great Basin Nat. 38:203-207.

Langdon, R. W. 1979. Food habits of the tui chub, Gila
bicolor, in Pyramid Lake, Nevada. Unpublished
thesis. Humboldt State Univ., Areata, California.
45 pp.

La Rivers, I. 1962. Fishes and fisheries of Nevada. Ne-
vada State Fish and Game Comm. 782 pp.

Miller, R. G. 1951. The natural history of Lake Tahoe
fishes. Unpublished dissertation. Stanford Univer-
sity. 160 pp.

MoYLE, P. B. 1976. Inland fishes of California. Univ. of
California Press, Berkeley. 405 pp.

Pennak, R. W. 1978. Freshwater invertebrates of the
United States. Wiley-lnterscience, New York. 803
pp.

Sharp, J. H. 1974. Improved analysis for "particulate"
organic carbon and nitrogen from seawater. Lim-
nol. Oceanogr. 19:984-989.

Snyder, D. E. 1976. Terminologies for intervals of larval
fish development. Pages 41-60 in J. Boreman,
ed.. Great Lakes fish egg and larval identi-
fication: proceedings of a workshop. National
Power Plant Team, U.S. Fish and Wildl. Serv.
FWS/OBS-76-23.

Snyder, J. O. 1917. The fishes of the Lahontan system of
Nevada and northwestern California. U.S. Bur.
Fish. Bull. 35(1915- 16):33-86.

Vice, S. 1978. Vertical distribution of adult fish in Pyra-
mid Lake, Nevada. Great Basin Nat. 38:417-428.

1981. Species composition and relative abun-
dance of adult fish in Pyramid Lake. Great Basin
Nat. 41:395-408.

VuciNiCH, N., D. L. Galat, and M. C. Coleman. 1981.
Size-class structure and food habits of young-of-
year tui chubs (Gila bicolor, Girard), in Pyramid
Lake, Nevada. Cooperative Fishery Research
Unit, Colorado State University, Fort Collins. 64
pp.

Williams, J. E., and C. D. Williams. 1980. Feeding
ecology of Gila boraxobius (Osteichthyes: Cyprin-
idae) endemic to a thermal lake in southeastern
Oregon. Great Basin Nat. 40:101-114.

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TABLE OF CONTENTS

Life history of the Lahontan cutthroat trout, Salmo clarki henshawi, in Pyramid
Lake, Nevada. William F. Sigler, WilHam T. Helm, Paul A. Kucera, Steven
Vigg, and Gar W. Workman 1

A review of the genus Soliperla (Plecoptera: Peltoperlidae). Bill P. Stark 30

A bibhography of Colorado vegetation description. William L. Baker 45

Evaluation of a program to control hydatid disease in central Utah. Ferron L. Ander-
sen, John R. Crellin, Craig R. Nichols, and Peter M. Schantz 65

Influence of cryptogamic crusts on moisture relationships of soils in Navajo National

Monument, Arizona. Jack D. Brotherson and Samuel R. Rushforth 73

A vascular flora of the San Rafael Swell, Utah. James G. Harris 79

Pronghom responses to hunting coyotes. Timothy D. Reynolds 88

Floristics of the upper Walker River, California and Nevada.. Matt Lavin 93

Agropyron arizonicum (Gramineae: Triticeae) and a natural hybrid from Arizona.

Grant L. Pyrah 131

Species composition, distribution, and phytosociology of Kalsow Prairie, a mesic tall-
grass prairie in Iowa. Jack D. Brotherson 137

Deer mouse, Peromyscus maniculatus, and associated rodent fleas (Siphonaptera) in
the arctic-alpine life zone of Rocky Mountain National Park, Colorado. R. B.
Eads and E. G. Campos 168

Food of larval Tui chubs, Gila bicolor, in Pyramid Lake, Nevada. David L. Galat and

Nancy Vucinich 1'^

HE GREAT BASIN NATURALIST

ume 43 No. 2

April 30, 1983

Brigham Young University

BRARY

1P«

GREAT BASIN NATURALIST

Editor. Stephen L. Wood, Department of Zoology, 290 Life Science Museum, Brigham Youing

Game, 407 West Line Street, Bishop, California 93514 (Fish Biology).
Dr. Wayne N. Mathis, Chairman, Department of Entomology, National Museum of

Natural History, Smithsonian Institution, Washington, D.C. 20560 (Entomology).
Dr. Theodore W. Weaver III, Department of Botany, Montana State University, Boze-
man, Montana 59715 (Plant Ecology).
The Great Basin Naturalist was founded in 1939 and has been published from one to four
times a year since then by Brigham Young University. Previously unpublished manuscripts in
Enghsh of less than 100 printed pages in length and pertaining to the biological natural his-
tory of western North America are accepted. Western North America is considered to be west
of the Mississippi River from Alaska to Panama. The Great Basin Naturalist Memoirs was es-
tablished in 1976 for scholarly works in biological natural history longer than can be accom-
modated in the parent publication. The Memoirs appears irregularly and bears no geographi-
cal restriction in subject matter. Manuscripts are subject to the approval of the editor.

6-83 65066305

ISSN 017-3614

Hie Great Basin Naturalist

Published at Provo, Utah, by
Brigham Young University

ISSN 0017-3614

Volume 43 April 30, 1983 No. 2

UTAH FLORA: COMPOSITAE (ASTERACEAE)

Stanley L. Welsh'

Abstract.— A revision of the sunflower family, Compositae (Asteraceae), is presented for the state of Utah. In-
cluded are 613 taxa in 100 genera. Keys to genera, species, and infraspecific taxa are provided, along with detailed
descriptions, distributional data, and pertinent discussion. Many nomenclatural changes have been necessary to
bring Utah composites into line with contemporary interpretations of the family. They include: Artemisia tridentata
Nutt. var. wyomingensis (Beetle & Young) Welsh; Aster kingii D.C. Eaton var. barnebyana (Welsh & Goodrich)
Welsh; Brickellia microphylla (Nutt.) Gray var. ivatsonii (Robins.) Welsh; Chrysothamntis nauseosiis (Pallas) Britt. —
var. abbreviatiis (Jones) Welsh, var. arenarius (L.C. Anderson) Welsh, var. glareosus (Jones) Welsh, var. iridis (L.C.
Anderson) Welsh, and var. nitidus (L.C. Anderson) Welsh; Cirsium arizonicum (Gray) Petrak var. nidulum (Jones)
Welsh; Cirsium calcareum (Jones) Woot. & Standi. — var. bipinnatum (Eastw.) Welsh and var. pulchellum (Greene)
Welsh; Cirsium neomexicanum Gray var. utahense (Petrak) Welsh; Cirsium undulatum (Nutt.) Spreng. var. tracyi
(Rydb.) Welsh; Crepis runcinata (James) T. & G. var. ghiuca (Nutt.) Welsh; Erigeron speciosus (Lindl.) DC. — var.
mollis (Gray) Welsh and var. uintahensis (Cronq.) Welsh; Gutierrezia petradoria (Welsh & Goodrich) Welsh; Gutier-
rezia pomariensis (Welsh) Welsh; Haplopappiis racemosus (Nutt.) Torr. — var. sessiliflortts (Greene) Welsh and var.
prionophyllus (Greene) Welsh; Haplopappus watsonii Gray var. rydbergii (Blake) Welsh; Lygodesmia grandiflora
(Nutt.) T. & G. — var. arizonica (Tomb) Welsh and var. dianthopsis (D.C. Eaton) Welsh; Machaeranthera canescens
(Pursh) Gray — var. commixta (Greene) Welsh, var. leucanthemifolia (Greene) Welsh, and var. vacans (A. Nels.)
Welsh; Petradoria pumila (Nutt.) Greene var. graminea (Woot. & Standi.) Welsh; and Senecio spartioides T. & G. var.
multicapitatus (Greenm. in Rydb.) Welsh.

This paper is one of a series of works lead- ornamentals. Few actual crop or food plants

ing to a definitive treatment of the flora of are derived from members of this family.

Utah. Previous papers have dealt with the Only lettuce and sunflower are grown as

Brassicaceae, Fabaceae, Rosaceae, and mis- crops or as garden plants from the vast array

cellaneous smaller families. of species in this great family. Despite the

The sunflower family has long been recog- paucity of food plants, there are many oma-

nized for its great size and complexity both mental species. These have been included in

in Utah and elsewhere; it is probably the the present treatment only when they have

largest flowering plant family on earth. In escaped, or when they have been planted

Utah it consists of 100 genera and 613 taxa, routinely for many years, and when speci-

of which some 40, or about 7 percent, are in- mens have been preserved in the herbarium,

troduced. The 573 indigenous taxa comprise The cultivated flora requires a separate in-

about 20 percent of the flora native to the tensive effort not herein attempted,
state. This large family is apparently unique The importance of members of this family

in total numbers, but is also unique in having to wildlife, both as cover and as food, is well

such a small proportion of adventive taxa. known by range managers. Despite the pres-

The figures are misleading, to an extent, be- ence of chemical substances produced by the

cause of the omission of numerous cultivated plants, which impart unpleasant flavors or

'Life Science Museum and Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602.

179

180

Great Basin Naturalist

Vol. 43, No. 2

even poisonous substances, many of them are
eaten by both wildlife and by livestock. Sage-
brush and rabbitbrush species are well known
for their value in reclamation of sites requir-
ing rehabilitation.

Negative values are reflected in the weedy
species that occupy cultivated lands, in the
mechanically injurious species (such as
thistles and their relatives), and in the numer-
ous poisonous kinds. Livestock losses have
been reported in literature from utilization of
species of the family, i.e., Baileya, Hyme-
noxys (especially richardsonii), Oxytenia ace-
rosa, Psilostrophe, Senecio, and Tetradymia.
Other plants are rendered unpalatable by
their complex biochemical compounds, and
they tend to increase on range lands where
other more palatable plants are eaten selec-
tively. Gutierrezia and Chrysothamnus spe-
cies fit this latter category, although selected
phases of the same Chrysothamnus species
might be palatable or differentially palatable.
Certainly there is much room for in-
vestigation of members of this huge
assemblage.

The largest genus, and one of the most
complex taxonomically, is Erigeron, with
more than 60 taxa. Without the able mono-
graph by Cronquist, and lacking his coopera-
tive help, the species of Erigeron would have
been much more difficult to interpret. Be-
cause of his knowledge, his dedication, and
his helpful cooperation, this treatment is ded-
icated to him. The same kinds of problems,
made even more complicated by extensive
hybridization, is true for Senecio, which
Barkley has treated so competently. Fortu-
nately, monographs or revisions are available
for most of the larger genera. However, no
modem work is definitive for Aster, which
requires interpretation on a cosmopolitan,
rather than provincial, basis. Also, Artemisia
has not been treated in its entirety for several
decades. Much research on the nature of the
biochemical constituents of Artemisia has
shed light on the taxonomy of the group.

Difficulties in the composite genera and
species, aside from those involving the great
number of taxa, include those due to hybridi-
zation, ploidy level, and apomixis. Blending
of morphological forms due to hybridization,
subtle changes due to variations in chromo-
some sets, and the problems of interpretation

of apomictic races each lead to problems not
easily soluble. And the end results are subject
to a variety of preliminary conclusions, each
subject to change as additional information
becomes available. Further, generic lines in
the family are not definitive, with aggrega-
tions of species sometimes representing con-
venience rather than actual affinities. Recent
workers have tended to segregate traditional
genera into finer subunits or additional gen-
era. This seems to represent a continuing
trend, and it seems probable that some of the
genera treated herein will be broken up in
the future. The weight of evidence for doing
so lies with the future workers. Those who
find it modern to give "new" generic inter-
pretations are often merely opting previous
workers, whose interpretations were flawed.
Another later generation will opt for a differ-
ent set of names based on what they consider
to be "modern."

Some of the species are edaphic specialists,
occurring on definite substrates of restricted
aerial or elevational distribution. The woody
asters (Xylorhiza) are selenophytes and are re-
stricted to shales and silt- and mudstones of
fine-textured geological formations in eastern
to southwestern Utah.

The present work should be considered
tentative at best. Despite the large number of
specimens available for study (13,700), many
of the taxa are imperfectly known and distri-
butions are yet to be understood. Mono-
graphic work is required for practically all
groups, whether completed in the recent past
or not. It is hoped that this work will provide
a useful summary for those who work with
Utah composites.

To the extent possible the work presented
below contains the most modern names for
the entities involved, based on application of
contemporary International Rules of Bo-
tanical Nomenclature. Following the name of
the entities are the synonyms that apply to
Utah plants specifically. No attempt has been
made to list synonymy exhaustively. The
basionym has been included more or less uni-
formly, and an attempt has been made to cite
all synonyms based on Utah plants, including
a brief notation of type locality.

The discussion following the description of
each species, or the name of the infraspecific
taxon, includes the plant communities in

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

181

which the taxon occurs. They are arranged in
ascending order from dry low elevation com-
munities to mesic high elevation commu-
nities. Community data is followed by eleva-
tional range (given in meters), counties of
known occurrence (in alphabetical order),
and the distribution outside Utah. At the end
of the discussion are two numbers, the first,
in Arabic niunerals, indicates the number of
Utah specimens examined for the taxon and
the second, in Roman numerals, is the num-
ber collected in Utah by the author. The
numbers are given to provide the reader with
the basis of understanding of the taxon by the
writer and his familiarity with the plants in
the field. The plants are not equally well
known by this writer, and the user should be
able to make judgements when the taxonomy
presented herein does not adequately ap-
proach the conditions as noted in the field.
The approach to reality is always an approx-
imation, and much improvement will take
place in the future, as more information is
derived from specimens not now available for
the present study.

COMPOSITAE (ASTERACEAE)

Sunflower Family

Annual, biennial, or perennial herbs, or
shrubs; leaves alternate, opposite, or whorled,
simple, pinnatifid, or compound; in-

florescence of involucrate heads, these soli-
tary or several in corymbose, racemose, pan-
iculate, or cymose clusters; flowers few to
numerous on a common receptacle, sur-
rounded by green bracts forming a cup-
shaped, cylindrical, or urn-shaped involucre
enclosing the flowers in bud; heads entirely
of tubular (disk) corollas, entirely of ligulate
(ray) corollas, or with tubular corollas form-
ing a central disk and an outer radiating row
of ligulate corollas; receptacle flat, convex,
conic, or cylindric, naked or bearing chaffy
bracts, scales, or hairs; calyx lacking, or
crowning the summit of the ovary and modi-
fied as a pappus of capillary bristles, scales,
or awns; stamens alternate with corolla lobes;
filaments free (rarely connate); the anthers
united and forming a tube (rarely separate);
ovary inferior, of 2 carpels, 1-loculed and
with a single ovule; styles 1, 2-cleft, exserted
through the anther tube; fruit an achene.
Note: All involucral measurements are from
dried pressed herbarium specimens. The
width measurements are sometimes broader
than in fresh material.

Arnow, L., B. Albee, and A. Wyckoff.
1980. Flora of the central Wasatch
Front, Utah. Univ. of Utah Printing
Service, Salt Lake City. 663 pp.
Meyer, S. E. 1976. Annotated checklist of
the vascular plants of Washington
County, Utah. Unpublished thesis,
Univ. of Nevada, Las Vegas. 276 pp.

1. Corollas all raylike; plants usually with milky juice KEY 1

— Corollas not all raylike, some or all of them tubular; juice seldom if ever milky 2

2(1). Corollas all tubular; no ray flowers present, or the rays vestigial and minute

KEY II

— Corollas not all tubular; ray flowers present 3

3(2). Pappus of capillary bristles, at least in part KEY III

— Pappus of awns or scales, or lacking 4

4(3). Pappus lacking KEY IV

— Pappus present, of awns or scales KEY V

KEY I.

Corollas all ray like; plants usually with milky juice.

Pappus lacking 2

Pappus present 3

182 Great Basin Naturalist Vol. 43, No. 2

2(1). Rays 10-20 mm long; plants glabrous, with leaves in basal rosette Atrichoseris

— Rays 5-7 mm long; plants pubescent, with well-developed cauline leaves .... Lapsana

3(1). Pappus, at least in part, of plumose bristles 4

— Pappus of simple bristles, of awns, or of scales 7

4(3). Plants acaulescent, with merely bracteate stems Hypochaeris

— Plants caulescent 5

5(3). Achenes not beaked, truncate at apex; involucres usually less than 15 mm long

Stephanomeria

— Achenes tapering or beaked at apex; involucres usually more than 15 mm long 6

6(5). Leaves pinnatifid; corollas white or pinkish; involucre with an outer series of

short bractlets; southern Utah Rafinesquia

— Leaves not pinnatifid, entire; corollas yellow or purplish; involucre lacking
short outer bractlets; widespread Tragopogon

7(3). Pappus of 1-3 series of unawned or awned scales 8

— Pappus of capillary bristles 9

8(7). Pappus of 2 or 3 series of unawned scales; corollas blue, closing by mid-
morning Cichorium

— Pappus scales in a single series, awned; corollas yellow, not closing by mid-
morning Microseris

9(7). Achenes more or less flattened; stems leafy; heads in panicles or in umbellate

clusters 10

— Achenes not flattened; stems leafy or scapose; heads solitary or variously dis-
posed 11

10(9). Involucres cylindric or ovoid-cylindric; achenes beaked; flowers yellow or

blue Lactuca

— Involucres broadly campanulate to hemispheric; achenes not beaked; flowers
yellow Sonchus

11(9). Corollas pink or purplish 12

— Corollas yellow or yellowish, or white or cream colored 14

12(11). Plants annual; heads mainly 5-7 mm long (from base of involucre to tip of

pappus) Prenanthella

— Plants perennial; heads mainly 8-20 mm long or more 13

13(12). Plants with rigid spine-tipped branches Stephanomeria

— Plants unarmed, the branches soft Lygodesmia

14(11). Leaves all basal; heads solitary on scapose peduncles 15

— Leaves not all basal, the stems leafy; heads not on scapose peduncles 17

15(14). Achenes not beaked, truncate; pappus bristles barbellate Microseris

— Achenes beaked or tapering to apex; pappus not of barbellate bristles 16

16(15). Achenes 10-ribbed or 10-nerved, not spinulose; involucral bracts usually imbri-
cated in several series Agoseris

— Achenes 4- to 5-ribbed, spinulose, especially near apex; principal bracts in a
single series, the outer much shorter Taraxacum

April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 183

17(14). Achenes ridged or tuberculate between the angles; leaves either crustaceous

margined or peduncles stipitate-glandular; southwestern Utah 18

— Achenes striate between the angles; leaves and peduncles otherwise (rarely
glandular setose in some Crepis species); widely distributed 19

18(17). Plants depressed annuals with crustaceous-margined leaves, not stipitate-
glandular; achenes abruptly beaked, transversely ridged between the ribs

Glyptopleura

— Plants erect, lacking crustaceous-margined leaves, conspicuously stipitate-
glandular above; achenes tapering to a beak, not transversely ridged Calycoseris

19(17). Pappus bristles early deciduous, more or less united below and falling together,

only a few of the stout outer ones may be persistent Malacothrix

— Pappus bristles persistent or tardily deciduous, and then falling separately 20

20(19). Pappus tan to brown; involucral bracts not thickened Hieracium

— Pappus white or whitish; involucral bracts somewhat thickened at base or on
midrib Crepis

KEY II.

Corollas all tubular; no ray flowers present.

1. Heads unisexual, the pistillate heads with 1-4 flowers enclosed in involucre; in-
volucre burlike or nutlike, only style tips exserted 2

— Heads perfect or unisexual; involucre not burlike or nutlike 4

2(1). Involucral bracts of the staminate heads separate; fruiting involucres burlike,

covered with hooked appendages Xanthium

— Involucral bracts of the staminate heads united; fruiting involucres various but,

if burlike, lacking hooked appendages 3

3(2). Shrubs; fruiting involucre with several transverse, scarious wings; leaves or

their lobes linear-filiform Hymenoclea

— Shrubs or herbs; fruiting involucre lacking transverse wings; leaves and their
lobes not linear-filiform Ambrosia

4(1). Stamens not united by their anthers; flowers always unisexual, the pistillate co-
rollas none or much reduced 5

— Stamens with united anthers or rarely not united in some species with perfect
flowers, at least some flowers usually perfect 7

5(4). Achenes long-villous; leaves or their lobes linear-filiform Oxytenia

— Achenes not long-villous; leaves or their lobes not linear-filiform 6

6(5). Pistillate flowers subtended by large, chaffy scales simulating inner involucral

bracts; achenes with pectinate or winged margins Dicoria

— Pistillate flowers subtended by chaffy scales or these lacking; achenes without
pectinate or toothed wings Iva

7(4). Involucral bracts with translucent, usually yellow or orange dots Porophyllum

— Involucral bracts without distinct dots; pappus various, but not as above 8

8(7). Pappus of capillary bristles, at least in part, these smooth, scabrous, barbellate,

or plumose 9

— Pappus lacking or, if present, not of capillary bristles 41

184 Great Basin Naturalist Vol. 43, No. 2

9(8). Leaves opposite or whorled, some or all cauline 10

— Leaves alternate, at least basally, or basal and actually alternate 13

10(9). Corollas yellow; involucral bracts in 1 series or in 2 series, but all equal in

length Arnica

— Corollas white, ochroleucous, flesh colored, blue, or purple; involucral bracts

in 2 to several series 11

11(10). Pappus double— the outer series of short scales, the inner series of capillary

bristles; shrubs with white bark Hofmeistera

— Pappus single, or else plants herbaceous; shrubs or herbs 12

12(11). Achenes 5-angled or 5-ribbed; involucral bracts subequal or in 2 series ... Eupatorium

— Achenes 10-angled or 10-ribbed; involucral bracts imbricated in several series

of different lengths Brickellia

13(9). Leaves spinescent, usually with spiny teeth or lobes, rarely entire but then

with spine-tipped apex, thistlelike 14

— Leaves entire, denticulate or lobed, lacking spines, not thistlelike 18

14(13). Corollas of some or all flowers bilabiate; basal leaf axils woolly; leaves spin-

ulose-dentate; flowers pink; arid sites in Kane and Washington counties Perezia

— Corollas not bilabiate; leaves not or seldom spinulose-dentate; basal leaf axils
woolly; flowers pink-white or cream; various distribution 15

15(14). Pappus of 2 series of awns, the outer long and naked, the inner short and hispi-

dulous; flowers yellow Cniciis

— Pappus of plumose or barbellate capillary bristles; flowers not yellow 16

16(15). Pappus bristles plumose (rarely some otherwise); receptacle densely bristly .. Cirsium

— Pappus bristles merely barbellate 17

17(16). Receptacle densely bristly, not fleshy or honeycombed; heads nodding Carduus

— Receptacle not bristly or scarcely so, fleshy and honeycombed; heads not nod-
ding Onopardum

18(13). Receptacle with dense bristles or narrow, chaffy scales between disk flowers 19

— Receptacle naked or at most short-hairy, never with dense bristles or scales 21

19(18). Involucral bracts with hooked spines; lower leaves large (resembling rhubarb),

cordate at base Arctium

— Involucral bracts without spines, or spines not hooked; lower leaves not large

and cordate at base 20

20(19). Receptacle chaffy except in center; plants small, woolly Filago

— Receptacle chaffy throughout; plants not small and woolly Centaurea

21(20). Heads unisexual; plants dioecious (staminate flowers may have styles but ovary

does not develop) 22

— Heads with at least central flowers perfect 24

22(21). Plants shrubs or else woody at base, not tomentose; leaves sometimes toothed

or lobed; involucral bracts not strongly scarious margined Baccharis

— Plants herbaceous, more or less tomentose; leaves entire; involucral bracts
strongly scarious, at least along margins 23

23(22). Pappus bristles of pistillate flowers united at base and falling together; pappus
bristles of staminate flowers usually club shaped at apex; plants usually less
than 30 cm tall; basal leaves commonly in a rosette; cauline leaves reduced
and different in shape; leaves usually tomentose on both sides Antennaria

April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 185

— Pappus of pistillate flowers separate at base and falling separately; pappus
bristles of staminate flowers not club shaped at apex; plants mostly over 30 cm

tall; leaves all alike, usually green and glabrate above Anaphalis

24(21). Stems longitudinally brown-striate; involucral bracts imbricate, chartaceous,
the inner with scarious margins and broadly rounded apices; shrubs with yel-
low flowers, of western Millard County Lepidospartum

— Stems striate or not; involucral bracts scarious, hyaline, or herbaceous but not

as above; herbs, or shrubs with flowers and distribution various 25

25(24). Involucral bracts scarious or hyaline (only partly so in Pluchea) 26

— Involucral bracts herbaceous, at least in the center 28

26(25). Involucral bracts subscarious; corollas purplish; plants not tomentose, slender

woody shrubs Pluchea

— Involucral bracts scarious; corollas rarely purplish; plants tomentose, prostrate

to erect herbs 27

27(26). Plants perennial, subdioecious pistillate heads usually with a few central, per-
fect flowers Anaphalis

— Plants annual or perennial, not dioecious; heads all alike, the marginal flowers
pistillate and central ones perfect Gnaphalium

28(25). Involucral bracts in a single series, a few very short ones may be present at the

very base 29

— Involucral bracts of 2 or more series, these often of different lengths 32

29(28). Plants woody, shrubs; involucral bracts 4-6 per head Tedradymia

— Plants herbaceous; bracts more than 6 per head 30

30(29). Plants annual; heads with inner flowers perfect, the outer pistillate Conyza

— Plants perennial; heads with all flowers perfect 31

31(30). Style branches with a tuft of hairs near the truncate apex; involucral bracts in

1 series only (a few short bracts may be present) Senecio

— Style branches without a tuft of hairs near the truncate apex; involucral bracts
actually in 2 or more series Erigeron

32(28). Pappus double, the outer series of short scales, the inner ones of capillary

bristles; shrubs with white bark Hofmeistera

— Pappus simple or else the plants herbaceous 33

33(32). Plants annual 34

— Plants perennial 36

34(33). Plants low, depressed, scurfy pubescent herbs; leaves broadly ovate or round-
ish, entire or toothed Psathyrotes

— Plants not as above 35

35(34). Leaves all entire Aster

— Leaves toothed or lobed, at least the lower Conyza

36(33). Involucral bracts in more or less distinct vertical rows Chrysothamnus

— Involucral bracts not in vertical rows 37

37(36). Involucral bracts usually in 1 subequal series Erigeron

— Involucral bracts imbricate, in 2 or more series 38

186 Great Basin Naturalist Vol. 43, No. 2

38(37). Involucral bracts not longitudinally striate; flowers commonly yellow .. Haplopappus

— Involucral bracts longitudinally striate; flowers commonly cream to ofl-white,

or pink to purplish 39

39(38). Flowers pink to purplish; plants of northwestern Utah Eupatorium

— Flowers cream to white; plants of various distribution 40

40(39). Pappus plumose; plants perennial herbs Kuhnia

— Pappus scabrous or hispidulose; plants shrubs or herbs Brickellia

41(9). Receptacle with bristles or chaffy scales among the flowers 42

— Receptacle naked or merely short-hairy 50

42(41). Receptacle densely bristly Centaurea

— Receptacle with chaffy scales 43

43(42). Plants low woolly annuals; outer bracts boat shaped and enclosing the achenes 44

— Plants various, but not low and woolly; outer bracts various but not usually en-
closing the achenes 45

44(43). Stem leaves opposite; style lateral Psilocarpus

— Stem leaves alternate; style terminal Stylocline

45(43). Involucral bracts in 2 distinct sets — the outer herbaceous, the inner differing

in shape and texture; leaves opposite, at least below, or alternate 46

— Involucral bracts not in 2 unlike sets; leaves alternate or basal 47

46(45). Leaves alternate throughout; outer involucral bracts about 5, spreading, her-
baceous, the irmer (1-3 subtending pistillate flowers) larger and broader, be-
coming strongly accrescent and hooded in fruit Dicoria

— Leaves opposite, at least below; outer involucral bracts various, but not as
above, not accrescent and hooded in fruit Thelesperma

47(45). Involucral bracts in 1 series, boat shaped, each bract enclosing a marginal

flower; rays short, yellow Madia

— Involucral bracts in 1 or more series, not boat shaped and enclosing marginal
flowers; rays lacking 48

48(47). Plants woody shrubs; mostly along the canyons of the Colorado and Green riv-
ers Encelia

— Plants herbaceous; widely distributed 49

49(48). Receptacles high-conical, mostly over 3 cm long; stems leafy Rudbeckia

— Receptacles merely convex, much less than 3 cm long; leaves all basal Enceliopsis

50(41). Pappus none 51

— Pappus present 54

51(50). Leaves opposite, some cauline, somewhat connate at base; plants of Grand, San

Juan, and Tooele counties Flaveria

— Leaves alternate or basal 52

52(51). Heads numerous, in spikes, racemes, or panicles; anthers with acute tips; re-
ceptacles flat; plants woody or herbaceous Artemisia

— Heads solitary on ends of stems, or sometimes corymbose or capitate; anthers
with rounded tips; receptacles convex or conic; plants herbaceous, or woody
only at base 53

April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 187

53(52). Plants annual; heads solitary or paniculately arranged; leaves green and

glabrous Chamomilla

— Plants perennial; heads corymbose or capitate; leaves usually silvery-canescent
Chrysanthemum

54(50). Plants dioecious shmbs Baccharis

— Plants not dioecious herbs or shrubs 55

55(54). Pappus of 2-8 caducous awns; plants usually strongly glutinous Grindelia

— Pappus various, but not of 2-8 caducous awns 56

56(55). Leaves and involucre conspicuously punctate with translucent oil glands .... Dyssodia

— Leaves and involucre sometimes impressed-punctate, but without translucent

oil glands 57

57(56). Pappus of 12 or more scale or bristlelike segments, these nearly or quite as

long as achene 58

— Pappus of fewer than 12 scalelike segments or else much shorter than achene 59

58(57). Pappus of 12-16 linear, acuminate awns; involucres glutinous; leaves 3- to 5-

nerved Vanclevea

— Pappus of ca 35 flattened, silvery scales and bristles of different widths; in-
volucres not glutinous; leaves 1-nerved Acamptopappus

59(57). Achenes strongly compressed; pappus of 1 or 2 slender awns Laphamia

— Achenes not compressed or, if so, then pappus not of 1 or 2 slender awns 60

60(59). Pappus a crown with margins entire or of short scales united into a crown 61

— Pappus not as above 63

61(60). Plants annual; heads solitary or paniculately arranged; flowers all perfect;

leaves green and glabrous Chamomilla

— Plants perennial; heads corymbose or capitate, rarely solitary; some marginal
flowers pistillate only; leaves mostly silvery-canescent 62

62(61). Plants 0.5-1 m tall; leaves doubly pinnately dissected, mainly 10-20 cm long ...

Tanacetum

— Plants mainly less than 0.3 m tall; leaves entire, once pinnately dissected, ter-
nate, merely toothed apically, or entire, mainly less than 10 cm long ... Sphaeromeria

63(60). Involucral bracts with a thin, scarious, white, yellow, or purplish margin and

tip Hymenopappus

— Involucral bracts without a scarious, colored margin and tip 64

64(63). Plants scapose; leaves roundish, entire, or crenate Chamaechaenactis

— Plants leafy stemmed; leaves not roundish and entire or subentire 65

65(64). Pappus scales with a strong midrib; leaves lanceolate or linear, entire; southern

Utah Palafoxia

— Pappus scales nerveless or essentially so; leaves, at least in part, toothed to pin-
natifid; widely distributed Chaenactis

KEY III.

Corollas not all tubular; ray flowers present pappus of capillary bristles.

I. Rays white, pink, violet, or purple, not yellow 2

— Rays yellow or orange-yellow 9

188 Great Basin Naturalist Vol. 43, No. 2

2(1). Pappus of numerous unequal bristles, alternating with shorter, lacerate scales;

involucral bracts subequal; low winter annuals Monoptilon

— Pappus of numerous bristles; involucral bracts imbricate or subequal; plants
various, but seldom low winter annuals 3

3(2). Pappus, at least of disk flowers, of several to many rigid bristles; achenes pu-
bescent with 2-forked hairs or the hairs barbed at apex Townsendia

— Pappus, at least of disk flowers, of many capillary bristles, at least in part;
achenes glabrous or pubescent with simple hairs 4

4(3). Rays very inconspicuous, shorter than the tube and scarcely if at all exceeding

their pappus; central perfect flowers few; plants annual Conyza

— Rays usually conspicuous, longer than the tube and pappus; central perfect
flowers several to many; plants annual, biennial, or perennial 5

5(4). Involucres subequal, rarely somewhat graduated; rays usually narrow; style tips

very short, triangular, rounded, or obtuse Erigeron

— Involucres usually strongly graduated; rays comparatively broad; style tips
ovate and acute to subulate, usually lanceolate 6

6(5). Plants perennial, rhizomatous, or annual, or, if from a caudex, ordinarily less

than 10 cm tall (see also Aster kingii) 7

— Plants from a caudex or taproot 8

7(6). Low, white-rayed perennial herbs from spreading cordlike rootstocks, in arid

sites; flowering in springtime Leucelene

— Low to tall, white- to pink- or purple-rayed annual or perennial herbs from
rhizomes or fibrous roots (a caudex in A. kingii); mainly flowering in summer

and autumn Aster

8(6). Plants herbaceous, from a taproot, biennial or perennial; heads usually several

to numerous Machaeranthera

— Plants more or less woody, from a ligneous caudex; heads usually solitary and
large (primary selenophytes) Xylorhiza

9(1). Leaves opposite, at least below 10

— Leaves alternate throughout 12

10(9). Plants subshrubs Laphamia

— Plants herbaceous 11

11(10). Leaves with stiff marginal bristles; involucre and leaves with conspicuous oil

glands; plants annual Pectis

— Leaves without stiff marginal bristles; involucre and leaves without oil glands;
plants perennial Arnica

12(11). Plants 1-1.5 m tall, herbaceous; heads 3-5 cm wide; rays 1-2 cm long Inula

— Plants various, usually less than 1 m tall, or, if taller, woody; heads much small-
er; rays seldom to 1 cm long 13

13(12). Pappus of 2-8 stiff, caducous bristles; plants usually glutinous Grindelia

— Pappus of numerous, usually soft, persistent bristles 14

14(13). Pappus of about 20 twisted, flattish bristles Amphipappus

— Pappus of numerous, straight, capillary bristles 15

15(16). Pappus double, the inner of numerous bristles, the outer sometimes scalelike 16

— Pappus not double, of subequal capillary bristles only 17

April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 189

16(15). Leaves essentially filiform Conyza

— Leaves not filiform, linear-oblong or broader Heterotheca

17(15). Involucral bracts in distinct vertical ranks 18

— Involucral bracts not in distinct vertical ranks 19

18(19). Outer involucral bracts with loose herbaceous tips; erect stems perennial;

plants shrubs; leaves deciduous Chrysothamnus

— Outer involucral bracts without loose herbaceous tips; erect stems annual;
plants herbaceous; leaves persistent Petradoria

19(17). Involucral bracts in 1 series, frequently with some smaller bracts at base; style

branches truncate apically Senecio

— Involucral bracts neither in 1 series nor with smaller bracts at base; style
branches without truncate tips 20

20(19). Heads small, the involucres usually less than 6 mm high, usually very numerous
and densely paniculate, rarely racemose or corymbose; plants rhizomatous,
fibrous rooted Solidago

— Heads medium to large, the involucres usually more than 6 mm high, neither
very numerous nor densely paniculate; plants with taproots, occasionally also
rhizomatous Haplopappus

Key IV.

Corollas not all tubular; ray flowers present; pappus lacking.

1. Rays white, pink, or pink-purple, sometimes yellow at base 2

— Rays yellow, sometimes partly purplish or maroon 6

2(1). Receptacle naked 3

— Receptacle with chaffy scales 5

3(2). Leaves all basal; plants scapose Bellis

— Leaves not all basal, at least some cauline; plants caulescent 4

4(3). Receptacle broad and flattish; involucral bracts with a dark brown submarginal

line Chrysanthemum

— Receptacle convex, conic, or hemispheric; involucral bracts without a dark
brown submarginal line Chamomilla

5(3). Heads small, numerous, in dense, flattish or rounded cymose panicles; plants

perennial Achillea

— Heads comparatively large, solitary or few; plants annual or perennial Anthemis

6(1). Receptacles not chaffy 7

— Receptacles chaffy, at least toward the margin 12

7(6). Heads 1- or 2-flowered, in dense glomerate clusters, sessile in the forks of the

stem, or terminal and leafy involucrate Flaveria

— Heads several- to many-flowered, solitary on terminal peduncles 8

8(7). Plants woolly 9

— Plants not woolly 10

9(8). Rays persistent, becoming papery Baileya

— Rays not persistent Eriophyllum

190 Great Basin Naturalist Vol. 43, No. 2

10(8). Involucre and leaves with translucent oil glands Pedis

— Involucre and leaves without translucent oil glands 11

11(10). Rays conspicuous; involucral bracts acuminate, without scarious margins Bahia

— Rays minute; involucral bracts obtuse, with scarious margins Tanacetum

12(6). Ray achenes partly or wholly enfolded by their involucral bracts; plants an-
nual, glandular-viscid above Madia

— Ray achenes not conspicuously enfolded by their involucral bracts or, if so,
then plants perennial; plants perennial or, if annual, not glandular above 13

13(12). Involucre distinctly double, the outer bracts herbaceous, the inner ones broad-
er and united to about the middle Thelesperma

— Involucre not double, the bracts distinct to the base 14

14(13). Plants scapose perennials; leaves broad, silvery-pubescent, entire; heads very

broad Enceliopsis

— Plants leafy stemmed or subscapose; leaves various but not broad and silvery-
pubescent, or if so, then sagittate; heads broad or narrow 15

15(14). Plants subscapose; leaves variously dissected or sagittate; heads broad .. Balsamorhiza

— Plants with stems definitely leafy; leaves usually not dissected or sagittate 16

16(15). Plants shrubby; achenes conspicuously ciliate on the margins, notched at the

apex, very flat Encelia

— Plants herbaceous; achenes not conspicuously ciliate on the margins 17

17(16). Leaves doubly pinnately dissected; heads numerous in corymbose cymes Achillea

— Leaves simple, entire or toothed to lobed; heads few to several 18

18(17). Achenes 2-winged; disks 15-25 mm wide; leaves white-strigose beneath, green

above Verbesina

— Achenes not 2-winged; disks 6-15 mm wide; leaves green on both sides .... Heliomeris

Key V.
Corollas not all tubular; ray flowers present; pappus of awns or scales.

1. Receptacle chaffy 2

— Receptacle not chaffy, either naked or bristly 17

2(1). Pappus scales fimbriate; ray flowers 4 or 5, white, only slightly surpassing the

disk; introduced weedy plants, to be expected in Utah Galinsoga parviflora Cav.

— Pappus scales or awns not fimbriate; ray flowers various in size and color; in-
digenous or introduced 3

3(2). Receptacle bearing a row of chaffy scales between the ray flowers and the out-
er disk flowers, otherwise naked; pappus of 10-20 slender setiform scales Layia

— Receptacle chaffy throughout; pappus not of 10-20 slender scales 4

4(3). Ray achenes dorsiventrally compressed, the thickened margins attached to
a contiguous pair of infertile disk flowers and the subtending bract, and falling

as a unit; pulvinate herbs of eastern Utah and shrubs of southwestern Utah

Parthenium

— Ray achenes various, but not as above; herbs or shrubs 5

5(4). Pappus of awns only, without scales 6

— Pappus, at least in part, of scales 10

April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 191

6(5). Achenes flat and obcompressed; awns retrorsely hispid Bidens

— Achenes not obcompressed; awns not retrorsely hispid 7

7(6). Achenes plump; pappus of 2 to several caducous awns Helianthus

— Achenes flat, very strongly compressed; pappus various 8

8(7). Plants scapose; heads large, solitary Enceliopsis

— Plants leafy stemmed; heads medium sized, usually several 9

9(8). Plants shrubby; achenes narrowly white margined, the margin not continuous

between weak awns Encelia

— Plants herbaceous annuals; achenes strongly white margined, the margin con-
tinuous between stout awns Geraea

10(5). Achenes very flat, strongly compressed 11

— Achenes not very flat, usually much thickened 13

11(10). Leaves once to twice pinnatifid Anthemis

— Leaves not pinnatifid, entire or nearly so 12

12(11). Plants scapose Enceliopsis

— Plants leafy stemmed Helianthella

13(10). Pappus caducous (of 2 awns and rarely some scales) Helianthus

— Pappus persistent 14

14(13). Inner involucral bracts united to middle into a cup Thelesperma

— Inner involucral bracts not united into a cup 15

15(14). Receptacle merely convex; rays pistillate Wyethia

— Receptacle conic or cylindric; rays neuter 16

16(15). Involucral bracts subequal, in 2 or 3 series Rudbeckia

— Involucral bracts unequal, in 2 series, the inner ones shorter Ratibida

17(1). Rays white or purple 18

— Rays yellow, sometimes marked with purple 24

18(17). Pappus a short crown 19

— Pappus of awns or scales 21

19(18). Leaves entire or pinnately divided Chrysanthemum

— Leaves irregularly 2-3 times pinnately dissected 20

20(19). Plants annual; heads 1-2.5 cm wide; receptacle conic, hollow Chamomilh

— Plants biennial or perennial; heads 3-5 cm wide; receptacle hemispheric . Matricaria

21(18). Pappus of 1 plumose awn and a denticulate crown Monoptilon

— Pappus of 2 to several awns or scales 22

22(21). Plants dwarf woolly annuals Eriophyllum

— Plants armual or perennial, not woolly 23

23(22). Pappus of numerous awns or scales; involucral bracts conspicuously scarious-

margined Touinsendia

— Pappus of 4 or 5 stiff awns; involucral bracts obscurely scarious-margined

Rigiopappus

24(17). Receptacle densely bristly or hairy 25

— Receptacle naked 26

192 Great Basin Naturalist Vol. 43, No. 2

Heads very small; involucres less than 10 mm wide Gutierrezia

Heads medium sized; involucres more than 10 mm wide Gaillardia

Pappus of 4 hyaline scales united at the base; rays reddish purple to yellow ... Hulsea
Pappus a crown, or of cauducous or persistent awns or scales; rays mostly yel-
low 27

Pappus a mere crown or of caducous awns 28

Pappus persistent, of awns or scales 30

Pappus of 2-8 caducous awns; plants glutinous Grindelia

Pappus a short crown; plants seldom if ever glutinous 29

Leaves entire, bristly margined basally Pedis

Leaves 2- or 3-pinnate Tanacetum

Pappus of 1 or 2 awns or scales (rarely more) with or without a crown Perityle

Pappus of 4 to many awns or scales 31

Pappus of about 20 slender, twisted awns; rays 1 or 2 small Amphipappus

Pappus of 4-16 twisted or plane awns or scales; rays usually several 32

Pappus of 4 or 5 stiff, narrowly lanceolate awns; achenes linear, transversely

rugulose Rigiopappus

Pappus of scales, awn-tipped scales, or setose-dissected awns 33

Pappus of several scales dissected nearly to base; dwarf woolly annuals

Syntrichopappus

Pappus awns or scales not dissected or else plants perennial or woody 34

Pappus of several more or less united scales; rays broad, papery, and persistent

Psilostrophe

Pappus not of united scales; rays not papery and persistent (occasionally so in
Hymenoxys) 35

Leaves and involucre with conspicuous oil glands Dyssodia

Leaves and involucre without conspicuous oil glands 36

Achenes slender, elongate-clavate 37

Achenes stouter, oblong or obovoid 38

Plants woolly Eriophyllum

Plants merely strigose Platyschkuhria

Involucral bracts spreading or reflexed; receptacle convex to subglobose;

leaves decurrent Helenium

Involucral bracts appressed; receptacle almost flat; leaves not decurrent 39

Pappus of numerous scales; stems leafy; leaves linear or linear-spatulate, entire,

2.5 mm wide or less Gutierrezia

Pappus of about 5 scales; leaves lobed or, if entire, broader and mostly or en-
tirely basal Hymenoxys

AcAMPTOPAPPUs Gray strongly imbricate, the bracts broad,
Shrubs with white bark; leaves alternate, ^-ounded, the tip greenish, the margin
entire, 1-nerved; heads yellow, discoid, sub- scarious, erose; receptacle convex, fimbril-
globose, cymose at tips of branches; flowers late; style branches linear; achenes sub-
all fertile; involucral bracts ca 4-seriate, turbinate, densely villous; pappus persistent.

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

193

of ca 35 flattened silvery scales and bristles of
different widths.

Acamptopappus sphaerocephalus (Harv. &
Gray) Gray Goldenhead. [Haplopappus
sphaerocephalus Harv. & Gray]. Low
rounded shrubs to 1 m tall, much branched,
glabrous throughout or scabrous along some
leaf margins; leaves spatulate to almost lin-
ear, 4-28 mm long, 1-5 mm wide, obtuse to
acute, mucronulate, thick, sessile; heads sub-
globose, 6-10 mm high; involucre 4-6 mm
high. Blackbrush, indigobush, and creosote
bush communities at 850 to 1375 m in Kane,
San Juan, and Washington counties; Arizona,
Nevada, and California; 22 (iv).

Achillea L.

Perennial, rhizomatous, aromatic herbs,
with watery juice; stems erect or ascending;
leaves alternate, 1- to 3-pinnately dissected;
leaves several to many, borne in compact to
open corymbose cymes; involucral bracts im-
bricate in several series, chaffy, the margins
scarious and hyaline; receptacle chaffy; ray
flowers present, usually 3-12, pistillate, fer-
tile, yellow, white, pink, or pink-purple; disk
flowers mostly 10 or more, perfect, fertile;
pappus none; style branches flattened;
achenes compressed, callus margined,
glabrous, beakless.

1. Flowers yellow; leaves coarsely twice pinnately dissected; plants cultivated

A. filipendulina

— Flowers white, pink, or pink-purple; leaves finely 2-3 times dissected; plants

indigenous or cultivated A. millefolium

Achillea filipendulina Lam. Fernleaf Yar-
row. Herbs, the stems erect, 8-12 dm tall or
more, longitudinally furrowed and minutely
glandular; leaves 4-35 cm long, doubly pin-
natifid, the lateral lobes with one large lobe
on the upper side; heads numerous, borne in
hemispheric or flat-topped corymbose cymes;
involucre 3-4 mm high, the bracts with pale
scarous margins, villous; rays about 5, to 1
mm long, yellow; disk flowers 30-40, yellow;
achenes 1-2 mm long. Cultivated ornamen-
tal. Salt Lake and Utah counties, and to be
expected elsewhere; introduced from Asia; 2
(0).

Achillea millefolium L. Milfoil Yarrow.
Herbs, the rhizomes horizontal; stems ascend-
ing to erect, 0.5-10 dm tall, villous-tomen-
tose, simple or branched above; leaves 2-26
cm long, reduced upwards, pinnately once to
thrice dissected, the segments very slender;
heads numerous, borne in hemispheric or
flat-topped, corymbose cymes; involucres
4-6 mm high, the bracts dark to pale mar-
gined, villous to glabrate; rays usually about
5, 2-3.5 mm long, white to pink or pink-
purple; disk flowers 10-20; achenes 1-2 mm
long. Gravelly, sandy, and clayey soils in
sagebrush, pinyon-juniper, cottonwood, juni-
per, rabbitbrush, ponderosa pine, mountain
brush, aspen, Douglas fir, spruce-fir, and al-

pine tundra commtmities at 1070 to 3750 m
in all Utah counties; widely distributed in
North America; circumboreal. Two very sim-
ilar taxa are present in Utah; the common, in-
digenous ssp. lanulosum (Nutt.) Piper
(n=18), and the introduced, cultivated, ssp.
millefolium (n = 27). A trend is recognizable
within ssp. lanulosum; the high elevation
specimens tend to have dark involucral
bracts, fewer heads, and lower stature. These
alpine plants have been treated as var. alpi
cola (Rydb.) Garrett, but they intergrade
completely with specimens attributable to
var. lanulosa. Indeed, the two extremes can
be found mounted on the same herbarium
sheet, taken from the same locality; 133 (xv).

Agoseris Raf.

Perennial scapose herbs with milky juice,
from taproots; leaves all basal, entire to pin-
nately lobed or merely toothed; heads soli-
tary on a naked scape; involucral bracts in 2
to several series, herbaceous, or the inner
ones hyaline or nearly so; receptacle usually
naked; corollas all raylike, perfect, yellow to
orange, often drying pinkish or purplish; pap-
pus of capillary bristles; style branches semi-
cylindric; achenes angular or terete, prom-
inently nerved, usually beaked.

194

Great Basin Naturalist

Vol. 43, No. 2

1. Plants annual; achene beak 2-3 times as long as the body; rare in Utah

A. heterophylla

— Plants perennial; achene beak less than half to 2 or more times as long as the
body 2

2(1). Achene beak striate, mostly less than half as long as the body (longer in some

var. laciniata); flowers yellow, often drying bluish to pinkish A. glauca

— Achene beaks scarcely striate, more than half to 2 or more times as long as the
body 3

3(2). Flowers brownish orange to yellow-orange, often drying purplish; achene beak

less than twice as long as the body A. aurantiaca

— Flowers yellow, often drying bluish or pinkish; achene beak more than twice

as long as the body A. grandiflora

Agoseris aurantiaca (Hook.) Greene Orange
Agoseris. Plants 0.6-6.6 dm tall, from a
simple or branched caudex; leaves 3.5-36 cm
long, 0.5-3 cm broad, narrowly oblanceolate,
entire to toothed or lobed, villous to glabrate;
scapes villous-tomentose to nearly glabrous;
involucres 10-27 mm long, 10-42 mm wide,
the outer bracts villous to glabrate and cil-

iate, often purple spotted; corollas brownish
orange to yellow-orange, often drying pur-
plish; achene body 4-8 mm long, the slender
beak not striate, from more than half as long
to longer than the body. Two rather weak
and intergrading phases are recognized at va-
rietal rank.

1. Involucres with bracts subequal or nearly so, slender, tapering, some often over

20 mm long A. aurantiaca var. aurantiaca

— Involucres with bracts definitely imbricate, broad, and rounded apically or

abruptly tapering A. aurantiaca var. purpurea

Var. aurantiaca [Troximon aurantiacum
Hook.; A. arizonica Greene; A. gracilens
(Gray) Kuntze; A. longirostris Greene, type
from Fish Lake]. Sagebrush, mountain brush,
juniper, pinyon-juniper, and alpine meadow
communities at 1375 to 3355 m in Beaver,
Box Elder, Carbon, Duchesne, Juab, Salt
Lake, San Juan, Sevier, Tooele, Uintah, Utah,
and Wasatch counties; Alberta to British Co-
lumbia, south to California and New Mexico;
33 (v).

Var. purpurea (Gray) Cronq. [Troximon
aurantiacum var. purpureum Gray; A. pur-
purea (Gray) Greene; A. confinis Greene,
type from near Marysvale]. Mountain brush,
aspen, aspen-fir, and spruce-fir communities
at 1700 to 3425 m in Carbon, Emery, Grand,
Juab, Iron, Piute, Sanpete, Sevier, and

Wasatch counties; Montana to Arizona and
New Mexico; 20 (vi).

Agoseris glauca (Pursh) Raf. Pale Agoseris;
Mountain Dandelion. Plants perennial,
0.2-6.4 dm tall, from a simple or branched
caudex; leaves 2-26 cm long, 0.2-3 cm
broad, narrowly oblanceolate to linear or
spatulate to elliptic, entire or toothed to
lobed, villous to glabrate; involucres 12-28
mm high, 0.8-4 cm wide, the outer bracts vil-
lous to glabrous, ciliate or not, sometimes
purple spotted; corollas yellow, often drying
bluish to pinkish; achene body 4-10 mm
long, the striate beak stout, to half as long as
the body (slender and to as long as the body
in some var. laciniata). Three intergrading
and partially sympatric varieties are present
in Utah.

1. Leaves laciniately toothed or lobed; plants of broad distribution, common

A. glauca var. laciniata

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

195

— Leaves entire, rarely with a few teeth or lobes; plants variously distributed, lo-
cally common 2

2(1). Plants pubescent, at least below the heads; mainly of spruce-fir and alpine

communities, sometimes lower A. glauca var. dasycephala

— Plants glabrous throughout, mainly of lower elevation wet meadows, but some-
times of high elevation meadows A. glauca var. glauca

Var. dasycephala (T. & G.) Jeps. [Am-
mogeton scorzoneraefolius Shrad.; A. scorzo-
neraefolia (Shrad.) Greene; Troximon pumi-
lum Nutt.; A. pumila (Nutt.) Rydb.; A. glauca
var. pumila (Nutt.) Garrett; T. glaucum var.
dasycephalum T. & G.; A. villosa Rydb.].
Sagebrush, mountain brush, aspen, spruce-fir,
and alpine tundra communities at 1830 to
3385 m in Duchesne, Garfield, Iron, Kane,
Piute, Salt Lake, San Juan, Sanpete, Sevier,
Tooele, Uintah, and Utah counties; Alaska to
Manitoba and south to Colorado. Plants of
this variety pass by degree into each of the
following; 29 (v).

Var. glauca [Troximon glaucum Pursh; A.
isomeris Greene, type from the Uinta Moun-
tains]. Meadows at 2325 to 3660 m in Box El-
der, Duchesne, Iron, Juab, Sanpete, Sevier,
Svimmit, Uintah, and Wasatch counties; Brit-
ish Columbia to Manitoba and south to Cali-
fornia and Arizona; 22 (iv).

Var. laciniata (D.C. Eaton) Smiley [Troxi-
mon parviflorum Nutt.; A. parviflora (Nutt.)
D. Dietr.; A. glauca var. parviflora (Nutt.)
Rydb.; T. taracifolium Nutt.; A. taracifolia
(Nutt.) D. Dietr.; Macorhynchus glaucus var.
laciniatus D.C. Eaton; A. taraxacoides
Greene, type from near Marysvale; A. caud-
ata Greene, type from Salina Canyon; A. agr-
estis Osterh.; A. glauca var. agrestis (Osterh.)
Q. Jones]. Sagebrush, mountain brush, juni-
per, pinyon-juniper, Douglas fir, aspen, and
spruce-fir communities at 1300 to 3050 m in
Beaver, Box Elder, Carbon, Daggett, Du-
chesne, Emery, Garfield, Grand, Juab, Kane,
Millard, Piute, San Juan, Salt Lake, Sanpete,
Sevier, Summit, Tooele, Uintah, Utah, Wash-
ington, Wayne, and Weber counties; Wash-
ington to Montana and south to Arizona. The
phase designated as var. agrestis blends com-
pletely in our area with that treated herein as
var. laciniata; 84 (xii).

Agoseris grandiflora (Nutt.) Greene

[Stylopappus grandiflorus Nutt.]. Plants pe-
rennial, 1.5-4.5 (7) dm tall, from a simple or
branching caudex; leaves 8-25 cm long, 1-3
cm broad, narrowly oblanceolate, pinnatifid
to subentire, villous to glabrate; involucres
15-38 mm long, 20-43 mm wide, the outer
bracts villous-tomentose to glabrate, ciliate,
often suffused with purple; corolla yellow,
drying bluish to pinkish; achene body 4-7
mm long, the nerveless beak more than twice
as long as the body. Specimens tentatively as-
signed to this species are from Cache, Iron,
Tooele, and Washington counties, where
they occur in sagebrush and mountain brush
communities at 1830 to 2135 m; British Co-
lumbia to California and Nevada; 4 (i).

Agoseris heterophylla (Nutt.) Greene An-
nual Agoseris. [Macrorhynchus heterophyllus
Nutt.]. Plants annual, 0.3-2.5 (4) dm tall,
with 1 to several scapes from the base; leaves
1-20 (15) cm long, 0.3-1.5 cm wide, narrow-
ly oblanceolate, toothed or pinnatifid to en-
tire, all basal, or with some not strictly basal;
involucres 5-20 mm long, 4-10 mm wide,
sparingly villous with multicellular hairs, the
cross-walls purplish; corolla yellow, some-
times turning pinkish on drying; achene body
2-5 mm long, prominently ribbed or winged,
the beak 2-3 times as long as the body. Our
one collection (Diehl D29, 1899 BRY) is from
Salt Lake County at low elevation; British
Columbia to California and Arizona; 1 (0).

Ambrosia L

Annual or perennial herbs or shrubs; leaves
alternate or opposite, pinnately or palmately
lobed, toothed, or dissected; heads unisexual,
discoid; staminate heads in slender spicate,
bractless racemes; involucre 5- to 12-lobed;

196

Great Basin Naturalist

Vol. 43, No. 2

receptacle flat, bearing flattened filiform-se-
tose bracts; staminal filaments monadelphous,
the anthers scarcely united; pistillate heads
borne below the fertile ones, mostly axillary,
their involucres closed, nutlike, armed with

prickles arranged in one or more series; pistil
naked, the corolla lacking; pappus lacking.
Payne, W. W. 1964. A ree valuation of the

genus Ambrosia. J. Arnold Arboretum

45:401-438.

1. Plants woody shrubs of southwestern Utah 2

— Plants armual or perennial herbs, of various distribution 3

2(1). Leaves mainly less than 15 mm long, pinnately lobed, the lobes again toothed

or lobed, silvery-strigose overall A. dumosa

— Leaves mainly more than 20 mm long, merely toothed or lobed, the lobes not
again toothed or lobed, bicolored, the upper surface green, the lower surface
white-tomentose A. eriocentra

3(1). Leaves palmately lobed, the lobes serrate; plants tall coarse herbs A. trifida

— Leaves pinnatifid or pinnately lobed; plants slender herbs usually less than 5

dm tall 4

4(3). Leaves bicolored, the lower surface obscured by appressed white hairs; plants

low rhizomatous perennials A. tomentosa

— Leaves various, but not definitely bicolored; plants from taproots or rhizomes,

but, if the latter, not as above 5

5(4). Plants perennial, rhizomatous; leaves opposite A. psilostachya

— Plants annual; leaves mainly alternate 6

6(5). Lower stems and leaves with pustular-based, stiff, multicellular hairs; plants of-
ten with lower lateral branches decumbent-ascending; burs with spines in more
than one series A. acanthicarpa

— Lower stems lacking pustular-based hairs, all stems slender and curved ascend-
ing-appressed; burs with spines in one series A. artemisiifolia

Ambrosia acanthicarpa Hook. Bur Rag-
weed. [Franseria acanthicarpa (Hook.) Co-
ville]. Plants annual, 0.9-7.5 dm tall, often
branching from the base, the lower branches
commonly decumbent-ascending; pubescence
of stiff multicellular hairs, the bases pustular;
leaves mostly alternate, petiolate, the blades
0.9-4.5 cm long, 0.6-3.5 cm wide, bipinnati-
fid to pinnatifid; heads numerous in terminal
or axillary racemes; staminate above, pistil-
late below; staminate heads short-peduncu-
late, not bracteate; pistillate solitary or clus-
tered in upper axils, with 2-3 series of
flattened, curved spines. Blackbrush, salt
desert shrub, desert shrub, pinion-juniper,
and riparian communities, often in sandy sub-
strates, at 850 to 2000 m, in Beaver, Carbon,
Daggett, Duchesne, Emery, Garfield, Grand,
Iron, Juab, Kane, Millard, San Juan, Tooele,
Uintah, Utah, Washington, and Wayne coun-
ties; Washington to Saskatchewan, south to
California, Arizona, and Texas; 60 (xi).

Ambrosia artemisiifolia L. Common Rag-
weed. [A. elatior L.]. Plants annual, mostly
3-9 dm tall, branching from above the
middle; pubescence of lax multicellular hairs,
the bases not pustular; leaves alternate, or
the lower usually opposite, petiolate, the
blades 2.5-8.5 cm long, 1.9-7.5 cm wide, 1-
to 2-pinnatifid; heads numerous in terminal
or axillary racemes, the staminate above, pis-
tillate below, clustered or solitary, with 1
series of tuberculate spines. Moist disturbed
sites at 1375 to 1500 m in Juab and Utah
counties; widespread in North America; 11

(i)-

Ambrosia dumosa (Gray) Payne Bur-sage.
[Franseria dumosa Gray]. Shrubs, 2-6 dm
tall, rounded, much branched; branchlets
white, subspinescent; pubescence dense, stri-
gose; leaves alternate, petiolate, the blades
9-30 mm long, 5-15 mm wide, mostly 2-pin-
natifid, uniformly hairy on both sides; stami-
nate heads spicate, rather few; pistillate

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

197

heads often scattered among the staminate;
pistillate involucre maturing 20-35 lance-
subulate spines. Creosote bush, blackbrush,
and Joshua tree communities at 670 to 1000
m in Washington County; Arizona, Califor-
nia, and Mexico; 28 (i).

Ambrosia eriocentra (Gray) Payne [Fran-
seria eriocentra Gray]. Shrubs, 3-10 (12) dm
tall, aromatic, branchlets white, sub-
spinescent, pubescence of white tomentum
and coarse multicellular hairs; leaves alter-
nate, subsessile, sinuately toothed to lobed or
1-pinnatifid, 8-40 (50) mm long, 2-20 mm
wide; staminate heads more or less clustered;
pistillate heads 1-flowered; pistillate in-
volucre with 12-20 flattened, subulate spines.
Creosote bush, blackbrush, and Joshua tree
communities at 670 to 1000 m in Washington
County; Arizona, Nevada, and California; 12
(ii).

Ambrosia psilostachya DC. Western Rag-
weed. [A. coronopifolia T. & G.]. Perennial
herbs, mostly 3-6 dm tall, simple or branch-
ing above the middle; pubescence of harsh,
spreading, multicellular, pustular-based hairs
(at least in part); leaves opposite, at least be-
low, petiolate to subsessile, the blades 4-10
cm long, 2.5-4.5 cm wide, mostly once pin-
natifid; staminate heads in terminal or ax-
illary spicate racemes; pistillate involucres
merely tuberculate or quite unarmed. Mead-
ows, stream banks, and roadsides in sagebrush
and other communities at 1300 to 2100 m in
Davis, Juab, Millard, Salt Lake, Utah, and
Weber counties; Washington to Illinois,
south to Arizona and Mexico; 20 (ii).

Ambrosia tomentosa Nutt. [Franseria dis-
color Nutt.; F. tomentosa (Nutt.) A. Nels., not
A. tomentosa Gray]. Perennial rhizomatous
herbs, mostly 1-3.5 dm tall, branching from
above the base; pubescence of short, stiff, ap-
pressed hairs; leaves alternate, petiolate, the
blades 2-15 cm long, 0.4-3.5 cm wide, 1- to
3-pinnatifid; staminate heads racemose; pis-
tillate heads armed with 2 or 3 series of
coarse spines. Meadows and stream banks at
1300 to 1525 m in Davis, Duchesne, and
Grand counties (likely elsewhere); Wyoming
and Colorado; 5 (iii).

Ambrosia trifida L. Giant Ragweed. An-
nual, robust herbs, 10-15 dm tall or more;
pubescence spreading-hirsute to hispid, at
least above; leaves opposite, petiolate, the

blades palmately 3- to 5-lobed, or unlobed,
mainly 5-20 cm long, 4-15 cm wide,
scabrous on both surfaces, serrate; staminate
involucres 3-nerved; pistillate involucres
5-10 mm long, bearing short spines at the
tip. Uncommon (introduced?) weedy plants
of disturbed sites in Salt Lake County (likely
elsewhere); widely distributed in North
America; 1 (0).

Amphipappus T. & G.

Low shrubs; branches white-barked, divari-
cate; leaves alternate, entire, short-petiolate;
heads small, radiate, few flowered, clustered
at tips of branches; involucre in ca 3 series,
strongly imbricate, straw colored to greenish,
the bracts broad, rounded; receptacle fimbril-
late; ray flowers yellow, 1 or 2, small; disk
flowers 3-6, perfect; ray achenes hairy,
broadly oblanceolate, compressed, their pap-
pus of more or less united bristles, awns, or
scales; disk achenes undeveloped, glabrous or
sparingly pilose, their pappus of twisted, his-
pidulous bristles or scales.
Porter, C. L. 1943. The genus Amphipappus
Torr. & Gray. Amer. J. Bot. 30:
481-483.

Amphipappus fremontii T. & G. Chaff-
bush. Shrubs 3-8 dm tall, the herbage
scabrous-puberulent; leaves 5-12 mm long,
2-5 mm wide, oblanceolate to elliptic, cu-
neate basally, acute to obtuse and apiculate,
green; heads 4-6 mm high, the bracts green-
ish medially near the apex, the margins hyal-
ine and more or less erose. Joshua tree and
creosote bush communities at 700 to 900 m
in Washington County; Nevada, Arizona, and
California. Our material belongs to var. spin-
osus (A. Nels.) C. L. Porter [ssp. spinosus (A.
Nels.) Keck]; 4 (i).

Anaphalis DC.

Perennial, dioecious or polygamo-
dioecious, rhizomatous herbs, with watery
juice; stems ascending to erect, simple or
branched above; leaves simple, alternate, en-
tire; heads several to many, in hemispheric or
flat-topped corymbose cymes; involucral
bracts imbricate in several rows, chaffy,
scarious, white, or with a dark triangular bas-
al spot; receptacle naked; corollas of disk
flowers only, imperfect, whitish, the pistillate

198 Great Basin Naturalist Vol. 43, No. 2

heads sometimes bearing some central stami- chesne, Iron, Juab, Salt Lake, Summit,

nate flowers, the pistillate corollas tubular- Wasatch, and Washington counties; widely

filiform, the staminate corollas tubular-fun- distributed in North America; Asia; 33 (vi).
nelform; pappus of capillary bristles; style

branches somewhat flattened; achenes small, Antennaria Gaertn.

roughened, glabrous to sparingly hairy.

Anaphalis margaritacea (L.) Benth. & Perennial, dioecious herbs with stolons.
Hook. Pearly Everlasting. [Gnaphalium mar- caudices, or rhizomes, the juice watery; stems
garitaceum L.]. Plants 1.5-8 dm tall, the ascending to erect, usually simple; leaves
stems white villous-tomentose; leaves only simple, alternate and basal, the cauline gen-
gradually reduced upwards, 2.5-12 cm long, erally reduced upward; heads solitary to
0.5-2 cm wide, narrowly lanceolate to ob- many, borne in corymbose cymes; involucral
long, elliptic, or oblanceolate, sessile, entire, bracts imbricate in several rows, scarious (at
flat to slightly revolute, white-tomentose be- least marginally), often colored; receptacle
low, commonly less pubescent and greenish naked; corollas of disk flowers only, imper-
above; heads showy, the involucres 4-7 mm feet, whitish or tawny; pistillate corollas
high, 5-10 mm broad, the bracts pearly- tubular-filiform, the pappus of numerous
white, with a dark triangular base, glabrous; capillary bristles; staminate corollas tubular-
achenes about 1 mm long. Meadows, stream- funnelform, the pappus of few clavate to bar-
banks, and openings in ponderosa pine, bellate, usually flattened bristles; style
lodgepole pine, box elder, and aspen commu- branches slightly flattened; achenes terete to
nities at 1150 to 2700 m in Box Elder, Du- slightly compressed, glabrous or papillose.

1. Heads solitary; flowering stems usually less than 5 cm tall A. dimorpha

— Heads (1) 2 to many (see A. rosulata); flowering stems often more than 5 cm

tall 2

2(1). Upper leaf surface green; leaf blades broadly spatulate, rounded to obtuse

A. neglecta

— Upper leaf surface not notably different from the lower; blades seldom both
spatulate and rounded to obtuse 3

3(2). Plants not forming mats, lacking leafy stolons, some caespitose from caudex or

rhizomes 4

— Plants mat forming, with leafy stolons 6

4(3). Involucral bracts glabrous or nearly so, scarious near the base, white-opaque

apically A. luzuloides

— Involucral bracts densely tomentose in the lower half, opaque to dark with
pale scarious apices 5

5(4). Involucral bracts blackish in aspect, the tips pale and scarious A. pulcherrima

— Involucral bracts opaque white, somewhat darkened at the middle .. A. anaphaloides

6(3). Terminal scarious portion of involucral bracts dirty brownish to blackish green

on at least the middle and outer ones 7

— Terminal scarious portion of involucral bracts white to pink, with a dark basal
spot on some only 8

7(6). Terminal scarious portion of involucral bracts blackish green; plants usually al-
pine in Uinta, Wasatch, and Tushar Mountains, and on the Markagunt Plateau
A. alpina

— Terminal scarious portion of bracts merely discolored and pale brown, or the
inner bracts whitish at the tips; plants usually of lower elevations A. umbrinella

8(6). Flowering stems less than 5 cm tall; heads 1 or 2; plants of Garfield, Kane, and

Wayne counties A. rosulata

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

199

— Flowering stems commonly more than 5 cm tall; heads usually 3 or more;
plants of broad, or other, distribution 9

9(8). Involucral bracts with a black spot between the tomentose greenish base and

the opaque white-scarious apex A. corymbosa

— Involucral bracts lacking a conspicuous black spot 10

10(9). Involucres mostly 4-7 mm high, often bright pink; pistillate corollas mostly
2-4.5 mm long A. microphylla

— Involucres mostly 7-11 mm high, seldom pink; pistillate corollas mostly 5-8

mm long A. parvifolia

Antennaria alpina (L.) Gaertn. Alpine
Pussytoes. Plants caespitose from a caudex,
mat forming and stoloniferous, 2-13 cm tall;
basal leaves 0.6-2.2 cm long, 2-6 mm wide,
cuneate-oblanceolate to spatulate, acute to
obtuse or rounded apically, grayish tomen-
tose on both surfaces or greenish and sub-
glabrous above on some leaves; heads 3-5,
borne in subcapitate cymes; pistillate in-
volucres 5-7 mm high, villous-tomentose be-
low, the scarious tips of bracts uniformly
blackish or brownish green, all rather blunt
apically, often erose; staminate involucres
mostly 4-5 mm high, the scarious tips of
bracts often pale apically; achenes glabrous.
Lodgepole pine, spruce-fir, and alpine tundra
communities at 3050 to 3550 m in Daggett,
Duchesne, Piute, Salt Lake, Uintah, and Utah
counties; north to Alaska and east to Lab-
rador; circumboreal. Our material belongs to
var. media (Greene) Jeps. [A. media Greene;
A. austromontana E. Nels., type from Tushar
Mountains]. There is a tendency for some
specimens to approach A. parvifolia in the
Uinta Mountains and A. umbrinella else-
where; 27 (vii).

Antennaria anaphaloides Rydb. Pearly
Pussytoes. Plants from a caudex, not mat
forming or stoloniferous, 1.5-3.5 (5) dm tall;
basal leaves 2.5-19 cm long, 4-18 mm wide,
narrowly oblanceolate to elliptic, tomentose
on both surfaces; heads several to many in
branching or compact cymes; pistillate in-
volucres 5-8 mm high, villous-tomentose be-
low, the scarious tips opaque-white, all
rounded or obtuse, often erose; staminate in-
volucres 5-8 mm high, similar to the pistil-
late; achenes glabrous. Aspen, spruce-fir,
sagebrush, and mountain brush communities
at 2440 to 3325 m in Daggett, Summit, and
Uintah coimties; British Columbia to Mon-
tana and south to Nevada and Colorado; 7 (i).

Antennaria corymbosa E. Nels. Plains Pus-
sytoes. [A. nardina Greene]. Plants caespi-
tose, mat forming and stoloniferous, 5-26 cm
tall; basal leaves 0.6-3.7 cm long, 2-6 mm
wide, narrowly oblanceolate to spatulate,
acute to obtuse apically, gray to greenish and
tomentose on both surfaces; heads commonly
3-8, in compact to branching cymes; pistil-
late and staminate involucres 4.5-6 mm high,
the bracts green and tomentose basally, with
a dark spot at the base of the white or sordid
terminal portion; achenes puberulent. Alpine
tundra, krumholz, spruce-fir, lodgepole pine,
and willow-alder communities, often along
stream banks and in wet meadows or bogs, at
2240 to 3355 m in Beaver, Duchesne, Gar-
field, Summit, and Uintah counties; Montana
and Idaho to Colorado and California(?). The
main body of the species in Utah lies in the
Uinta Mountains, with outliers in the Stans-
bury and Tushar mountains, and in the Mark-
agunt Plateau; 22 (iii).

Antennaria dimorpha (Nutt.) T. & G. Low
Pussytoes. [Gnaphalium dimorphum Nutt.; A.
dimorpha var. macrocephala D.C. Eaton,
type from Salt Lake City]. Plants caespitose,
mat forming, rooting from short caudex
branches, not truly stoloniferous, 1-5 (7) cm
tall; basal leaves narrowly oblanceolate,
0.6-4 cm long, 1-14 mm wide, acute api-
cally, grayish tomentose on both sides; heads
solitary, terminal on short leafy stems; pistil-
late involucres (7) 10-18 mm long, the bracts
strongly imbricated, slender, attenuate, green
at base, suffused with brown above the base,
the apical portions yellowish to brownish
scarious; staminate involucres 6-9 mm long,
tomentose at the base, brown above the base,
the broad apical portion hyaline to scarious;
achenes puberulent. Mat-saltbush, sagebrush,
juniper, oak-serviceberry, ponderosa pine,
and spruce-fir-lodgepole pine communities at

200

Great Basin Naturalist

Vol. 43, No. 2

1430 to 3050 m in Beaver, Daggett, Du-
chesne, Iron, Juab, Millard, Salt Lake, San-
pete, Sevier, Summit, Tooele, Utah, and
Washington counties (and certainly else-
where); British Columbia to Montana, south
to California, Nevada, Colorado, and Nebras-
ka; 37 (viii).

Antennaria luzuloides T. & G. Rush Pus-
sytoes. [A. oblanceolata Rydb.]. Caespitose
from a caudex, 1.1-5 (7) dm tall; basal leaves
2-5 (8) cm long, 2-8 mm wide, greenish, to-
mentose on both surfaces; heads numerous in
a compact or more often branched corym-
bose inflorescence; pistillate and staminate
involucres similar, 4-5 mm high, glabrous to
the base, the bracts brownish scarious and
more or less hyaline below, opaque whitish
above; achenes puberulent. Openings in as-
pen-conifer and lodgepole pine-spruce com-
munities at 2950 to 3050 m in Duchesne and
Summit counties; British Columbia to Mon-
tana, south to California, Nevada, and Colo-
rado; 7 (i).

Antennaria microphylla Rydb. Rosy Pus-
sytoes. [A. rosea Rydb.; A. concinna E. Nels.;
A. arida A. Nels.]. Plants caespitose, stoloni-
ferous and mat forming, 0.4-3 (4) dm tall;
basal leaves 0.5-3 cm long, 2-8 mm wide,
oblanceolate to spatulate; heads 2-13 (or
more), in congested to open cymes; pistillate
involucres 4-7 mm high, the bracts tomen-
tose below, greenish or scarious below the
middle, often somewhat brownish below the
scarious, whitish or pinkish, terminal portion;
mainly known from pistillate individuals;
achenes glabrous or sparingly hispidulous.
Sagebrush, juniper, ponderosa pine, Douglas
fir, lodgepole pine, spruce-fir, and alpine
meadow communities at 1830 to 3450 m in
Beaver, Box Elder, Carbon, Daggett, Du-
chesne, Emery, Garfield, Iron, Juab, Kane,
Millard, Piute, Rich, Salt Lake, Sanpete, Se-
vier, Summit, Tooele, Uintah, Utah, Wasatch,
Wayne, and Weber counties; Alaska to On-
tario, south to California and New Mexico;
105 (xxiii).

Antennaria neglecta Greene Field Pus-
sytoes. [A. marginata Greene]. Plants caespi-
tose, stoloniferous and mat forming, 5-15 (25)
cm tall; basal leaves 1.8-3.5 cm long, 3-15
mm wide, spatulate, thinly tomentose to
glabrous and green above, white-tomentose
beneath; heads mainly 3-5, in compact

cymes; pistillate involucres 6-11 mm high,
the bracts tomentose on the greenish base,
the apical scarious portion white or suffused
with pink; staminate plants rare; achenes
glabrous or minutely pubescent. Pinyon-juni-
per and shrub communities at 1525 to 1900
m in San Juan, Utah, and Washington coun-
ties; Alaska to Newfoundland, south to Cali-
fornia, Arizona, and Virginia; 3 (i). Our few
specimens are hardly adequate to represent
this species clearly in Utah.

Antennaria parvifolia Nutt. [A. aprica
Greene; A. obtusita Greene]. Plants caespi-
tose, stoloniferous and mat forming, 3-15 cm
tall; basal leaves 0.8-3.5 cm long, 3-8 mm
wide, spatulate, obtuse to acute apically, to-
mentose on both sides; heads 2-6 or more;
pistillate involucres 7-11 mm high, the bracts
more or less tomentose on the greenish base,
the scarious portion white, sordid, or pink;
staminate plants rarely collected; achenes
glabrous. Mountain brush, pinyon-juniper,
sagebrush, ponderosa pine, aspen, lodgepole,
and spruce-fir communities at 1650 to 3250
m in Beaver, Carbon, Daggett, Duchesne,
Emery, Garfield, Juab, Kane, Piute, San Juan,
Summit, Tooele, Uintah, Utah, and Wayne
counties; British Columbia to Manitoba,
south to Arizona and New Mexico; 46 (iv).

Antennaria pulcherrima (Hook.) Greene
Showy Pussy toes. [A. carpathica var. pulcher-
rima Hook.]. Plants from a caudex, not mat
forming or stoloniferous, 23-40 cm tall; basal
leaves 4-19 cm long, 5-23 mm wide, narrow-
ly to broadly oblanceolate to elliptic, tomen-
tose on both surfaces; heads several to many
in branching or compact cymes; pistillate
and staminate involucres both 6.5-8 mm
long, the bracts tomentose at the greenish
base, the terminal scarious portion blackish
to brownish or the apex whitish; achenes
glabrous. Sedge-rush meadows, streamsides,
and bogs at 2440 to 2800 m in Duchesne,
Garfield, and Summit counties; Alaska to
Newfoundland, south to Colorado; 5 (i).

Antennaria rosulata Rydb. Plants caespi-
tose, stoloniferous and mat forming, 1-3 cm
tall; basal leaves 0.5-1.1 cm long, 2-5 mm
broad, spatulate, obtuse to rounded apically,
tomentose on both surfaces; heads 1 or 2, ter-
minating short erect branches; pistillate in-
volucres 5-9 mm high, the outer bracts

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

201

greenish and tomentose to the apex, the inner
ones green at base, with scarious slender
white tips; staminate involucres 4-5 mm
high, the bracts densely tomentose at base,
the broad scarious tips white-opaque;
achenes puberulent. Ponderosa pine, aspen,
Douglas fir, limber pine, sagebrush, and
spruce communities, and in alpine meadows,
at 2600 to 3350 m in Garfield, Kane, and
Wayne counties; Colorado, New Mexico, and
Arizona; 14 (i).

Antennaria umbrinella Rydb. [A. dioica
authors, not (L.) Gaertn.]. Plants caespitose,
mat forming and stoloniferous, 2-14 mm tall;
basal leaves 0.7-2 cm long, 0.2-1.5 cm wide,
cuneate-oblanceolate to spatulate, acute to
obtuse apically, tomentose on both sides;
heads 2-6, borne in subcapitate cymes; pistil-
late involucres 5-8 mm long, the bracts
greenish and tomentose at the base, the
scarious tips dirty brownish to pale tan, or
the innermost almost white, acute to
rounded, usually erose; staminate plants un-
known in our region; achenes glabrous. As-
pen communities and alpine meadows at

2745 to 3500 m in Duchesne, Juab, and Sum-
mit counties; Alaska to Hudson Bay, south to
California, Arizona, and Colorado; 4 (0).
Specimens assigned here are more or less in-
termediate between A. alpina and A. micro-
phylla. Many more specimens are required to
provide definitive information on this entity
in Utah.

Anthemis L.

Annual or short-lived perennial, aromatic
herbs from taproots, the juice watery; stems
erect, commonly branched; leaves alternate,
1-3 pinnately dissected; heads solitary on the
uppermost branches; involucral bracts imbri-
cated in several series, chaffy, the margins
scarious or hyaline; receptacle hemispheric,
chaffy at least near the middle; ray flowers
present, white or yellow, usually 10 or more,
sterile; disk flowers numerous, perfect, fer-
tile; pappus none or a short crown; style
branches flattened; achenes subterete or com-
pressed, not callous-margined, glabrous,
beakless.

Rays white; pappus lacking; disk commonly less than 10 mm broad A. cotula

Rays yellow; pappus a short crown; disk commonly more than 12 mm broad
A. tinctoria

Anthemis cotula L. Mayweed. Plants an-
nual, 1-7.5 dm tall; stems simple or
branched, ill scented; leaves 1-6 cm long,
twice pinnatifid, the ultimate segments
lance-oblong, sparsely villous and glandular-
dotted; heads borne solitary at the upper
ends of the uppermost branches; ray flowers
commonly 10-20, white, sterile, 5-10 mm
long; disk flowers numerous; disk 4-10 (12)
mm wide; receptacle chaffy only in the
middle, the bracts narrowly subulate; achenes
slightly flattened, glandular, the pappus lack-
ing. Introduced Old World weeds of fields,
roadsides, revegetated woodlands, and other
disturbed sites at 1280 to 1400 m in Du-
chesne, Morgan, Salt Lake, Utah, and Weber
counties (likely elsewhere); widespread in
North America; 7 (0).

Anthemis tinctoria L. Yellow Camomile.
Plants short-lived perennials, 2.5-6 dm tall;
stems simple or branched; leaves 1.5-7 cm
long, 1- to 2-pirmatifid, the segments oblong

in outline, merely toothed or lobed, villous-
tomentose below, glabrous or glabrate above,
sparsely glandular-dotted; heads borne soli-
tary at ends of the uppermost branches; ray
flowers 20-35, yellow, fertile, 7-14 mm long;
disk flowers numerous; disk 12-15 mm wide
or more; receptacle chaffy throughout, the
bracts narrow and with yellow awn-tips;
achenes compressed; pappus a short crown.
Old World cultivated ornamentals; widely
planted and occasionally escaping (Salt Lake
County, Garrett 8865 BRY); widespread in
North America; 1 (0).

Arctium L.

Biennial, coarse herbs with watery juice,
from a taproot; leaves rhubarblike, basal and
alternate, entire or toothed; heads few to nu-
merous in axillary or terminal corymbose or
racemose inflorescences; flowers all tubular,
perfect, the corollas pink to purplish; in-

202 Great Basin Naturalist Vol. 43, No. 2

volucres urn shaped, the bracts imbricate in achenes sHghtly compressed, more or less 3-
many series, the tips slender and inwardly angled, many nerved, truncate apically; pap-
hooked; receptacle flat, densely bristly; pus of numerous, scaly, deciduous bristles.

1. Heads mainly 1.5-2.5 cm thick, arranged in racemelike axillary clusters, the

terminal also racemelike A. minus

— Heads commonly over 2.5 cm thick, arranged in corymbose clusters, especially

the terminal A. lappa

Arctium lappa L. Great Burdock. Plants Sevier, Summit, Tooele, Uintah, Utah,
8-15 dm tall; basal leaves long-petiolate, the Wasatch, Wayne, and Weber counties, and
blades commonly 2-5 dm long, 1-3 dm probably cosmopolitan; widespread in North
broad, cordate-ovate, obtuse, thinly tomen- America; Eurasia; 23 (i).
tose beneath, glabrous or nearly so above; in-
florescence corymbosely disposed, the pe- Arnica L
duncles glandular or glandular-hairy; heads

2.5-4 cm broad, the involucre greenish stra- Perennial herbs from rhizomes or caudices,
mineus, glabrous or glandular, often sparingly the juice watery; stems erect, simple or
arachnoid-tomentose. Cultivated for its branched above; leaves opposite or the up-
edible roots, and persisting; introduced from permost alternate, simple, entire or toothed;
Eurasia; 1 (0). heads solitary, or 3-9 (11) in corymbose clus-

Arctium minus (Hill) Bernh. Burdock, ters; involucral bracts subequal or evidently

Plants 5-15 dm tall; basal leaves long- biseriate, herbaceous; receptacle naked, con-

petiolate, the blades commonly 1-3.5 (4) dm vex; ray flowers present, yellow or orange,

long, 1-3 dm wide, cordate-ovate, obtuse, several to many, fertile, or lacking (in A. par-

thinly tomentose to glabrous beneath, ryi); disk flowers numerous, perfect, fertile;

glabrous above or nearly so; inflorescence pappus of barbellate or subplumose capillary

racemosely disposed, the peduncles short or bristles; style branches flattened; achenes cy-

lacking; heads 1-2.5 cm thick (rarely more), lindrical, 5- to 10-nerved, pubescent to gla-

the bracts glabrous or glandular to definitely brate or glabrous, often glandular,

arachnoid. Introduced Old World weed of Maguire, B. 1943. A monograph of the genus

consequence in Cache, Juab, Millard, Piute, Arnica. Brittonia 4:386-510.

1. Heads discoid (rarely some with rays), the lateral (lower) ones spreading or re-
flexed, the uppermost one erect A. parryi

— Heads radiate, the lateral ones (if any) erect like the uppermost 2

2(1). Cauline leaves (4) 5-9 pairs; pappus brownish; heads often 5 or more per main

stem 3

— Cauline leaves 1-4 (5) pairs; pappus white or brownish; heads mainly 1-4 per
stem 4

3(2). Involucral bracts merely acute to abruptly rounded (rarely acuminate), bearing

an apical or subapical tuft of hairs A. chamissonis

— Involucral bracts acuminate to attenuate, not especially more hairy at the
apex A. longifolia

4(2). Leaves (at least the lower) cordate, ovate, or broadly ovate-lanceolate, often

cordate, truncate, or obtuse basally, seldom cuneate 5

— Leaves narrowly lanceolate to lance-oblong or lanceolate, usually cuneate ba-
sally 7

5(4). Pappus brownish, subplumose; main cauline leaves obtuse to subcuneate ba-
sally A. diversifolia

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

203

— Pappus white, merely barbellate; main cauline leaves usually cordate, truncate,

or obtuse basally 6

6(5). Blades of main cauline leaves much longer than the petiole, or sessile; achenes

glabrous throughout, or at least near the base A. latifolia

— Blades of main cauline leaves subequal to or shorter than the petioles; achenes
uniformly, though sometimes sparingly, hairy A. cordifolia

7(4). Pappus brownish, subplumose A. mollis

— Pappus usually white or tawny, merely barbellate 8

8(7). Heads turbinate-campanulate, commonly with 7-10 rays; lower cauline leaves

sessile or nearly so A. rydbergii

— Heads hemispheric, commonly with 10-20 rays; lower cauline leaves often
petiolate 9

9(8). Old leaf bases bearing dense brown wool in the axils; disk corollas both spread-
ing hairy and stipitate-glandular A. fulgens

— Old leaf bases lacking axillary tufts of hair, or with white hair only; disk
corollas merely stipitate-glandular A. sororia

Arnica chamissonis Less. [A. foliosa Nutt.;
A. chamissonis ssp. foliosa (Nutt.) Maguire;
A. foliosa var. incana Gray]. Plants 1-6 (8)
dm tall, the stems erect or ascending, simple
or more commonly branched in the in-
florescence, sparsely to densely villous with
multicellular hairs and often glandular as
well; basal leaves 3-11 (15) cm long, 3-16
(20) mm wide, lanceolate to oblong or oblan-
ceolate, with 3-5 main veins, pilose to villous
or tomentose, tapering to a slender petiole,
entire to distinctly toothed, smaller than the
cauline ones and often withered by flowering
time; cauline leaves (4) 5-8 (9) pairs, lanceo-
late to lance-elliptic, the largest near the
middle of stem or slightly below, the lower
ones petiolate and with membranous con-
nate-sheathing bases, the upper sessile, entire
to distinctly toothed; heads (1) 3-9, the pe-
duncle apex sparingly to densely villous with
whitish hairs often intermixed with glands;
involucres 9-15 mm high, the bracts lanceo-
late, obtuse, acute, or" less commonly acumi-
nate, sparsely to densely pilose, ciliate, the
tips with a conspicuous tuft of whitish hairs;
rays usually 10-16, yellow; achenes 4-6 mm
long, hairy to glandular or glabrate; pappus
brownish to straw colored, barbellate. Stream
banks, gravel bars, and lake shores in aspen,
willow, and spruce-fir communities at 2300
to 3350 m in Duchesne, Emery, Garfield,
Iron, Kane, Salt Lake, Sanpete, Sevier, Sum-
mit, Tooele, Wasatch, Washington, and
Wayne counties; Alaska to Hudson Bay,

south to California and New Mexico; 29 (v).
Maguire (1943) treated all Utah material as
ssp. foliosa (Nutt.) Maguire. Cronquist (Univ.
Washington Publ. Biol. 17(5): 45-54. 1955)
cited var. incana (Gray) Hulten [ssp. incana
(Gray) Maguire] from Utah. Our specimens
are only arbitrarily separable into two
phases, differing mainly in degree of pu-
bescence. The correct name at varietal level
for our gray hairy plants appears to be A.
chamissonis var. andina (Nutt.) Ediger &
Barkley.

Arnica cordifolia Hook. Plants 1.5-4 dm
tall, the stems erect or ascending, simple or
branched above, sparsely villous with multi-
cellular hairs and often glandular as well;
basal leaves smaller than the cauline, often
withered at anthesis; petioles of main leaves
(at least) often longer than the blades; cauline
leaves 2-4 (5) pairs, the blades 2-9 cm long
(from sinus to apex), 1-9 cm wide, cordate-
ovate to orbicular or reniform, or the up-
permost lanceolate, the largest below the
middle of the stem, the lower leaves petio-
late, the upper ones sessile or subsessile, ser-
rate-dentate to subentire; heads 1 (3), rarely
more, the peduncle apex villous with whitish
hairs often intermixed with glands; involucres
14-20 mm high, the bracts lanceolate to ob-
long, acuminate to acute, sparsely to densely
pilose and often glandular-ciliate, the tip
with a moderate tuft of hair; rays usually
10-15, yellow; achenes 4-5.5 mm long, uni-
formly hairy and often glandular; pappus

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Great Basin Naturalist

Vol. 43, No. 2

white, barbellate. Sagebrush, Douglas fir,
white fir, lodgepole pine, ponderosa pine, as-
pen, and spruce-fir communities at 1525 to
3355 m in Beaver, Box Elder, Cache, Carbon,
Daggett, Duchesne, Garfield, Iron, Juab,
Piute, Salt Lake, San Juan, Sanpete, Sevier,
Summit, Uintah, Utah, Wasatch, Washington,
Wayne, and Weber counties (likely univer-
sal); Alaska to Michigan, south to California,
Arizona, New Mexico, and Nebraska; 102
(xii). The white pappus and cordate long-
petiolate leaves are diagnostic for this
species.

Arnica diversifolia Greene Plants 1.5-4.2
dm tall, the stems erect or ascending, simple
or branched above, sparsely villous with mul-
ticellular hairs and often glandular, or almost
glabrous; basal leaves smaller than the cau-
line and often withered by flowering time,
borne on slender to broadly winged petioles
shorter than or subequal to the blades; cau-
line leaves 2-4 (5) pairs, blades 2-8 cm long,
0.8-4 (6) cm wide, ovate or the uppermost
lanceolate, the largest at the middle or be-
low, becoming sessile to subsessile above,
subentire or irregularly serrate; heads 1-3 or
more, the peduncle apex sparsely to moder-
ately villous with whitish hairs and often
with glands; involucre 10-16 mm high;
bracts lanceolate, acuminate to acute, sparse-
ly to densely pilose and often glandular, cil-
iate, the tip lacking a tuft of hairs; rays usual-
ly 10-15, yellow; achenes 5-7 mm long,
glabrous or sparsely and uniformly hairy;
pappus brownish, subplumose. Stream sides,
meadows, and scree slopes in spruce-fir and
alpine tundra communities at 2560 to 3400 m
in Duchesne, Grand, Piute, San Juan, San-
pete, Summit, Uintah, and Utah counties;
Alaska and Yukon, south to Oregon and Colo-
rado; 19 (ii). This taxon is not well collected
in Utah. The broad leaves and brownish sub-
plumose pappus are diagnostic for these
plants that might be regarded as consisting of
a series of hybrid derivatives between A. mol-
lis and A. cordifolia, A. latifolia, or A. ryd-
bergii. More work is necessary.

Arnica fulgens Pursh [A. pedunculata
Rydb.]. Plants 1.5-6 (7) dm tall, the stems
erect, the basal leaf axils with tufts of long
brown woolly hair, otherwise stipitate-
glandular and often hairy as well; basal

leaves smaller than the cauline, often with-
ered at anthesis, with broadly winged pet-
ioles or subsessile; cauline leaves 2-4 pair,
the blades oblanceolate to elliptic (often nar-
rowly so), mostly 3-12 cm long, 0.6-4 cm
wide, the largest ones near the base, becom-
ing sessile upward, subentire to entire; heads
1-3, the peduncle apex yellowish villous; in-
volucre 10-15 (18) mm high, the bracts nar-
rowly elliptic to lance-elliptic, attenuate to
an obtuse or acute apex, villous, the tips pu-
bescent within; rays mostly 10-20, yellow to
yellow-orange; achenes 4-5.5 mm long,
densely hairy; pappus whitish to cream col-
ored, barbellate. Dry sagebrush community
at 2000 m in Daggett County; British Colum-
bia to Saskatchewan, south to California, Ne-
vada, and Colorado; 1 (0).

Arnica latifolia Bong. [A. gracilis Rydb.;
A. jonesii Rydb.]. Plants 1-4 (6) dm tall, the
stems erect or ascending, simple or branched
above, sparsely villous with multicellular
hairs and often glandular; basal leaves
smaller than the cauline, usually withered by
flowering time, the petioles (if any) usually
shorter than the blades; cauline leaves 2-5
pairs, the blades 2-4.5 (7) cm broad, cordate-
ovate to lanceolate, the largest ones at the
middle or below, the lower ones with petioles
shorter than the blades, the upper ones sessile
or subsessile, serrate-dentate, less commonly
entire or nearly so; heads 1-5 or rarely more,
the peduncle apex sparsely to moderately vil-
lous with whitish or yellowish hairs and often
glandular; involucres 9-17 mm high, the
bracts lanceolate, acuminate to acute, sparse-
ly pilose and often glandular, ciliate, lacking
an apical tuft of hair; rays usually 8-12, yel-
low; achenes 5-8 mm long, glabrous or
sparsely hairy, or glabrous in the lower por
tion; pappus white, barbellate. Lodgepole
pine, spruce-fir, and alpine tundra commu-
nities at 2240 to 3400 m in Duchesne, Salt
Lake, Summit, and Utah counties; Alaska and
Yukon to California and Colorado; 24 (v).
Specimens available for study are variable.
TTiey occur in the Uinta and Wasatch moun-
tains and on the Tavaputs Plateau. The var.
gracilis (Rydb.) Cronq. was reported from
Utah by Maguire (I.e., as A. gracilis Rydb.),
but has not been seen by me. It differs from
the bulk of our material in its small size (1-3
dm), more numerous heads (3-9), and narrow

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

205

small involucre 9-13 mm high. The single
collection cited by Maguire is from Salt Lake
or Utah County. More material is necessary.

Arnica longifolia D.C. Eaton in Wats. [A.
caudata Rydb., type from Big Cottonwood
Canyon]. Plants 3-10.5 dm tall; stems erect
or sprawling, tufted from caudexlike short-
ened rhizomes, simple or branched above,
shortly villous to puberulent and often some-
what glandular-viscid; basal leaves lacking or
soon withering, the cauline ones 5.5-20 cm
long, 0.6-3 cm wide, lanceolate to elliptic,
with 3-5 main veins, puberulent, all sessile,
5-7 pairs, the largest near the middle of the
stem, the lower ones connate-sheathing, en-
tire or nearly so; heads 1-9, the peduncle
apex sparingly yellowish villous; involucres
6-13 mm high, the bracts lanceolate to lance-
oblong, acute to aciiminate, sparingly pilose
and glandular, ciliate, the tips sparingly
white-hairy; rays mainly 8-13, yellow;
achenes 4.5-5.5 mm long, glabrate, or uni-
formly stipitate-glandular; pappus brownish
to straw colored, barbellate. Snow flushes,
talus, and stream banks in lodgepole pine, as-
pen, ponderosa pine, Douglas fir, white fir,
and spruce-fir communities at 1890 to 3325
m in Box Elder, Cache, Garfield, Grand,
Juab, Rich, Salt Lake, Sanpete, Summit,
Tooele, and Washington counties; Washing-
ton to Alberta, south to California, Nevada,
and Colorado; 15 (i). Our material belongs to
var. longifolia.

Arnica mollis Hook. [A. arachnoidea
Rydb., type form Big Cottonwood Canyon;
A. chamissonis var. longinodosa A. Nels.,
type from near Marysvale; A. ovata Greene,
type from Alta]. Plants 1.5-6.5 dm tall, the
stems erect or ascending, loosely to com-
pactly clump-forming, simple, or branched in
inflorescence, puberulent to villous and
glandular; basal leaves smaller than the cau-
line ones, often withered at anthesis, the cau-
line ones 4.5-18 cm long, 0.8-4 cm wide, ob-
lanceolate to obovate, lanceolate or elliptic,
the lower slenderly to broadly petiolate, be-
coming sessile upwards, 3-4 pairs, the largest
below the middle, the lower connate-sheath-
ing, entire to irregularly denticulate; heads
1-5 (7), the peduncle apex sparingly yellow-
ish villous; involucres 10-17 mm high, the
bracts lanceolate to lance-elliptic, acute to
attenuate, sparingly to densely villous-pilose
and more or less glandular, lacking a sub-

apical tuft of hair; rays mainly 12-18, yellow;
pappus brownish, subplumose; achenes pu-
bescent to stipitate-glandular. Meadows,
bogs, stream banks, seeps, talus slopes, and
rock stripes in sagebrush, ponderosa pine,
lodgepole pine, Douglas fir, white fir, aspen,
spruce-fir, and alpine tundra communities at
1950 to 3550 m in Box Elder, Cache, Dag-
gett, Duchesne, Garfield, Juab, Kane, Piute,
Rich, Salt Lake, Sanpete, Summit, Tooele,
Uintah, and Utah counties; British Columbia
to California, Nevada, and Colorado; 50 (xii).

Arnica parryi Gray. Plants 1.5-5 (6) dm
tall, erect or ascending, from elongate rhi-
zomes, simple or branched in inflorescence,
villous and more or less glandular; basal
leaves smaller than the cauline ones, often
withered at anthesis, the cauline ones long
petioled below, becoming sessile upwards,
the blades 2-9.5 cm long, 0.4-4 cm wide, lan-
ceolate to ovate, the base obtuse to truncate
or cuneate, 2-4 (5) pairs, the largest near the
stem base, the lower connate-sheathing, en-
tire to denticulate; heads 3-12, nodding in
bud, the peduncle apex glandular-villous; in-
volucres 10-16 mm high, the bracts narrowly
lanceolate, acute to attenuate, glandular-vil-
lous, lacking a subapical tuft of hairs; rays
lacking, or rarely present, yellow; pappus
brownish, barbellate to subplumose; achenes
glabrous to glandular or hairy. Aspen and
spruce-fir communities at 2415 to 3175 m in
Carbon, Daggett (?), Garfield, Iron, Salt
Lake, Sanpete, and Summit counties; British
Columbia and Alberta to California and Col-
orado; 9 (ii). A specimen from Piute County
(Welsh et al. 14018 BRY) has ray flowers well
developed.

Arnica rydbergii Greene. Plants 1-2.6 dm
tall, erect or ascending, from elongate rhi-
zomes, sparingly villous and shortly stipitate-
glandular; basal leaves smaller than the cau-
line, sometimes bladeless, often withered at
anthesis, the cauline ones short to long pet-
ioled below, becoming sessile upwards, the
blades 2-5 cm long, 0.4-1.5 cm wide, lan-
ceolate to elliptic, ovate, or obovate, the base
obtuse to cuneate, 2 or 3 (4) pair, the largest
at or near the middle of the stem, the lower
connate-sheathing, entire or denticulate;
heads 1-5, the peduncle apex yellowish vil-
lous, glandular; involucres 9-13 mm high, the

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Great Basin Naturalist

Vol. 43, No. 2

bracts narrowly lanceolate, acute to attenu-
ate, stipitate-glandular, ciliate, lacking a sub-
apical tuft of hair; rays mainly 7-10, yellow;
pappus white, barbellate; achenes shortly pi-
lose. Spruce-fir and lodgepole pine forests in
Duchesne, Summit, and Utah counties; Brit-
ish Columbia and Alberta to Oregon and Col-
orado; 12 (iii).

Arnica sororia Greene Plants 1.5-6 dm
tall, the stems erect, the basal axils lacking
tufts of hair, otherwise more or less villous
and glandular; basal leaves smaller than the
cauline, often withered at anthesis, with
winged to narrow petioles or subsessile; cau-
line leaves 2-4 pair, the blades lanceolate to
elliptic, mostly 3-10 cm long, 0.5-2 cm wide,
the largest ones near the base, becoming ses-
sile upward, mainly entire; heads 1-3, the pe-
duncle apex sparingly villous; involucres
10-15 mm high, the bracts narrowly oblong-
lanceolate, attenuate, villous, the tips more
or less hairy within; rays mainly 9-15, yel-
low; achenes 4-6 mm long, densely short-
hairy; pappus white, barbellate. Meadows
and foothills in sagebrush and aspen commu-
nities at 1675 to 2100 m in Cache and Rich
counties; Alberta and British Columbia to
Wyoming, Nevada, and California; 2 (0).

Artemisia L.

Annual, biennial, or perennial herbs, sub-
shrubs, or shrubs from taproots, caudices, or
rhizomes, the juice watery; stems decumbent
to ascending or erect, simple or branched;
leaves alternate or basal, entire or toothed,
lobed, or divided; heads several to numerous,
borne in spicate, racemose, or paniculate
clusters; involucral bracts imbricate in sever-
al series, dry, at least the inner with scarious
margins; receptacle naked or beset with long
hairs, often glandular; corollas of disk flowers
only (rarely with minute bilabiate ray flowers
in A. bigelovii), perfect, or sometimes the
central ones sterile, the marginal merely pis-
tillate; marginal corollas tubular (or bila-
biate), the central ones tubular- funnelform;
pappus lacking, or a short crown; style
branches flattened; achenes subterete or an-
gular, glabrous.

Beetle, A. A. 1960. A study of sagebrush —
The section Tridentate of Artemisia.

Univ. Wyoming Agr. Expt. Sta. Bull.

368. 83 pp.
Keck, D. D. 1946. A revision of the Artemisia

vulgaris complex in North America.

Proc. Calif. Acad. 25:421-468.

1. Plants shrubs or subshrubs 2

— Plants herbs 13

2(1). Heads with both ray and disk flowers, the ray flowers 2-lipped; branchlets of
inflorescence spreading to reflexed; plants of rimrock areas in Colorado drain-
age, rarely in southern Great Basin A. bigelovii

— Heads discoid; branchlets of inflorescence variously disposed; plants seldom of
rimrock, the distribution various 3

3(2). Leaves 1 to 3-pinnately or ternately dissected, the segments linear 4

— Leaves entire or toothed, or if lobed, the lobes oblong or broader, or if linear

(see A. filifolia), tall shrubs of sandy areas at low elevations 6

4(3). Plants silvery-canescent; receptacle hairy; growing commonly on windswept

ridges, but not always so restricted A. frigida

— Plants green to gray-green; receptacle glabrous, or, if hairy, plants of low ele-
vations 5

5(4). Shrubs with spreading branches, spinescent, flowering in springtime .... A. spinescens

— Shrubs with erect or ascending branches, not spinescent, flowering in late sum-
mer and autumn A. pygmaea

6(3). Leaves linear-filiform, less than 1 mm wide, entire, or 3-parted; tall plants of

sandy low elevation sites A. filifolia

— Leaves broader, entire, or the segments broader than 1 mm wide; plants of var-
ious habitats and elevations 7

April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 207

7(6). Leaves entire or with 1 or 2 teeth; heads borne in slender panicles; plants of high

elevations A. cana

— Leaves toothed or lobed at the apex; heads borne in slender spicate to broad
panicles 8

8(7). Plants usually less than 3 dm tall; leaves usually less than 1 cm long; foliage

dull yellow- to lead-gray or rarely silvery A. nova

— Plants mainly more than 3 dm tall; leaves usually more than 1 cm long (at least
some); foliage silvery-canescent 9

9(8). Leaves all, or many of them, deeply cleft into narrowly oblong lobes which

may be further divided; flowers commonly 5-8 per head A. tripartita

— Leaves mainly merely toothed apically; flowers various 10

10(9). Leaves coarsely and deeply 3-lobed, the lobes broad and rounded apically; in-
florescence narrow, seldom over 1.5 cm wide; plants of Rich and Summit
counties A. longiloba

— Leaves variously 3- to 5-toothed, seldom lobed; inflorescence various; plants
variously distributed 11

11(10). Inflorescence open, paniculate, commonly more than 2 cm wide; plants of

broad distribution, our common sagebrush species A. tridentata

— Inflorescence narrow, spicate, commonly less than 1.5 cm wide; plants less
broadly distributed 12

12(11). Plants often less than 4 dm tall; leaves mainly less than 1.5 cm long; heads small;

plants usually of middle elevations A. arbuscula

— Plants often over 4 dm tall; leaves mostly over 2 cm long; heads large; plants

of high elevations A. spiciformis

13(1). Leaves all entire, or the lower ones toothed or lobed, glabrous and green above
and beneath, or white-hairy on both surfaces (see also A. carruthii and A. mich-
auxiana), usually much longer than broad 14

— Leaves deeply incised, pinnatifid, or temately divided, variously pubescent,
various in length-width proportions 15

14(13 ) Leaves green above and beneath; central flowers of heads with normal ovaries

A. dracunculus

— Leaves white-hairy above and beneath or green above; central flowers of head
with abortive ovaries A. ludoviciana

15(14). Plants annual or biennial from a taproot; leaves green, essentially glabrous; ad-

ventive 16

— Plants perennial from a rhizome or caudex; leaves tomentose, strigose, or pilose ... 17

16(15). Inflorescence paniculate, loose and open; heads borne on short peduncles; in-
volucres 1-2 mm high A. annua

— Inflorescence a spicate panicle, the branches appressed-ascending; heads ses-
sile or nearly so; involucres more than 2 mm high A. biennis

17(15). Cauline leaves reduced upwards, the largest leaves in a basal rosette, silvery-
villous to strigulose, scarcely tomentose and uniformly colored above and be-
neath; plants from caudices, only occasionally rhizomatous 18

— Cauline leaves not especially reduced upwards, seldom with a basal rosette,
variously tomentose and often bicolored; plants often rhizomatous (except in

A. absinthium) 20

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18(17). Pubescence of leaves loosely villous to glabrous; corollas hairy, the receptacle

glabrous; plants of high elevations in the Uinta and La Sal mountains 19

— Pubescence of leaves appressed strigose or villosulose; corollas glabrous or
hairy, but, if hairy, the receptacle long- villous; plants variously distributed 20

19(18). Involucres 3-4 mm high; plants of the La Sal Mountains A. parryi

— Involucres 4-5.3 mm high; plants of the Uinta Mountains A. norvegica

20(19). Inflorescence a spicate raceme; receptacle and corolla long-villous; plants of

high elevations A. scopulorum

— Inflorescence a slender panicle; receptacle and corollas glabrous; plants of low
elevations, seldom of high elevations A. campestris

21(17). Receptacle beset with numerous long hairs between the flowers; leaves about

equally hairy above as below; plants introduced, weedy, of low elevations

A. absinthium

— Receptacle naked; leaves more or less tomentose below, usually green or
greenish above, or equally tomentose on both sides; plants indigenous, not
weedy, of mid-to-high elevations 22

22(21). Leaves entire or with entire lobes; plants of moderate elevations in central and

southern Utah A. carruthii

— Leaves bipinnatifid, the lobes again toothed; plants of high elevations in the
Uinta, Wasatch, and La Sal mountains A. michauxiana

Artemisia absinthium L. Absinthe. Pe-
rennial fragrant herbs from a rhizomatous
caudex, 5-10 (12) dm tall, appressed sericeus;
leaves bi- or tripinnatifid, the main lobes
again lobed or toothed, silvery-sericeus on
both surfaces, with very short tangled hairs,
hardly tomentose, 1.5-5.5 cm long on flower-
ing stems (2-10 cm long on sterile stems) the
main ultimate segments mostly 2-4 mm
wide, petiolate below, shortly petiolate and
less commonly divided above; involucres 2-3
mm high, the bracts scarious over the green-
ish center, the margins brownish hyaline;
flowers all fertile, the marginal ones pistil-
late; receptacles with numerous long slender
hairs; achenes glabrous. Roadsides, stream-
banks, and abandoned fields in Garfield,
Rich, and Utah counties; widely established
in North America; adventive from Europe; 6
(ii).

Artemisia annua L. Sweet Wormwood.
Annual fragrant herbs, mainly 0.3-1.5 (3) m
tall; stems sparingly glandular; leaves 2- or 3-
pinnatifid, the main lobes again lobed, green
and minutely glandular on both surfaces,
1.5-8 (10) cm long, the main ultimate seg-
ments 0.5-3 mm wide, petiolate below, sub-
sessile or shortly petiolate above; involucres
1.3-2 mm high; involucral bracts with green

centers minutely glandular, the margins hya-
line; receptacles naked; achenes glabrous. In-
troduced weedy species of disturbed sites in
Washington County; adventive from Eurasia,
now widely naturalized in North America; 1 (0).
Artemisia arbuscula Nutt. Low Sagebrush.
[A. tridentata ssp. arbuscula (Nutt.) H. & C.;
A. tridentata var. arbuscula (Nutt.) McMinn].
Shrubs, commonly 2-4 (5) dm tall, the vege-
tative stems 1.5-10 cm long, the flowering
stems erect, 8-30 cm long; leaves 0.4-1.6 cm
long, shallowly 3- to 5-dentate to deeply
lobed, cuneate basally, appressed canescent;
inflorescence spicate, mostly less than 2 cm
wide; involucres 4-6 mm long, campanulate;
involucral bracts 4-8, canescent, the margins
brownish-scarious; flowers 4-9, all perfect;
receptacle naked; achenes glabrous. Pinyon-
juniper, mountain brush, sagebrush, white fir,
aspen, and spruce-fir communities at 1375 to
2550 m in Box Elder, Cache, Millard, Rich,
Salt Lake, Summit, and Tooele counties;
Washington to Montana, south to California
and Nevada; 14 (0). A. arbuscula, or low
sagebrush, has been confused with both A.
tridentata and A. nova. It can be distin-
guished from the former by its narrow in-
florescence, and from the latter by its canes-
cent involucres. Beetle (Ic.) reports

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

209

intermediates with A. longiloha, a taxon with
broadly campanulate heads and bluntly lobed
leaves.

Artemisia biennis Willd. Biennial Worm-
wood. Plants annual or biennial, with tap-
roots, the stems 0.3-9 (10) dm tall or more,
glabrous; basal leaves often withered by an-
thesis; cauline leaves well developed, 1.5-10
(15) cm long, once pinnately divided, the seg-
ments oblong to oblanceolate, again toothed,
essentially glabrous, green; inflorescence spi-
cate or in spicate panicles; heads numerous,
crowded, sessile or subsessile, erect or nearly
so; involucres 2-3 mm high, 2-4 mm broad,
the bracts glabrous, greenish to yellowish, the
margins hyaline; marginal flowers perfect,
fertile, the corollas glabrous; receptacle and
achenes glabrous. Floodplains, lake beds and
shores, mud flats, and pond margins at 1375
to 2900 m in Cache, Duchesne, Garfield,
Grand, Iron, Millard, Salt Lake, Sanpete,
Tooele, Uintah, and Utah counties; wide-
spread in North America, where presumably
indigenous in the western portion; Europe;
20 (v).

Artemisia bigelovii Gray Bigelow Sage-
brush. Shrubs, commonly 2-7 (10) dm tall or
more, the vegetative stems 1-3 dm long, the
flowering stems erect, 3-4.5 dm tall; leaves
0.3-2.3 cm long, 1-7 mm wide, entire or
shallowly 3-toothed, basally cuneate, ap-
pressed to loosely canescent-tomentose; in-
florescence narrowly paniculate, mostly less
than 4 cm wide, the branches often lax and
with heads tending to be pendulous; in-
volucres mainly 2.5-3.5 mm high, sub-
cylindric to narrowly campanulate, the
bracts 5-10, silvery canescent, with narrow
scarious margins; flowers 3 or 4, imperfect or
some perfect, the marginal pistillate (ray)
flowers bilaterally symmetrical; receptacle
naked; achenes glabrous. Rimrock areas in
pinyon-juniper and mixed desert shrub com-
munities at 975 to 2135 m in Duchesne,
Emery, Garfield, Grand, Kane, Millard, San
Juan, Sevier, Uintah, and Wayne counties;
California and Nevada east to Colorado, New
Mexico, and Texas; 45 (xvii).

Artemisia campestris L. Field Wormwood.
Perennial herbs from a caudex and taproot,
the stems (1.5) 2.5-7 dm tall (rarely taller),
tomentose or glabrous; basal leaves well-de-
veloped (often withered at anthesis), 2-12 cm

long, 2- to 3-pinnatifid or ternate, the seg-
ments linear to narrowly oblong or spatulate,
villous or pilose to glabrous on both sides;
cauline leaves reduced upwards, once pin-
natifid, ternate, or entire; inflorescence of
narrow to lax panicles; heads numerous,
shortly pedunculate on contracted to lax
branchlets, finally pendulous; involucres
2.5-3.8 mm high, 2-2.3 mm wide, the bracts
glabrous, greenish to yellowish, the margin
hyaline; marginal flowers pistillate, fertile,
the corollas glabrous; disk flowers sterile, the
ovaries abortive; receptacle and achenes
glabrous. Saltbush, greasewood, sagebrush,
mountain brush, and pinyon-juniper commu-
nities, mainly in dunes and other sandy sites
at 1250 to 2075 m in Emery, Garfield,
Grand, Kane, San Juan, Sevier, Washington,
and Wayne counties; Arizona, New Mexico,
Colorado, Wyoming, and west to the Pacific;
24 (vii). Our material is assignable to ssp. ho-
realis (Pallas) H. & C, in a broad sense, and
belongs to var. scouleriana (Benth.) Cronq.
[A. pacifica Nutt.; A. campestris ssp. pacifica
(Nutt.) H. & C; A. forwoodii authors, not
Wats.; A. caudata authors, not Michx.] in a
more narrow sense.

Artemisia cana Pursh Silver Sagebrush.
Shrubs, commonly 2.5-12 (15) dm tall, the
vegetative branches 1-3 (5) dm long; flower-
ing stems erect, 1-3 dm tall; leaves 0.8-5.3
(7) cm long, linear to narrowly elliptic or ob-
long, entire, or some of them toothed or
deeply lobed, usually acute basally, acute to
obtuse apically, appressed tomentose; in-
florescence narrowly spicate or glomerate-
paniculate, mostly less than 5 cm wide, often
conspicuously bracteate, the branches, when
present, erect, the heads erect; involucres
3.3-6.1 mm high, 3.5-6 mm wide, cam-
panulate; bracts numerous, the outer silvery-
canescent, with greenish median, the margins
brownish scarious, rounded-erose; flowers
10-20, perfect; receptacle naked; achenes
glabrous. Meadows and stream terraces, less
commonly on moist slopes away from mead-
ows and streams at 2270 to 3050 m in Cache,
Carbon, Daggett, Duchesne, Emery, Gar-
field, Iron, Juab, Kane, Piute, Rich, Sanpete,
Sevier, Summit, Utah, Wasatch, and Wash-
ington counties; British Columbia to Sas-
katchewan, south to California, Nevada, and
New Mexico. Our materials are assigned to

210

Great Basin Naturalist

Vol. 43, No. 2

var. viscidula Osterh. [A. cana ssp. viscidula
(Osterh.) Beetle], which differs from typical
var. cana in its smaller, narrower leaves and
less canescent herbage. Silver sagebrush
forms intermediates with both A. tridentata
var. vaseyana and A. spiciformis, within
whose altitudinal range it occurs, but whose
habitats are ordinarily separate; 42 (viii).

Artemisia carruthii Wood ex Carruth Car-
ruth Wormwood. [A. wrightii Gray; A. vul-
garis ssp. wrightii (Gray) H. & C.]. Plants pe-
rennial herbs, with well-developed rhizomes,
the stems 2-7 dm tall, sparingly to densely
tomentose; basal leaves not well developed;
cauline leaves various but usually pinnatifid
with linear lobes, those of innovations and
sometimes the primary ones at base of flow-
ering stems entire or merely lobed, 0.6-3 cm
long, the lobes 0.5-1.5 (2) mm wide, linear or
narrowly oblong, tomentose on both sides, or
less so above; inflorescence paniculate (nar-
rowly so) or spicate; heads numerous, shortly
pedunculate to sessile, erect; involucres 2.3-3
mm high, 2-2.5 mm wide, the bracts spar-
ingly tomentose, pale greenish with hyaline
margins; marginal flowers pistillate, fertile;
central flowers perfect, fertile, the corollas
glabrous; receptacle and achenes glabrous.
Canyon bottoms, slopes, and rock outcrops in
sagebrush, mountain brush, aspen, and
spruce- fir communities at 1890 to 3050 m in
Emery, Iron, Piute, San Juan, Utah, and
Washington counties, and likely elsewhere;
east to Kansas and south to Arizona, New
Mexico, and Texas. This taxon is allied to A.
ludoviciana, and some specimens appear to
be intermediate between them. There is justi-
fication for inclusion of A. carruthii within an
expanded A. ludoviciana, but no formal pro-
posal is intended or implied herein. The
deeply pinnatisect main foliage leaves are
thought to be diagnostic. The species has not
been collected in sufficient numbers as to un-
derstand its distribution in any definitive
manner; 16 (ii).

Artemisia dracunculus L. Terragon. [A.
glauca Pallas; A. dracunculus ssp. glauca
(Pallas) H. & C; A. aromatica A. Nels.; A.
dracunculoides Pursh]. Plants shortly rhizo-
matous, perermial herbs, the stems (2) 5-12
(15) dm tall, glabrous (rarely tomentose?);
leaves primarily cauline, entire or rarely a

few of them cleft, 1.2-7.5 cm long, 1-6 mm
wide, glabrous, green on both surfaces; in-
florescence paniculate; heads numerous,
short-pedunculate to subsessile, more or less
pendulous; involucres 2-2.8 mm high, 2.2-3
mm wide, the bracts glabrous, greenish, with
broad hyaline margins; marginal flowers pis-
tillate, fertile; central flowers sterile, the
ovaries abortive, the corolla glabrous (often
glandular); receptacle and achenes glabrous.
Rabbitbrush, sagebrush, skunkbush, wildrye,
salt desert shrub, pinyon-juniper, ponderosa
pine, aspen, spruce-fir, and hanging garden
communities at 1220 to 3200 m in Duchesne,
Emery, Garfield, Grand, Iron, Juab, Kane,
Millard, Piute, San Juan, Sanpete, Sevier,
Tooele, Uintah, Utah, Wasatch, Washington,
and Wayne counties; Yukon southeast to Il-
linois and south to Mexico. Our material fits
within the concept of var. glauca (Pallas)
Bess, in Hook., which is probably not sepa-
rable from var. dracunculus of the Old
World; 63 (xvii).

Artemisia filifolia Torr. Sand Sagebrush;
Old-man Sagebrush. Shrubs commonly 5-15
dm tall, the vegetative branches 1-3 dm
long; flowering branches erect, 1.5-6 dm
long; leaves 0.6-8 cm long, 0.3-1.5 mm wide,
revolute (appearing terete) or somewhat flat-
tened (and still revolute), entire or the lower
ternate, appressed villous-tomentose; in-
florescence paniculate, mostly more than 3
cm wide, conspicuously bracteate, the
branches erect, the heads pendulous; in-
volucres 1.6-2.2 mm long 1.5-2.2 mm wide,
campanulate to subglobose; bracts 5-9,
densely silvery canescent; flowers 3-9, the
marginal ones pistillate, fertile, the central
ones sterile; receptacle naked; achenes
glabrous. Sandy sites in blackbrush, creosote
bush, ephedra, Poliomintha, Eriogonum, rab-
bitbrush, and pinyon-juniper communities at
825 to 2290 m in Emery, Garfield, Grand,
Iron, Kane, San Juan, Washington, and
Wayne counties; Colorado and South Da-
kota, south to Arizona, Texas, and Mexico; 43
(vi).

Artemisia frigida Willd. Fringed Sage-
brush; Prairie Sagewort. Shrubs 0.5-4.5 dm
tall, white-tomentose to strigulose; flowering
stems arising from short prostrate or ascend-
ing woody offsets; leaves of basal offsets

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

211

much like the stem leaves, 0.5-1.5 (2.5) cm
long, 2- to 3-temately (or subpinnately) di-
vided into linear segments mainly 0.3-0.8
mm wide, often with stipulelike divisions
near the base, whitish pilose-tomentose
throughout (fading brownish); inflorescence
paniculate or less commonly borne sessile or
on very short pedimcles; involucres 2-3.5
mm high, 4-6 mm broad, the bracts pilose-to-
mentose, with brownish scarious margins;
marginal flowers pistillate, fertile; central
flowers perfect, fertile, the corolla glabrous
(often glandular), yellow or tinged reddish;
receptacle long-hairy; achenes glabrous.
Shadscale, sagebrush, pinyon-juniper, ponde-
rosa pine, mountain brush, aspen, spruce, and
alpine (often on windswept ridge crests) com-
munities at 900 to 3480 m in Box Elder, Car-
bon, Duchesne, Emery, Garfield, Grand,
Juab, Kane, Millard, Piute, San Juan, San-
pete, Sevier, Summit, Uintah, Utah, Wasatch,
and Wayne counties (likely elsewhere);
Alaska to Quebec, south to Arizona and Kan-
sas; Asia; 78 (x).

Artemisia longiloba (Osterh.) Beetle
Longleaf Sagebrush. [A. spiciformis (?) long-
iloba Osterh.]. Shrubs, mainly 2-5 dm tall,
appressed villous-tomentose; flowering stems
1-2 dm long; leaves 0.4-2 cm long, broadly
cuneate, deeply 3-lobed, the lobes obtuse, ap-
pressed villous-tomentose; inflorescence spi-
cate, the heads several, shortly pedunculate
to sessile, erect; involucres 4-6 mm high, 3-5
mm wide, the 4-12 bracts villous-tomentose,
green, with brownish scarious margins; mar-
ginal flowers perfect, fertile, central flowers

perfect, fertile, the corolla glabrous (glandu-
lar), cream colored; receptacle and achenes
glabrous. Sagebrush and grass communities at
1675 to 2440 m in Rich and Summit counties;
Oregon to Montana, south to Nevada and
Colorado. This entity is reported to grow in
tight to heavy soils (Beetle 1960) and matures
seed in July and August. The plants have
large heads similar to those of A. cana and
the low habit of A. nova. Possibly they would
best be treated within an expanded A. triden-
tata, but no combination is proposed herein;
4 (0).

Artemisia ludoviciana Nutt. Perennial
rhizomatous herbs, the stems 2-10 dm tall (or
white-tomentose or glabrate to

glabrous; leaves mainly cauline, entire, lobed,
or pinnately incised, white-tomentose below,
green and glabrous or tomentose above
(rarely glabrous throughout), 0.8-9 cm long,
0.1-1 (2) cm wide; inflorescence spicate to
paniculate; heads numerous, shortly pe-
dunculate to sessile, more or less pendulous;
involucres 2.5-4.5 mm high, 3-7 mm wide
(or more), the bracts tomentose to glabrous,
with broad scarious margins; marginal flow-
ers pistillate, fertile; central flowers perfect,
fertile, the corolla glabrous, yellow; recep-
tacle and achenes glabrous. This is a wide
spread species of many phases and habitats.
In Utah there are five more or less distinctive
varieties. Two of the varieties, ludoviciana
and incompta are especially abundant, the re-
maining three less so. Not all specimens are
readily separable into the named varieties,
and the following key is arbitrary.

1. Inflorescence an open panicle, often more than 8 cm wide; plants of southern

and southeastern Utah 2

— Inflorescence a spicate panicle, usually less than 6 cm wide; plants of various
distribution 3

2(1). Leaves mainly less than 2.5 cm long, the margin often narrowly revolute

A. ludoviciana var. alhula

— Leaves mainly over 2 cm long, the margins not revolute

A. ludoviciana var. mexicana

3(1). Leaves entire or less commonly some of them toothed or lobed

A. ludoviciana var. ludoviciana

— Leaves more or less deeply parted or divided 4

4(3). Involucres 3.5-4.2 mm high, 4-7 mm wide A. ludoviciana var. latiloba

— Involucres 2.5-3.5 (3.8) mm high, 2.5-5 mm wide A. ludoviciana var. incompta

212

Great Basin Naturalist

Vol. 43, No. 2

Var. albula (Wooton) Shinners [A. albula
Wooton, nomen novum pro A. microcephalu
Wooton]. This distinctive short-leaved vari-
ety has open inflorescences; it occurs in ri-
parian areas with rabbitbrush, cottonwood,
and copperweed at 880 to 1680 m in Emery,
Garfield, Kane, San Juan, and Washington
counties; Nevada and Colorado south to Mex-
ico; 4 (ii).

Var. incompta (Nutt.) Cronq. [A. incompta
Nutt.; A. ludoviciana ssp. incompta (Nutt.)
Keck]. The deeply lobed or cleft leaves and
compactly spicate inflorescence are diagnos-
tic. The plants occur at moderate to high ele-
vations (2135 to 3500 m) in aspen, spruce-fir,
willow-wet meadow, and riparian commu-
nities in Cache, Carbon, Duchesne, Juab,
Millard, Piute, Salt Lake, San Juan, Sanpete,
Sevier, Summit, Tooele, Utah, and Wasatch
counties; Oregon to Montana, south to Cali-
fornia, Nevada, and Colorado; 65 (vi). This
variety passes by degree into A. michauxiana
at high elevations.

Var. latiloba Nutt. [A. candicans Rydb.; A.
ludoviciana ssp. candicans (Rydb.) Keck].
This variety is similar to var. incompta,
differing in larger (higher and wider) heads.
It is poorly understood in Utah, where it was
reported by Keck (1946) from Cache County.
Our one specimen is from Utah County,
without locality data; 1 (0).

Var. ludoviciana [A. gnaphaloides Nutt.;
A. ludoviciana var. gnaphaloides (Nutt.) T. &
G.; A. purshianus Bess, in Hook.]. The typi-
cal variety is a plant with entire or cleft
(rarely deeply cleft or parted) leaves and
loose, but not open, inflorescences. Rabbit-
brush, sagebrush-grass, mountain brush, pin-
yon-juniper, ponderosa pine, and hanging
garden communities at 880 to 2750 m in all
Utah counties, except for Morgan, Piute, and
Summit (and likely there also); British Co-
lumbia to Ontario, south to California, Ari-
zona, New Mexico, Texas, and Indiana; 86
(xvi).

Var. mexicana (Willd.) Fern. [A. mexicana
Willd.; A. ludoviciana ssp. mexicana (Willd.)
Keck]. This is the long-leaved plant with
open inflorescences; it forms the counterpart
of var. albula. It is a component of riparian,
pinyon-juniper, ponderosa pine, and aspen
communities at 750 to 2600 m in Garfield,
Grand, Kane, San Juan, Washington, and

Wayne counties; Colorado to Missouri, south
to Mexico; 17 (viii).

Artemisia michauxiana Bess. Michaux
Wormwood. [A. discolor Dougl. ex Bess.]. Pe-
rennial herbs, the stems 0.8-4 dm tall (rarely
more), white tomentose to glabrate or
glabrous; leaves mainly cauline, 0.5-4 (5) cm
long and about as broad, bipinnately dis-
sected, the secondary segments again toothed
or lobed, acute, the uppermost seldom entire,
commonly green above and tomentose be-
neath, but often green beneath also; in-
florescence spicate; heads several to numer-
ous, commonly pedicellate, erect or nodding;
involucres 3.4-4.4 mm high, 3-6 mm wide,
the bracts glabrous or sparingly tomentose,
green, the broad margins brownish scarious
and erose-ciliate; marginal flowers pistillate,
fertile; central flowers perfect, fertile, the co-
rolla glabrous (glandular), yellow; receptacle
and achenes glabrous. Spruce-lodgepole pine
and alpine tundra communities, often in
boulder stripes and talus, at 2950 to 3500 m
in Duchesne, San Juan, Summit, and Utah
counties (Uinta, La Sal, and Wasatch moun-
tains); British Columbia and Alberta south to
Nevada and Wyoming; 11 (ii). Keck (1946)
notes that A. michauxiana is connected
through a series of intermediates with A. lu-
doviciana var. incompta in Nevada speci-
mens. This is true for ours also. There ap-
pears to be some justification for treating A.
michauxiana within an enlarged A. ludovi-
ciana, but such a combination is not implied
herein.

Artemisia norvegica Fries Spruce Worm-
wood. Perennial herbs, 2-4.1 dm tall, from a
simple or branched caudex and stout taproot,
the caudex branches short, clothed with per-
sistent leaf bases, the flowering stems arising
directly from the caudex, villous, often red-
dish; leaves of basal rosettes 2-19 cm long,
bi- or tripinnatifid, the segments lance-atten-
uate to acute, villous on both surfaces; cau-
line leaves becoming smaller upwards, often
with stipulelike divisions near the base; in-
florescence racemose; heads several to nu-
merous, finally nodding, the peduncles to 4.5
cm long; involucres 4-5.3 mm high, 6-11
mm wide, the bracts sparingly to densely vil-
lous-pilose, more or less green, the margins
broadly dark brownish scarious; marginal

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

213

flowers pistillate, fertile; central flowers per-
fect, fertile, the corollas long-hairy from near
the base, cream colored; receptacle and ach-
enes glabrous. Spruce-fir, lodgepole pine, and
alpine tundra communities in Duchesne and
Summit counties; Alaska east to Mackenzie,
and south to California and Colorado. Our
material belongs to var. piceetorum Welsh &
Goodrich in Welsh; 4 (iii).

Artemisia nova A. Nels. Black Sagebrush.
[A. tridentata ssp. nova (A. Nels.) H. & C; A.
tridentata var. nova (A, Nels.) McMinn; A.
arbuscula ssp. nova (A. Nels.) Ward]. Shrubs,
1-3 (5) dm tall, the main branches spreading,
the vegetative stems 1-3 dm long (rarely
more); flowering stems mainly 1.5-3 (4) dm
long; leaves 0.3-2.1 cm long, shallowly to
deeply 3- to 5-lobed or -toothed, the lobes or
teeth rounded, often lead-gray or gray green,
cuneate basally, appressed canescent and of-
ten minutely punctate; inflorescence narrow-
ly paniculate, seldom more than 3 cm wide;
involucres 3.1-5.8 mm long, 1.4-3.4 mm
wide, cylindric to narrowly campanulate;
bracts 8-12, canescent to glabrous, green to
yellowish, the margin hyaline; flowers 3-8,
all perfect; receptacle glabrous; achenes
glabrous. Horsebrush, greasewood, shadscale,
ephedra, juniper, sagebrush, rabbitbrush,
winterfat, pinyon-juniper, and mountain
brush communities at 1400 to 2600 m in Bea-
ver, Box Elder, Cache, Carbon, Daggett,
Duchesne, Emery, Garfield, Grand, Iron,
Juab, Kane, Millard, Piute, Rich, San Juan,
Sanpete, Sevier, Summit, Tooele, Uintah, and
Weber counties; Oregon to Montana, south
to California, Arizona, and New Mexico; 57
(x). Black sagebrush forms intermediates with
all other members of the section Tridentatae
that it contacts. The intermediates form nar-
row bands along lines of contact, but general-
ly the habitats are mutually exclusive. There
is little justification for considering black
sagebrush in an expanded A. tridentata unless
one is willing to accept most of the remain-
der of the section as portions of that species
also.

Artemisia parryi Gray Parry Wormwood.
Perennial herbs, 0.8-2 (4) dm tall, from a
simple or branched caudex and stout taproot,
the caudex branches short, clothed with per-
sistent leaf bases, the flowering stems arising

directly from the caudex, sparingly and loose-
ly villous to glabrous, often reddish; leaves of
basal rosettes 2-4 (8) cm long, twice pinnati-
fid, the segments oblong to lance-oblong,
sparingly and loosely villous (to glabrous?);
cauline leaves becoming smaller upwards; in-
florescence racemose to subspicate; heads
several to numerous, commonly nodding, the
peduncles 1-5 mm long; involucres 3-4 mm
long, 3-5 mm wide, the bracts sparingly vil-
lous to glabrate, with green to brownish
middle and brownish scarious margins; mar-
ginal flowers pistillate, fertile; central flowers
perfect, fertile, the corollas long-hairy (to
glabrous?); receptacle and achenes glabrous.
Alpine sites in the La Sal Mountains (Grand
and San Juan counties); Colorado; 0 (0). The
species is reported for Utah by Hall and Cle-
ments (I.e.), but no specimens have been seen
from the state by me. Possibly it is only a
phase of A. norvegica.

Artemisia pygmaea Gray Pygmy Sage-
brush. Shrubs 0.5-2 dm tall, from superficial
woody caudexlike branches and stout tap-
roots, the vegetative stems to 0.5 dm long;
flowering stems erect to 2 dm tall; leaves
0.3-1 cm long, pinnately (or subbipinnately)
3- to 10-lobed, the lobes acute, yellow- to
gray-green, sparingly villous to glabrous; in-
florescence spicate or narrowly paniculate,
less than 2 cm wide; involucres 5.2-6.3 mm
high, 3-4.5 mm wide, cylindric or becoming
campanulate upon drying; involucral bracts
oblong, 15 or more, sparingly villous to
glabrous, green, the margins stramineous-
hyaline; marginal flowers lacking; central
flowers 3-5, perfect, fertile, the corollas
cream colored, glandular; receptacle and
achenes glabrous. Black sagebrush, rabbit-
brush, shadscale, greasebush, juniper, pinyon-
juniper, and ponderosa pine communities at
1600 to 2300 m in Beaver, Duchesne, Emery,
Garfield, Iron, Millard, Piute, Sevier, and
Uintah counties (likely elsewhere); Arizona
and Nevada; 27 (viii). This dwarf sagebrush
occurs in peculiar edaphic situations on
Green River Shale, in clay soils forming the
matrix in igneous gravels, on calcareous
gravels, and on dolomitic outcrops and
gravels. It is often a component of commu-
nities that support rare plant species.

Artemisia scopulorum Gray Dwarf Sage-
wort. Perennial herbs, 0.5-3.7 dm tall, from a

214

Great Basin Naturalist

Vol. 43, No. 2

simple or branched caudex and stout taproot,
the caudex branches short, clothed with per-
sistent leaf bases, the flowering stems arising
directly from the caudex, appressed pilose to
loosely and sparingly villous, often reddish or
purplish; leaves of basal rosettes 1.5-9 cm
long, twice pinnatifid, the segments oblong
to elliptic, pubescent like the stems; in-
florescence spicate to racemose; heads sever-
al to numerous, erect or nodding, the pe-
duncles lacking, or to 2.3 cm long; involucres
3-5.2 mm high, 3-8 mm wide, the bracts vil-
lous, green to brownish in the middle, the
margins brown-scarious; marginal flowers
pistillate, fertile; central flowers perfect, fer-
tile, the corollas cream colored, long-hairy;
receptacle copiously long-villous; achenes
glabrous. Talus slopes, moraines, and outwash
plains and terraces in alpine tundra and
meadows in spruce, lodgepole pine, and
Douglas fir communities at 3050 to 4000 m
in Boulder, Tushar, La Sal, and Uinta moun-
tains; Beaver, Duchesne, Garfield, Grand,
Piute, Summit, and Uintah counties; Mon-
tana, Wyoming, Colorado, and New Mexico;
30 (xi). The hairy corollas and long-villous re-
ceptacles are diagnostic for this distinctive
species.

Artemisia spiciformis Osterh. Osterhout
Sagebrush. Shrubs, mainly 5-8 dm tall, the
vegetative stems 0.4-1 dm long, the flower-
ing stems erect, 1.5-3.4 dm long; leaves
1.7-5.7 cm long, shallowly to deeply 3- to 5-
lobed or -toothed, often widest below the
teeth, the lobes acute to obtuse (or rounded)
or lacking, gray-green, long-cuneate basally,
appressed villous-canescent; inflorescences
narrowly paniculate, usually less than 4 cm
wide; involucres 5-6.3 mm long, 3.5-7 mm
wide, cylindric to campanulate; involucral
bracts 8-12 or more, canescent to glabrate,
green, with broad yellowish brown scarious
margins; flowers 6-10 or more, all perfect;
receptacle and achenes glabrous. Ridge mar-
gins and snow-flushes in sagebrush-grass,
snowberry, aspen, spruce-fir, and Douglas fir
commimities at 2680 to 3050 m in Cache (?),
Duchesne, Emery, Juab, Sanpete, Summit,
Tooele, and Wasatch counties (likely else-
where); Colorado and Wyoming; 16 (ii). This
is the material which has long passed under
the name of A. rothrockii Gray in Utah.

Resemblance to that species appears to be
superficial, with relationships running to both
A. cana and A. tridentata var. vaseyana. Its
habitat is intermediate between the high ele-
vation, moderately xeric conditions of var.
vaseyana, and the more mesic stream terrace
and valley bottoms of A. cana.

Artemisia spinescens D.C. Eaton in Wats.
Budsage. Shrubs, flowering in springtime, the
branches spreading and often prostrate, 0.5-3
dm long or more, the vegetative stems
mainly 0.3-0.8 dm long, commonly surpass-
ing the flowering stems; leaves 0.4-2 cm
long, petiolate, the blade palmately 3- to 5-
cleft, the main divisions again cleft, sub-
orbicular in outline, villous; inflorescence of
short leafy-bracted racemose or spicate
branches, or of solitary heads, the rachis per-
sistent as a thorn; involucres 2-3.5 mm high,
3.5-5 mm wide; involucral bracts 4-8, vil-
lous, green, with narrow hyaline margins;
flowers 6-20 or more, the marginal ones pis-
tillate, fertile, the central ones sterile; co-
rollas copiously long-hairy; receptacle naked;
achenes long-hairy. Silty, clayey, or gravelly,
often saline, substrates in black sagebrush,
shadscale, tetradymia, greasewood, black-
brush, juniper, and winterfat communities at
1200 to 1925 m in Carbon, Duchesne,
Emery, Garfield, Juab, Kane, Millard, Piute,
San Juan, Sevier, Tooele, Uintah, and Utah
counties; Oregon to Montana, south to Cali-
fornia and New Mexico; 92 (vii). This low
shrub is a principal browse plant for domes-
tic livestock on the spring ranges of western
and southern Utah.

Artemisia tridentata Nutt. Big or Com-
mon Sagebrush. Shrubs 4-20 (30) dm tall;
branches spreading to erect, the vegetative
branchlets 0.5-2 dm long; flowering stems
mostly 1.5-4 dm long, usually much surpass-
ing the vegetative ones; leaves 0.5-5 cm
long, 3- to 5-toothed apically, or the upper
ones entire, long-cuneate; inflorescence pan-
iculate, 3-20 (15) cm wide; involucres 3-5
mm long, 2-4 mm wide, the bracts 10-20,
green, canescent, the margins scarious; flow-
ers 3-8, all perfect, the corollas cream col-
ored, glandular; receptacle and achenes
glabrous. Three more or less completely in-
tergrading varieties are known from Utah;
they tend to occupy distinctive habitats, but

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

215

intermediates form wherever they meet. Fur-
ther, this taxon is known to hybridize with
most if not all other members of the section
Tridentatae. The help of Durrant MacArthur
and Sherel Goodrich of the U.S. Forest Ser-
vice is here gratefully acknowledged. They

sorted our materials into their respective va-
rieties following my initial attempt and gen-
eral failure. While it is not possible to segre-
gate all specimens, the following key will
prove useful to those who must manage the
sagebrush lands of Utah and the west.

2(1).

Vegetative stems short, standing at about the same height, the inflorescence

rather uniformly overtopping them; plants of middle and higher elevations

A. tridentata var. vaseyana

Vegetative stems short to long, the inflorescence not uniformly overtopping
them; plants of low to moderate elevations 2

Leaves mainly to 2 cm long or more, narrowly cuneate; plants of low to mod-
erate elevations A. tridentata var. tridentata

Leaves mainly less than 1.2 cm long, cuneate to cuneate-flabellate; plants
mainly of moderate elevations, in drier sites A. tridentata var. wyomingensis

Var. tridentata Big Sagebrush. Sagebrush,
juniper, pinyon-juniper, and rabbitbrush
communities at 1220 to 2410 m in most, if
not all, Utah counties; Washington to Mon-
tana, south to California, Arizona, and New
Mexico; 57 (xviii).

Var. vaseyana (Rydb.) B. Boi. Vasey Sage-
brush. [A. tridentata ssp. vaseyana (Rydb.)
Beetle]. Sagebrush, rabbitbrush, mountain
brush, pinyon-juniper, aspen, Douglas fir,
ponderosa pine, and spruce-fir communities
at 1830 to 3050 m in all, or nearly all, Utah
coimties; Idaho to the Dakotas, south to Col-
orado; 55 (viii).

Var. wyomingensis (Beetle & Young)
Welsh Stat. nov. [based on Artemisia triden-
tata ssp. wyomingensis Beetle & Young Rho-
dora 67: 405. 1965]. Wyoming Sagebrush.
Shadscale, rabbitbrush, sagebrush, juniper,
bitterbrush, and mountain mahogany com-
mimities at 1525 to 1980 m in Box Elder,
Garfield, Emery, Rich, Tooele, and Uintah
counties; Wyoming and Idaho to Colorado.
This is the sagebrush of drier sites at middle
elevations. Its distribution is poorly under-
stood; likely it is widespread. Its recognition
allows management considerations by profes-
sionals in the various state and federal
agencies; 9 (0).

Artemisia tripartita Rydb. Threetip Sage-
brush. Shrubs 2-20 dm tall, the branches
erect, the vegetative ones 0.3-1.5 dm long,
the flowering stems 0.6-3.5 dm long; leaves
1-4 cm long, deeply 3-cleft, the linear lobes

0.5-0.8 mm wide, canescent, the lobes some-
times again divided, or the upper ones entire;
inflorescence paniculate, commonly 2-5 cm
wide; involucres campanulate, 3-4 mm long,
1.5-4 mm wide; bracts many, imbricate, ca-
nescent and more or less green, the inner
with broad brownish scarious margins; flow-
ers 4-8, all perfect, the corollas stramineus to
cream-colored, more or less glandular; ach
enes and receptacle glabrous. Sagebrush and
mountain brush communities at ca 1525 to
1830 m in Box Elder and Cache counties;
British Columbia to Montana, south to Cali-
fornia and Colorado; 1 (0).

Aster L.

Annual or perennial herbs from rhizomes
(suffrutescent in A. spinosus), with watery
juice; stems decumbent to ascending or erect,
simple or branched; leaves alternate, simple,
entire or toothed; heads solitary or few to
several in corymbose clusters; involucral
bracts strongly imbricate to subequal (or the
outer surpassing the inner), herbaceous
throughout, or with scarious margins near the
base; receptacle flat or merely convex,
naked; rays blue, purple, pink, or white, few
to numerous, pistillate; disk flowers numer-
ous, perfect, fertile, yellow or tinged reddish
or purplish; pappus of capillary bristles; style
branches flattened, oblong to lanceolate,
mostly more than 0.5 mm long; achenes
mostly several nerved.

216 Great Basin Naturalist Vol. 43, No. 2

1. Plants suffrutescent, rushlike, armed with axillary or subaxillary thorns, from a
deep-seated rhizome; known from Garfield County (possibly Washington also)
A. spinosus

— Plants herbaceous, annual or perennial, unarmed, from a taproot or rhizome; distri-
bution various 2

2(1). Plants annual, from taproots 3

— Plants perennial, from rhizomes or subrhizomatous caudices, or from branch-
ing caudices 4

3(2). Involucral bracts definitely acute; rays wanting or nearly so, the pistillate

corollas tubular, shorter than the style A. brachyactis

— Involucral bracts obtuse to obtusish; rays to 2 mm long, longer than the style ...

A. frondosus

4(2). Plants with a well developed caudex; involucral bracts reflexed, at least the

outer; plants of rock crevices in the Wasatch and Canyon mountains A. kingii

— Plants with caudex lacking or poorly developed, rhizomatous; involucral bracts

not reflexed; plants of various habitats and localities 5

5(4). Leaves all erect-ascending, thickened, to about 4 mm wide; pappus double, the
outer series of very short bristles; heads solitary; plants known from Box Elder
County A. scopiiloriim

— Leaves various, seldom as above; pappus in one series, or rarely double; heads
solitary to numerous; distribution various 6

6(5). Involucral bracts dry, chartaceous, with scarious tips (at least the innermost),

with a distinctive midvein, not herbaceous (the outer sometimes so) 7

— Involucral bracts herbaceous at the tips or throughout, lacking a distinctive
midvein 9

7(6). Involucral bracts (at least the outer) bluntly obtuse apically; herbage strongly

glaucous; plants often of open calcareous sites A. glaucodes

— Involucral bracts acute; herbage green, not glaucous; plants of various habitats 8

8(7). Rays white (drying pinkish); main leaves often over 20 mm wide; plants 6-15

dm tall, of montane areas in central northern Utah A. engelmannii

— Rays purple or violet; leaves mainly less than 15 mm wide; plants 2-6 dm tall,

of central northern and western Utah A. perelegans

9(5). Involucres and peduncles glandular 10

— Involucres and peduncles lacking glands or apparently so 13

10(9), Stems glabrous; leaves linear to linear-oblanceolate, 2-5 mm wide, 1.5-7 cm

long; plants of saline or hot water seeps and springs A. pauciflorus

— Stems puberulent to villous with multicellular hairs, or glabrous, but, if so,
differing in other respects 11

11(10). Rays white; leaves glaucous; plants of central to south central Utah ..A. wasatchensis

— Rays blue to purple, lavender, or violet 12

12(11). Cauline leaves clasping the stem, mainly 15-40 mm wide; involucres 8-15

mm high; plants of central northern Utah A. integrifolius

— Cauline leaves not or only slightly clasping, 2-10 mm wide; involucres 5-8

mm high A. campestris

April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 217

13(9). Pubescence occurring in decurrent lines below leaf bases, commonly not uni-
form below the heads, or only in the inflorescence; inflorescence often conic,
mostly large and leafy A. hesperius

— Pubescence of stem uniform, or, if in decurrent lines, uniform below the heads
and confined to the inflorescence; inflorescence few to many flowered and not
usually leafy to large and leafy (see also A. eatonii) 14

14(13). Rays white; involucral bracts strigulose dorsally (rarely glabrous), with spread-
ing to squarrose minutely spinulose tips; heads numerous 15

— Rays pink to purple, or less commonly white; involucral bracts mucronate at

the tip; heads few to numerous 16

15(14). Rhizomes well developed, creeping; involucres 4.6-6.5 mm high, 7-9.5 mm

wide (when pressed); plants of western Utah A. falcatus

— Rhizomes mainly poorly developed, or reduced and caudexlike; involucres
3.8-4.9 mm high, 4.5-6 mm wide; plants of eastern Utah A. pansus

16(14). Achenes glabrous or nearly so; herbage glabrous except for lines of pu-
berulence in the inflorescence, tending to be glaucous; rare plants in
southeastern Utah A. laevis

— Achenes pubescent, except in some A. foliaceus; herbage pubescent to almost
glabrous, scarcely glaucous 17

17(16). Involucral bracts strongly imbricate, the outer ones at least obtuse or obtusish

(sometimes acute), not foliaceous; pubescence below the heads harsh A. chilensis

— Involucral bracts not strongly imbricate, or, if so, the bracts sharply acute,
the outer ones acute, or, if obtuse, foliaceous; pubescence below heads soft or
minute 18

18(17). Inflorescence a long slender leafy panicle; heads numerous; stem pubescence
short, uniform; leaves mostly more than 7 times longer than wide; rays usually
pink to white A. eatonii

— Inflorescence an open or congested panicle; heads solitary to several;
pubescence various; rays usually blue to violet 19

19(18). Involucral bracts slender, never foliaceous; leaves at midstem mostly less than

1 cm wide, mostly over 7 times longer than broad A. occidentalis

— Involucral bracts various, but some of them usually enlarged and foliaceous;
leaves at midstem mostly less than 7 times longer than broad A. foliaceus

Aster brachyactis Blake in Tidestr. [Tripo- tubular filiform, lacking rays, much shorter
Hum angustum Lindl. in Hook.; A. angustus than the styles; pappus abundant, white,
(Lindl.) T. & G., not Nees; Brachyactis ang- longer than the corollas. Sandbars, terraces,
ustus (Lindl.) Britt. in Britt. & Brown]. An- stream banks, marshes and pond margins, of-
nual herbs, with taproots, glabrous through- ten where saline, in tamarix, rush, rabbit-
out, except for leaf margins and involucral brush, and cottonwood commimities at 1220
bracts; stems 0.9-5.3 (7) dm tall; leaves 1.3-8 to 1525 m in Box Elder, Carbon, Duchesne,
(12) cm long, 1-7 (9) mm wide, linear to nar- Emery, Grand, Garfield, Salt Lake, Uintah,
rowly oblong, entire, the lower ones soon de- and Utah counties; British Columbia to Min-
ciduous; heads few to numerous, in pan- nesota, south to Washington and Colorado;
iculate to spicate inflorescences; involucres 14 (i).

5.5-9.4 (11) mm high, 7-15 (17) mm wide. Aster campestris Nutt. Meadow Aster. Pe-

the bracts linear-oblong, acute to attenuate, rennial rhizomatous herbs, glandular, at least

herbaceous, subequal to somewhat imbricate, in inflorescence; stems puberulent to

or some outer ones often surpassing the in- glabrous, mainly 1-5 dm tall; leaves 2-8 cm

ner; marginal flowers pistillate, the corollas long, 2-8 mm wide, linear to oblong, entire.

218

Great Basin Naturalist

Vol. 43, No. 2

sessile, sometimes clasping, the lower ones
larger and more or less petiolate, or smaller,
soon deciduous; heads solitary or several to
many; involucres 5-8 mm high, glandular,
the bracts subequal to definitely imbricate,
acute or attenuate, with long herbaceous tips;
rays 15-20, violet to purple, 6-12 mm long.
Meadows at 1525 to 2475 m, reported for
Utah (Univ. Washington Publ. Biol. 17(5): 77.
1955), but I have seen no specimens from the
state.

Aster chilensis Nees Pacific Aster. Pe-
rennial rhizomatous to subrhizomatous herbs,
imiformly harshly strigose to strigulose, at
least above; stems (0.8) 1.2-10.5 dm tall;
leaves 0.6-16.5 cm long, 2-16 (20) mm wide,
entire or nearly so, pubescent to glabrous, cil-
iate, the lower ones more or less petiolate, of-
ten deciduous at anthesis in taller plants, be-
coming smaller and sessile upwards,
sometimes markedly reduced-bracteate in in-
florescence; inflorescence of 1 to many heads,
narrow, corymbose, or open paniculate; in-
volucres 5-8 mm high, 6-15 mm broad, the
bracts imbricate, green tipped (machaeran-
theroid), the chartaceous bases white to straw
colored, the outer ones abruptly pointed but
mucronate; rays commonly 15-40, purplish
to violet (rarely white) or pink, 5-15 mm
long; achenes pubescent. Alluvial fans, ter-
races, and slopes along stream and canal
banks, in hanging gardens, rabbitbmsh, sage-
brush, grass-sedge, cottonwood-willow, pon-
derosa pine, juniper-pinyon, mountain brush,
aspen, and spruce-fir communities at 850 to
3200 m in all Utah counties; Washington to
Saskatchewan, south to California and New
Mexico; 189 (xxviii). The Pacific aster is a
generalized taxon with no clearly diagnostic
features. It is separated from its near con-
geners by a group of intangible character-
istics. Involucral bracts are definitely imbri-
cate, with the greenish portion usually
glabrous, and margins ciliate. The tips of out-
er bracts are often but not always obtuse, and
the tip, even when abruptly contracted is
mucronate. These features, which I designate
as "machaerantheroid," are shared to a great-
er or lesser extent with A. eatonii, A. occiden-
talis, and A. foliaceus. The harsh pubescence
below the heads appears to be diagnostic, but
is difficult to distinguish from the soft or
merely puberulent vesture of closely related

species. Not all specimens can be assigned
with certainty to any of the taxa. There are
two intergrading morphological phases of the
Pacific aster, which are striking in their ex-
tremes, but which probably represent noth-
ing more than developmental gradients.
There are plants with few flowers that lack
distinctive reduced bracteate leaves in the in-
florescence, and taller plants with more nu-
merous heads and distinctively bracteate in-
florescences. The inflorescences of the taller
plants are mainly corymbiform, and not cy-
lindroid as in A. eatonii. More work is in-
dicated. Our material belongs to ssp. adscen-
dens (Lindl.) Cronq.

Aster eatonii (Gray) Howell Eaton Aster.
[A. foliaceus var. eatonii Gray; A. oregonus
authors, not (Nutt.) T. & G.]. Perennial rhizo-
matous to subrhizomatous herbs, uniformly
puberulent, at least above (below the heads
and sometimes on upper leaves), the stems
(2.7) 6-10.5 dm tall, often reddish; leaves
0.8-15 cm long, 2-25 mm wide, entire or ser-
rate, puberulent to glabrous, ciliate, the
lowermost shortly petiolate, often deciduous
in anthesis, becoming smaller and sessile up-
wards, linear to narrowly elliptic or lanceo-
late to oblanceolate; inflorescence of few to
numerous heads, commonly open-cylindric to
conic in form; involucres 4.5-8 (10) mm high,
6-10 mm wide, the bracts more or less sub-
equal to indistinctly imbricate, green tipped
(but not especially machaerantheroid), the
chartaceous bases white to straw colored, all
or most of them mucronate; rays 20-40, com-
monly pink (sometimes white), 5-12 mm
long; achenes pubescent. Gravel bars, stream
terraces, meadows, canal banks, hanging gar-
dens, and marshes at 1370 to 2325 m in
Cache, Garfield, Grand, Iron, Juab, Kane,
Salt Lake, Summit, Uintah, Utah, Wasatch,
and Washington counties; British Columbia
to Saskatchewan, south to California, Ari-
zona, and New Mexico; 48 (ix). The pink or
white rays, uniform upper stem puberulence
and leaves many times longer than broad are
diagnostic for most specimens. Reports of A.
junciformis Rydb. for Utah appear to be
based on slender phases of the Eaton aster
with linear leaves and slender rhizomes.

Aster engelmannii (D.C. Eaton) Gray
Engelmann Aster. [A. elegans var. engel-
mannii D.C. Eaton]. Perennial rhizomatous

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

219

or subrhizomatous herbs, puberulent to spar-
ingly villous with multicellular hairs, or
somewhat glandular, the stems 2-15.2 dm
tall, reddish at the base; leaves 2-13.5 cm
long, 3-46 mm wide, elliptic to lanceolate,
entire (or nearly so), sparingly puberulent to
glabrous or sparsely villous, sessile, largest
near midstem, the lowermost reduced to
scales; inflorescence of 1 to numerous large
heads, corymbose or conic; involucres 8-13
mm high, 11-25 mm wide, the bracts mainly
strongly imbricate, with a definite midvein,
commonly purplish (at least the inner), the
outer sometimes green and more or less fo-
liaceous, sometimes all greenish or straw col-
ored to the tip, glabrous dorsally, ciliate; rays
8-23, white (drying pinkish), 12-25 mm long;
achenes pubescent. Mountain brush, juniper,
Douglas fir, aspen, white fir, lodgepole pine,
and spruce-fir communities at 1950 to 3200
m in Cache, Carbon, Davis, Duchesne, Juab,
Salt Lake, Sanpete, Summit, Utah, and
Wasatch counties; British Columbia and Al-
berta, south to Nevada and Colorado; 57 (vi).
Aster falcatus Lindl. [A. miiltiflorus var.
commutatus T. & C; A. commutatus (T. &
G.) Gray]. Perennial rhizomatous herbs, vil-
lous or villous-hirsute with multicellular
hairs, the stems 2.8-7.5 dm tall; leaves 1.2-6
(8) cm long, 2-8 mm wide, entire, antrorsely
scaberulous on both surfaces (or glabrous),
sessile, linear to narrowly oblong, often spin-
ulose-mucronate, the lowermost often lacking
at anthesis; inflorescences several- to many-
headed, cylindroid; involucres 4.6-6.5 mm
high, 7-9.5 mm wide, the bracts strongly to
only somewhat imbricate, with a green tip,
scaberulous to glabrous dorsally and ciliate;
rays mainly 17-25, white (drying pale laven-
der in some), 6-8 mm long; achenes pu-
bescent. Oak, sagebrush, and ponderosa pine
communities at 1525 to 2135 m in Box Elder,

Kane, Utah, and Washington counties; Alaska
to Minnesota, south to California, New Mexi-
co, and Kansas; 7 (i). The species is closely al-
lied to A. pansus (q.v.), which has smaller
heads.

Aster foliaceus Lindl. in DC. Leafybract
Aster. Perennial rhizomatous or sub-
rhizomatous herbs, uniformly and shortly
soft-villous below the heads, uniformly vil-
lous to glabrous below, or in lines below leaf
bases, the stems 1.3-7 dm tall; leaves 1,8-16
cm long, 3-34 mm wide, entire or nearly so,
strigose to glabrous, ciliate, the lower ones
petiolate (often lacking at anthesis), becom-
ing smaller and sessile (and more or less
clasping) upwards; inflorescence of 1-19 (50)
corymbosely arranged large and showy
heads; involucres 6-12 mm high, 10-20 mm
wide, the bracts imbricate to slightly so, fo-
liaceous or slender, green with pale white to
yellowish or brownish chartaceous bases (at
least the inner), acute to obtuse or rounded,
mucronate; rays mainly 15-50, pink to
purple, blue, or violet, 9-16 (20) mm long;
achenes hairy. The leafybract aster is a por-
tion of an assemblage that includes the con-
cept of A. subspicatus Nees. Both A. fo-
liaceus and A. subspicatus were described
from coastal Alaska (Unalaska and Yakutat
Bay, respectively). Brownish bases of in-
volucral bracts, commonly serrate leaves, and
reddish pappus are supposedly diagnostic for
A. subspicatus, which is not known from
Utah, but some specimens of A. foliaceus
have one or more of these features. In the
Alaska Flora (Welsh 1974), I treated both
species under the older name of A. sub-
spicatus. Now, I follow tradition so as to
avoid creation of synonyms should further
study indicate a better course of action.
Three more or less distinctive infraspecific
taxa are present in Utah.

1. Involucral bracts foliaceous, 2-6 mm broad; plants uncommon

A. foliaceus var. canbyi

— Involucral bracts not especially foliaceous, mainly less than 2 (2.5 ) mm wide;
plants common to uncommon 2

2(1). Plants mainly 0.5-2.5 dm tall, decumbent or ascending; bracts often purple

margined or tipped; known from high elevations, rare A. foliaceus var. apricus

— Plants often more than 2 dm tall, erect; bracts seldom as above; known from

low to high elevations, common A. foliaceus var. parryi

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Var. apricus Gray Meadows in spruce-fir
forest at 3050 to 3660 m in Summit County;
British Columbia to Montana, south to Cali-
fornia and Colorado; 1 (0).

Var. canbyi Gray Mountain brush, aspen,
and spruce-fir communities at 1950 to 2900
m in Iron, Juab and Salt Lake counties;
Washington to Wyoming, south to California
and New Mexico; 5 (0).

Var. parryi (D.C. Eaton) Gray [A. adscen-
dens var. parryi D.C. Eaton; A. foliaceus var.
frondeus Gray]. Meadows and openings in as-
pen, spruce, lodgepole pine, and Douglas fir
communities at 1890 to 3265 m in Cache,
Duchesne, Garfield, Iron, Juab, Piute, Salt
Lake, Sanpete, Summit, Tooele, Uintah, and
Utah counties; Washington to Wyoming,
south to California and New Mexico; 36
(viii). This is the phase of the leafybract aster
that simulates A. occidentalis (q.v.), but
which seldom has long peduncles, dark blue-
purple ray corollas, and much reduced upper
stem leaves of that species.

Aster frondosus (Nutt.) T. & G. Leafy As-
ter. [Tripolium frondosum Nutt.]. Annual
herbs from taproots; stems 0.2-3.6 cm tall;
leaves 1-6 cm long, 2-12 mm wide, linear to
oblong or oblanceolate, entire, the lower
ones sometimes deciduous; heads few to nu-
merous, in a narrow paniculate to spicate in-
florescence; involucres 5-9 mm high, 6-13

mm wide, the bracts oblong to narrowly ob-
lanceolate, obtuse or obtusish, herbaceous,
subequal to moderately imbricate; marginal
flowers pistillate, the rays developed, pink, to
2 mm long; pappus abundant, white, longer
than the disk corollas. Lake shores, seep mar-
gins, wet meadows, and stream banks in salt-
grass, tamarix, Russian olive, rabbitbrush, and
greasewood communities at 1250 to 2270 m
in Beaver, Duchesne, Garfield, Grand, Juab,
Kane, Salt Lake, San Juan, Utah, and Wayne
counties; Washington to Wyoming, south to
California and New Mexico; 18 (ii).

Aster glaucodes Blake Blueleaf Aster. Pe-
rennial rhizomatous herbs, glabrous and
glaucous, or puberulent to glandular in the
inflorescence; stems 1.1-7 dm tall; leaves
1.4-12.5 cm long, 4-25 mm wide, entire,
lance-oblong to oblong or elliptic, glaucous,
glabrous, sessile and clasping, the lower often
lacking at anthesis, reduced upwards; heads
few to numerous in corymbose in-
florescences; involucres 6-9 mm tall, 7-9 mm
wide, the bracts imbricate, dry, chartaceous
throughout or sometimes some of them
greenish, the midvein prominent, commonly
suffused with pink or purple, mainly obtuse
to less commonly acute apically; rays 10-20,
white or pink, 11-17 mm long. There are
two varieties within our specimens.

Peduncles and/ or involucres glandular-pubescent; plants of Washington and

adjacent western Kane counties A. glaucodes var. pulcher

Pedimcles and involucres lacking glandular pubescence; plants widespread

A. glaucodes var. glaucodes

Var. glaucodes This is the common phase
of the species, often on calcareous substrates
at higher elevations and in saline seeps at
moderate to lower elevations in sagebrush,
pinyon-juniper, mountain brush, ponderosa
pine, ryegrass, spruce-fir, Douglas fir, lodge-
pole pine, and hanging garden communities
at 1220 to 3050 m in Cache, Carbon, Du-
chesne, Emery, Garfield, Grand, Kane, San
Juan, Sanpete, Sevier, Summit, Tooele, Uin-
tah, Utah, Washington, and Wayne counties;
Idaho and Wyoming, south to Arizona and
Colorado; 56 (ix).

Var. pulcher (Blake) Kearney & Peebles
Note: This taxon was published at subspecific

rank by Blake, and was inadvertantly accept-
ed at varietal rank by Kearney and Peebles
(Arizona Flora 872. 1951), without citation of
the basionym, A. glaucodes ssp. pulcher
Blake, Proc. Biol. Soc. Washington 35: 174.
1922. Salt desert shrub, sagebrush, pinyon-
juniper, and ponderosa pine communities at
825 to 2136 m in Washington and adjacent
western Kane counties; Arizona; 6 (0).

Aster hesperius Gray Siskiyou Aster. [A.
laetivirens Greene]. Perennial rhizomatous
herbs, villous with multicellular hairs in de-
current lines from leaf bases, or less com-
monly almost glabrous and with decurrent
lines below the heads; stems 3.6-9.5 (15) dm

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

221

tall; leaves 3-17 (21) cm long, 5-27 mm
wide, entire or serrate, glabrous or scabrous,
ciliate, the lower ones commonly petiolate,
often deciduous at an thesis, becoming small-
er, sessile and more or less clasping upwards,
sometimes much reduced in inflorescence;
heads few to numerous in open to narrow
subcorymbose inflorescences; involucres
4.5-7 (8) mm high, 7-12 mm wide, the bracts
imbricate to subequal, green tipped, the
chartaceous base white to straw colored, all
acute and mucronate; rays commonly 20-50,
pink to blue or white, 6-14 mm long; ach-
enes hairy. Wet meadows, canal banks, and
stream sides with sedges, rabbitbrush, willow,
and other riparian communities at 850 to
2135 m in Box Elder, Cache, Duchesne, Gar-
field, Grand, Kane, Millard, Summit, Utah,
Wasatch, and Washington counties; Alberta
to Saskatchewan, south to California, Ari-
zona, New Mexico, and Missouri; 32 (vii).
This plant occurs at lower elevations in Utah
and has been confused with A. foliaceus, with
which some plants share the subequal bracts.
It has also been mistaken for A. chilensis,
with which it is partially sympatric. The lack
of uniformly disposed hair in the in-
florescence appears to be diagnostic.

Aster integrifolius Nutt. Thickstem Aster.
Perennial subrhizomatous herbs, glandular
villous with multicellular hairs, at least
above; stems 2.3-6.4 (7) dm tall; leaves
2.5-19 cm long, 8-50 mm wide, entire, ob-
lanceolate to elliptic or lanceolate, glandular-
villous, ciliate, the lower ones petiolate, be-
coming smaller, sessile and clasping upward;
heads few to several (numerous), large and
showy, in elongate to subcorymbose clusters;
involucres 8-13 (14) mm high, 12-23 mm
wide, the bracts mainly subequal, green or
suffused with purple, glandular dorsally, fo-
liaceous or not; rays commonly 10-25, dark
purple, 10-15 mm long. Meadows and moist
woods in sedge-willow, sagebrush, Douglas
fir, and spruce communities at 2275 to 3125
m in Rich, Salt Lake, Summit, and Wasatch
counties; Washington and Montana, south to
California and Colorado; 8 (0).

Aster kingii D.C. Eaton [Machaeranthera
kingii (D.C. Eaton) Cronq. & Keck]. Pe-
rennial herbs from a caudex and taproot, the
caudex branches clothed with blackish or
dark brown marcescent leaf bases, these

scarious and ashy when young; stems 3-12
(15) cm long, more or less villous below,
stipitate-glandular above; basal leaves 0.8-12
cm long, 3-22 mm wide, petiolate, the pet-
iole bases expanded and scarious, the blades
oblanceolate or spatulate, glabrous or glandu-
lar, or less commonly hispidulous or merely
puberulent on one or both sides; heads 1-5,
racemosely or corymbosely arranged; in-
volucres 8-11 mm high, 10-16 mm wide;
bracts glandular to shortly stipitate-glandu-
lar, herbaceous above the middle, scarious
below, often suffused purplish, especially the
inner, the tips of at least the outer reflexed;
rays 15-27, white (often fading pale pink),
8-17 mm long, 1.5-2.8 mm wide; achenes ca
3.5 mm long. Douglas fir-white fir, mountain
brush, and Cottonwood communities at 1839
to 3050 m in Juab, Millard, Salt Lake, and
Utah counties; endemic; 21 (i). The southern
populations have at least some toothed leaves
and stems with longer stipitate-glandular
hairs; they belong to var. barnebyana (Welsh
& Goodrich) Welsh comb. nov. [based on:
Machaeranthera kingii var. barnebyana
Welsh & Goodrich Brittonia 33: 299. 1981];
6 (0). Attempts to segregate genera within
the Astereae are often fraught with diffi-
culties. This is especially true of that core of
genera involving Haplopappus, Machaeran-
thera, Xylorhiza, and Aster. Cronquist and
Keck (1957. Brittonia 9: 231-329) reconsti-
tuted the genus Machaeranthera, and includ-
ed within that expanded generic definition
those species treated elsewhere herein as
Machaeranthera and Xylorhiza. Included
within the series Integrifoliae of section Xy-
lorhiza was Aster kingii. Watson (1978.
Madrono 25: 205-210) has shown the
chromosome number to be 2n=18 for Aster
kingii, and he notes that its placement within
Machaeranthera section Xylorhiza "is pheno-
logically, ecologically, morphologically, and
chromosomally anomalous. ..." The chromo-
some numbers reported for Xylorhiza are
2n = 12 or 24; that of Machaeranthera, in a
restricted sense, is 2n = 8, 10, or 16; that of
Aster is mainly 2n = 18. The taproots and
squarrose involucral bracts suggest an al-
liance with Machaeranthera, shorn of Xylo-
rhiza, but the similarity seems superficial, es-
pecially in light of different chromosome
numbers. Some asters in a strict sense, i.e.,

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Great Basin Naturalist

Vol. 43, No. 2

A. alpigenns Rydb., have a caudex, with the
rhizome attenuated. The logical conclusion
of such an attenuation is the caudex of A.
kingii, and the squarrose bracts seem to have
been secondarily derived, being present to a
greater or lesser degree in other Aster species
as well as in Machaeranthera. Hence, it seems
best to treat this taxon within Aster.

Aster laevis L. Smooth Aster. Sub-
rhizomatous perennial herbs, glabrous or
nearly so; stems mainly 5-12 dm tall; leaves
0.8-14 cm long, 2-30 mm wide, entire or ser-
rate, linear-subulate to lanceolate or elliptic,
the lower ones petiolate, often lacking at an-
thesis, becoming smaller, sessile, and more or
less clasping upwards; heads numerous, in co-
rymbose inflorescences; involucres 5-8 mm
high, 7-12 mm broad (when pressed), the
bracts slender, green tipped, the chartaceous
bases straw colored to brownish or white,
acute and mucronate; rays 15-30, blue or
purple, 6-9 mm long; achenes glabrous. Ri-
parian communities at ca 1400 m in Grand
(and San Juan?) County; Yukon to Maine,
south to Oregon, New Mexico, and Georgia;
1 (i). This plant is rare in collections from
Utah, due presumably to the paucity of late
season collections from southeastern Utah.

Aster occidentalis (Nutt.) T. & G. Western
Aster. [Tripolium occidentale Nutt.]. Rhizo-
matous or subrhizomatous perennial herbs,
uniformly, softly, and often loosely villous (at
least above); stems 0.9-8.5 dm tall; leaves
1-15 cm long, 1-20 mm wide, entire or
toothed, glabrous or nearly so, ciliate, the
lower ones petiolate, sometimes lacking at
anthesis, rather abruptly smaller and finally
sessile upwards; inflorescence mainly of 1-7
(rarely to 15), corymbosely arranged large
and showy heads; involucres 5-12 mm high,
7-20 mm wide, the bracts imbricate to sub-
equal, slender, green, with pale yellowish to
white or brownish chartaceous bases (at least
the inner), mainly acute, mucronate; rays
20-50, blue to purple, 6-15 mm long; ach-
enes hairy. Meadows and stream sides in
lodgepole pine, cottonwood, willow, aspen,
and spruce-fir communities at 2175 to 3175
m in Carbon, Daggett, Duchesne, Emery,
Garfield, Grand, Piute, Sanpete, Sevier, Sum-
mit, and Wasatch counties; Mackenzie to
Colorado and California; 43 (ix). This species

shares the features of soft loose pubescence
and general aspect with the partially sympa-
tric A. foliaceus. The very slender and
abruptly reduced cauline leaves are diagnos-
tic in most instances.

Aster pansus (Blake) Cronq. Elongate Aster.
Subrhizomatous herbs, villous or villous-hir-
sute with multicellular hairs, the stems 3-12
(or more) dm tall; leaves 1-6 cm long, 2-8
mm wide, entire, antrorsely scaberulous on
both surfaces, sessile, linear to narrowly ob-
long, often spinulose-mucronate, the lower-
most commonly lacking at anthesis; in-
florescence paniculate to secund-paniculate,
narrow; involucres 3.8-4.9 mm high, 4.5-6
mm wide, the bracts strongly imbricate,
green tipped, scaberulous dorsally and cil-
iate; rays mainly 15-25, white, 3-8 mm long;
achenes hairy. Drainages, meadows, seeps,
and hanging gardens at 1220 to 1890 m in
Daggett, Grand, San Juan, and Uintah coun-
ties; British Columbia to Montana, south to
Colorado and Nebraska; 6 (iv). This species
forms the basis for inclusion in previous bo-
tanical works of the name A. ericoides L. in
the Utah flora. It is closely allied to A. fal-
catus, but differs in the smaller heads, taller
stature, and eastern distribution.

Aster pauciflorus Nutt. Alkali Aster. [A.
thermalis Jones, type from Monroe Hot
Springs]. Subrhizomatous perennial herbs,
glabrous below, stipitate-glandular above and
in inflorescence; stems 2-7.5 dm tall; leaves
1.1-12.5 cm long, 1-4 mm wide, entire,
acicular to lance-linear or linear, glaucous,
glabrous, all sessile or the lowermost petio-
late, reduced upwards; heads few to several
in corymbose inflorescences; involucres 4.3-7
mm long, 7-10 mm wide, the bracts imbri-
cate to subequal, glandular dorsally, green
throughout, narrow and acute; rays mainly
20-35, blue to purple, 5-12 mm long; ach-
enes hairy. Hot springs, stream terraces, and
salt grass meadows, often in saline or alkaline
substrates at 1300 to 2135 m in Box Elder,
Duchesne, Emery, Juab, Kane, Millard, San-
pete, Sevier, Salt Lake, and Utah counties;
Saskatchewan to Nevada, Arizona, and Mexi-
co. This distinctive glandular aster has been
collected in full anthesis on 27 April growing
in hot water at Monroe Hot Springs in Sevier
County. It continues to flower into October;
21 (vi).

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Welsh: Utah Flora: Compositae (Asteraceae)

223

Aster perelegans Nels. & Macbr. Nuttall
Aster. [EucepJialus elegans Nutt.; A. elegans
(Nutt.) T. & G., not Willd.]. Subrhizomatous
perennial herb, puberulent to glabrate (some.-
times glandular); stems 3-7 dm tall; leaves
1.3-6.5 cm long, 3-14 mm wide, entire, ob-
long to oblong-lanceolate or elliptic,
scabrous, firm, sessile, the lowermost reduced
in size; heads 3-16, in corymbose in-
florescences; involucres 7-10 mm high, 7-12
mm wide, the bracts chartaceous, imbricate,
with prominent midvein, and acute to obtuse
apex, the margins hyaline and ciliate, more
or less puberulent dorsally; rays 5-16, dark
purple, 7-13 mm long; achenes hairy. Sage-
brush, mountain brush, Douglas fir, aspen,
and limber pine communities at 1725 to 3050
m in Carbon, Duchesne, Juab, Millard, Salt
Lake, Wasatch, and Weber counties; Oregon
to Montana, south to Nevada; 21 (iv).

Aster scopulorum Gray Crag Aster. [Chry-
sopsis alpina Nutt., not A. alpinus L.]. Pe-
rennial subrhizomatous herbs with a woody
caudex, villous on stems and peduncles; stems
4-12 cm tall; leaves 5-12 (15) mm long, 1-3
mm wide, overlapping, elliptic to oblong or
linear, firm, scabrous or puberulent, often
with some villous hairs above, spinulose-
mucronate; heads solitary, pedunculate; in-
volucre 7-11 mm high, 8-12 mm wide, the
bracts imbricate, sparingly villous-hirsute and
glandular, with a prominent midvein in the
lower half, greenish, with chartaceous border
and hyaline margins, acute; rays mainly 8-15,
blue or purplish, 6-15 mm long; achenes
hairy. Sagebrush community at 2440 to 2745
m in Box Elder County; Oregon to Montana,
south to California and Nevada; 8 (0).

Aster spinosus Benth. Mexican Devilweed.
Suffrutescent, rushlike plants from a deeply
placed rhizome, glabrous; stems 6-12 (or
more) dm tall, with axillary or supraxillary
thorns to 1.5 cm long; leaves 2-4 cm long,
2-5 mm wide, firm, entire to toothed, re-
duced above to scales; heads solitary at ends
of branches, or some axillary; involucres 4-6
mm high, 6-8 mm wide, the bracts imbricate,
slender, acute to acuminate, green, with
prominent scarious margin; rays 15-30,
white, very short; achenes glabrous. Riparian
communities at below 1130 m in Garfield
and probably Washington counties; Califor-
nia to Texas, south to Central America; 1 (0).

The plant was collected at the mouth of Ti-
caboo Canyon, along the Colorado River in
Glen Canyon (Lindsay 20, 1958 UT), at a site
now inundated by Lake Powell. This is one
of a series of extirpations related to construc-
tion of Glen Canyon Dam. The plant should
be sought in the St. George vicinity.

Aster wasatchensis (Jones) Blake Mark-
agunt Aster. Subrhizomatous perennial,
glandular-puberulent; stems 3.5-6.5 dm tall;
leaves 1.8-8.5 cm long, 6-24 mm wide, en-
tire, lanceolate to oblong, or oblanceolate,
glandular-puberulent to glabrous, firm, more
or less glaucous, the lowermost often smaller
and commonly lacking at an thesis; heads sev-
eral to numerous, more or less corymbosely
arranged; involucres 8-11.5 mm long, 10-20
mm wide, the bracts herbaceous throughout
or the inner with scarious bases, glandular
dorsally, abruptly acute to attenuate, api-
cally; rays 15-25, white or pink, 10-20 mm
long; achenes hairy. Pinyon-juniper, aspen,
limber pine, and spruce-fir communities at
1890 to 3050 m in Garfield, Iron, Millard,
and Piute counties; endemic. This remarkable
aster is unique in Utah in having foliaceous
or subfoliaceous glandular involucral bracts
and glaucous leaves; 14 (vi).

Atrichoseris Gray

Annual scapose herbs, with milky juice,
from taproots; leaves all basal, sinuate-den-
tate, often spotted; heads on slender pe-
duncles, few to numerous, corymbosely ar-
ranged; involucre of about 12-15 subequal
but biseriate, lance-linear scarious-margined
bracts and some shorter outer bracts; recep-
tacle naked; corollas all raylike, perfect,
white; pappus lacking; achenes oblong, with
corky-thickened ribs.

Atrichoseris platyphylla Gray Tobacco-
weed; Gravel Ghost. Plants 3-10 dm tall (or
more), from slender taproots; leaves 1.2-10.5
cm long, 0.5-6 cm wide, obovate to broadly
spatulate, tapering abruptly to a broad pet-
iole, sinuate-dentate, the teeth mucronate-
cuspidate, glabrous, often mottled, more or
less glaucous; involucres 6-8 mm high, 12-16
mm wide, the outer bracts ovate-lanceolate,
hyaline, more or less scurfy, the inner ones
lance-acuminate, with broad hyaline margins;

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Great Basin Naturalist

Vol. 43, No. 2

corollas white, 8-20 mm long; achenes white,
with corky ridges. Joshua tree, ambrosia,
yucca, cholla communities at 670 to 750 m in
Washington County; California and Arizona;

4(i).

Baccharis L.

Dioecious shrubs; leaves alternate, entire
or toothed; heads discoid, many flowered, the

corollas white, turbinate, borne in corymbose
or paniculate clusters; involucres imbricate,
the bracts chartaceous, whitish; pistillate
heads with tubular-filiform obscurely toothed
or truncate corollas, the pappus of copious
capillary bristles; staminate heads of tubular
5-toothed corollas, the pappus (often scanty)
of usually twisted clavellate scales; recep-
tacle naked; style branches flattened; achenes
subcylindric, 5- to 10-ribbed.

1. Branches fastigiate, deeply sulcate and more or less ridged, the leaves com-
monly deciduous at anthesis; achenes 10-ridged 2

— Branches not especially fastigiate, commonly spreading to ascending; leaves
commonly persistent at flowering time; achenes 5- or 10-ribbed 3

2(1). Main leaves linear; pistillate pappus to 10 mm long or more in fruit ... B. sarothroides

— Main leaves obovate-spatulate; pistillate pappus to 3 mm long in fruit. ..B. sergilloides

3(1). Leaves long-cuneate basally, thickened, entire or few toothed toward apex;
branches often subfastigiate; achenes 10-nerved; plants of Virgin and Colorado
drainages 4

— Leaves not especially long-cuneate basally, commonly thin, entire, or toothed
from below the middle; achenes 5-nerved 5

4(3). Staminate involucres 3.5-5.3 mm long, 3.7-4.8 mm wide; pistillate involucres
7.3-8.5 mm long; pappus 11-13 mm long; plants of Washington and Kane
counties B. emoryi

— Staminate involucres 5.3-6 mm long, 5-10 mm wide; pistillate involucres
6-6.5 mm long; pappus 8-9.5 mm long; plants of Emery, Grand, Garfield,

and San Juan counties B. salicina

5(3). Leaves mainly entire; panicles terminating short lateral branches; plants of

Washington County B. viminea

— Leaves usually serrate; panicles terminating main stems; plants of Washington

and Kane counties B. glutinosa

Baccharis emoryi Gray in Torr. Shrubs,
mainly 1-2 (3) m tall, the branches green to
olive or brownish, ascending, subfastigiate,
more or less glutinous; leaves 1.2-8.5 cm
long, 3-20 mm wide, spatulate-oblanceolate
to elliptic or linear, cuneate to a slender pet-
iole, thick, entire or sparingly and irregularly
toothed, obtuse to acute apically; heads nu-
merous in a conic to pyramidal panicle; pis-
tillate involucres 7.3-8.3 mm high, 4.5-7 mm
wide, the bracts in several series, scarious, of-
ten glutinous, with thickened green or brown
to reddish tips and hyaline margins; stami-
nate involucres 3.7-5.3 mm high, 3.7-4.8 mm
wide; pistillate corollas 4.5-5.5 mm long, the
pappus 11-13 mm long; achenes 10-ribbed.
Stream and canal banks and hanging gar-

dens at 825 to 1220 m in Kane and Washing-
ton counties; Arizona, Texas, and California;
Mexico; 19 (iv).

Baccharis glutinosa Pers. Shrubs, mainly
1-3 m tall, the branches straw colored to
brownish or greenish, ascending-spreading,
not fastigiate, glutinous; leaves 1.2-12.5 (15)
cm long, 4-18 mm wide, elliptic to narrowly
lanceolate, acuminate to attenuate, cuneate
to a short petiole, evenly serrate to entire;
heads numerous in terminal cymose panicles
(less commonly in lateral ones) with pistillate
and staminate heads about the same size; in-
volucres 3.5-4.5 mm high, 4-5.5 (7.5) mm
wide; corollas 2.2-3 mm long, the pistillate
pappus 3.5-4.5 mm long; involucral bracts in
several series, chartaceous, greenish in the

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

225

center, the margins scarious, not glutinous;
achenes 5-ribbed. Stream bars and banks, and
in seeps, at 670 to 1130 m in Kane and Wash-
ington counties; Colorado and Nevada to
Texas and California; South America; 6 (ii).

Baccharis salicina T. & G. Shrubs, mainly
1.5-3 m tall, the branches green to brownish,
subfastigiate, glutinous; leaves 1.4-8 cm long,
4-18 mm wide, elliptic to oblanceolate or
linear, cuneate to a short petiole, thick or
thin, entire or sparingly toothed or lobed
mainly near the apex, acute to rounded api-
cally; heads few to numerous in axillary
and/ or terminal panicles; pistillate involucres
6-6.5 mm high, 4-6 mm wide, the bracts in
several series, scarious, often glutinous, with
thickened greenish to reddish tips and hya-
line margins; staminate involucres 5.3-6 mm
high, 5-10 mm wide; pistillate corollas
2.5-3.5 mm long, the pappus 8-10 mm long;
achenes 10-ribbed. Stream banks and hanging
gardens at 1220 to 1525 m in Emery, Gar-
field, Grand, and San Juan counties; Colo-
rado to Kansas, south to New Mexico and
Texas; 7 (iv). Our material of B. salicina has
long been mistaken for B. emoryi, to which it
is allied. The shorter pistillate involucres and
broader staminate involucres are diagnostic.

Baccharis sarothroides Gray Broom Bac-
charis. Shrubs, mainly 1-3 m tall, the
branches green to brown, fastigiate, glu-
tinous, finally almost leafless; leaves 1-3.5 cm
long, 2-5 mm wide, linear to oblong, entire,
ridged; heads solitary at tips of fastigiate
branches forming a hemispheric panicle; pis-
tillate involucres 6-8 mm high, 5-10 mm
wide, the bracts in several series, cream col-
ored; staminate involucres 3-4 mm high, 4-8
mm wide, the bracts green apically; pappus
6-11 mm long; achenes 10-ribbed. Reported
for Washington County (UT), where presum-
ably it occurs along streams; California to
New Mexico; 0 (0).

Baccharis sergilloides Gray Squaw Water-
weed. Shrubs, mainly 0.3-2 m tall, the
branches green to brown, fastigiate, glu-
tinous, finally almost leafless; leaves 0.5-2.5
cm long, 1-10 mm wide, spatulate to obo-
vate, entire or few toothed, thick; heads nu-
merous, borne in conic to pyramidal panicles;
involucres 2.5-3.5 mm high, 2.5-3.5 mm
wide, the bracts in several series, straw col-
ored, or with thickened brownish centers;
pappus 2.5-3 mm long; achenes 10-ribbed.

Stream bars and banks at 670 to 825 m in
Washington County; California and Arizona;
3(i).

Baccharis viminea DC. Mule-fat. Shrubs,
mainly 2-3 m tall, the branches green to
straw colored or brownish, spreading-ascend-
ing, not fastigiate, glutinous; leaves 0.8-9.5
cm long, 2-9 mm wide, elliptic to lance-el-
liptic or narrowly oblong, attenuate to acute,
cuneate to a short petiole, entire to evenly
serrate; heads few to many in terminal cy-
mose panicles on short lateral branches, with
pistillate and staminate heads about the same
size; involucres 3-5.7 mm high, 6-9 mm
wide; corollas 2.5-3.8 mm long; pistillate
pappus 5-6 mm long; involucral bracts in
several series, chartaceous, commonly with
reddish centers, the margins scarious, not glu-
tinous; achenes 5-ribbed. Stream bars and
banks at 650 to 900 m in Washington Coun-
ty; California and Arizona; 10 (ii).

Bah I A Lag.

Biennial or short-lived perennial herbs
with watery juice, arising from taproots;
stems erect or ascending, puberulent; leaves
alternate, once to twice ternately divided;
heads few to numerous, in corymbose pan-
icles; involucral bracts subequal, in 1 or 2
series, greenish; ray flowers present, yellow,
pistillate, fertile; disk flowers perfect, fertile;
pappus none; style branches flattened; ach
enes 4-angled, 12-nerved.

Bahia dissecta (Gray) Britt. [Amauria dis-
secta Gray]. Biennial or short-lived perennial
herbs, the stems 2-8 dm tall, minutely pu-
berulent; leaves 1-10 cm long, the blade 1- to
3-ternately divided, oval to cordate in out-
line, strigulose; peduncles glandular hairy; in-
volucres hemispheric, 3.4-6 mm high, 8-12
mm wide, the bracts more or less glandular
hairy (or merely villous), greenish, abruptly
contracted to a broadened apex; rays mainly
10-15, yellow, 4.5-9 mm long; achenes
glabrous. Sagebrush, pinyon-juniper, moun-
tain brush, aspen, lodgepole pine, ponderosa
pine, and spruce communities at 1700 to
2930 m in Beaver, Garfield, Grand, Kane, Se-
vier, Uintah, Washington, and Wayne coun-
ties; Nevada to Wyoming, south to Califor-
nia, Arizona, and Mexico; 24 (iii). Those
species treated elsewhere in this work as Pla-
tyschkuhria belong to Bahia in a broad sense

226

Great Basin Naturalist

Vol. 43, No. 2

and are probably best treated in the latter
genus, but their combination is not implied
here.

Baileya Harv. & Gray

Annual, biennial, or perennial herbs from
taproots, with watery juice; stem erect,

white-tomentose; leaves alternate, 1- or 2-
pinnatifid to entire; heads solitary or few in
cymose clusters; involucral bracts subequal,
white-tomentose; receptacle naked; ray flow-
ers persistent, yellow, pistillate, fertile; disk
flowers perfect, fertile; pappus none; style
branches short, truncate; achenes oblong or
clavate, striate.

1. Ray flowers 7 or fewer; plants slender annuals with involucres less than 8 mm

wide B. pauciradiata

— Ray flowers 20 or more; plants annual, biennial, or perennial, with involucres
10-26 mm wide 2

2(1). Rays 11-22 mm long; peduncles (4.5) 12-32 cm long in anthesis; involucres

5.7-7.5 mm high, 13-26 mm wide B. multiradiata

— Rays 8-10 mm long; peduncles 1-8 (11) cm long in anthesis; involucres 3-5.5

mm high, 10-16 mm wide B. pleniradiata

0.8-5 dm tall, white-tomentose; leaves 0.8-12

Baileya multiradiata Harv. & Gray Bien- cm long, the blades 1- to 2-pinnately lobed to

nial or short-lived perennial herbs; stems entire, obovate to linear, white-tomentose;

1.9-5 (5.2) dm tall, white-tomentose; leaves peduncles 1-8 (11) cm long in anthesis,

0.8-10 cm long, the blade 1- to 2-pinnately white-tomentose; involucres 3.5-5.5 mm

lobed to entire, ovate-oval to linear, white- high, 6-13 mm wide, the bracts slender,

tomentose; peduncles (4.5) 13-32 cm long in greenish, white-tomentose; rays 18-58, yel-

anthesis, white-tomentose; involucres 5-7.5 low, 8-10 mm long; achenes glabrous. Creo-

mm high, 13-26 mm wide, the bracts slender, sote bush, blackbrush, shadscale, mesquite,

greenish, white-tomentose; rays 25-40 or sagebrush, and pinyon-juniper communities

more, yellow, 11-22 mm long; achenes at 820 to 1100 m in Washington County; Ne-

glabrous. Creosote bush, Joshua tree, bur- vada and California to Texas; Mexico; 29 (ii).
robush, blackbrush, and sagebrush commu-
nities at 670 to 1320 m in western Kane and

Washington counties; Nevada and California Balsamorhiza Nutt.
south to Mexico; 36 (iii).

Baileya pauciradiata Harv. & Gray An- Perennial scapose or subscapose herbs from
nual herbs; stems mainly 1.5-4 (6) dm tall, taproots, the juice watery; leaves mainly bas-
densely floccose-lanate; leaves 3-10 cm long, al, simple and entire or variously pinnatifid,
the blades entire or the lower irregularly pin- reduced and bractlike upward; heads solitary,
natifid (or bipinnatifid), linear or linear-Ian- or few to several; involucral bracts in several
ceolate, white-tomentose; peduncles 2-5 cm series, imbricate or subequal, herbaceous; re-
long in anthesis, tomentose; involucres 5-6 ceptacle chaffy, convex, the bracts enclosing
mm high, 5-8 mm broad, the bracts slender, the achenes; ray flowers present, pistillate,
greenish, loosely tomentose; rays 5-7, yellow, fertile, usually yellow; disk flowers numerous,
5-8 mm long; achenes glabrous. Reported perfect, fertile, yellow; pappus none; style
from Washington County by Meyer (1976), branches slender; achenes compressed. Note:
where it was collected at Warner Valley The genus is notorious for the lack of genetic
Spring; California, Arizona, and Mexico; 0 (0). barriers to hybridization. Any two taxa can

Baileya pleniradiata Harv. & Gray An- intergrade where they occur together,
nual to short-lived perermial herbs; stems

Leaves sagittate, with entire margins B. sagittate

Leaves pinnatifid or variously cleft 2

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

227

2(1). Leaves mainly 3-6 dm long, with segments mainly 5-12 cm long, these entire

or few lobed or toothed B. macrophylla

— Leaves mainly 1-3 dm long, with segments mostly 1-5 cm long, these entire

or variously lobed or toothed 3

3(2). Involucral bracts abruptly tapering to a long-attenuate apex; stem leaves rela-
tively well developed, pinnatifid or bipinnatifid; reported for northern Utah,
but no specimens have been seen B. hirsuta Nutt.

— Involucral bracts gradually tapering to an attenuate apex; stem leaves lacking

or small and inconspicuous B. hookeri

Balsamorhiza hookeri Nutt. Hooker Bal- ^^^ ^^^^t^ lance-linear, evenly tapering to the
samroot. Perennial scapose herbs from a ^P^^ °' somewhat enlarged at the base, long-
thick taproot, mainly 0.9-4.5 (5.2) dm tall; ^^1^^*^' glandular to tomentose dorsally; rays
leaves 6-30 cm long, (0.3) 1.5-11 cm wide, "^^'^^ly ^^-l^' Y^^^^w, 16-40 mm long; ach-
pinnatifid or bipinnatifid, the segments to 5.5 ^"^^ glabrous. Phases of this taxon are known
cm long; peduncles naked or with a few in- *« ^^^ mtermediates with B. sagittata, and
conspicuous, linear, entire or pinnatifid Presumably with B. macrophyllum. Ours are
bracts near the base; heads solitary; in- separable into two modestly distinctive
volucres 13-24 mm high, 21-47 mm wide, varieties.

Involucres densely villous-tomentose dorsally; plants of Daggett, Duchesne,

and Uintah counties B. hookeri var. neglecta

Involucres glandular to glabrous dorsally; plants of broad distribution, occa-
sionally of Daggett and Duchesne counties B. hookeri var. hispidula

Var. hispidula (Sharp) Cronq. [B. hispidula
Sharp]. This is the common phase of the spe-
cies in Utah, and it has been confused with B.
hirsuta Nutt., with which it is compared in
the key. Bunchgrass, sagebrush, mountain
brush, juniper, pinyon-juniper, and salt desert
shrub communities at 1240 to 2745 m in Bea-
ver, Box Elder, Daggett, Duchesne, Juab, Salt
Lake, Tooele, Utah, Wasatch, and Washing-
ton counties; Nevada, Idaho, and Wyoming
(?); 31 (ii).

Var. neglecta (Sharp) Cronq. [B. hirsuta
var. neglecta Sharp] .Salt desert shrub, sage-
brush, pinyon-juniper, and ponderosa pine
communities at 1640 to 2625 m in Daggett,
Duchesne, and Uintah counties; Nevada,
Idaho, and Wyoming (?); 18 (ii). Plants of this
variety form hybrids with B. sagittata.

Balsamorhiza macrophylla Nutt. Cutleaf
Balsamroot. Perermial scapose herbs from a
thick taproot, mainly 3-7 dm tall; leaves
15-60 cm long, 3.7-25 cm wide, pinnatifid,
the segments entire, few toothed or lobed, up
to 12.5 cm long; peduncles sparingly long
shaggy-villous, naked, or with one to few re-

duced leaves near the base; heads solitary; in-
volucres 23-35 mm high, 30-60 mm wide,
the bracts lance-linear, attenuate, long-
ciliate, glandular and more or less long-vil-
lous dorsally; rays 9-14, yellow, 30-55 mm
long; achenes glabrous. Mountain brush and
sagebrush or bunchgrass communities at 1525
to 2290 m in Box Elder, Cache, Salt Lake,
Summit, Utah, and Weber counties; Idaho to
Montana and Wyoming; 7 (0).

Balsamorhiza sagittata (Pursh) Nutt. Ar-
rowleaf Balsamroot. [Bupthalmium sagitta-
tum Pursh]. Perennial scapose herbs, from
thick taproot, mainly 1.5-8 dm tall; leaves
(including long slender petioles) 5-45 cm
long, 1.5-15 cm wide, sagittate, entire, or the
cauline ones from near the summit to near
the middle of the subscapose stem and linear
to elliptic; peduncles villous-tomentose;
heads solitary (or with additional reduced
ones); involucre 15-30 mm long, 20-50 mm
wide, the bracts lance-linear, attenuate, vil-
lous-tomentose; rays 8-25, yellow, 25-60 mm
long; achenes glabrous. Sagebrush, mountain
brush, pinyon-juniper, ponderosa pine.

228

Great Basin Naturalist

Vol. 43, No. 2

Douglas fir, aspen, and fir communities at
1340 to 3020 m in Beaver, Box Elder, Cache,
Davis, Garfield, Iron, Juab, Kane, Millard,
San Juan, Salt Lake, Sanpete, Sevier, Summit,
Tooele, Uintah, Utah, and Washington coun-
ties; British Columbia to Montana and South
Dakota, south to California, Nevada, and
Colorado; 43 (vii).

Bellis L.

Scapose perennial herbs, with fibrous roots
and short stolons, the juice watery; stems
leafless, simple; leaves all basal, simple, pet-
iolate, toothed to entire; heads solitary; in-
volucral bracts in 2 subequal series, her-
baceous; receptacle conic to hemispheric,
naked; rays white, pink, or purple, numerous,
pistillate; disk flowers numerous, perfect, yel-
low; pappus lacking; style branches flattened;
achenes flattened, usually 2-nerved,
pubescent.

Bellis perennis L. European Daisy. Plants
0.2-2 dm tall; leaves all basal, with short to
long petioles, the blades 0.7-3 (4) cm long,
5-25 mm wide, obovate to oval or orbicular,
dentate to entire, obtuse to rounded or
emarginate apically, pubescent on both sides
with coarse spreading hairs; scapes pubescent
with ascending hairs; heads solitary; in-

volucres 4-7 mm high, 9-15 mm wide, the
bracts ovate to broadly lanceolate, rounded
to obtuse apically, sparsely hairy dorsally, of-
ten suffused with purple, mostly 8-10 mm
long, 1.5-2.5 mm wide; pappus lacking; ach-
enes flattened. Cultivated ornamental, escap-
ing and persisting in lawns of lower valleys in
Salt Lake and Utah counties; adventive from
Europe; 4 (0).

BiDENS L.

Annual herbs with fibrous roots, or rooting
along the lower stem, the juice watery; stems
decumbent to erect, commonly branched;
leaves opposite, simple or pinnately com-
pound; heads few to several in cymose in-
florescences; involucral bracts in 2 series, the
outer herbaceous, the inner somewhat pet-
aloid and striate; receptacle flat or slightly
convex, chaffy throughout, the chaff similar
to the inner involucral bracts; ray flowers
present, yellow, neutral or pistillate, or lack
ing; disk flowers numerous, perfect, fertile,
yellow; pappus of (1) 2-4 awns or teeth,
these retrorsely barbed, persistent; style
branches flattened; achenes flattened, pu-
bescent, usually 2- to 4-awned.
Sherff, E. E. 1937. The genus Bidens. Field
Mus. Pub. Bot. 16:1-709.

Leaves simple, the middle and upper ones (at least) sessile or subsessile B. cernua

Leaves pinnately compound, with 3-5 leaflets, all petiolate B. frondosa

Bidens cernua L. Bur-marigold. Plants
1-13 dm tall, the stems sparingly sp reading-
hairy to glabrous; leaves simple, 1.5-15 cm
long, 0.5-4 cm wide, narrowly lanceolate to
lance-ovate, coarsely serrate to subentire,
glabrous; heads nodding in age; outer in-
volucral bracts 5-8, green, foliaceous, un-
equal, spreading or reflexed, the inner bracts
erect, mostly 6-15 mm long; rays 6-8, yel-
low, or lacking; achenes mainly 5-7 mm
long, tan, the 2-4 awns retrorsely barbed.
Wet meadows, bogs, stream banks, bars, and
shores, at 1300 to 2380 m in Cache, Garfield,
Juab, Kane, Salt Lake, Sevier, Summit, Uin-
tah, and Tooele coimties; widely distributed
in the Northern Hemisphere; 27 (iii).

Bidens frondosa L. Devil's Beggarticks.
Plants 2-12 dm tall, the stems short-hairy to
glabrous; leaves petiolate, pinnately com-
pound with 3-5 leaflets, these 2-10 cm long,

0.5-3 cm wide, lanceolate, serrate; heads
erect in age; outer involucral bracts 5-8,
green, subfoliaceous, subequal, erect or
spreading, the inner bracts erect, mostly 5-8
mm long; rays usually lacking; achenes 5-9
mm long, dark brown to black, the 2 awns
barbed. Marshes, pond and lake shores, bars,
wet meadows, and irrigation canals at 1190
to 1650 m in Davis, Grand, Salt Lake, and
Utah counties; widespread in North America;
12 (ii). Note: The panboreal weed, Bidens tri-
partita L., might occur in our area. It is dis-
tinguished from B. frondosa in its simple but
trifid leaves, and from B. cernua in its petio-
late trifid leaves.

Brickellia Ell. Nom. Cons.

Perennial herbs, subshrubs or shrubs; leaves
alternate or opposite, simple; heads cam-

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

229

panulate or cylindric, cymose or paniculate,
discoid; flowers all perfect, fertile; involucral
bracts imbricate in several series, striate; re-
ceptacle almost flat, naked; style branches
flattened, with long-papillate appendage;

achenes 10-ribbed; pappus of barbellate,

smooth, or subplumose bristles.

Robinson, B. L. 1917. A monograph of the

genus Brickellia. Mem. Gray. Herb. 1:

1-151.

1. Leaves spinulose-serrate, or spinulose tipped; low rounded shrubs of Washing-
ton and San Juan counties B. atractyloides

— Leaves entire or toothed, not spinulose; herbs, subshrubs, or tall shrubs of vari-
ous distribution 2

2(1). Plants herbaceous; heads reflexed, broadly campanulate; leaves sagittate- to

cordate-ovate, longer than broad B. grandiflora

— Plants, shrubs, or subshrubs; heads narrowly cylindric, or, if campanulate, erect;
leaves ovate to linear, if cordate, about as broad as long or broader 3

3(2). Leaves petiolate, the blades cordate-ovate to ovate or suborbicular, 1-5 cm

broad B. calif ornica

— Leaves sessile or subsessile, linear to narrowly lanceolate, or, if broader, mainly

less than 1 cm broad 4

4(3). Leaves linear to lanceolate or narrowly elliptic; shrubs 6-15 dm tall or more;

flowers 3-5 per head B. longifolia

— Leaves ovate to oval or oblong to linear; shrubs or subshrubs less than 5 dm

tall; flowers many per head 5

5(4). Leaves 5-10 times longer than broad or more, entire or nearly so, sessile; in-
volucres 10-20 mm high B. oblongifolia

— Leaves only somewhat longer than broad, often toothed or lobed, at least some
evidently petiolate; involucres 8-12 mm high B. microphylla

Brickellia atractyloides Gray Shrubs,
much branched, mostly 3-5 dm tall, the
branchlets greenish to straw colored, soon
gray; leaves alternate, short-petiolate, the
blades 0.6-3.2 cm long, 0.3-2.2 cm wide,
lance-ovate to ovate, obtuse to rounded ba-
sally, spinulose-serrate to entire, acuminate
and spinulose tipped apically, thick and
prominently veined, glabrous or minutely
glandular puberulent; heads solitary, termi-
nating the branches; peduncles 1-5.2 cm
long, glandular-puberulent; involucres
10-13.5 mm high, 8-16 mm wide, the outer
bracts ovate-lanceolate, acuminate apically,
many veined; the inner narrower, glandular-
puberulent dorsally; flowers 50-75 or more;
achenes black, 3.8-4.2 mm long, hirtellous on
the ribs. Rock crevices and talus slopes,
creosote bush, blackbrush, and indigo bush
communities at 820 to 1130 m in San Juan
(confluence of San Juan and Glen Canyon
arms of Lake Powell) and Washington coun-
ties; Nevada and Arizona; 9 (i). The type is

from the Colorado River (Utah?), Palmer sn,
1870 (US!).

Brickellia californica Gray [Bulbostylis
californica T.& G.; Coleosanthus californicus
(T. & G.) Kuntze]. Subshrubs, mainly 5-10
dm tall, the branchlets whitish to brownish;
leaves alternate, petiolate, the blades 1.7-5.2
cm long, 1.3-4.5 cm wide, cordate-ovate to
ovate or orbicular, truncate to cordate ba-
sally, crenate-serrate, rounded to obtuse api-
cally, the veins not prominent, glandular-
scabrous; heads clustered in a leafy-bracteate
panicle; sessile or shortly pedunculate; in vol
ucres 5.5-8 mm high, 4-7 mm wide, the out-
er bracts very short, rounded apically, few
veined, the inner long and slender, often suf-
fused with red or purple, glabrous; flowers
8-18; achenes straw colored, 2.5-3.5 mm
long. Canyons and rock outcrops at 825 to
2135 m in Garfield, Kane, San Juan, Utah,
and Washington counties; Colorado to
California and south to Texas and Mexico; 15

230

Great Basin Naturalist

Vol. 43, No. 2

Brickellia grandiflora (Hook.) Nutt. [Eu-
patorium grandiflorum Hook.]. Perennial
herb, from a caudex and taproot, the stems
green to straw colored, 2.5-9.5 dm tall;
leaves alternate, petiolate, the blades sagit-
tate to cordate-ovate, 1.5-9 (11) cm long,
0.6-6.5 cm wide, cordate to truncate basally,
serrate to doubly so, attenuate to acuminate
apically, the veins not prominent, minutely
puberulent or hirtellous; heads several to nu-
merous in short corymbose panicle, com-
monly reflexed; involucres 7-12 mm high,
6-10 mm wide, the outer bracts lance-acumi-
nate, the inner abruptly acuminate, pu-
berulent dorsally; flowers mostly 20-40 (70);
achenes brown to black, 3.5-4.5 mm long,
hirtellous. Pinyon-juniper, mountain brush,
ponderosa pine, aspen, Douglas fir-white fir,
spruce, and bristlecone pine communities at
1640 to 3200 m in Beaver, Duchesne, Gar-
field, Iron, Juab, Kane, Salt Lake, San Juan,
Tooele, Utah, and Washington counties;
Washington east to Missouri, south to Mexi-
co; 34 (vii).

Brickellia hngifolia Wats. [Coleosanthus
longifolia (Wats.) Kuntze]. Shrubs, with stems
and white to tan bark, mainly 10-15 dm tall;
leaves alternate, sessile or subsessile, 1.2-13.5
cm long, 3-8 mm broad, lance-linear to
lance-elliptic, obtuse to acute basally, attenu-
ate apically, the veins not prominent.

glabrous, glandular-resinous; heads numerous
in panicles; involucres 3.4-6.2 mm high,
2.3-4 mm wide, the outer bracts ovate,
acute, the inner, longer and slender, glabrous;
flowers 3-5; achenes 1.8-2.4 mm long,
brown, glabrous. Canyon bottoms, stream
margins, seeps, and hanging gardens at 750 to
1590 m in Emery, Garfield, Grand, Kane, San
Juan, Washington, and Wayne counties; Cali-
fornia, Nevada, Arizona; 23 (viii).

Brickellia microphylla (Nutt.) Gray [Bul-
hostylis microphyllus Nutt.]. Shrubs or sub-
shrubs, with tan to whitish bark, mainly 2-7
dm tall; leaves alternate, shortly petiolate to
subsessile or sessile, 3-14 (20) mm long, 1-9
(12) mm wide, ovate to suborbicular, toothed
to entire, commonly glandular-villous or -his-
pidulose, the veins not especially prominent,
rounded to acute apically; heads solitary or
few at tips of branches, racemosely arranged
in leafy-bracteate panicles; involucres 7-10.3
mm high, 4-8.5 mm wide, the outer bracts
oval to ovate, with thickened glandular tips,
the inner often lacking glands and more or
less 3-lobed or 3-veined; flowers 8-18; ach-
enes 3.5-4.3 mm long, blackish, hirtellous or
glabrous. Two distinctive phases, which have
been treated at specific level, are present in
Utah. There is justification for treating them
at specific rank, but they are similar in vege-
tative features and general aspect.

Flowers 8-11 per head; involucres 7-10 mm long; 4-7.5 mm wide; plants of

the Green, Colorado, and Virgin river systems B. microphylla var. scabra

Flowers (12) 17-18 per head; involucres 8.5-10.3 mm long, 6.5-10 mm wide;
plants of the Great Basin B. microphylla var. watsonii

Var. scabra Gray Blackbrush, rabbitbrush,
sagebrush, shadscale, Grayia, greasewood,
juniper, and pinyon-juniper communities
mainly on sandstone outcrops at 885 to 2170
m in Daggett, Duchesne, Emery, Garfield,
Grand, Kane, San Juan, Uintah, and Wash-
ington counties; Colorado, Nevada, Arizona.
Our material is uniformly hispidulose-glandu-
lar along upper stems at least, and has 8-11
flowers per head; 35 (xi). Note: A peculiar
specimen from San Juan County (Anderson
A-6 BRY) has heads nearly all clustered at
branch tips.

Var. watsonii (Robins.) Welsh comb. nov.
[based on Brickellia watsonii Robins. Mem.
Gray Herb. 1:42. 1917]. Sagebrush, shadscale.

mountain brush, and juniper communities at
1525 to 2440 m in Juab, Millard, Sevier,
Tooele, and Utah counties; Nevada and Cali-
fornia. All modern floras distinguish B. micro-
phylla by its heads "about 22-flowered." Our
material fits well within the concept of B.
watsonii Robins., which has heads "18-flow-
ered." Specimens from the Great Basin of
Utah are uniformly 18-flowered, except in
depauperate heads that vary downward to 12
flowers per head. Stems are villous to glandu-
lar-villous, with the type of B. watsonii Rob-
ins. (Watson 494 US!) at the villous end of a
cline; 7 (ii).

Brickellia oblongifolia Nutt. Subshrubs or
subherbaceous, with green to tan branches.

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

231

mainly 1-5.5 dm tall; leaves alternate, sessile
or nearly so, 0.9-4 cm long, 1-11 (15) mm
wide, elliptic to oblong, or lance-oblong, en-
tire or essentially so, glandular-hispidulous,
the veins not especially prominent, acute to
attenuate or obtuse apically; heads solitary
and terminating branches, or corymbosely ar-
ranged; involucres 10.8-15 mm long, 12-22
mm wide, the bracts all acute to acuminate,
glabrous or glandular to glandular-
puberulent; flowers (11) 26-40 (50); achenes
4.8-5.8 mm long, blackish, hispidulous. Gray-
ia, shadscale, rabbitbrush, blackbnish, desert
almond, juniper, pinyon-juniper, and pon-
derosa pine communities at 1280 to 2500 m
in Beaver, Duchesne, Emery, Garfield, Juab,
Kane, Millard, San Juan, Sevier, Uintah,
Utah, Wasatch, Washington, and Wasatch
coimties; British Columbia to Montana, south
to California, Arizona, and New Mexico. Our
material is assignable to var. linifolia (D.C.

Eaton) Robins. [B. linifolia D.C. Eaton, type
from Jordan Valley, American Fork] which is
distinguished by its achenes being his-
pidulous, not glandular-hispidulous or
glandular. The segregation is tenuous at best;
41 (x).

Calycoseris Gray

Annual subscapose or caulescent herbs,
with milky juice, from taproots, beset with
tacklike stipitate glands above; leaves mostly
basal, pinnately parted; heads solitary or few
on leafy-bracteate peduncles; involucral
bracts in 2 series, herbaceous, the inner with
hyaline margins; receptacle with capillary
bristles; corollas all raylike, yellow or white
tipped; achenes fusiform, 5- or 6-ribbed, ta-
pering to a short beak, this produced apically
into a low denticulate cup; pappus abundant,
white, of barbellate capillary bristles falling
attached.

1. Rays white, with pink or purple dots or streaks dorsally; stipitate glands pale ...

C. wrightii

— Rays yellow; stipitate glands purple C. parryi

Calycoseris parryi Gray Annual herbs,
mainly 0.7-3 dm tall, the stems simple or
with spreading branches; leaves basal and al-
ternate along stem, pinnately parted, the
lobes linear, reduced and entire above,
glabrous except for a few tangled long hairs
on lower surface; peduncles mainly 0.5-4 cm
long, clad with tacklike, long-stipitate,
purple or purplish-black glands; involucres
11-15 mm high, 8-14 mm wide (when
pressed), the bracts linear-subulate to lance-
subulate, more or less stipitate-glandular, at-
tenuate apically; rays yellow, 10-20 (25) mm
long; pappus surpassing the achene. Creosote
bush and Joshua tree communities, reported
for Utah by Munz (1959. Calif. Flora p.
1300); to be expected in Washington County;
California and Arizona. Measurements are
from Arizona and California materials; 0 (0).

Calycoseris wrightii Gray Annual herbs,
mainly 1.4-4 dm tall, the stems commonly
with spreading branches; leaves basal and al-
ternate along the stem, pinnately parted, the
lobes linear, reduced and subentire upward,
glabrous except for a few long tangled hairs
on lower surface; peduncles mainly 0.3-5 cm
long, clad with tacklike long-stipitate pale

glands; involucres 12-17 mm long, 12-20
mm wide, the bracts linear-subulate to lance-
subulate, more or less stipitate-glandular, at-
tenuate apically, rays 10-25 mm long, white,
with pink or purple markings dorsally; pap-
pus shorter than achene. Creosote bush and
Joshua tree communities in Washington
County; California, Nevada, Arizona; 1 (0).

Carduus L.

Biennial or annual herbs with taproots, the
juice watery; stems erect, simple or
branched; leaves alternate simple, pinnatifid
to bipinnatifid or merely pinnately lobed, of-
ten decurrent, spiny; heads solitary or few,
borne in corymbose cymes; involucral bracts
imbricated in several series, spine tipped; re-
ceptacle hemispheric, densely bristly; disk
flowers only present, perfect, red-purple,
with long slender lobes; pappus of barbellate
bristles; style branches connate, shortly hairy
at base of branches; achenes compressed.

Carduus nutans L. Nodding Thistle; Musk
Thistle. Rank biennial or annual herbs, most-
ly 0.6-20 (25) dm tall; stems arachnoid-to-
mentose to glabrate; leaves alternate, decur-
rent, 3-40 cm long, 0.5-20 cm wide (or

232

Great Basin Naturalist

Vol. 43, No. 2

more), lance-linear to elliptic, glabrous, or to-
mentose along veins beneath; heads com-
monly solitary, nodding; involucres 20-30
mm long, 30-80 mm wide, the bracts 2-8
mm wide, ovate-lanceolate to lanceolate,
glabrous or nearly so, spinose tipped, at least
the outermost reflexed near the middle, the
midrib prominent; flowers red-purple; ach-
enes 3.5-4.5 mm long, smooth, marked with
vertical lines, umbonate. Disturbed sites
along roads and in fields and pastureland at
1340 to 2440 m in Daggett, Juab, Salt Lake,
Sanpete, and Utah counties, and probably
universal; introduced Old World plants, now
widely established in the United States; 15

Centaurea L.

Annual, biennial, or perennial herbs with
taproots or rhizomes, the juice watery; stems
erect or ascending; leaves alternate, entire to

pinnatifid; heads solitary, or few to numer-
ous, discoid; involucral bracts imbricate in
several series, spine tipped or some of them
enlarged and with scarious or hyaline erose
to lacerate or pectinate appendages; recep-
tacle bristly; flowers all tubular, perfect, or
the marginal ones sterile and falsely sub-
radiate; purple, blue, yellow, pink, or white;
pappus of bristles, scales, or none; style
branches more or less connate, with a thick-
ened often hairy ring at the base; achenes
obliquely or laterally attached to receptacle.
Note: This is a large genus, mainly of the
Mediterranean region of the Old World, but
with some indigenous to North America, Aus-
tralia, and South America. All of ours are in-
troduced, and the potential for other in-
troductions in this remarkable genus is great.
In Flora Europaea, our species are treated
within three genera: Amberboa (Pers.) Less.
(C. moschata L.), Acroptilon Cass (C. repens
L.), and Centaurea for the others.

1. Involucral bracts definitely spine tipped, at least some with spines 1-20 mm

long 2

— Involucral bracts definitely not spine tipped, or, if shortly spinose as in C.
maculosa and C. scabiosa, the heads 6-25 mm wide .' 5

2(1). Stem definitely winged, the leaf bases decurrent; pappus present (central flow-
ers, at least) 3

— Stems angled, not winged; pappus none 4

3(2). Apical spine of involucral bract 5-9 mm long; plants arachnoid when young;

flowers all with evident pappus C melitensis

— Apical spine of involucral bract 11-20 mm long; plants persistently tomentose;
flowers in center only with a pappus C. solstitialis

4(2). Apical spine of bracts 5-15 mm long or more C. calcitrapa

— Apical spine of bracts 1-4 mm long C. virgata

5(1). Leaves entire or merely toothed, not pinnatifid 6

— Leaves pirmatifid or deeply pinnately lobed 8

6(5). Leaves linear to lance-linear, entire or nearly so, less than 1 cm wide C. cyanus

— Leaves various, but, if as above, plants rhizomatous 7

7(6). Plants rhizomatous; leaves mainly 2-10 mm wide; pappus evident, 6-11 mm

long C. repens

— Plants not rhizomatous; leaves 6-15 mm wide; pappus 2-5 mm long C. jacea

8(5). Leaves merely pinnately lobed; involucral bracts entire or nearly so

C. moschata

— Leaves pinnately divided, the lobes linear to narrowly oblong; involucral
bracts pectinately lobed 9

April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 233

9(8). Involucres 15-25 mm wide; lobes of leaves often again toothed or lobed

C. scabiosa

— Involucres mainly 6-10 mm wide; lobes of leaves usually entire C. maculosa

Centaurea calcitrapa L. Star-thistle. Bien-
nial herbs, from taproots, the stems usually
branched, 1-8 dm tall, arachnoid-villous to
glabrate; leaves 0.5-4.5 cm long, pinnatifid,
the lobes linear to oblong, attenuate, or the
upper ones entire; heads few to numerous;
involucres urn shaped, 10-18 mm high,
mainly 8-12 mm wide, the bracts weakly
spinose-ciliate, with a stout apical spine
mainly 5-30 mm long; flowers few, purple;
pappus none. Roadside weeds, Utah County
(Wadley & Holmgren 381 UT); introduced
from Eurasia; 1 (0).

Centaurea cyanus L. Bachelor's Button;
cornflower. Annual or biennial herbs from
taproots, the stem usually branched, mostly
1-8 (12) dm tall, arachnoid-tomentose; leaves
2-10 (13) cm long, 1-8 mm wide, entire or
some with slender lobes, attenuate; heads few
to numerous; involucres hemispheric, 10-16
mm high, 10-23 mm wide, the bracts with a
tapering pectinate or fringed tip, often
purplish suffused, the central apical tooth not
especially spinose; flowers several, blue,
purple, pink, or white, the marginal ones en-
larged, irregular; pappus 2-3 mm long. Culti-
vated ornamental, now established in dis-
turbed sites in Cache, Salt Lake, Tooele,
Utah, Wasatch, and Washington counties; ad-
ventive from Europe; 6 (0).

Centaurea jacea L. Perennial herbs from
taproots, the stems simple or branched from
the middle, mostly 5-12 dm tall, glabrous or
somewhat arachnoid; leaves entire or toothed
to shallowly lobed, the basal ovate to lanceo-
late, petiolate, becoming smaller upward;
heads few to numerous; involucre 12-18 mm
high, 12-15 mm wide, ovoid, the bracts with
orbicular appendages, scarious, brown, dark-
er in middle, the outer denticulate to pecti-
nate-lacerate, the inner less so and often bi-
fid; flowers purple or white, the outer more
or less radiate; pappus none or very short.
Cultivated ornamental, now established in
Salt Lake County; adventive from Europe; 0
(0). Note: The large headed C. montana L., is
cultivated in Utah. It has wedge-shaped in-
volucral bracts and decurrent large leaves.

Centaurea maculosa Lam. Biennial or
short-lived perennial, the stems simple or
commonly branched above the middle,
mainly 3-10 (15) dm tall, tomentose and
sparingly scabrous-puberulent; leaves 1-9 cm
long, pinnatifid, the lobes linear to lanceolate
or oblong, entire or variously toothed or
lobed, reduced and bracteate in the in-
florescence; heads few to many, hemispheric
to vase shaped; involucres 10-13 mm high,
10-13 mm wide, the bracts with short dark
pectinate tip, the central tooth produced as a
spine to 0.5 mm long; flowers pink or pur-
plish, rarely white, the marginal ones radiate;
pappus to 2 mm long, rarely lacking. Road-
sides in Beaver, Juab, and Tooele counties;
adventive from Europe; 3 (i).

Centaurea melitensis L. Annual or bien-
nial, the stems sparingly branched from
middle or below, 1.5-8 dm tall, winged by
decurrent leaf bases; basal and lower cauline
leaves oblanceolate, toothed to lyrate-pin-
natifid or sinuately lobed, reduced upward,
finally entire; heads solitary, terminating
branches, or 2 or 3 in clusters; involucres
8-15 mm high, 8-12 mm wide, tapering api-
cally, the middle and outer bracts spine
tipped, the spines 5-8 mm long; flowers yel-
low, all alike; pappus 1.5-3 mm long. Adven-
tive Old World species of disturbed sites in
Salt Lake County (Without collector UT); 1
(0).

Centaurea moschata L. [Amberboa mos-
chata (L.) DC.]. Annual herbs; simple or
sparingly branched, mainly 3-7 dm long,
sparingly tomentose; leaves 1-9.5 cm long,
1-3 cm wide, pinnatifid, the lowermost pet-
iolate, becoming sessile upward; heads soli-
tary, on peduncles 8-15 cm long or more; in-
volucres vase shaped, 12-14 mm high, 18-22
mm wide, the bracts oval, with purplish mar-
gins, only the inner with broad, reflexed, en-
tire appendage; flowers pink; pappus shorter
to about equaling the achenes. Cultivated or-
namental, escaping and persisting in Wash-
ington County; adventive from Asia; 1 (0).

Centaurea repens L. Russian Knapweed.
[C. picris Pallas ex Willd.; Acroptilon repens

234

Great Basin Naturalist

Vol. 43, No. 2

(L.) DC.]. Perennial rhizomatous herbs, most-
ly 3-8 dm tall, arachnoid-tomentose to gla-
brate; leaves in a basal rosette and cauline,
the basal leaves often withered by flowering
time, the cauline mainly 1-6 cm long, 2-12
mm wide, entire or serrate; heads few to nu-
merous, terminating branches; involucre
9-15 mm high, 5-12 mm wide, more or less
um shaped, middle and outer bracts broad,
glabrous, with broader rounded, subentire
hyaline tips, the inner bracts narrow, taper-
ing, and with plumose hairy tips; flowers
pink to purplish, all alike; pappus bristle sub-
plumose, 6-11 mm long. Introduced Old
World primary noxious weed, now widely es-
tablished at 1220 to 2380 m in Cache, Dag-
gett, Duchesne, Emery, Garfield, Grand,
Kane, San Juan, Salt Lake, Tooele, Uintah,
and Utah counties; widespread in North
America; adventive from Eurasia; 28 (ii).

Centaurea scabiosa L. Perennial herbs,
mostly 5-15 dm tall, scabrous-puberulent;
leaves 4-20 cm long or more, the lowermost
long-petiolate, once to twice pinnatisect, the
segments linear to oblong, entire or dentate-
serrate to lobed, the upper pinnately divided,
sessile; heads few to several, terminating
branches; involucres 13-20 mm high, 18-25
mm wide, ovoid-globose; bracts ovate,
glabrous or arachnoid, the appendages
triangular-ovate, brown or black, with pale
brown teeth; flowers purple, alike or nearly
so; pappus 4-5 mm long. Cultivated orna-
mental, persisting and escaping, Salt Lake
County and probably elsewhere; adventive
from Europe; 2 (0).

Centaurea solstitialis L. Annual or bien-
nial, grayish tomentose, the stems 1-6 (10)
dm tall, evidently winged; leaves mainly
1-12 (20) cm long, 0.1-3 (5) cm wide, the
basal ones lyrate to pinnatifid, the cauline
ones progressively smaller and entire up-
wards, linear to linear-subulate; heads few to

numerous, terminating branches; involucres
8-15 mm high, 7-15 mm wide, urn shaped,
the middle and outer bracts with central
apical spines 10-20 (30) mm long, the inner
with a small hyaline appendage; flowers yel-
low, all alike; pappus of marginal flowers
none, that of the central ones 3-5 mm long.
Roadsides and abandoned fields at 915 to
1900 m in Wasatch, Washington, and Weber
counties; adventive from Europe; 4 (i).

Centaurea virgata Lam. Perennial, from a
caudex, more or less grayish tomentose, the
stems 4-9 dm tall, branched above; leaves
mainly 0.5-15 cm long, 0.1-6 cm wide, the
basal ones petiolate, once to twice pinnately
divided, the lobes linear, these often again
toothed or lobed; cauline leaves smaller, ses-
sile, and lobed to entire; heads several to nu-
merous, terminating short branches; in-
volucre 7-10 mm high, 3-5 mm wide, the
bracts pale or suffused with red or purple,
with a slender apical spine 1-2 mm long;
flowers pink; pappus about 1.5 mm long.
Roadsides and other disturbed sites in Grand,
Juab, and Utah counties at 1525 to 1830 m;
adventive from Eurasia; 5 (i).

Chaenactis DC.

Annual, biennial, or perennial herbs, from
taproots; leaves alternate or mainly basal,
pinnately dissected to entire; heads solitary
or few to several, borne in corymbose cymes,
discoid, the flowers white, or cream to pink,
all perfect, the marginal ones sometimes en-
larged and raylike; involucral bracts in 1-3
series, herbaceous; receptacle flat, naked;
pappus of 4-20 hyaline scales; style branches
slightly compressed; achenes clavate, terete
or more or less compressed.
Stockwell, p. 1940. A revision of the genus

Chaenactis. Contr. Dudley Herb.

3:89-168.

2(1).

Plants perennial from a simple or branching caudex; 2-9 cm tall; stemless or

with few short internodes; of high elevations C. alpina

Plants annual or biennial, rarely perennial, the caudex seldom developed;
stems mainly 10-30 cm tall, or, if less, plants definitely not perennial; dis-
tribution usually of middle and lower elevations 2

Basal rosette well developed; plants biennial or short-lived perennials; pappus
scales 10-16 C. douglasii

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

235

— Basal rosettes poorly, if at all, developed; plants annual; pappus scales 4 or 5
(rarely 8) 3

3(2). Lower and upper cauline leaves simple, the middle ones few lobed;

Washington County C. fretnontii

— Lower, middle, and upper leaves pinnately divided, or only the uppermost
simple 4

4(3). Heads mostly 15-22 mm high; flowers pink, much surpassing the involucre;

anthers included C. macrantha

— Heads mostly 8-10 mm high; flowers white or cream, only slightly surpassing

the involucre; anthers exserted 5

5(4). Involucral bracts blunt or nearly acute apically; plants widely distributed

C. stevioides

— Involucral bracts long-attenuate and bristle tipped apically; plants of
Washington and Millard counties C. carphoclina

Chaenactis alpina (Gray) Jones Alpine
Dusty-maiden. [C. douglasii var. alpina
Gray]. Perermial, from a simple or branched,
sometimes soboliferous caudex, 3.5-9 cm tall;
stems with few contracted internodes, very
short, or not developed; leaves 1.3-5 cm
long, pinnately divided, the lobes again
toothed or lobed, 1-7 mm long, gray tomen-
tose to glabrate; heads solitary or sometimes
2, the peduncles tomentose or glandular,
0.5-6 cm long; involucres (7.5) 10-13 mm
long, (8) 10-17 mm wide, the bracts often
suffused with purple, glandular or tomentose;
corolla purplish to white, glandular or spar-
ingly tomentose; pappus of 10 oblong-spatu-
late rounded hyaline scales, in 2 series; ach-
enes 6-8 mm long, hairy. Boulder stripes and
talus in alpine tundra or upper montane com-
munities at 2980 to 3965 m in Duchesne, Salt
Lake, Summit, and Utah counties; Oregon to
Montana, California and Colorado. Our ma-
terials are separable into two more or less
distinctive phases; a glandular phase, with
distribution mainly in the Wasatch Moun-
tains, which is var. alpina [including C. ru-
bella Greene; C. alpina var. rubella (Greene)
Stockwell], and a tomentose phase, mainly
from the Uinta Mountains, which might be
assignable to var. leucopsis (Greene) Cock-
erell [C. leucopsis Greene]. More work is nec-
essary, including evaluation of the type speci-
men of var. leucopsis; 10 (i).

Chaenactis carphoclina Gray Annual,
from a taproot, 6-28 (40) cm tall; stems well
developed, more or less flexuous; leaves
0.8-5.6 cm long, mealy-puberulent, 1- to 2-

pinnatifid, the segments linear-filiform, 1-20
mm long; heads few to numerous, on slender
farinose to glandular peduncles 0.4-3 cm
long; involucres 6-9 mm high, 6-15 mm
wide, the bracts lance-attenuate into slender,
bristlelike tips, glandular; flowers white to
cream; pappus of central flowers usually of 4
lance-acuminate scales, those of marginal
flowers sometimes shorter; achenes 3.5-4.5
mm long, hairy. Larrea community at 850 to
1000 m in Washington County; California,
Nevada, Arizona; 10 (0).

Chaenactis douglasii (Hook.) H. & A.
Douglas Dusty-maiden. [Hymenopappus
douglasii Hook.; C. achilleaefolia H. & A.; C.
douglasii var. achilleaefolia (H. & A.) A.
Nels.; C. douglasii var. montana Jones; C.
brachiata Greene, type from Springdale; C.
brachiata var. stansburyi Stockwell, type
from Stansbury Island]. Biennial or short-
lived perennial, from a taproot, seldom with
a caudex, mainly 5-50 (60) cm tall, sparsely
to densely tomentose; stems with few to
many well developed internodes; leaves
0.6-12 (15) cm long, 1-3 pinnatifid, the lobes
1-3 cm long, tomentose to glabrate; heads
solitary or several in a corymbose cyme; in-
volucre 7-16 mm high, 8-25 mm wide, the
bracts glandular to glandular-tomentose, ob-
long to narrowly oblanceolate or linear, blunt
apically; flowers white to pink; pappus of
10-16 scales in 2 series; achenes 6-8 mm
long, hairy. Shadscale, sagebrush, pinyon-
juniper, mountain brush, ponderosa pine,
white fir, Douglas fir, aspen, and limber pine
communities at 1340 to 3050 m in all Utah

236

Great Basin Naturalist

Vol. 43, No. 2

counties; British Columbia to Montana, south
to California, Arizona, and Colorado. It does
not seem reasonable to attempt to segregate
our materials into varieties. The variability
apparently does not demonstrate geographic
correlation; 132 (xx).

Chaenactis fremontii Gray Annual or win-
ter annual, from a taproot, 10-25 (40) cm
tall, glabrate or sparingly tomentose when
young; leaves 0.6-6.5 cm long, the lower and
upper simple, linear, the middle few lobed,
glabrous; heads solitary to several on tomen-
tose to glabrate (glandular?) peduncles 1-5
cm long; involucres 8-10 mm high, 10-12
mm wide, glabrous or tomentose, attenuate
but not caudate; flowers white to pinkish, the
outer ones enlarged; pappus of central flow-
ers of 4 scales; achenes hairy. Creosote bush
and Joshua tree communities at 670 to 885 m
in Washington County; Arizona, Nevada,
California; 2 (0).

Chaenactis macrantha D.C. Eaton Annual
or winter annual, from a taproot, mainly
6-25 cm tall, branching from the base or
simple, floccose-tomentose to glabrate; leaves
0.5-5 cm long, 1- to 2-pinnatifid, the lobes to
1 cm long, broad, floccose to glabrate; heads
solitary to several, on tomentose peduncles
0.5-5 cm long; involucres 12-17 mm high,
8-22 mm wide, the bracts oblong-lanceolate,
rather abruptly short-acuminate, tomentose;
corollas pink to white, all about alike; an-
thers included; pappus of 4 linear-oblong
scales and 2-4 short outer ones or these lack-
ing; achenes hairy. Shadscale, pinyon-juniper,
creosote bush, and blackbrush communities
at 885 to 2135 m in Beaver, Juab, Kane, Mil-
lard, Tooele, and Washington counties; Cali-
fornia, Nevada, Arizona; 17 (iii).

Chaenactis stevioides H. & A. Annual or
winter annual, from a taproot, mainly 4-42
cm tall, branching from the base or simple,
more or less tomentose; leaves 0.3-10 cm
long, 1-2 pinnatifid, the lobes to 2.5 cm long,
linear to oblong, sometimes all or nearly all
simple in depauperate specimens; heads soli-
tary to several on glandular peduncles 0.3-3
cm long; involucres 6-11 mm high, 8-22 mm
wide, the bracts oblong-lanceolate to linear,
acute to shortly acuminate apically, glandu-
lar; corollas white to cream, the outer ones
enlarged; pappus of 4 oblong-lanceolate
scales; achenes hairy. Creosote bush, black-
brush, mat-atriplex, shadscale, indigo bush,

and juniper communities at 915 to 1891 m in
Beaver, Carbon, Duchesne, Emery, Garfield,
Grand, Juab, Kane, San Juan, Washington,
and Weber counties; Wyoming south to Ne-
vada, west to California; 63 (vi).

Chamaechaenactis Rydb.

Perennial scapose herbs from a long-pilose
caudex, clothed with marcescent leaf bases,
and taproot, with watery juice; leaves all bas-
al, petiolate, simple; heads solitary; in-
volucres turbinate, the bracts subequal or the
outer shorter; receptacle naked; rays none;
disk flowers perfect, fertile, cream colored to
pink; pappus of hyaline scales; style branches
flattened, papillate; achenes 4-angled, hairy.

Chamaechaenactis scaposa (Eastw.) Rydb.
[Chaenactis scaposa Eastw.] Plants 2-9 cm
tall, the scapes long-villous; leaves petiolate,
the blades 0.4-1.8 cm long, 3-13 (15) mm
wide, lance-oblong to ovate, to oval or or-
bicular, obtuse to rounded apically, obtuse to
truncate basally, villous beneath, strigose to
strigulous or villous above; heads solitary; in-
volucre 7-17 mm high, 10-23 mm wide, the
bracts oblong or linear-oblong, the outer
densely villous, green or suffused with red-
purple, the margin hyaline; corollas cream to
pink; pappus scales oblanceolate-spatulate,
rounded; achenes black, hirsute-pilose. Shad-
scale, galleta, pygmy sagebrush, mountain
brush, pinyon-juniper, and ponderosa pine
communities at 1580 to 2565 m in Carbon,
Duchesne, Emery, Garfield, Grand (?), and
San Juan counties; Arizona and Colorado; 40
(V).

Chamomilla S.F. Gray

Annual herbs, aromatic in some; leaves al-
ternate, 2- or 3-pinnatifid, with linear fili-
form ultimate segments; heads radiate or dis-
coid, solitary or corymbose; involucral bracts
greenish-chartaceous, the margins hyaline, in
2 or 3 series, subequal to imbricate; recep-
tacle conic, hollow, naked; marginal flowers
pistillate; rays (when present) white, the cen-
tral disk flowers perfect and fertile, the style
branches truncate, tufted-hairy apically; pap-
pus a short crown of minute scales, or vesti-
gial or lacking; achenes subcylindric, the ven-
tral face with 3-5 narrow ribs, the dorsal face
smooth and convex.

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

237

1. Heads radiate; disk corollas 5-lobed; involucre 11-25 mm in diameter C. recutita

— Heads discoid; disk corollas 4-lobed; involucre 4-10 mm in diameter ... C. suaveolens

Chamomilla recutita (L.) Rauschert
Chamomile. [Matricaria chamomilla L.]. An-
nual herbs; stems 0.2-4 (6) dm tall, erect or
ascending, branched above; herbage glabrous
or puberulent; leaves 2-6 cm long; heads soli-
tary or more commonly few to many and
corymbosely arranged; involucres saucer
shaped, 3-4 mm high, 11-25 mm wide, the
bracts subequal, the margins broadly hyaline,
the midstripe greenish to brownish; rays
10-20, white, 4-10 mm long. Moist disturbed
soils at low to moderate elevations in Salt
Lake and Wasatch counties; adventive from
Europe; 2 (0).

Chamomilla suaveolens (Pursh) Rydb.
[Matricaria inatricarioides (Less.) Porter]. An-
nual herbs; stems 0.4-4 dm tall, erect or as-
cending, branched from the base or simple;
herbage glabrous or pubescent; leaves 1-5 (9)
cm long; heads few to many, paniculately ar-
ranged; involucres saucer shaped, 2-6 mm

high, 4-10 mm wide, the bracts subequal to
somewhat imbricate, the margins hyaline, the
midstripe greenish; rays lacking; disk flowers
4-lobed. Disturbed sites at 1310 to 2810 m in
Box Elder, Cache, Carbon, Rich, Salt Lake,
Sevier, Utah, and Weber counties; adventive
from Europe; 16 (0).

Chrysanthemum L.

Perennial herbs from a rhizome or a cau-
dex, with watery juice; stems erect or nearly
SO; leaves alternate, serrate to pinnatifid;
heads solitary or few to numerous in open
corymbose clusters; involucral bracts imbri-
cate, in 2-4 series, greenish or straw colored,
the margins brownish-scarious; receptacle
naked; ray flowers white, numerous, pistil-
late, fertile, or lacking; disk flowers numer-
ous, perfect, fertile, yellow; pappus lacking
or a short crown; style branches flattened;
achenes several nerved, beakless, glabrous.

1. Leaves finely serrate; heads usually numerous, small, commonly rayless..C. balsamita

— Leaves coarsely serrate or pinnatifid; heads larger, fewer, commonly with rays 2

2(1). Heads solitary or few; involucres 7-10 mm high; rays 1-2 cm long; leaves ser-
rate to more or less once pinnatifid C. leucanthemum

— Heads several to numerous; involucres 3-4.5 mm high; rays 2-6 mm long

C. parthenium

Chrysanthemum balsamita L. Costmary.
[Balsamita major Desf.]. Perennial herbs,
from a caudex, commonly 5-10 (12) dm tall;
stems strigose, at least above; leaves petiolate
below, sessile or subsessile above, the blades
0.9-10 (15) cm long, 0.6-5 (8) cm wide, ellip-
tic to oblanceolate, finely serrate, strigose;
heads numerous, corymbose; involucres
3.7-4.6 mm high, 6-8 mm wide, the bracts
oblong, sparingly strigose, the tip hyaline;
ray flowers (when present) 4-6 mm long.
Fields, roadsides, and cemetaries at 1370 to
2135 m Salt Lake, Summit, Tooele, and Utah
counties; escaped from cultivation, now
widely established in the United States; 5 (i).

Chrysanthemum leucanthemum L. Oxeye-
daisy. [Leucanthemum vulgare Lam.] Pe-
rennial rhizomatous or subrhizomatous herbs,
commonly 2-8 (10) dm tall; stems glabrous or

nearly so, mainly simple; leaves petiolate be-
low, becoming smaller and sessile above, the
blades 0.8-5 cm long, oblanceolate to obo-
vate or linear, serrate, crenate, or pinnately
lobed, glabrous or villosulose; heads solitary;
involucres 7-10 mm high, 15-23 mm wide,
the bracts lance-ovate to oblong-linear, with
brown margins, hyaline apically; rays mainly
15-30, white, 10-22 mm long; pappus none.
Roadsides, fields, and other disturbed sites at
1525 to 2135 m in Salt Lake, Utah, Wasatch,
and Weber counties; widespread in North
America; adventive from Eurasia; 6 (0).

Chrysanthemum parthenium (L.) Bemh.
[Matricaria parthenium L.; Leucanthemum
parthenium (L.) Gren. & Godron; Pyrethrum
parthenium (L.) Sm.; Tanacetum parthenium
(L.) Schultz-Bip.]. Perennial herbs with cau-
dex and taproot; commonly 3-9 dm tall;

238 Great Basin Naturalist Vol. 43, No. 2

stems glabrous, or puberulent above; leaves Chrysothamnus Nutt.

petiolate, becoming smaller, but still petio- ^ i -.l i.-. u i .u r i.

r , , 1 1 ? r> ,- o 1 nV. A c Shrubs with white bark, or the suriace ob-

late above, the blades 0.5-8 cm long, 0.6-4.5 J, ^ ^ ^V.- U 1 J 1

.-. , 1 , , 1 1 scured by a tomentum, this orten giandular-

(6) cm wide, pinnatirid or doubly so; heads . \ ,. . ,. ^ ui ^

^ ' ' ^ ■' resinous; leaves alternate, linear to oblong, or

several to numerous, the inflorescence cor- ^^^^^^^^^^ ^^^^^^e, entire; heads white or yel-

ymbose; involucres 3-4.5 mm high, 7-10 mm j^^ ^^^^^^^ ^^^^.^.^^ ^^ contracted to open

wide, the bracts oblong, with a dark center, paniculate inflorescences; flowers perfect,

otherwise scarious except the tip hyaline; £gj.tile; involucral bracts imbricate, more or

rays 10-20, white, 4-8 mm long; pappus a \q^^ keeled, in 4 or 5 vertical or obscure

crown or none. Cultivated ornamental, es- ranks, chartaceous or coriaceous, or the tip

caping and persisting at 1525 to 1950 m in herbaceous; receptacle naked; style branches

Carbon, Salt Lake, Utah, and Weber coun- flattened; achenes slender, flattened, angled,

ties; widely established in the United States; or terete, hairy or glabrous; pappus of nu-

adventive from Europe; 5 (0). merous capillary bristles.

1. Flowers white; leaves terete; plants of western tier of counties (except Iron and

Washington) C. alhidus

— Flowers yellow; leaves various, but, if terete, of Washington County or rarely
elsewhere 2

2(1). Leaves terete, resinous punctate; stems more or less fastigiate; plants of Wash-
ington County C. paniculatus

— Leaves commonly more or less flattened, resinous-punctate or not; stems not
especially fastigiate; plants of broad or other distribution 3

3(2). Stems obscured by a tomentum, this often impregnated with resinous-glandular

material 4

— Stems glabrous or puberulent, the surface readily apparent 5

4(3). Involucral bracts long-attenuate, membranous; inflorescence more or less race-
mose C. parryi

— Involucral bracts obtuse to acute, rarely attenuate, but, if so, chartaceous; in-
florescence cymose C. nauseosus

5(3). Leaves lanceolate to lance-oblong, not contorted; shrubs mainly 6-20 dm tall;

plants of the Uinta and Navajo basins C. linifolius

— Leaves linear, oblong, or lanceolate, but, if lanceolate, twisted and shrubs
mainly less than 6 dm tall; distribution various 6

6(5). Achenes hairy 7

— Achenes lacking hairs, sometimes glandular, or, if sparingly hairy, the in-
volucre over 10 mm long 8

7(6). Involucral bracts acuminate-cuspidate; leaves 1-2 mm wide C greenei

— Involucral bracts acute to obtuse; leaves various C viscidiflorus

8(6). Flowers 10-12 mm long, surpassed by the pappus; plants of Emery, Wayne,

and San Juan counties C. pulchellus

— Flowers 7-9 mm long, surpassing or subequal to the pappus; distribution
various 9

9(8). Involucral bracts strongly ranked; involucres 9.2-13 mm long C. depressus

— Involucral bracts not strongly ranked; involucres 6.2-7.5 mm long C. vaseyi

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

239

Chrysothamnus albidus (Jones) Greene

Alkali Rabbitbrush; White Rabbitbush. [Bige-
lovia alhida Jones]. Shrubs, mainly 5-10 dm
tall, more or less fastigiately branched, white
barked, glabrous, resinous-viscid, aromatic;
leaves 0.5-3.5 cm long, terete, 0.5-1 mm
thick, glandular-punctate, mucronate,
crowded, often with axillary fascicles; heads
clustered at branchlet apices; involucres
6.8-9 mm high, 3-7 mm wide, the bracts ob-
scurely 4- to 5-ranked, the outer ones lance-
ovate, thickened in lower half, abruptly sub-
ulate-attenuate, the inner oblong, acuminate
to acute, the margin hyaline, glandular to to-
mentose; corollas white, 6-7.5 mm long; ach-
enes 4-4.5 mm long, pilose and glandular;
pappus abundant. Local in salt grass, pickle-
weed, and alkali-saccaton communities at
1450 to 1650 m in Beaver, Box Elder, Juab,
Millard, and Tooele counties; California, Ne-
vada; 8 (iii).

Chrysothamnus depressus Nutt. Dwarf
Rabbitbrush. Low, spreading shrubs, the as-
cending to erect, subherbaceous stems 0.6-3
dm tall, white barked, scabrous-puberulent or
glandular-puberulent; leaves 0.4-2 cm long,
1-4 (5) mm wide, flat, narrowly lanceolate to
oblanceolate or spatulate, flat, scabrous-
puberulent, obtuse, rounded or sharply apicu-
late; heads clustered at branch apices; in-
volucres 9.2-13 mm high, 4.5-7 mm wide,
the bracts in 4 or 5 definite vertical ranks,
keeled, lance-attenuate, the subulate tip soft,
the outer more or less herbaceous (sometimes
suffused with purple) and the inner with
broad hyaline margins; corollas yellow, 7.5-9
mm long; achenes (5) 6-7 mm long, glabrous
or sparingly stipitate-glandular; pappus off-
white to brownish, abundant. Sagebrush, salt
desert shrub, juniper, pinyon-juniper, moun-
tain brush, ponderosa pine and alpine fir
communities at 1550 to 2900 m in Carbon,
Duchesne, Emery, Garfield, Iron, Juab, Kane,
Millard, Piute, San Juan, Sanpete, Sevier,
Summit, Uintah, Utah, Wasatch, Washington,
and Wayne counties; Colorado, New Mexico,
Arizona, and Nevada; 34 (iv).

Chrysothamnus greenei (Gray) Greene
Greene Rabbitbrush. Low, ascending to erect
shrubs, with subherbaceous stems from a
woody crown, mainly 1-3.5 dm tall, white-
barked, glabrous; leaves 0.3-3.5 cm long,
0.8-1.2 mm wide, flat, linear, glabrous or

scabrous-ciliate; heads numerous, corym-
bosely clustered at branch tips; involucres
5-7.1 mm high, 2.5-4 mm wide, the bracts
obscurely ranked, the outer ones herbaceous-
thickened near the tip, gradually acuminate-
cuspidate, the inner ones abruptly narrowed,
glabrous or more or less tomentose, narrowly
if at all hyaline-margined; corollas yellow,
3.5-4.8 mm long; achenes 3.3-4 mm long, pi-
lose. Rabbitbrush, black sagebrush, shadscale,
winterfat, sagebrush, and pinyon-juniper
communities at 1280 to 2745 m in Carbon,
Duchesne, Emery, Garfield, Grand, Juab,
Millard, Piute, Tooele, Uintah, Utah, and
Wayne counties; Colorado, New Mexico, Ari-
zona, and Nevada; 53 (vi). This entity forms
intermediates with phases of C. viscidiflorus.

Chrysothamnus Unifolius Greene Spread-
ing Rabbitbrush. Tall shrubs, the branches
erect-ascending, mainly 8-20 (35) dm tall,
white barked, glabrous; leaves 0.9-7.7 cm
long, 1-9 mm wide, flat, plane (not contorted
or rarely somewhat so), thick, oblong to ellip-
tic or narrowly lanceolate, glabrous,
scabrous-ciliate, attenuate to acute; heads nu-
merous, corymbosely arranged at branch tips;
involucres 4.3-7.2 mm long, 1.8-3 mm wide,
the bracts indistinctly ranked, the outer dis-
tinctly herbaceous at tip, the inner often
merely glandular thickened at midrib, all ob-
tuse to rounded, glabrous; corollas yellow,
4.5-5.8 mm long; achenes 2.1-2.8 mm long,
pilose. Stream banks and terraces, irrigation
canals, seeps and springs in riparian commu-
nities at 1130 to 2535 m in Carbon, Daggett,
Duchesne, Emery, Garfield, Grand, Kane,
San Juan, Sanpete, Sevier, Uintah, and
Wayne counties; Montana to Arizona and
New Mexico; 54 (xvii).

Chrysothamnus nauseosus (Pallas) Britt.
Rubber Rabbitbrush. Low to tall shrubs, the
branches erect-ascending, mainly 2-20 (30)
dm tall, the bark obscured by a tomentum,
this often resinous-glandular impregnated;
leaves 0.6-7 (10) cm long, 0.5-5 (10) mm
wide, 1- to 3-nerved, tomentose to glabrate
or glabrous, subcylindric to flat, if the latter
then commonly plane, linear to narrowly ob-
long, acute to apiculate apically; heads nu-
merous, in terminal paniculate cymes; in-
volucres (6) 6.5-11.5 (13) mm high, 1.5-7.2
mm wide, the bracts obscurely to definitely
ranked, the outer ones sparingly tomentose to

240 Great Basin Naturalist Vol. 43, No. 2

glabrous, the inner commonly glabrous, ob- complex in Utah is represented by a diverse

long, chartaceous to more or less herbaceous- assemblage of more or less geographically

thickened, obtuse to acute or shortly acumi- and ecologically segregated races, which are

nate apically; corollas yellow or yellow-or- placed in some 14 varieties. The following

ange, 6-10.3 (12) mm long; achenes 2.5-5.5 arbitrary key will serve to identify most

mm long, glabrous or hairy. The nauseosus specimens.

1. Shrubs usually 3 dm tall or lower; plants local endemics in Piute, Sanpete,

Sevier, Carbon, Emery, Daggett, and Duchesne counties 2

— Shrubs usually more than 3 dm tall, seldom lower, but then of different distri-
bution 4

2(1). Involucres glabrous, 8.5-9.5 mm high; plants of Emery, Carbon, Wasatch and

Duchesne counties C. nauseosus var. psilocarpus

— Involucres tomentose or glabrous, 10-12 (13.5) mm high; plants of Sanpete,
Sevier, and Piute counties 3

3(2). Involucres glabrous; corollas 7.8-9 mm long; plants local on Arapien shale in

Sanpete and Sevier counties C. nauseosus var. iridis

— Involucres tomentose; corollas 10-12 mm high; plants local in Piute County ....

C. nauseosus var. glareosus

4(1). Achenes and ovaries glabrous 5

— Achenes and ovaries pilose 8

5(4). Flowers 5-8 mm long; involucres 7-8.5 (9) mm long, 1.5-3 mm wide (when

pressed) C. nauseosus var. abbreviata

— Flowers 8.3-10 mm long; involucres 9-11 mm long, 3.7-7 mm wide (when
pressed) 6

6(5). Involucres subcylindric; plants of dunes and deep sands of western Utah and in

the Uinta Basin C. nauseosus var. turbinatus

— Involucres tapering to the base; plants of south central and southeastern Utah 7

7(6). Achenes 5-5.5 mm long; plants low, commonly less than 5 dm tall; known

from San Juan and Emery counties C. nauseosus var. bigelovii

— Achenes 2.5-4 mm long; plants taller, commonly over 5 dm tall; known from
Kane County C. nauseosus var. nitidus

8(4). Involucres over 10 mm long; corollas 9.5-10.5 mm long 9

— Involucres 6.5-8.6 (9.5) mm long (to 11 mm long in var. junceus); corollas
5-8.6 (10) mm long 10

9(8). Involucres cylindric, the bracts neither strongly keeled nor ranked; plants of

dime areas in western and northeastern Utah C. nauseosus var. turbinatus

— Involucres tapering, clavate, the bracts strongly keeled and aligned; plants of
Kane County C. nauseosus var. arenarius

10(8). Leaves 3-5 (10) mm wide; plants of central to north central Utah

C. nauseosus var. salicifolius

— Leaves 0.5-3 mm wide; plants of various distribution 11

11(10). Corolla lobes commonly long-pilose (glabrate in age); leaves often deciduous

by anthesis; plants of southeastern Utah C. nauseosus var. junceus

— Corolla lobes glabrous; leaves present or absent at anthesis; distribution vari-
ous 12

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

241

12(11). Corolla lobes 0.4-0.9 mm long C. nauseosus var. gnaphaloides

— Corolla lobes 1-2 mm long 13

13(12). Leaves (1) 3- to 5-nerved, commonly 1-3 mm wide C. nauseosus var. glabratus

— Leaves 1-nerved, commonly 0.5-1.5 mm wide 14

14(13). Leaves and/ or stems usually grayish or whitish tomentose or green, not

especially yellow-green; involucres more or less tomentose

C. nauseosus var. albicaulis

— Leaves and/or stems usually yellowish-green, the tomentum commonly
resinous-matted; involucres glabrous C. nauseosus var. consimilis

Var. abbreviatus (Jones) Welsh comb. nov.
[based on: Bigelovia leiosperma var. abbre-
viata Jones Proc. Calif. Acad. II, 5: 693.
1895; type from Clear Creek Canyon, Sevier
County; C. nauseosus var. leiosperma (Gray)
Hall; C. nauseosus ssp. leiospermus (Gray) H.
& C; Bigelovia leiosperma Gray, type from
St. George.] Blackbrush, Grayia, shadscale,
black sagebrush, Vanclevea, pinyon-juniper,
and ponderosa pine communities at 1070 to
2745 m in Emery, Garfield, Grand, Kane,
Millard, Piute, Sevier and Washington coun-
ties; Nevada, California; 15 (v). The materials
from Emery and Grand counties have leaves
that are very slender and subterete. The con-
dition is presumably derived from in-
trogression with var. bigelovii.

Var. albicaulis (Nutt.) Rydb. [C. nauseosus
var. albicaulis Nutt.]. Saltgrass, sagebrush,
pinyon-juniper, and ponderosa pine commu-
nities at 1310 to 2290 m in Box Elder, Cache,
Carbon, Juab, Kane, Millard, Morgan, Salt
Lake, San Juan, Uintah, Utah, Wasatch, and
Weber counties; Oregon to Wyoming, south
to California, Nevada, and New Mexico; 24
(i). This taxon forms intermediates with var.
glabratus. In low elevation phases of saline
substrates the stems are white-pannose.

Var. arenarius (L.C. Anderson) Welsh
comb. nov. [based on: C. nauseosus ssp. are-
narius L.C. Anderson Phytologia 38: 311.
1978.]. Sagebrush, juniper, and pinyon-juni-
per communities at 1675 to 1830 m in Kane
County; Arizona; 3 (i). This is a plant of deep
sandy alluvium.

Var. bigelovii (Gray) Hall [C. nauseosus
ssp. bigelovii (Gray) H. & C.; Linosyris
(Chrysothamnus) bigelovii Gray]. Grayia and
pinyon-juniper communities 1460 to 1950 m
in Emery and San Juan (Lavender Mesa)
counties; Arizona, Colorado, New Mexico; 2
(i). More collections of this entity are
required.

Var. consimilis (Greene) Hall [C. nau-
seosus ssp. consimilis (Greene) H. & C; C.
consimilis Greene]. Saline meadows, riparian
zones, and terraces in saltgrass-alkali sacca-
ton, shadscale, sagebrush, rabbitbrush, moun-
tain brush, pinyon-juniper, and ponderosa
pine communities at 1280 to 3000 m in all
Utah counties except Grand and San Juan;
Oregon to Wyoming, south to California,
Arizona and New Mexico; 100 (xxv). This is
the common narrow-leaved phase with cone-
shaped panicles. They occur frequently in sa-
line moist sites, such as the travertine mounds
at Monroe Hot Springs.

Var. glabratus (Gray) Cronq. [Bigelovia
graveolens var. glabrata Gray; C. nauseosus
ssp. graveolens (Gray) Piper; C. nauseosus
var. graveolens (Gray) Hall]. Desert willow-
baccharis, willow-cottonwood, greasewood-
tamarix, sagebrush, shadscale, mountain
brush, and ponderosa pine communities at
750 to 2475 m in Summit, Wasatch, Utah,
Sanpete, Sevier, Piute, Iron, and Washington
counties, and in all counties east of those;
Idaho to North Dakota, south to Arizona, and
New Mexico; 88 (xxiii).

Var. glareosus (Jones) Welsh stat. nov.
[based on: Bigelovia glareosa Jones Zoe 2:
247. 1891, type from Marysvale; C. nau-
seosus ssp. glareosa (Jones) H. & C.]. The
type specimen is lost, and the ultimate dis-
position of this taxon is uncertain; it should
be sought in the canyon north of Marysvale,
on Tertiary igneous substrates; endemic; 0
(0).

Var. gnaphaloides (Greene) Hall [C. speci-
osus var. gnaphaloides Greene; C. nauseosus
ssp. hololeucus (Gray) H. & C, in part].
Shadscale, pigmy sagebrush, rabbitbrush,
sagebrush, and pinyon-juniper communities
at 1070 to 2380 m; known in all Utah coun-
ties except Box Elder, Daggett, Duchesne,
Kane, Morgan, Rich, Summit, and Wayne,

242

Great Basin Naturalist

Vol. 43, No. 2

and likely in them also; California, Nevada,
and Arizona (?); 73 (vii). This taxon is a near
ally of ssp. hololeucus (Gray) H. & C., and
should that taxon be placed within a quad-
rinomial, then the var. gnaphaloides would
be placed within it. However, no such com-
bination is implied or proposed herein.

Var. iridis (L.C. Anderson) Welsh stat.
nov. [based on: C. nauseosus ssp. iridis L.C.
Anderson Great Basin Nat. 41:311. 1981,
type from Rainbow Hills, Sevier County].
Rabbitbrush-sagebrush community on an in-
cipient seep in Arapien shale at ca 1980 m in
Sevier Co.; endemic; 2 (i).

Var. junceus (Greene) Hall [C. nauseosus
ssp. junceus (Greene) H. & C; Bigelovia jun-
cea Greene]. Blackbrush, shadscale, rabbit-
brush, matchweed, and pinyon-juniper com-
munities at 1220 to 1800 m in Emery,
Garfield, Grand, Kane, San Juan and Wayne
counties; Arizona; 18 (iv). The nonglandular,
clear straw-colored, long involucres with
bracts usually aligned are distinctive of this
variety.

Var. nitidus (L.C. Anderson) Welsh stat.
nov. [based on: C. nauseosus ssp. nitidus L.C.
Anderson Phytologia 38: 313. 1978]. Van-
clevea-ephedra community at about 1250 m
in Kane County; Arizona; 1 (0). This variety
has the general aspect of vars. bigelovii and
abbreviata. It is a taller plant than either, and
differs otherwise as set forth in the key.

Var. psilocarpus Blake [C. nauseosus ssp.
psilocarpus (Blake) L.C. Anderson]. Sage-
brush and salina wildrye communities at
1925 to 2290 m in Carbon, Duchesne,
Emery, and Wasatch counties; endemic; 5
(0). These peculiar low shrubs occasionally
produce taller intermediates with var. gla-
bratus (qv.)

Var. salicifolius (Rydb.) Hall [C. salici-
folius Rydb., type from Strawberry Valley; C.
nauseosus ssp. salicifolius (Rydb.) H. & C.].
Sagebrush, pinyon-juniper, mountain brush,
and aspen communities at 1310 to 2870 m in
Box Elder, Carbon, Duchesne, Emery, Juab,
Salt Lake, Sanpete, Sevier, Summit, Tooele,
Utah, and Wasatch counties; endemic; 19
(iii). This entity forms intermediates with var.
glabratus, and might represent nothing more
than a broad-leaved extension of that taxon.

Var. turbinatus (Jones) Blake [Bigelovia
turbinata Jones, type from Kane County; C.

nauseosus ssp. turbinatus (Jones) H. & C.].
Rabbitbrush, saltbush, ephedra, juniper, and
greasewood communities at 1370 to 1710 m
in Beaver, Iron, Juab, Kane, Millard, and Uin-
tah counties; Nevada(?); 10 (iii). Both
glabrous and pilose achenes occur in this dis-
tinctive taxon. It shares the feature of villous
corolla lobes with the sand-loving var. jun-
ceus of the Navajo Basin. The Uintah Basin
materials differ in the more keeled and atten-
uate involucres and flowers that are more ex-
serted from the involucre.

Chrysothamnus paniculatus (Gray)
Greene [Bigelovia paniculata Gray]. Tall
shrubs, the branches subfastigiate, mainly
6-20 dm tall, the bark green, becoming tan
to gray in age, resinous-punctate; leaves
0.4-3 cm long, about 0.5 mm wide, linear-
filiform, terete, mucronate apically; heads
numerous, in usually conic panicles; in-
volucres 4.8-6.5 mm high, 2-3 mm wide, the
bracts indistinctly ranked, chartaceous-in-
durate, scarcely if at all glandular, thickened
at midrib, obtuse, glandular; corollas yellow,
5.5-6 mm long; achenes 1.8-3.4 mm long, pi-
lose. Roadsides, stream banks, terraces, and
slopes in creosote bush, Joshua tree, and bac-
charis communities at 670 to 1220 m in
Washington County; Nevada, Arizona, Cali-
fornia; 9 (iii). The plants begin to flower in
October and continue into November.

Chrysothamnus parryi (Gray) Greene
Low to moderate shrubs, the branches not es-
pecially fastigiate, mainly 2-6 dm tall, the
bark pannose-tomentose or the tomentum
glandular-resinous; leaves 0.6-6 (8) cm long,
1-2 mm wide, 1- to 3-nerved, green, viscid or
sometimes tomentulose, flat, usually plane,
linear to narrowly oblong; heads several to
many, the inflorescences tending to be elon-
gate and subracemose; involucres 9-14.5 mm
high, 4-8 mm wide, the bracts obscurely to
definitely ranked, puberulent to glabrous, the
outer usually with elongate herbaceous tips,
the inner chartaceous, with glandular-thick-
ened midrib, abruptly to gradually acumi-
nate-attenuate or attenuate; corollas yellow
or creamy yellow, 8-10 mm long; achenes
3.3-7.5 mm long, pilose. Plants of the parryi
complex form hybrid derivatives with phases
of C. nauseosus, and with other named segre-
gates within the complex. Except for varie-
ties parryi and nevadensis, only arbitrary

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

243

segregation appears possible. Thus, the con- low seems to best reflect the nature of C. par-
servative treatment as outlined be- ryi in Utah.

1. Flowers usually more than 10 per head C. parryi var. parryi

— Flowers commonly 5-9 per head 2

2(1). Involucral bracts mainly 24-28; plants of southwestern Utah

C. parryi var. nevadensis

— Involucral bracts mainly 12-22; plants of south central, central, and north-
eastern Utah C. parryi var. attenuatus

Var. attenuatus (Jones) Kittell in Tidestr.
& Kittell [Bigelovia howardii var. attenitata
Jones, type from near Marysvale; C. parryi
ssp. attenuatus (Jones) H. & C; C. affinis. A.
Nels.; C. parryi ssp. affinis (A. Nels.) L.C.
Anderson; Linosyris howardii Parry in Gray;
C. parryi ssp. howardii (Parry) H. & C; C.
parryi var. howardii (Parry) Kittell in Tidestr.
& Kittell]. Meadows, sagebrush, juniper, pin-
yon-juniper, mountain brush, ponderosa pine,
and aspen communities at 1740 to 2930 m in
Beaver, Carbon, Daggett, Duchesne, Gar-
field, Grand, Iron, Kane, Piute, Sanpete, Se-
vier, Uintah, Utah, Wasatch, and Wayne
counties; Wyoming and Nebraska, south to
Arizona and New Mexico; 55 (xv). The how-
ardii phase differs supposedly in the brac-
teate leaves overtopping the inflorescence
and in the pale colored flowers; both charac-
ters fail as diagnostic features.

Var. nevadensis (Gray) Kittell in Tidestr.
& Kittell [Linosyris howardii var. nevadensis
Gray; C. parryi ssp. nevadensis (Gray) H. &
C.]. Sagebrush, juniper, pinyon-juniper,
mountain brush, and ponderosa pine commu-
nities at 1830 to 2565 m in Beaver, Iron, Mil-
lard, and Washington counties; Arizona; 10
(ii). The var. nevadensis differs only in degree
from var. attenuatus, with which it is con-
tiguous, if not partially sympatric, to the east.
Should the two be combined, then the cor-
rect name will be var. nevadensis, since that
name has priority in rank. Plants with leaves
overtopping the inflorescence occur; techni-
cally they would key to the howardii phase of
var. attenuatus.

Var. parryi [Linosyris parryi Gray]. Pon-
derosa pine and spruce-fir communities at
2075 to 2625 m in Beaver, Emery (?), Gar-
field, Kane, Millard, and Washington coun-
ties; Wyoming, Colorado, New Mexico, and
Nevada; 9 (ii).

Chrysothamnus pulchellus (Gray) Greene

Low to moderately tall shrubs, the branches
not fastigiate, mainly 5-10 dm tall, the bark
white, becoming tan or brown in age,
glabrous or puberulent above; leaves 0.4-3
cm long, 1-2 mm wide, linear to narrowly
oblanceolate, glabrous or puberulent, flat or
revolute, mucronate; heads few to many, in
corymbose panicles; involucres 11.5-15 mm
high, 4.5-6 mm wide, the bracts distinctly
aligned, more or less herbaceous toward the
apex, glandular, attenuate to sharply acute;
corollas yellow, 9-10 (14) mm long; achenes
3.8-4.5 mm long, sparingly hirsute and
glandular. Shadscale, blackbrush, ephedra,
pinyon-juniper, and ponderosa pine commu-
nities at 1370 to 2350 m in Emery, Wayne,
and San Juan counties; Arizona to Kansas,
south to Mexico; 4 (i). Our material belongs
to var. baileyi (Woot. & Standi.) Blake [ssp.
baileyi (Woot. & Standi.) H. & C.].

Chrysothamnus vaseyi (Gray) Greene
[Bigelovia vaseyi Gray]. Low shrubs, mainly
1-3 dm tall, the branches not especially fasti-
giate, the bark green, becoming whitish tan
or finally gray in age, puberulent; leaves
0.3-3.7 cm long, 0.8-3 mm wide, linear to
oblong or narrowly oblanceolate, glabrous or
glandular, flat, plane, mucronate; heads nu-
merous in compact terminal cymes; in-
volucres 6.2-7.5 mm high, 3-6 mm wide, the
bracts more or less aligned, commonly her-
baceous or thickened near the apex, glandu-
lar, obtuse, the margins fimbriate-hyaline; co-
rolla yellow, 4.8-7 mm long; achenes 2.6-4
mm long, glabrous. Meadows, sagebrush, rab-
bitbrush, juniper, mountain brush, and pon-
derosa pine communities at 1675 to 2900 m
in Beaver, Carbon, Emery, Garfield, Juab,
Kane, Iron, Piute, San Juan, Sanpete, Sevier,
and Utah counties; Nevada, Wyoming, Colo-
rado, New Mexico; 21 (ii).

244

Great Basin Naturalist

Vol. 43, No. 2

Chrysothamnus viscidiflorus (Hook.)
Nutt. Low to moderate shrubs, mainly 2-10
dm tall, the branches fastigiate or not, the
bark green to tan or white, finally gray in
age, glabrous or puberulent; leaves 0.3-4.5
(6) cm long, 0.5-4 (10) mm wide, 1- to 5-
nerved, linear to oblong, elliptic or oblan-
ceolate, often twisted, mucronate; heads nu-
merous, in compact to open terminal cymes;
involucres 5-7.5 mm high, 2-4 mm wide, the
bracts not well aligned, commonly her-
baceous or thickened near the apex (at least
the outer), glandular or puberulent, obtuse,
or abruptly acute, the margin narrow, hya-
line; corollas yellow, 3.8-6 mm long; achenes
3-4 mm long, pilose. The viscidiflorus com-
plex is separable into two groups on the basis
of pubescence of upper stems or the lack of
pubescence. The segregation is not complete,
because pubescence or its absence is not an
absolute criterion. There is a cline in the
amount of pubescence from abundant to few
(or none), and the adoption of a position that

one hair equals pubescence and, therefore
one part of the complex and not the other,
will lead to absurdity. Within the hairy phase
of the complex are two more or less dis-
tinctive but largely sympatric varieties. The
"glabrous" portion of the species is more dif-
ficult to separate into its constituent entities.
Anderson (Great Basin Nat. 40: 117-20,
1980) reviewed this portion of the complex;
concluding that there are three taxa involved,
i.e. ssp. axillaris, ssp. viscidiflorus var. vis-
cidiflorus, and ssp. viscidiflorus var. steno-
phyllus. Only arbitrary separation of the
three is possible, and segregation of the ax-
illaris phase is problematical. In my view it is
not practical to attempt recognition of more
than two taxa, i.e. var. stenophyllus (includ-
ing axillaris) and var. viscidiflorus. They are
aH recognized herein at varietal level, but
probably would best fit within an expanded
ssp. viscidiflorus as varieties (a course not in-
tended or implied herein). The following key
will allow for identification of most
specimens.

1. Stems (at least above) and/or leaves puberulent to hispidulous 2

— Stems and leaves glabrous, or the leaves ciliate, or rarely with a few short hairs

on stems or with glandular excrescences in the inflorescence 3

2(1). Leaves 0.5-2 mm wide; stems finely puberulent above

C. viscidiflorus var. puberulus

— Leaves 2-5 mm wide; stems hispidulous-puberulent above

C. viscidiflorus var. lanceolatus

3(1). Leaves 0.5-1.5 mm wide; plants mainly 2-3 dm tall

C. viscidiflorus var. stenophyllus

— Leaves mainly 1-4 mm wide (or more); plants mainly 3-10 dm tall

C. viscidiflorus var. viscidiflorus

Var. lanceolatus (Nutt.) Greene [C. lan-
ceolatus Nutt.; C. viscidiflorus ssp. lanceo-
latus (Nutt.) H. & C.]. Sagebrush, pinyon-
juniper, mountain brush, aspen, Douglas fir,
lodgepole pine, spruce-fir, and alpine mead-
ow communities at 1375 to 3200 m in all
Utah counties except Kane and Washington,
and likely there also; British Columbia to
South Dakota, and south to California, Ne-
vada, Arizona, and New Mexico; 112 (xii).

Var. puberulus (D.C. Eaton) Jepson [Lino-
syris viscidiflora var. puberula D.C. Eaton;
C. viscidiflorus ssp. puberulus (D.C. Eaton)
H. & C.]. Rabbitbrush, black sagebrush,

shadscale, sagebrush, pinyon-juniper and pon-
derosa pine communities at 1460 to 2200 m
in the western tier of counties, east to Piute,
Sevier, Emery, Carbon, Utah, and Salt Lake
counties; Oregon and Idaho south to Califor-
nia, Nevada, and Arizona; 44 (vii).

Var. stenophyllus (Gray) Hall [Bigelovia
douglasii var. stenophylla Gray; C. vis-
cidiflorus ssp. stenophylla (Gray) H. & C; C.
axillaris Keck; C. viscidiflorus ssp. axillaris
(Keck) L.C. Anderson]. Ephedra, blackbrush,
rabbitbrush, sagebrush, galleta, shadscale, and
pinyon-juniper communities at 1280 to 2075
m in all Utah counties except Piute, Sevier,

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

245

Sanpete, Carbon, Duchesne, Wasatch, Utah, Cirsium Mill,

Salt Lake, Davis, Weber, Morgan, Summit,

and Cache; Oregon to Wyoming and south to Annual, biennial, or perennial, caulescent

California,, Nevada, Arizona, and Colorado; ""' acaulescent, spiny herbs from taproots,

04 / -jx with caudices or rhizomes in some, the juice

Var.' viscidifhrus [Crinitaria viscidiflora ^^^^^y' ^^^^^^ ^^^^^ ^"^ ^^"^i"^' alternate;
Hook.; C. viscidiflorus var. pumilus authors, ^^^^' fo^it^^X *« ^^^^^f 1' involucral bracts in
not (Nutt.) Jeps. (?). Rabbitbrush, shadscale, ^^^^^^^ ^^"^^' subequal to imbricate, some or
sagebmsh, pinyon-juniper, mountain brush, "^^f ^^ ^^^"^ ^P^^e tipped; receptacle dense-
white fir, ponderosa pine, and aspen commu- ^X bristly; corollas all discoid, pink, purple,
nities at 1460 to 2900 m in all or nearly all '^^' ""' ^^^^"^y ^^^^e, perfect or imperfect;
Utah counties; Washington to Nebraska, P^PP"^ ^^ plumose bristles (or those of the
south to California, Nevada, Arizona, and outermost flowers merely barbellate); style
Colorado; 100 (xx). The var. viscidiflorus ^^^^ ^ thickened minutely hairy ring below
forms intermediates with all other taxa in the ^^^ ^^^^^X ^o""^*^ 1°^^^' ^^^enes glabrous,
species, and with C.greend also. ?^"^^!^ °' ^-angled, 4- to many-nerved.

Note: This is a particularly complex genus

taxonomically, with both introduced and in-

CiCHORiuM L. digenous species. The indigenous members

Perennial herbs, with milky juice, from ^'^ especially difficult, due in part to hy-
taproots; leaves alternate, toothed to pinnati- bridization, mainly within species groups,
fid; heads sessile or subsessile, numerous, ^^ following treatment is tentative, but rep-
borne in clusters at nodes of a spicate, simple, resents an attempt to categorize the variation
or branched inflorescence; involucral bracts P^'^^^"* ^^ Utah plants and to provide a le-
biseriate, the outer shorter; corollas all ray- gitimate name for each. Several taxa pre-
Uke, perfect; pappus of 2 or 3 series of scales, ^^o^^^X reported from the state are excluded,
sometimes minute; achenes angular or some- ""' ^^y ^'^ ^'^^^^"^ ^^t^*'^ *e constituent
what compressed, glabrous. t^^a. All involucral measurements are in

Cichorium intybus L. Chickory. Plants pressed condition!

3-10 dm tall or more, hirsute or glabrous; ^o^^^, R. J. and C. Frankton. 1963a. Cyto-

lower leaves petiolate, the blades 6-20 cm taxonomic notes on some Cirsium spe-

long, 1-5 (7) cm wide, sinuate-dentate to run- ^^^j ?f J^^ ;^^^tf " V^l*. '

■I ■ \c^ u ■ u A nad. J. Bot. 41: 1553-1567.

cmate-pmnatirid, becommg smaller and ses- ^r:r>r^,\ * i r- ^- c ^- ■

., ^ J £• 11 1 ..• 11 1963b. A clarification of Ctrstum

sue upward, some finally subentire; heads r ;• j /-• • j j- o
, 11 ir, ^ r ■ foliosum and Cirstum drummondii. Ca-
large and showy, 1-3 per node of in- ^^^^ . g^^ 42: 451-461.
florescence; flowers pure blue, rarely white; ^ggg Cytotaxonomy of Cirsium hook-
involucre 9-15 mm high, the outer bracts erianum and related species. Canad. J.
chartaceous at base, herbaceous apically; gQj 43. 597_613

achenes 2-3 mm long. Roadsides and dis- I973. xhe Cirsium arizonicum com-

turbed sites at 1340 to 2135 m in Duchesne, plex of the southwestern United States.

Iron, Kane, Salt Lake, Tooele, and Utah Canad. J. Bot. 52: 543-551.

counties; widespread in North America; na- Petrak, F. 1917. Die nordamerikanischen

tive of Eurasia; 8 (i). The herb C. endiva L. is Arten der Gattung Cirsium. Beih. Bot.

grown in Utah; the extent is not known. Centralbl. (Abt. 2), 35: 223-567.

1. Flowers mainly imperfect; heads unisexual; plants perennial, from rhizomes;

introduced weed of consequence C. arvense

— Flowers perfect; plants biennial or perennial, seldom if ever with rhizomes 2

2(1). Leaves roughly hispid above, green; stems conspicuously winged decurrent;

plants biennial, introduced C. vulgare

— Leaves villous, floccose, arachnoid, tomentose, or glabrous, white to gray or
green; stems not winged-decurrent, except in some species; plants indigenous
biennials or perennials 3

246 Great Basin Naturalist Vol. 43, No. 2

3(2). Basal rosettes to 10 dm across, the mature leaves commonly 10-30 cm wide,
green, glabrate or glabrous on both sides; heads small, with long, tapering, re-
curved spines; plants of hanging gardens in southeastern Utah, rarely below
them C. rydbergii

— Basal rosettes rarely to 5 dm across, the mature leaves usually less than 8 cm
wide, floccose, tomentose, arachnoid, or glabrous on one or both sides; plants
seldom of hanging gardens in southeastern Utah 4

4(3). Bracts, at least the innermost, conspicuously dilated (but not lacerate), or
definitely tan to silvery in appearance, contrasting with the overall aspect of
the bracts; plants commonly of meadows C. scariosum

— Bracts all spinose, or the innermost occasionally twisted to contorted at the
tips, but not especially dilated or conspicuously different in color or texture
from the overall aspect of bracts (see C. centaureae); plants of various habitats 5

5(4). Involucral bracts (at least the outer) pinnately spinose; plants green, with yel-
lowish spines, of high elevations in the Wasatch, Tushar, and Uinta mountains
C. eatonii

— Involucral bracts not, or rarely, pinnately spinose (except in C. clavatum, C.
scopuhrum, and C. ownbeyi); plants of low to high elevations, but, if pinnately
spinose, of other distribution or of low elevations 6

6(5). Heads 1.8-2.7 cm high, and about as wide; inner bracts with coarsely lacerate

margins; plants of lower middle elevation meadows C. centaureae

— Heads 1.5-3 cm high, 1.5-4.5 (6) cm wide; inner bracts not lacerate; leaves
thinly textured, finely to coarsely spined, definitely tomentose or glabrous;
plants of various distribution 7

7(6). Herbage definitely white- to gray-tomentose (or rarely green); involucres 1.5-2
cm high, 1.5-2.5 cm wide; known from white shale outcrops in the
Uinta Basin C. bamebyi

— Herbage green, or white- to gray-tomentose; involucres mainly longer and
broader, but if not, then of different distribution 8

8(7). Stems definitely winged-decurrent; heads mainly 1.3-2 cm high, 1.2-3.2 cm
wide; herbage white- to gray-tomentose; plants of Sanpete and Washington
counties 9

— Stems not winged, or if so, the herbage green and glabrous or nearly so, or the
heads commonly larger; plants of various distribution 10

9(8). Leaves of upper stem merely spinose-toothed, tapering from base to apex;

plants of Washington County only C. virginensis

— Leaves of upper stem definitely lobed, the lobes spinose-toothed, with parallel
sides from base to near apex; plants not of Washington County C. subniveum

10(8). Herbage glabrous or glabrate, green 11

^ Herbage tomentose, floccose-tomentose, gray or white, or only the upper leaf

surfaces green 16

11(10). Flowers bright red or carmine; corolla lobes 15-18 mm long; spines of middle

involucral bracts 7-11 mm long or more; plants of San Juan County

C. rothrockii

— Flowers pink, pink-purple, or white; corolla lobes less than 15 mm long; spines

of middle involucral bracts 1-6 mm long; plants of various distribution 12

April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 247

12(11). Outer bracts not pinnately spinose; mainly low elevation plants, usually in

gypsiferous soils, in the Navajo Basin C. calcareum

— Outer bracts more or less pinnately spinose; plants of the Navajo and Uinta ba-
sins 13

13(12). Stems strongly winged almost or quite the length of upper intemodes; main
upper leaves tripinnatifid; plants of lower elevations in northern Uintah and
Daggett counties C. ownbeyi

— Stems not winged, or rarely some intemodes with incipient wings; main upper
leaves pinnatifid to bipinnatifid; plants of moderate to high elevations in the
southern end of the Uinta Basin and southward 14

14(13). Involucral bracts ciliate with long yellowish or brownish multicellular hairs;
spines of bracts 6-15 mm long or more; plants of the east Tavaputs Plateau
and La Sal Mountains C. scopulorum

— Involucral bracts more or less ciliate with whitish hairs or a tomentum; spines
of bracts mainly 3-7 mm long; plants from the Tavaputs Plateau and south
westward 15

15(14). Involucral bracts scabrous dorsally, at least the innermost; herbage not at all

tomentose; plants of the Henry Mountains C. calcareum

— Involucral bracts not scabrous dorsally; herbage more or less tomentose; plants

not of the Henry Mountains C. clavatum

16(11). Heads campanulate, mainly 3.5-6.5 cm wide at anthesis, or, if narrower,

bracts commonly glandular-thickened dorsally 17

— Heads turbinate to subcylindric, mainly 2-3.5 cm wide at anthesis; involucral
bracts seldom glandular-thickened dorsally 18

17(16). Involucral bracts appearing brown to gray-brown, the spines arising from the
body of the bract, not from spreading long-attenuate herbaceous terminal por-
tions; bracts of inflorescence usually prominent; plants of broad distribution

C. undulatum

— Involucral bracts appearing green or fresh green or at least herbaceous, the
spines arising from the apex of spreading long-attenuate terminal portions;

bracts of inflorescence much reduced; plants of various distribution

C. neomexicanum

18(16). Corollas bright red or carmine; plants from Garfield and Iron counties south-
ward C. arizonicum

— Corollas pale pink, pink, rose-purple, or white; plants from Garfield and Iron
counties northward 19

19(18). Involucral bracts (at least the inner) tapering, wedge-shaped, definitely

scabrous roughened on dorsal surface, often suffused with red or purple

C. calcareum

— Involucral bracts smooth dorsally, seldom only somewhat scabrous, not con-
spicuously tapering, and seldom conspicuously suffused with red or purple

C. wheeleri

Cirsium arizonicum (Gray) Petrak Ari- lobed or toothed, the main spines 1-6 mm

zona Thistle. [Cnicus arizonicus Gray]. Bien- long, white to grayish tomentose below, more

nial or short-lived perennial herbs from a or less tomentose and greenish to green

taproot, the caudex sometimes developed; above; stems 4-7.5 dm tall, more or less floc-

leaves of basal rosettes 7-36 cm long, bi- cose-tomentose; cauline leaves 3-35 cm long,

pinnately lobed or parted, the lobes again 1-8 cm wide, with lobing and vesture similar

248

Great Basin Naturalist

Vol. 43, No. 2

to the basal, reduced and less deeply lobed
upward; involucres 22-30 mm high, 20-50
mm wide, subcylindric to turbinate, the
bracts tomentose at margins, and over back,
smooth and often shiny medially, rarely
glandular-thickened, the apical portions, es-

pecially of the inner definitely scabrous;
spines yellowish, 3-10 (15) mm long; corollas
crimson to carmine, 25-34 mm long, the tube
8-13 mm long, throat 1.5-11 mm long, the
lobes 10-19 mm long. Two more or less dis-
tinctive but intergrading phases are present.

Heads subcylindric to turbinate; spines 3-10 mm long; plants mainly of the

Colorado drainage system (also in western Garfield, and in Iron counties)

C. arizonicum var. arizonicum

Heads turbinate to broadly so; spines 3-15 mm long or more; plants mainly of
the Great Basin and Virgin drainages (also in eastern Iron and western Garfield
counties) C. arizonicum var. nidulum

Var. arizonicum Salt desert shrub, pinyon-
juniper, ponderosa pine, spruce-fir, and hang-
ing garden communities at 1220 to 3050 m in
Garfield, Iron, Kane, Piute, San Juan, and
Washington counties; Arizona; 26 (iv).

Var. nidulum (Jones) Welsh comb, no v.
[based on: Cnicus nidulus Jones Proc. Calif.
Acad. II. 5: 705. 1895]. Pinyon-juniper,
mountain brush, aspen, ponderosa pine,
Douglas fir, white fir, and spruce-fir commu-
nities at 1890 to 3200 m in Beaver, Garfield,
Iron, Kane, San Juan and Washington coun-
ties; Arizona, Nevada; 37 (iii). Relationships
apparently lie with C. rothrockii, C. cal-
careum, and, to a lesser extent, with C.
wheeleri.

Cirsium arvense (L.) Scop. Creeping or
Canada Thistle. [Serratula arvensis L.]. Pe-
rennial rhizomatous herbs, the stems mostly
5-10 dm tall, glabrous or sparingly tomen-
tose; leaves 3-15 cm long, 1-6 cm broad,
deeply pinnatifid or lobed to merely toothed,
glabrous to tomentose above and beneath;
heads several to many, mainly unisexual; in-
volucres 10-20 (25) mm high, 10-25 mm
wide, the bracts lance-ovate, at least the out-
er ones and often all of them spine tipped, to-
mentose to glabrous; corollas pink-purple to
white; pappus of pistillate heads longer than
the corollas, that of staminate heads shorter
than the corollas; achenes 3-5 mm long.
Roadsides, fields, and other disturbed sites,
but also invading native plant communities,
at 1280 to 2535 m, probably in all Utah
counties; widespread in North America; ad-
ventive from Eurasia; 42 (iii). We have two
phases of creeping thistle in Utah; the one
with merely toothed (unlobed) leaves is var.
mite Wimm. & Grab., and the common one

with deeply lobed leaves is var. horridum
Wimm. & Grab. This common weed and the
bull thistle are our only two introduced
thistles in the genus Cirsium, which makes up
a huge assemblage in the Old World. We can
expect more introductions.

Cirsium barnebyi Welsh & Neese in
Welsh Barneby Thistle. Perennial herbs from
a caudex and taproot, the caudex clothed
with black marcescent leaf bases; leaves of
basal rosettes 11-25 cm long, bipinnately
lobed or parted, the lobes again lobed or
toothed, the main spines 3-5 mm long, whit-
ish- to grayish tomentose on both sides; stems
3-5 dm tall, whitish tomentose (rarely green);
cauline leaves 2-30 cm long, 1-8 cm wide,
with lobing and vesture similar to the basal,
reduced and less deeply lobed upwards; in-
volucres 15-22 mm high, 20-30 mm wide,
turbinate, the bracts glabrate or sparingly
arachnoid on margins, glutinous dorsal ridge
inconspicuous, smooth medially, the apical
portions of the inner often contorted, not
scabrous dorsally; spines 2-7 mm long, flat-
tened apically, more or less spreading; co-
rollas bluish pink. Sagebrush, juniper, cryp-
tantha, ephedra, wildrye, and rabbitbrush
communities at 1525 to 2257 m in Uintah
County; endemic; 7 (iii). The Barneby thistle
is apparently related to the undulatum
complex.

Cirsium calcareum (Jones) Woot. &
Standi. Cainville Thistle. [Cnicus calcareus
Jones, type from Cainville]. Perennial herbs
from a caudex and taproot, the caudex with
brownish black to castaneous marcescent leaf
bases; leaves of basal rosettes 6-35 cm long,
pinnatifid to bipinnatifid, glabrous and green
or tomentose on one or both surfaces, the

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

249

main spines 3-8 mm long; stems mainly 2-5
dm tall, glabrous or more or less floccose-to-
mentose, winged-decurrent or not; cauline
leaves 3-28 cm long, 0.8-7 cm wide, bipin-
natifid, with lobing and vesture like the basal,
reduced upward, the main spines 3-8 mm
long; involucres 19-34 mm long, 15-45 mm
wide, the bracts ovate-lanceolate to linear,
more or less tomentose at the margins,
smooth and often shiny medially, the dorsal

ridge glandular-thickened or not, the apical
portions of at least the inner scabrous; spines
straw colored, 1.5-6 mm long; corollas pink
to blue-pink. The calcareum complex is a
portion of the arizonicum group of thistles,
and has long been misinterpreted. There are
three more or less confluent varieties present
in Utah. Specimens collected are few, espe-
cially in the critical southeastern portion of
the state. More work is indicated.

2(1).

Herbage permanently tomentose, the leaves grayish tomentose beneath

C. calcareum var. pulchellum

Herbage green, the leaves rarely sparingly tomentose along the midveins be-
neath 2

Leaves definitely decurrent, the stems winged 2-6 cm below leaf base; plants

of San Juan and Wayne counties C. calcareum var. calcareum

Leaves not or scarcely decurrent; plants of other distribution

C. calcareum var. bipinnatum

Var. bipinnatum (Eastw.) Welsh stat. nov.
[based on: Cnicus drummondii var. bipinna-
tum Eastw. Zoe 4: 8. 1893]. Aspen, Douglas
fir, and riparian communities at 1130 to 3150
m in Garfield, Kane, and San Juan counties;
Colorado, New Mexico, and Arizona; 8 (i).

Var. calcareum [Cirsium pulchellum var.
glabrescens Petrak type from Elk Mountains,
San Juan County]. Riparian communities at
1460 to 2200 m in Carbon, San Juan, and
Wayne counties; endemic (?); 4 (i). Jones
(Proc. Calif. Acad. II., 5: 704. 1895) cited two
collections with the protologue; i.e., Jones
5695bh from Bromide Pass in the Henry
Mountains and Jones 5696 from Cainville.
His description best fits the Cainville mate-
rials, and that collection is here selected as
lectotype. The material from Bromide Pass
seems best to fit var. bipinnatum (q.v.). A pe-
culiar plant with thin leaves that are glabrous
on both sides and subentire is known from
Cedar Canyon (Atwood and Higgins 5918
BRY). How it fits into the scheme of Utah
thistles is not known, but the plant appears to
be intermediate between this and some other
thistles. The status of the Cainville thistle, as
strictly interpreted, beyond Utah is unknown;
it seems likely that it does not occur outside
the state.

Var. pulchellum (Greene) Welsh comb,
nov. (based on: Carduus pulchellus Greene ex
Rydb. Fl. Colorado 400, 401. 1906). Rabbit-

brush, sagebrush, tamarix, rabbitbrush, pin-
yon-juniper, and aspen communities at 1340
to 2745 m in Carbon, Emery, Garfield,
Grand, Kane, San Juan, Uintah, Utah, and
Wayne counties; Colorado, New Mexico, Ari-
zona; 41 (vii). Both winged and wingless
stems are present within our material. There
are plants from the San Rafael Swell with
winged stems and they are similar to C. och-
rocentrum Gray of New Mexico, but they ap-
pear to be transitional in every way with the
wingless plants. And it seems probable that
they are not conspecific with that plant as it
occurs beyond Utah. Possibly they do war-
rant taxonomic recognition. Further collec-
tions are necessary.

Cirsium centaureae (Rydb.) K. Schum.
[Carduus centaureae Rydb.]. Fringed Thistle.
Perennial herbs from a simple caudex and
taproot, the caudex with chestnut leaf bases;
leaves of basal rosette 2-28 cm long, 1-8 cm
wide, pinnatifid, the lobes often again
toothed, tomentose below, thinly tomentose
to glabrous above, the main spines 1-5 mm
long; stems 3-12 dm tall, not succulent,
arachnoid or glabrous; cauline leaves with
lobing and vesture like the basal, the spines
3-8 mm long; involucres 18-27 mm high,
and about as wide, the outer bracts lance-
ovate, the inner with coarsely lacerate mar-
gins, usually dilated in the upper half, tomen-
tose to glabrous on the margins, the dorsal

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Great Basin Naturalist

Vol. 43, No. 2

ridge not well developed, the longest spines
2-5 mm long, straw colored; flowers white to
pink or purple. Montane communities at
3355 m in San Juan Co.; Wyoming and Colo-
rado; 2 (0).

Cirsium clavatum (Jones) Petrak Fish
Lake Thistle. [Cnicus clavatus Jones, type
from Fish Lake]. Perennial or biennial herbs
from a taproot, and often with a caudex, the
caudex clothed with marcescent chestnut-
brown leaf bases; leaves of basal rosettes
2.5-22 cm long, bipinnately parted to merely
toothed, green on both sides or more or less
tomentose below, the main spines 1-6 mm
long; stems 3-10 dm tall, glabrous or thinly
tomentose; cauline leaves 3-26 cm long,
0.5-7 cm wide, with lobing and vesture like
the basal, reduced and less lobed above; in-
volucres 18-23 (32) mm high, 22-30 (55) mm
wide, the bracts more or less villous-tomen-
tose on margins, the outer ones usually pin-
nately spiny, smooth medially, the dorsal
ridge not especially glandular, apical por-
tions of the inner ones often scabrous, some-
times slightly dilated-erose; spines yellowish,
3-8 (18) mm long; corollas white or less com-
monly pink. Sagebrush, meadow, aspen,
Douglas fir, and spruce-fir communities at
2135 to 3200 m in Beaver, Carbon, Emery,
Garfield, Grand, Kane, Piute, Sanpete, Se-
vier, Uintah, and Wayne counties; endemic;
27 (viii). The Fish Lake thistle is apparently
related to the allopatric C. eatonii. It is more
or less transitional to C wheeleri, and prob-
ably other taxa, especially those with

scabrous inner bracts. Rarely some have de-
current leaf bases, and when the pinnately
spinose bracts are poorly developed, this
thistle approaches C. calcareum. Moore and
Frankton (1965) proposed that C clavatum
was a hybrid between C. eatonii and C. cen-
taureae. However, despite its possible origin
from hybridization, the taxon seems to be or-
ganized on about the same basis as other
thistles. Further, its distribution is distinct
from that of the putative parents. There does
not seem to be justification for recognition of
this entity as a hybrid.

Cirsium eatonii (Gray) Robins. Eaton
Thistle. [Carduus eatonii Gray]. Perennial
herbs from a simple or rarely branched cau-
dex and taproot, the caudex clothed with
brownish black to brown marcescent leaf
bases; leaves of basal rosette 4-20 cm long,
more or less bipinnatifid, green and glabrous
or nearly so on both sides, the main spines
1.5-4 mm long; stems 1.5-5 dm tall, glabrous
or nearly so; cauline leaves 3-25 cm long,
0.6-5.5 cm wide, with lobing like the basal,
reduced upward; involucres 20-37 mm high,
25-50 mm wide, the bracts ovate-lanceolate
to lance-linear, tomentose to long-villous
marginally (rarely overall), the outer ones
usually pinnately spiny, smooth to roughened
medially, the dorsal ridge not developed, the
apical portions of the inner ones sometimes
contorted; spines 5-18 mm long, straw col-
ored; corollas pink to white. Three more or
less distinctive varieties are present.

2(1).

Involucral bracts copiously gray- to brown-villous with multicellular hairs; co-
rollas ocroleucous; plants of the Uinta Mountains from Lake Fork eastward

C. eatonii var. murdockii

Involucral bracts merely white-tomentose or rarely with short multicellular
hairs; corollas mainly pink or rose; plants of western Uinta Mountains, and
elsewhere 2

Involucral bracts commonly suffused with dark purple; involucres not ob-
scured by outer spinose bracts; plants of the Tushar Mountains

C. eatonii var. harrisonii

Involucral bracts green or variously purplish; involucres with copious pinnate
spines, mainly obscuring the surface of inner bractlets; plants of western Uinta
and Wasatch mountains, and Great Basin ranges C. eatonii var. eatonii

Var. eatonii [C. eriocephalum var. leio-
cephalum D.C. Eaton; this is the basionym
for C. eatonii in a strict sense, which was re-
named by Gray in honor of D.C. Eaton who

collected with Sereno Watson in 1869]. The
lectotype came from the head of the Bear
River, in Summit County (Watson 691, 1869
US!), with syntypical material being taken

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

251

under the same number in Cottonwood Can-
yon (now Salt Lake County). Lodgepole pine
and spruce communities upwards into alpine
tundra at 2375 to 3420 m in Duchesne, Juab,
Salt Lake, Summit, Tooele, and Weber coun-
ties; Nevada and Colorado; 31 (iv). Speci-
mens from the Deep Creek Mountains have
few lateral spines on the outer bracts, and ap-
proach C. clavatum in technical features.
More material is needed to determine their
status and relationships.

Var. harrisonii Welsh Talus slopes and al-
pine meadows at 2975 to 3450 m in Beaver
and Piute counties; endemic; 6 (v). This low
phase of the Eaton thistle stands geographi-
cally apart from the remainder of the species,
isolated on the islandlike Tushar Mountains.

Var. murdockii Welsh The plants grow in
talus slopes and on rock stripes at 3230 to
3660 m in Daggett, Duchesne, and Uintah
counties; endemic; 7 (iii). This variety has
been regarded as constituting a portion of C.
tweedyi (Rydb.) Petrak. That entity was re-
viewed by Moore and Frankton (1965) and
was mapped to include northeastern Utah in
its range. However, no specimens were cited

from Utah. I have seen the type of that taxon,
and other material within its range in north-
western Wyoming, and they differ in pu-
bescence of involucral bracts being merely
white tomentose along the margins.

Cirsium neomexicanum Gray Biennial
herbs from taproots; leaves of basal rosette
5-25 cm long (or more), pinnatifid, the lobes
again toothed or lobed, white tomentose be-
low and less so above, the main spines 1-6
mm long; stems 6-15 dm tall, whitish tomen-
tose; cauline leaves 1.5-35 cm long, 0.5-7 cm
wide, tomentose, appearing filmy greenish
white, lobed like the basal ones, rather
abruptly reduced upward, finally minute
spiny bracts; involucres 20-30 mm high,
40-65 mm wide, the bracts green or greenish,
narrowly lanceolate, tomentose marginally
(or overall), the outer ones often reflexed, the
inner minutely serrulate-ciliate, long-attenu-
ate apically, the spine a continuation of the
attenuation, smooth medially, the glandular
dorsal ridge more or less well developed, the
apical portions of the inner often contorted;
spines 1-9 mm long, yellowish; corollas
creamy white.

1. Involucral bracts green throughout, the attenuate apex not differing in texture

from the body of the bract C. neomexicanum var. neomexicanum

— Involucral bracts not green throughout, the attenuate apex differing in texture

from the body of the bract C. neomexicanum var. utahense

Var. neomexicanum Creosote bush, Joshua
tree, blackbrush, shadscale, sagebrush, and
pinyon-juniper communities at 915 to 2050
m in Beaver, Garfield, Grand, Juab, Kane,
Millard, San Juan, Tooele, and Washington
counties; Nevada, Arizona, New Mexico; 26
(vii). This is one of the most distinctive spe-
cies of thistle in Utah. The tall slender stems,
with one or few large heads with creamy
white flowers, stand in candelabra form in
the arid portions of western and southern
Utah. Ghostlike stalks of previous years per-
sist for a time, reminding one of the regime
which allowed their growth.

Var. utahense (Petrak) Welsh comb. nov.
[based on: C. utahense Petrak Beih. Bot.
Centr. 35(2): 470. 1917.] Salt desert shrub,
sagebrush, pinyon-juniper, and mountain
brush communities at 1220 to 2300 m in
Cache, Carbon, Emery, Millard, Rich, Salt
Lake, Tooele, and Utah counties; Colorado

(?); 24 (ii). This taxon has long been confused
with C. undulatum with which it shares the
grayish tomentum, large heads, and tall stat-
ure. They have been separated previously on
the basis of glandular development of the
dorsal ridge; a feature which is, unfortu-
nately, not diagnostic. The long-attenuate
bract apices from which the spines arise are
apparently distinctive for this taxon. It is es-
sentially intermediate between undulatum
and neomexicanum in a strict sense. The type
is from Silver Reef, Washington County, but
the main area of distribution for this variety
is apparently along the Wasatch Mountains
in northern Utah.

Cirsium ownbeyi Welsh Ownbey Thistle.
Perennial herbs from caudex and taproot, the
caudex with marcescent dark brown leaf
bases; leaves of basal rosettes 5-13 cm long,
1.5-3 cm wide, tripinnatifid, green on both
sides, sparingly tomentose along lower side of

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Great Basin Naturalist

Vol. 43, No. 2

midrib; cauline leaves with vesture and lob-
ing like the basal; stems 5-7 dm tall, winged-
decmrent, sparingly tomentose; involucres
1.8-2.5 cm high, 1.5-2.5 cm wide, the out-
ermost bracts more or less pinnately spinose,
lance-attenuate, smooth medially, the dorsal
ridge not well developed, not scabrous, spar-
ingly tomentose along margins, the inner
more or less contorted apically; spines 3-8
mm long; corollas rose-pink. Juniper, sage-
brush, and riparian communities at 1678 to
1891 m in Daggett and Uintah counties; en-
demic; 2 (i). Relationships of the Ownbey
thistle apparently lie with C. eatonii.

Cirsium rothrockii (Gray) Petrak Rothrock
Thistle. [Cnicus rothrockii Gray; Cnicus roth-
rockii var. diffusus Eastw., type from Willow
Creek, San Juan County]. Perennial or bien-
nial herbs from a caudex and taproot, the
stems 5-8 dm tall, sparingly tomentose or
glabrate to glabrous; cauline leaves 3.5-30
cm long, 2-9 cm wide, bipinnatifid, green
and glabrous or nearly so on both sides, car-
ried well to the inflorescence; involucres (19)
23-28 (34) mm long, 20-35 mm wide, the
bracts lanceolate to lance-linear, more or less
tomentose along the margins, smooth me-
dially, the dorsal ridge not or only somewhat
glandular, sometimes purplish apically, the
apical portions of the inner definitely
scabrous, the spines 7-17 mm long; corollas
red to carmine. Mixed shrubs and ponderosa
pine woods at 1830 to 2560 m in San Juan
County; Arizona; 3 (0). This entity is poorly
known in Utah; its relationship is with both
C. calcareum and C. arizonicum. It is a green
subglabrous plant with red flowers and long
involucral spines.

Cirsium rydbergii Petrak Rydberg Thistle.
[Cirsium lactucinum Rydb., type from Bluff].
Perennial herbs from a definite caudex and
taproot, the caudex clothed with blackish
brown leaf bases; leaves of basal rosette
mainly 30-90 cm long, 15-40 cm wide, bi-
pinnatifid, the lobes narrow to very broad,
glabrous to glabrate on both surfaces, the
main spines 2-11 mm long; stems 6-12 dm
tall or more, glabrous; cauline leaves

glabrous, less lobed and much reduced up-
wards; involucres 10-17 mm high (not mea-
suring the reflexed outer bracts), 13-26 mm
wide, the outer bracts lance-ovate, rather
abruptly contracted into recurved spines
3-25 mm long, sparingly tomentose mar-
ginally; dorsal glandular ridge lacking, the in-
ner attenuate, not scabrous; flowers pink.
Hanging gardens, or rarely in canyons below
them, at 1125 to 1525 m in Grand, Kane, San
Juan, Wayne (and probably in Garfield)
counties; Arizona (?).; 15 (v). Both C ryd-
bergii Petrak and C. lactucinum Rydberg are
based on the same type collection from the
hanging gardens near Bluff. The Rydberg
thistle is a plant with huge basal rosettes, tall
slender flowering stems, and small heads.

Cirsium scariosum Nutt. Meadow Thistle.
[Carduus lacerus Rydb., type from near Mid-
way; Carduus olivescens Rydb., type from
the Aquarius Plateau; Cirsium acaule var.
americanum Gray; Cnicus drummondii var.
acaulescens Gray; C foliosum authors, not T.
& G.; C. drummondii authors, not T. & G.].
Perennial herbs from a simple caudex and
taproot, the caudex with chestnut leaf bases;
leaves of basal rosette 2-28 cm long, 1-8 cm
wide, merely spiny toothed to bipinnatifid,
the lobes often again toothed, tomentose to
glabrate below, thinly tomentose to glabrous
above, the main spines 1-5 mm long; stems
lacking, or 1-12 dm tall or more, often suc-
culent and edible, arachnoid to glabrous;
cauline leaves (when stems present) bipin-
natifid or merely pinnatifid, the spines 3-35
mm long, with vesture like the basal; in-
volucre 22-35 mm high, 20-65 mm wide, the
outer bracts lance-ovate, the inner progres-
sively more lance-attenuate, smooth me-
dially, the margins smooth to minutely
scabrous, tomentose to glabrous on margins,
the dorsal ridge not well developed, the long-
est spines mainly 2-5 mm long, straw col-
ored, the inner with tips more or less con-
torted, dilated, or fimbriate, usually whitish
or silvery; flowers white to pink or pink-
purple. Our specimens fall into two rather
distinctive varieties.

1. Heads 25-35 mm high, 35-80 mm wide; inner bracts slender, sometimes con-
torted, not especially dilated; plants mainly 6-12 dm tall

C. scariosum var. thorneae

— Heads 22-30 mm high, 20-40 mm wide; inner bracts often dilated or con-
torted, sometimes fimbriate; plants 0-6 dm tall C. scariosum var. scariosum

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

253

Var. scariosum [Cirsium acaule var. ameri-
canum Gray]. This taxon, as here interpreted,
consists of an amazingly diverse assemblage
that has passed under a series of names in-
cluding those cited above; and, if it is demon-
strated that C. foliosum (Hook.) DC. is ac-
tually conspecific, that name has priority.
Saline seeps and salt marshes, stream sides,
terraces, and other meadowlands at 1310 to
3175 m in Carbon, Duchesne, Emery, Gar-
field, Juab, Millard, Salt Lake, Sanpete, Se-
vier, Summit, Tooele, and Utah counties;
British Columbia to Montana, south to Cali-
fornia, Arizona, and Colorado; 43 (x). This
phase of C. scariosum has passed under the
names C. acaulescens (Gray) Schum., C. colo-
radoense (Rydb.) Cockerell; C tioganum
(Congdon) Petrak, C drummondii T. & G.,
and C. foliosum. Nomenclature is still un-
clear, and more work is indicated. Our highly
variable material is transitional from acaules-
cent to caulescent within populations, with
stems, when present, fleshy and edible. This
is our common thistle of meadowlands, and it
is unfortimate that nomenclatural entangle-
ments have not allowed selection of an
unequivocal name. Reported for the state is
C. parryi (Harrington, Flora of Colorado,
1952), but I have seen no specimens of that
entity from Utah. It would key to C. scario-
sum in the present work. It has densely
arachnoid involucral bracts, with at least the
innermost dilated-fringed at the tips; flowers
are greenish yellow and the leaves are gla-
brate on both surfaces.

Var. thorneae Welsh Stream terraces and
seeps or springs at 1650 to 2475 m in Beaver,
Garfield, Iron, Kane, Millard, and Piute
counties; endemic (?); 10 (vi). In addition to
the features noted above, the cauline leaves
are thick, with coarse veins, and spines 8-35
mm long.

Cirsium scopulorum (Greene) Cockerell in
Daniels [Carduus scopulorum Greene]. Pe-
rennial herbs from taproots; leaves of basal
rosettes 3-28 cm long, 0.8-8 cm wide, with
spines 2-6 mm long, unlobed to bipinnatifid,
tomentose below, glabrate to glabrous and
green above; stems mainly 3-7 dm tall, spar-
ingly arachnoid, not winged-decurrent; cau-
line leaves mainly bipinnatifid, or the upper

ones merely pinnatifid, green above, glabrous
to sparingly tomentose below, rather gradu-
ally reduced upward; heads in a compact
subglobose terminal cluster; involucres 30-35
mm high, 30-55 mm wide, the bracts lance-
attenuate, abundantly villous marginally,
with long yellowish to brownish multicellular
hairs, the outer ones usually pinnately spiny,
the dorsal crest not glandular, smooth me-
dially, the apical portions of the inner ones
often contorted; spines 10-18 mm long, yel-
lowish; corollas pale yellow to cream. Sage-
brush, aspen, and spruce-fir communities at
2135 to 3000 m in Grand, San Juan (?), and
Uintah counties; Colorado; 3 (0).

Cirsium subniveum Rydb. Perennial herbs
from taproots; basal rosettes not seen; stems
mainly 6-10 (13) dm tall, tomentose, winged-
decurrent; cauline leaves 3-25 cm long or
more, 1-6 cm wide, pinnatifid, tomentose on
both sides, or less so above, the bases decur-
rent; involucres 17-25 mm high, 20-30 mm
wide, the bracts ovate-lanceolate, smooth
medially, the glandular dorsal ridge more or
less developed, none scabrous, tomentose
marginally; spines 3-5 mm long; corollas ap-
parently white to cream. Pinyon-juniper
community at 1890 m in Rich and Sanpete
counties; Oregon to Montana; 2 (0).

Cirsium undulatum (Nutt.) Spreng. Gray
Thistle. [Carduus undulatus Nutt.]. Perennial
herbs from a simple caudex and taproot, the
caudex more or less clothed with persistent
leaf bases; leaves of basal rosette mainly 7-25
cm long, 1.5-6 cm wide, merely toothed to
bipinnatifid, tomentose on both sides, white-
tomentose below, white to greenish above,
the main spines 1-6 mm long; stems 2-10
(12) dm tall, tomentose; cauline leaves bipin-
natifid or the upper ones merely pinnatifid,
with vesture as in the basal ones, rather grad-
ually reduced upward; involucres (15) 20-30
mm high, 20-60 mm wide, the bracts brown
or brownish, lance-ovate to lanceolate, to-
mentose on margins or overall, the dorsal
ridge strongly glutinous to undeveloped, the
spinose tips spreading, with yellowish spines
mainly 2-5 (10) mm long, smooth medially,
the apical portion of the innermost more or
less contorted; corollas pink, pink-purple, or
creamy white.

254
1.

Great Basin Naturalist

Vol. 43, No. 2

Heads mainly less than 2.5 cm wide, even the largest, commonly (1) 3-10 or

more per stem C undulatiim var. tracyi

Heads mainly more than 2.5 cm wide, at least the largest, commonly 1-3 per
stem C. undulatum var. undulatum

Var. tracyi (Rydb.) Welsh comb. nov.
[based on: Carduus tracyi Rydb. Bull Torrey
Bot. Club. 32: 133. 1905]. Sagebrush, moun-
tain brush, juniper, aspen, and Douglas fir
communities at 1525 to 2900 m in Duchesne,
Emery, Grand, Juab, Summit, and Uintah
counties; Colorado; 26 (iii). This variety
grades with the type variety, and separation
is at least partially arbitrary.

Var. undulatum Desert shrub, sagebrtish,
pinyon-juniper, mountain brush, ponderosa
pine, and aspen communities at 1400 to 2600
m in Cache, Carbon, Daggett, Duchesne,
Emery, Garfield, Juab, Grand, Rich, San
Juan, Salt Lake, Sanpete, Sevier, Summit,
Tooele, Uintah, Utah, Wasatch, and Weber
counties; British Columbia to Minnesota,
south to Arizona, New Mexico, and Missouri;
59 (vi).

Cirsium virginensis Welsh Virgin Thistle.
Perennial(?) herbs from taproots; leaves of
basal rosettes 6-35 cm long, 1-5 cm wide,
unlobed, pubescent like the cauline ones,
with spines 1-4 mm long; stems 6-15 dm tall,
tomentose, winged by definitely decurrent
leaf bases; cauline leaves 1.5-15 cm long or
more, sinuate-dentate to pinnatifid, whitish
tomentose on both sides, or greenish above,
often reduced to spiny bracts upwards; in-
volucres 13-20 mm tall, 12-32 mm wide, the
bracts ovate-lanceolate to narrowly lanceo-
late, brownish to straw colored, or often suf-
fused with purple, tomentose marginally (or
overall), the outer not especially reflexed, the
inner serrulate or entire, smooth medially,
the glandular dorsal ridge more or less devel-
oped, the apical portions of the inner often
contorted; spines 2-6 (8) mm long, yellowish;
corollas pink to lavender (or white?). Saline
seeps and stream terraces at 850 to 950 m in
Washington Co.; Arizona; 9 (i). The small
heads and long decurrent leaf bases are diag-
nostic. The relationships of the Virgin thistle
are unknown. It does not appear to be closely
related to other species groups represented in
our area.

Cirsium vulgare (Savi) Ten. Bull Thistle.
[Carduus vulgaris Savi]. Biennial herbs from

taproots; leaves of basal rosette mainly 5-25
cm long, 2-8 cm wide, merely doubly ser-
rate-dentate to doubly pinnatifid, tomentose
beneath, coarsely hispid above; stems mainly
3-12 (15) dm tall, spiny-winged by decurrent
leaf bases; cauline leaves mainly bipinnatifid,
with vesture as in the basal ones; involucres
28-40 mm high, 35-70 mm wide, the bracts
narrowly lanceolate, with spreading spine-
tips, tomentose marginally, the dorsal ridge
not developed, the inner sometimes con-
torted apically; spines 1-4 mm long, yellow-
ish; corollas rose-purple. Meadows, fields,
roadsides, and other disturbed sites from 1340
to 2745 m in most, if not all, Utah counties;
widespread in North America; 52 (i).

Cirsium wheeleri (Gray) Petrak Wheeler
Thistle. [Cnicus wheeleri Gray]. Perennial or
biennial herbs from a simple or branched
caudex and taproot, the caudex clothed with
persistent brown to dark brown leaf bases;
leaves of basal rosettes mainly 7-20 cm long,
1-5 cm wide, once to twice pinnatifid, or
merely toothed or spinose-serrate, grayish or
whitish tomentose below, thinly so to
glabrous and green above, the main spines
0.5-4 mm long; stems 2.5-7 dm tall; cauline
leaves 2-25 (32) cm long, 0.5-5 (7) cm wide,
with lobing and vesture similar to the basal,
carried well to the inflorescence, though re-
duced above; involucres 20-27 mm high,
20-35 mm wide, the bracts lance-ovate to
lance-linear, more or less tomentose along
the margins, smooth medially, the dorsal
ridge not or only somewhat glandular, some-
times purplish tipped, the apical portions of
at least the inner more or less scabrous; co-
rollas pink to pink-purple, or less commonly
white. Mountain brush, pinyon-juniper, white
fir, aspen, and spruce-fir communities at
(1980) 2165 to 3150 m in Beaver, Emery,
Garfield, Iron, Juab, Kane, Millard, San Juan,
Sanpete, and Sevier counties; Colorado, New
Mexico, and Arizona. Our materials appar-
ently intergrade with C. undulatum, C. nidu-
lum, and possibly C. scariosum. The moder-
ate sized heads, usually pink or pink-purple
flowers, low stature, essentially nonglandular

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

255

bracts, and usually green upper leaf surface
appears to be diagnostic. The phases from
Cedar Canyon (Iron County), with merely
spinose unlobed leaves, are striking, but
probably not more than minor variants; 39
(iv).

Cnicus L.

Annual caulescent spiny herbs from tap-
roots, the juice watery; leaves alternate;
heads solitary, terminating branches; in-
volucral bracts in several series, spine tipped,
the inner ones pinnately spiny; receptacle
densely bristly; corollas all discoid, yellow,
perfect; pappus in 2 series, the outer smooth,
long, alternating with short sparingly pecti-
nate ones; style with a ring of hairs at base of
divergent branches; achenes terete, strongly
ribbed, glabrous.

Cnicus benedictus L. Blessed Thistle.
Plants 1-5 dm tall or more, branching from
near the base; stems villous; leaves mainly
8-15 cm long, pinnatifid, more or less
glandular and sparingly villous, the spines
0.5-1.5 mm long, the lower ones petiolate,
becoming sessile above; involucres 3-4 cm
high, closely subtended and obscured by the
foliose bracteate upper leaves; corollas yel-
low. Waste places and gardens at 885 m in
Washington County; widespread in the U.S.;
adventive from Europe; 1 (0).

CoNYZA Less. Nom. Cons.

Annual herbs from taproots, with watery
juice; stems erect, commonly branched;

leaves alternate, simple; heads numerous, in
cylindric to conic panicles; involucral bracts
more or less imbricate, herbaceous medially;
receptacle flat or nearly so, naked; rays min-
ute, white or purplish, scarcely surpassing the
pappus; disk flowers seldom more than 20,
perfect, fertile; pappus of capillary bristles;
achenes 1- or 2-nerved or nerveless.

Conyza canadensis (L.) Cronq. Horse-
weed. [Erigeron canadensis L.]. Annuals,
mainly 0.5-10 dm tall, glabrous or spreading-
hairy; leaves 2-8 (10) cm long, 2-8 cm wide,
linear to oblanceolate, ciliate-serrate, often
deciduous by late anthesis; heads numerous,
inconspicuous; involucres 2-3.5 (4) mm high,
(2.5) 3-7 mm wide, the bracts lance-subulate,
the midvein glandular-thickened, herbaceous
medially, glabrous or strigose; rays white or
purplish. Weedy species, often in riparian or
other moist disturbed sites at 850 to 2135 m
in all (?) Utah counties; widespread in North
America; Europe; 30 (vi). Our material be-
longs to var. glabrata (Gray) Cronq.

Crepis L.

Annual, biennial, or perennial caulescent
or subacaulescent herbs, from taproots, with
milky juice; leaves basal and cauline, alter-
nate, pinnatifid to toothed or entire; heads
few to numerous, in corymbose or paniculate
clusters; involucral bracts in 1 or 2 series,
herbaceous; receptacle naked; corollas all
raylike, perfect, yellow or yellowish; pappus
of numerous white capillary bristles; achenes
terete or nearly so, 10- to 20-ribbed, often
beaked.

1. Plants annual, adventive, of disturbed sites C. capillaris

— Plants perennial, indigenous, neither weedy nor of disturbed sites 2

2(1). Leaves and stems glabrous (or glandular-hispid only above); plants sub-
acaulescent or subscapose 3

— Leaves and stems more or less tomentose or puberulent to setose or glandular
hispid; plants caulescent 4

3(2). Plants less than 10 cm tall, soboliferous, of high elevations C. nana

— Plants mainly 15-40 cm tall, never soboliferous, of lower-elevation meadows ...
C. runcinata

4(2). Heads narrowly cylindric; involucral bracts 5-7 (8), the inner commonly

glabrous; flowers mostly 5-10 C. acuminata

— Heads narrowly to broadly campanulate; involucral bracts 8-15, tomentose

and often setose-hispid; flowers mostly 8-60 5

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Great Basin Naturalist

Vol. 43, No. 2

5(4). Leaf segments linear to narrowly lanceolate, entire or nearly so, the terminal

lobe more than 5 cm long; achenes commonly green C. atrabarba

— Leaf segments narrowly lanceolate to triangular, some usually toothed, the ter-
minal lobe less than 5 cm long; achenes mainly yellowish to brownish 6

6(5). Involucres more than twice longer than broad; leaves usually green, runcinate-

pinnatifid C intermedia

— Involucres less than twice longer than broad, or leaves not green or not run-
cinate-pinnatifid 7

7(6). Involucre and stems not or sparingly setose, but, if setose, the setae gland-
tipped C. occidentalis

— Involucre and/or stems conspicuously setose, the setae not glandular

C. modocensis

Crepis acuminata Nutt. Perennial herbs, communities at 1890 to 2870 m in Daggett,

2.5-8.5 dm tall, with 1-several stems from a Garfield, Salt Lake, Summit, Tooele, Uintah,

caudex, the caudex clothed with dark brown and Utah counties; British Columbia and Al-

marcescent leaf bases; herbage more or less berta, south to Nevada and Colorado; 8 (i).

tomentose to glabrate; basal and lowermost The species is evidently uncommon in Utah,

cauline leaves 8-33 (40) cm long, 2-12 cm It is known to form apparent hybrids with C.

wide, petiolate, the blade elliptic to oblan- acuminata, and probably with other taxa as

ceolate in outline, pinnatifid to runcinate- well.

pinnatifid, the lobes triangular to narrowly Crepis capillaris (L.) Wallr. Slender
subulate, sometimes toothed or lobed; heads Hawksbeard. Annual or biennial herbs, the
mainly 20-75 or more, cylindric, 5- to 10- stems erect, simple or branched, mostly 1-6
flowered; involucres (8) 9-13.5 (16) mm high, dm tall, sparingly spreading-hairy; basal
3-7 mm wide, the inner ones 5-8, glabrous or leaves 3-20 cm long, 0.5-3 cm broad, lan-
sometimes shortly villous-tomentose, the out- ceolate to oblanceolate, denticulate to pin-
er bracts much shorter, commonly tomen- natifid or bipinnatifid, glabrous or pubescent
tose; corollas 10-18 mm long, yellow; ach- with stiff spreading hairs, especially along the
enes yellow to brown, narrowed above, lower midvein, petiolate; cauline leaves re-
Sagebrush, mountain brush, white fir, aspen, duced upward, sessile and auriculate-clasp-
and spruce-fir communities at 1430 to 2900 ing; heads (1) several to numerous, mostly 20-
m in most if not all Utah counties; Washing- to 60-flowered, borne in an open in-
ton to Montana, south to California, Arizona, florescence; involucres 5-8 mm high, 5-14
and New Mexico; 69 (viii). mm wide, the inner bracts lance-attenuate,

Crepis atrabarba Heller [C. occidentalis 8-16, tomentose, often glandular-hairy,

var. gracilis D.C. Eaton]. Perennial herbs, glabrous within, the outer bracts lance-linear;

2-4.5 dm tall, with 1-several stems from a achenes 2-5 mm long, pale brown to straw

caudex, the caudex with dark brown to colored; achenes 2-5 mm long, pale brown to

purplish marcescent leaf bases; herbage gray straw colored, not beaked. Ruderal weed of

villous-tomentose to glabrate, basal and Salt Lake County; widely scattered in North

lowermost cauline leaves 6-22 cm long, America; adventive from Europe; 1 (0).
1.5-4 cm wide, petiolate, the blade lance- Crepis intermedia Gray Gray Hawksbeard.

elliptic in outline, pirmatifid, the lobes linear [C. barbigera Leiberg, in part]. Perennial

or linear-subulate, the terminal lobe 5-9 cm herbs, 2.5-7 dm tall, with 1-several stems

long, entire; heads mainly 2-15, cam- from a caudex, the caudex clothed with pale

panulate, commonly 10- to 40-flowered; in- to dark brown marcescent leaf bases; herbage

volucres 9-15 mm long, 7-13 mm wide, the more or less tomentose or villous; basal and

inner ones 8-10, usually grayish tomentulose lowermost cauline leaves 15-30 cm long,

and often with few glandless black setae; co- 2-10 cm wide, petiolate, the blade elliptic to

rollas 10-18 mm long, yellow; achenes usual- oblanceolate in outline, pinnatifid to run-

ly greenish, attenuate at the apex. Sagebrush, cinate-pinnatifid, the lobes triangular to lin-

ponderosa pine, Douglas fir, and white fir ear-subulate, sometimes toothed or lobed, the

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

257

terminal lobe less than 5 cm long; heads
mainly 10-60, campanulate, 7- to 16-flow-
ered; involucres 11-20 mm high, 6-12 mm
wide, the inner ones 7-12, tomentulose
(rarely glabrate), sometimes setose with non-
glandular setae, the outer bracts much short-
er; corollas 13-20 mm long, yellow; achenes
mainly yellowish or brownish, narrowed
above. Sagebrush, pinyon-juniper, and moun-
tain brush communities at 1525 to 2575 m in
Beaver, Cache, Duchesne, Garfield, Salt
Lake, Sevier, Utah, Wasatch, and Washing-
ton counties; Washington to Alberta, south to
California, Nevada, and Colorado; 16 (iii).
The intermedia assemblage consists of a series
of apomictic intermediates involving C.
acuminata as one of the parental types, and
one or more of the other taxa (i.e., occiden-
talis or modocensis) to complete the complex.
Included here is the concept of C. barbigera
as it has been applied in Utah; it consists of a
similar hybrid sequence of polyploid apo-
micts from outside our area.

Crepis modocensis Greene Modoc Hawks-
beard. Perennial herbs, 1.5-3.7 dm tall, with
1 -several stems from a caudex, the caudex
clothed with pale to brown marcescent leaf
bases (the stem base often yellow); herbage
more or less tomentose; basal and lowermost
cauline leaves 9-25 cm long, 2-5 cm wide,
petiolate, the blade elliptic to oblanceolate in
outline, bipinnatifid, the lobes linear to
lance-subulate, again toothed or lobed, the
terminal lobe less than 5 cm long; heads 1-9,
10- to 60-flowered; involucres 11-16 mm
high, 11-23 mm wide, the inner bracts
10-15, tomentulose, commonly setose, the
setae not glandular, the outer bracts much
shorter; corollas 13-22 mm long, yellow; ach-
enes greenish black to reddish brown, attenu-
ate. Sagebrush, pinyon-juniper, and mountain
brush communities at 1640 to 3175 m in Bea-
ver, Box Elder, Cache, Daggett, Juab, Mil-
lard, Rich, Salt Lake, Sanpete, Sevier,
Tooele, Uintah, and Utah counties; British

Columbia to California, Nevada, and Colo-
rado; 24 (0). The peculiar numerous slender
lateral lobes of the deeply dissected or parted
leaf blades are diagnostic.

Crepis nana Richards. Dwarf Hawksbeard.
Perennial caespitose herbs, the stems much
branched, often soboliferous, mostly 0.2-1.1
dm tall, contracted, usually obscured by the
leaves, glabrous; basal leaves mainly 1-7.5
cm long, 0.2-1.8 cm wide, the blades spatu-
late to orbicular, elliptic, or ovate, glabrous,
petiolate; cauline leaves similar to the basal,
not clasping; heads few to numerous, mostly
4- to 12-flowered, borne in a compact cush-
ionlike inflorescence; involucre 7-12 mm
high, 3-6 mm wide, the inner bracts narrow-
ly oblong, 8-12, greenish or blackish,
glabrous, the outer much shorter; achenes
brownish, ribbed, shortly beaked. Alpine
communities, mainly in talus, at 3050 to 3425
m in Juab, Piute, and Utah or Salt Lake
counties; Alaska to Labrador, south to Cali-
fornia and Utah; 10 (iii).

Crepis occidentalis Nutt. Western Hawks-
beard. Perennial herbs, 1-4 dm tall, with 1-
several stems from a caudex, the caudex
clothed with brown marcescent leaf bases
(the stem base often yellow); herbage tomen-
tose; basal leaves mainly 6-30 cm long, 1-5
cm wide, petiolate, the blade lanceolate to
elliptic in outline, pinnatifid to bipinnatifid,
the lobes triangular to oblong or linear-sub-
ulate, usually again toothed or lobed, the ter-
minal lobe less than 5 cm long; heads 2-25,
12- to 30-flowered; involucres 10-20 mm
high, 6-15 mm wide, the inner bracts (7)
8-13 (18), tomentose, the outer ones much
shorter; corollas 10-22 mm long, yellow; ach-
enes pale to dark brown, not much attenuate
apically. There are three rather weak and ar-
bitrarily recognizable varieties of this species,
with some geographical correlation, in Utah.
Intermediates occur between the varieties
and with other taxa as well.

2(1).

Largest heads 12- to 14-flowered, with 8 or 9 involucral bracts; plants mainly

of the Great Basin C. occidentalis var. costata

Largest heads with more than 15 flowers, with 10-13 involucral bracts; plants

of various distribution 2

Involucres with few glandular setae, or none; plants mainly 2-3 dm tall, of the
Great Basin C. occidentalis var. pumila

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Great Basin Naturalist

Vol. 43, No. 2

Involucres with few to numerous glandular setae; plants mainly 1-2 dm tall, of

the Colorado drainage system, less commonly in the southern Great Basin

C. occidentalis var. occidentalis

Var. costata Gray Sagebrush, pinyon-
juniper, mountain brush, and aspen commu-
nities at 1525 to 2200 m in Box Elder, Juab,
Millard, Salt Lake, Tooele (type from Stans-
bury Island), Utah, and Washington counties;
British Columbia to California and Colorado;
23 (0).

Var. occidentalis Shadscale, rabbitbrush,
sagebrush, pinyon-juniper, and ponderosa
pine communities at 1280 to 2565 m in Bea-
ver, Daggett, Duchesne, Garfield, Kane,
Piute, Sanpete, Sevier, San Juan, and Wash-
ington counties; Oregon to Wyoming, south
to California and New Mexico; 25 (v).

Var. pumila (Rydb.) Babcock & Stebbins
[Crepis pumila Rydb.]. Sagebrush, pinyon-
juniper, and mountain brush communities at
1700 to 2100 m in Millard and Tooele coun-
ties; 7 (0).

Crepis runcinata (James) T. & G. [Hier-
acium runcinatum James]. Perennial herbs,
1.5-5 (7) dm tall, with 1-several stems from a
caudex, the short caudex clothed with brown
marcescent leaf bases; herbage glabrous or
hispid above (puberulent in some), not to-
mentose; basal leaves mainly 2-25 cm long,
1-6 (8) cm wide, petiolate or not, spatulate
to oblanceolate, or the blades ovate to oval,
oblong, or oblanceolate, more or less pinnati-
fid to lobed or entire, commonly glaucous;
heads 1-30, with 20-50 flowers; involucres
campanulate, 8-16 mm high, 6-15 mm wide
or more, the inner bracts mainly 10-16, pu-
berulent or hispid, the outer ones much
shorter; corollas 9-18 mm long, yellow; ach-
enes light to dark brown, attenuate, or short-
ly beaked. Three distinctive varieties are
present.

2(1).

Involucres merely puberulent; plants mainly of saline meadows

C. runcinata var. glauca

Involucres hispid with black hairs (resembling species of Hieracium); plants of
saline or nonsaline sites 2

Basal leaves definitely petiolate, the blade 2-4 times longer than broad

C runcinata var. hispidulosa

Basal leaves broadly winged-petiolate, the blade 4-8 times longer than broad ...
C. runcinata var. runcinata

Var. glauca (Nutt.) Welsh stat. nov. [based
on: Crepidium glaucum Nutt., Trans. Amer.
Phil. Soc. II. 7: 436. 1841]. Meadows, lake
shores, seeps, and hot springs in salt grass,
rush, alkali sacaton, and common reed com-
munities at 1220 to 2200 m in Carbon, Dag-
gett, Duchesne, Emery, Grand, Juab, Kane,
Millard, Piute, San Juan, Sevier, Tooele, Uin-
tah, Utah, and Wayne counties; Idaho to Sas-
katchewan, south to Arizona and New Mexi-
co; 34 (xii). This variety has been collected in
full flower on 27 April at Monroe Hot
Springs.

Var. hispidulosa Howell ex Rydb. Sedge-
willow and meadow communities at 1370 to
2535 m in Box Elder, Duchesne, Kane, Piute,
Rich, Sanpete, Sevier, Summit, and Utah
counties; Washington to Montana, south to
California and Colorado; 14 (ii). The mead-
ows are seldom saline where this plant
occurs.

Var. runcinata [C. runcinata var. alpicola
Rydb.]. Bogs in Salt Lake and Utah counties;
Manitoba to Minnesota, south to Idaho and
New Mexico; 1 (0). This variety is evidently
uncommon in Utah.

DicoRiA T. & G.

Annual herbs; leaves alternate or the lower
ones opposite, simple, entire or toothed;
heads unisexual or perfect, discoid; involucral
bracts strongly dimorphic, the ca 5 outer
ones small, herbaceous, the inner subtending
the 1 or 2 pistillate flowers, subscarious, ac-
crescent, much larger than the outer at matu-
rity; chaff narrow, tardily deciduous; pis-
tillate flowers without corolla; staminate
flowers with funnelform corolla, the anthers
distinct; achenes plano-convex, black,
toothed to pectinately wing margined; pap-
pus lacking.

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

259

Foliose bracts of inflorescence orbicular to broadly ovate; plants of Washing-
ton County D. canescens

Foliose bracts of inflorescence lance-ovate to lanceolate; plants not of Wash-
ington County D. brandegei

Dicoria brandegei Gray [D. paniculata
Eastw.; D. wetherillii Eastw., a monstrous
form]. Plants branched from the base up-
wards, 1.5-5.5 dm tall, the herbage white-pi-
losulose to strigose, the hairs multicellular;
lower cauline leaves linear to lanceolate,
more or less hastately lobed, toothed, or sub-
entire, 1-7 cm long (including petiole),
0.2-1.5 cm wide; foliose bracts linear to ob-
long, lanceolate or ovate, rarely if ever or-
bicular, the blades 0.6-4 cm long; outer in-
volucral bracts oblong, 1.5-3 mm long, the
inner ones suborbicular, glandular-pu-
berulent, accrescent in fruit; achenes 5-8 mm
long, the winged margin toothed to pecti-
nate, black like the body or pale. In dunes
and other sandy sites, in wavy-leaf oak,
eriogonum, amsonia, old-man sagebrush, rab-
bitbrush, ephedra, and vanclevea commu-
nities at 1130 to 1830 m in Emery, Garfield,
Grand, Kane, San Juan, and Wayne counties;
Arizona, New Mexico, and Colorado (?); 21

(V).

Dicoria canescens Gray in Torr. Plants
branched from base upward, 2.5-9 dm high,
the herbage white-pilosulose to strigose and
glandular, the hairs multicellular; lower cau-
line leaves deltoid-lanceolate, dentate, 1-5
cm long; foliose bracts ovate to orbicular, the
blades 0.6-1.5 cm long; outer involucral

bracts oblong, 2-3 mm long, the inner ones
suborbicular, glandular-puberulent, accres-
cent in fruit, to 10 mm long or more; achenes
5-6 mm long, the winged margin toothed to
pectinate, black like the body, or pale. Dunes
and other sandy sites in blackbrush and creo-
sote bush communities at 825 to 1000 m in
Washington County; Arizona, Nevada, and
California; 4 (1). Our material belongs to ssp.
clarkae (Kennedy) Keck.

Dyssodia Cav.

Annual or perennial herbs or subshrubs
from taproots, the juice watery; herbage with
conspicuous translucent oil glands; stems
striate, numerous; leaves opposite or alter-
nate, entire to pinnatisect; heads solitary at
branch ends, or few to several in cymose
clusters; involucral bracts in 2 series, distinct
or united, and usually with a much shorter
outer set; receptacle flat or convex, pu-
berulent; ray flowers yellow, pistillate, fer-
tile; disk flowers fertile; pappus of 10-15
bristle-tipped scales, or these dissected into 3
or more bristles; style branches with a short,
conic appendage.
Strother J. L. 1969. Systematics of Dyssodia

Cavanilles (Compositae: Tageteae).

Univ. Calif. Publ. Bot. 48: 1-88.

1. Plants annual; leaves bipinnatisect; stems villosulous D. papposa

— Plants perennial, herbs or subshrubs; leaves simple or merely pinnatisect; stems
hispidulous 2

2(1). Heads borne on elongate merely bracteate peduncles; leaves pinnately 5-lobed,

shortly hispid; pappus scales tipped with usually a solitary bristle

D. pentachaeta

— Heads sessile or essentially so; leaves simple, entire or rarely irregularly few
lobed; glabrous or merely ciliate; pappus scales with 3-5 bristles D. acerosa

Dyssodia acerosa DC. Dogweed. Plants glands; ray flowers 7-8, lemon-yellow; disk

suffruticose, 10-25 cm tall, forming compact flowers 18-25, pale yellow; pappus of ca 20

clumps, from taproots; herbage glabrous or scales, each dissected into 3-5 bristles; ach-

villosulous; leaves opposite (or alternate enes dark brown, 3-3.5 mm long, strigose.

above), simple or irregularly lobed, 3-18 mm Blackbrush communities at 1130 to 1350 m

long, 0.5-1 (2) mm wide, glandular, ciliate or in Garfield, Washington, and San Juan coun-

glabrous; heads sessile or subsessile; in- ties; Arizona and New Mexico, south to Mex-

volucres turbinate-cylindric, 5-7 mm high, ico; 5 (ii).

3-4 mm wide; involucral bracts ca 13, con- Dyssodia papposa (Vent.) A.S. Hitchc.

nate, each bract with conspicuous orange [Tagetes papposa Vent.]. Plants annual, 1.5-4

260

Great Basin Naturalist

Vol. 43, No. 2

dm tall; herbage glabrous to sparingly pu-
berulent; leaves opposite below, alternate
above, 1.5-3 (5) cm long, pinnatisect into
11-15 lobes, these sometimes again lobed;
heads shortly pedunculate to subsessile; in-
volucres turbinate to campanulate, 6-10 mm
high, and about as wide; involucral bracts
6-12, oblanceolate, with yellowish oil glands,
connate only at the base; ray flowers 8 or
fewer, yellow-orange; disk flowers mainly
20-40, dull yellow; pappus of ca 20 scales,
each dissected into 5-10 bristles; achenes
black, 8-35 mm long. Sandy roadsides at
1450 to 1500 m in Carbon, Duchesne, San-
pete, and Tooele counties; through much of
the United States and Mexico; 4 (0).

Dyssodia pentachaeta (DC.) Robins.
[Hymenatherum pentachaetum DC] Plants
suffruticose, 8-28 cm tall, forming rounded
clumps, from taproots; leaves opposite, pin-
nately parted into 3-5 rigid linear lobes,
0.5-2 cm long, sparingly hirtellous; pe-
duncles 1-8 cm long; involucres turbinate,
4.8-6 mm high, 5-10 mm wide; involucral
bracts in 2 series, connate for much of their
length, with distinctive yellow oil glands; ray
flowers usually 13, bright yellow; disk flow-
ers 50-70, dull yellow; pappus usually of 10

scales, these awnless or with 1-3 awns;
achenes brown, 2.2-3 mm long, hispid to
glabrous. Blackbrush, ephedra, shadscale,
creosote bush, and Joshua tree communities
at 700 to 1220 m in Garfield, Kane, San Juan,
and Washington counties; Nevada and Cali-
fornia to Texas and Mexico; South America;
30 (vi). Our material has been assigned to var.
belinidium (DC.) Strother [D. thurberi (Gray)
Woot. & Standi.].

Encelia Adams

Shrubs; stems ascending to erect, grayish
or whitish, the branchlets commonly pu-
bescent; leaves alternate, simple, petiolate,
entire or toothed; heads solitary or in cymose
clusters, radiate or discoid; involucral bracts
in 2 or 3 series; receptacle convex to flat,
chaffy, the scales clasping the achenes and
falling with them; ray flowers (when present)
sterile, yellow; disk flowers perfect, yellow;
pappus lacking (or of 2 awns); achenes flat,
obovate, villous-ciliate and pubescent on the
surfaces.
Blake, S. F. 1913. A recision of Encelia and

some related genera. Proc. Amer. Acad.

49: 358-376.

1. Leaves white-tomentulose; peduncles glabrous; heads in branching cymes;

plants rare in Washington County E. farinosa

— Leaves strigose to hispid, green; peduncles scabrous to strigose; heads soli-
tary at branch ends; plants of Washington County, and elsewhere E. frutescens

Encelia farinosa Gray Incienso. Plants
mainly 3-10 dm tall, aromatic; leaves clus-
tered at apex of current stems, 2-8 cm long,
ovate, entire or toothed, silvery tomentose,
petiolate; peduncles elongate, cymosely
branched or simple; heads showy, the disk
1-1.5 cm wide; involucres 4-7 mm high, vil-
lous and glandular dotted; rays 8-12 mm
long, orange-yellow; achenes narrowly obo-
vate. Blackbrush community at 1280 m in
Washington County; Nevada, Arizona, and
California; Mexico; 1 (0).

Encelia frutescens Gray Bush Encelia.
[Simsia frutescens Gray]. Plants mainly 3-12
(15) dm tall; leaves scattered along current
stems, the blades commonly 0.5-2.5 cm long,
0.3-2 dm wide, ovate to orbicular or lanceo-
late, entire or toothed, strigose to hispid with
pustular-based hairs; heads showy or not, the
disk 1-3 cm wide; involucres 6-10 mm high,
strigose or glandular; rays lacking or 1-16 (or
more), 2-12 mm long, yellow; achenes obo-
vate. Two distinctive varieties are present in
Utah.

Herbage strigose, also with some pustular-based hairs; involucral bracts abrupt-
ly caudate-acuminate, strigose; plants of Washington County

£. frutescens var. virginensis

Herbage hispid with pustular-based hairs; involucral bracts gradually attenu-
ate, more or less glandular (sometimes strigose) E. frutescens vslt. frutescens

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

261

Var. frutescens [E. frutescens var. resinosa
Jones in Blake] Talus and slickrock in black-
brush and shadscale communities at 1130 to
1830 m in Emery, Grand, Kane, and San Juan
counties; Arizona, California; 15 (vii). There
is a cline of glandularity in leaves from defi-
nitely glandular in the southern portion of
the range in Utah to no glands at all in the
northern material. Also, our plants vary from
discoid to radiate.

Var. virginensis (A. Nels.) Blake [E. virgi-
nensis A. Nels.]. Creosote bush, Joshua tree,
and blackbrush communities at 760 to 1325
m in Washington County; Nevada, Arizona,
and California; 23 (i).

Enceliopsis (Gray) A. Nels.

Perennial scapose or subscapose herbs,
from tuberous roots or taproots and sub-
terranean to superficial caudex; herbage pi-
losulose to velutinous; leaves all basal (rarely
some reduced bracteate ones along the
scape), the blades spatulate, lanceolate, ob-
lanceolate, ovate, or orbicular; heads solitary;
involucral bracts in 2 or 3 series, herbaceous
throughout; receptacle flat to convex, chaffy,
the scales clasping the achenes; rays yellow,
sterile (but apparently pistillate), or lacking;
disk flowers numerous, perfect, fertile, yel-
low; pappus of 2 awns and with or without
small scales between, or none; achenes flat-
tened, blackish.

1. Heads discoid; herbage pilose-hirsutulose; plants arising from a subterranean

tuberous root E. nutans

— Heads radiate; herbage tomentulose; plants arising from a superficial caudex 2

2(1). Petioles broadly winged, mainly shorter than the blades; plants reported from
the Virgin Narrows section of Washington County, but none have been seen by
me E. argophyllus (D. C. Eaton) A. Nels.

— Petioles slender, not or only narrowly winged, mainly longer than the blades;
plants commonly in eastern and west central portions of the state E. nudicaulis

Enceliopsis nudicaulis (Gray) A. Nels.
[Encelia nudicaulis Gray]. Scapose, caespi-
tose perennials from a superficial, branching
caudex, 10-43 cm tall, the herbage tomentu-
lose, silvery white; petioles 0.7-17 cm long,
narrowly if at all winged; leaf blades 2-9 cm
long, 1.3-10 cm wide, ovate to elliptic, or-
bicular or spatulate, cuneate to subcordate
basally, obtuse to rounded apically; scapes of-
ten with a reduced foliose bract; involucres
1.3-2.2 cm high, 3-5.6 cm wide, the bracts
ovate-lanceolate to lanceolate or linear-lan-
ceolate, attenuate to acuminate; rays 13-21,
yellow, 22-38 mm long; achenes 10-12 mm
long, long silky-pilose, cuneate, black or dark
brown; pappus commonly of 2 awns con-
nected by a crown of short connate scales (or
none). Commonly on gypsiferous semibarren
knolls in blackbrush, rabbitbrush, ephedra,
shadscale, grayia, and pinyon-juniper com-
munities in Beaver, Emery, Garfield, Grand,
Millard, Piute, San Juan, Sevier, Uintah, and
Wayne counties; Idaho, Nevada, Arizona,
and California; 66 (xiii).

Enceliopsis nutans (Eastw.) A. Nels. [En-
celia nutans Eastw.]. Scapose, discoid pe-
rermials, 10-25 cm tall, from a subterranean

caudex (2-15 cm long) and tuberous root to 4
cm thick, the herbage strigose to pilosulose
(antrorsely on the upper surface, retrorsely so
below), green; petioles 2-6.5 cm long, often
narrowly winged; leaf blades 2-7.5 cm long,
1.4-6 cm wide, ovate to orbicular or spatu-
late, cuneate basally, obtuse to rounded api-
cally; scapes not bracteate; involucres 0.9-1.5
cm high, 2.5-4 cm wide, the bracts lance-at-
tenuate; rays lacking; achenes 9-11 mm long,
oblanceolate, long silky-pilose, brown; pap-
pus lacking. Mainly in finely textured soils in
shadscale, budsage, galleta, and ephedra com-
munities at 1310 to 1830 m in Carbon, Du-
chesne, Emery, Grand, Uintah, and Wayne
counties; Colorado (a Colorado Basin endem-
ic); 35 (iii).

Erigeron L.

Annual, biennial, or perennial herbs from
caudices, rhizomes, stolons, or taproots, with
watery juice; stems decumbent to ascending
or erect, rarely prostrate; leaves alternate,
simple, entire, toothed, or pinnatifid to pal-
matifid; heads solitary or few to numerous in
corymbose or paniculate inflorescences; in-

262 Great Basin Naturalist Vol. 43, No. 2

volucral bracts equal, or slightly to definitely flattened, 2 (rarely 4-14) -nerved. Note: This

imbricate, slender, herbaceous (or scarcely is a large and complex genus. The species, al-

herbaceous) throughout; receptacle flat, though mainly distinctive, are distinguished

naked; rays white, pink, purple, bluish, or by minute features that can be interpreted

yellow, numerous, pistillate, or lacking; disk variously. The genus is a near congener of

flowers numerous, yellow or tinged reddish; both Aster and Conyza, and is not always sep-

pappus of capillary bristles, sometimes with arable from either.

an outer series of short bristles or scales; style Cronquist, A. C. 1947. Revision of the

branches with lanceolate and acute or North American species of Erigeron

triangular and obtuse appendages; achenes north of Mexico. Brittonia 6: 121-302.

1. Plants with yellow ray flowers, known from Box Elder County E. linearis

— Plants with ray flowers pink, pink-purple, blue, blue-purple, or white, but not
yellow, of various distribution 2

2(1). Plants annual, biennial, or short-lived perennials from usually slender taproots,
lacking rhizomes (except in some specimens of E. proselyticus) or woody
caudices KEY I

— Plants definitely perennial, often from rhizomes or caudices 3

3(2). Plants silvery pubescent; achenes with 4 or more nerves; involucral bracts defi-
nitely imbricate KEY II

— Plants green, or less commonly silvery pubescent; achenes with 2 nerves, or, if
with more nerves, involucral bracts subequal 4

4(3). Involucres wooUy-villous to spreading villous, or villous-hirsute with at least

some long spreading multicellular hairs KEY III

— Involucres merely glandular, glabrous, puberulent, or with appressed simple or
multicellular hairs, rarely with some spreading long hairs near the base KEY IV

Key I.

Plants armual, biennial, or short-lived perennials from slender taproots,
lacking rhizomes or woody caudices.

1. Pistillate corollas very numerous, filiform, the rays short, erect, not exceeding

the disk, or the inner ones tubular and lacking rays 2

— Pistillate corollas few to numerous (rarely lacking), the tube generally cylin-
dric, the rays well developed and spreading, rarely reduced or absent 3

2(1). Cauline leaves narrowly lanceolate to oblong, or less commonly linear; rayless
pistillate flowers present between the ray and disk flowers; inflorescence
corymbose, the peduncles curved-ascending, or the heads solitary E. acris

— Cauline leaves linear to oblong; rayless pistillate flowers lacking; inflorescence
racemose, the peduncles erect or nearly so, or the heads solitary E. lonchophyllus

3(1). Pappus of ray and disk flowers unlike, that of the disk flowers composed of
bristles and short outer setae, that of the ray flowers lacking bristles; plants
tall adventive weedy species 4

— Pappus of ray and disk flowers alike, consisting of bristles, sometimes also with
outer setae or scales; plants indigenous, low to tall 5

4(3). Foliage ample; plants mainly 6-12 (15) dm tall; pubescence of stem long and

spreading (at least below); plants introduced, weedy E. annuus

— Foliage sparse; plants mainly 3-7 dm tall; pubescence various; plants to be
sought in Utah E. strigosus Muhl.

April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 263

5(3). Plants diffusely branched, annual; leaves linear to linear-oblong; hairs of stem

short and incurved; pappus simple E. hellidiastrum

— Plants various, but seldom as above, or, if so, the pappus double 6

6(5). Disks mainly over 1 cm wide; stems commonly simple, with solitary or few

heads, and broad cauline leaves E. glabellus

— Disks mostly less than 1 cm wide; stems commonly branched, often with sever-
al to many heads 7

7(6). Stems with hairs all spreading E. divergens

— Stems with hairs appressed or ascending, or glabrous 8

8(7). Leaves entire; plants with sterile flagellate branches E. flagellaris

— Leaves pinnately lobate or toothed, or if entire then the plants lacking sterile
flagellate branches 9

9(8). Ray flowers commonly 40-80; plants psammophytes of eastern Washington

and western Kane counties E. religiosus

— Ray flowers commonly 25-40; plants of various substrates in eastern Washing-
ton and Kane counties 10

10(9). Involucres 2.5 mm high or less; stems 1-11 cm tall; peduncles sparingly villous;

plants known from seeps and moist sandstone in Zion National Park

E. sionis

— Involucres 2.5-3 mm high; stems 14-25 cm long; peduncles hirsute; plants
from limestone and sandstone outcrops in eastern Iron and adjacent Kane
counties E. proselyticus

KEY II.

Plants perennial, silvery pubescent; achenes 4 (or more) -nerved;
involucral bracts imbricate.

1. Achenes glabrous, with 8-14 nerves; caudex clothed with marcescent leaf
bases, the midribs evident in age; plants of higher elevations in southern Utah
£. canus

— Achenes more or less hairy, with 3-8 nerves; caudex lacking marcescent leaf
bases, or if these present then the midribs not evident; plants of low to
moderate elevations, more widely or otherwise distributed 2

2(1). Involucres villous-hirsute with multicellular spreading hairs, the bracts more or

less glandular apically; achenes with 3-5 nerves E. pulcherrimus

— Involucres more or less strigose with simple hairs; achenes 4- to 8-nerved 3

3(2). Basal leaves evident, tufted, persistent; heads one per stem; plants through

much of Utah E. argentatus

— Basal leaves mostly withered at anthesis, not forming a conspicuous tuft; plants
mainly of southeastern Utah E. utahensis

KEY III.

Plants perennial; achenes mostly 2-nerved; bracts mostly subequal,
villous with woolly or spreading multicellular hairs.

1. Plants with pinnatifid or palmatifid or merely lobed leaves, low-spreading,

more or less mat or clump forming, of high elevations 2

264 Great Basin Naturalist Vol. 43, No. 2

— Plants with entire leaves, or, if some of them lobed, otherwise differing; low to

tall, of various elevations 4

2(1). Plants soboliferous, the caudex divided into elongate spreading branches;

leaves merely toothed, or if lobed not as below E. vagus

— Plants not soboliferous, the caudex branches short; leaves pinnately to
palmately lobed or divided 3

3(2). Leaves pinnately lobed; plants of the La Sal Mountains E. mancus

— Leaves palmately lobed or divided; plants widespread E. compositus

4(1). Involucres long- and shaggy-villous, the hairs sometimes obscuring the bract

surface from middle to base 5

— Involucres hirsute to shortly villous, or, if long and shaggy-villous, the hairs not
obscuring the bract surface even in the lower portion 7

5(4). Plants 4-7 dm tall or more; known from southeastern Utah E. elatior

— Plants mainly 0.3-1 dm tall; distribution various 6

6(5). Hairs of involucre with black or dark purple cross-walls; basal leaves rounded

to refuse apically; plants of the La Sal mountains E. melanocephalus

— Hairs of involucre with pale cross-walls or some with bright reddish purple to
dark purple cross-walls; basal leaves acute to abruptly obtuse apically; plants

of the Uinta, Deep Creek, Tushar, and La Sal Mountains E. simplex

7(4). Cauline leaves ample, usually lanceolate or broader; plants tall, erect (more or

less asterlike) 8

— Cauline leaves usually much reduced, subulate, linear, oblong oblanceolate, or,

if broader, the plants not tall or not erect 11

8(7). Hairs of involucre with black cross-walls near their bases; rays white; plants

rare, known from the Wasatch Mountains E. coulteri

— Hairs of involucre with pale cross-walls; rays white, pink, or purple; abun-
dance and distribution various 9

9(8). Plants with cauline leaves well developed and equably distributed, only gradu-
ally reduced upward, the middle ones as large as or larger than the lower ones
E. speciosus

— Plants with cauline leaves rather abruptly reduced upward, those of the middle
smaller than the lower ones 10

10(9). Involucres glandular or viscid toward the apex; stems curved at base

E. formosissimus

— Involucres seldom if at all glandular or viscid; stems erect E. glabellus

11(7). Ray flowers lacking E. aphanactis

— Ray flowers present 12

12(11). Plants subscapose, the bracteate leaves very small; caudex branches with per-
sistent leaf bases 13

— Plants not subscapose, the leaves merely reduced upward; caudex branches
with or without persistent leaf bases 16

13(12). Stems and involucres with long, contorted villous hairs; plants of Box Elder and

Daggett counties E. nanus

— Stems and involucres strigose, pilosulose, or hispidulous, the hairs appressed or
ascending to spreading; distribution various 14

April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 265

14(13). Leaves linear; herbage strigose; rays 7-11 mm long; plants mainly of lower

elevations in the Navajo and Great basins E. compactus

— Leaves narrowly oblanceolate to spatulate; herbage strigose to pilosulose or
hispidulous; rays 4-8.2 mm long; plants of the Uinta Basin and Wasatch
Plateau 15

15(14). Involucres long-villous with spreading multicellular hairs; rays 6.8-8.2 mm

long; plants of the Wasatch Plateau E. carringtonae

— Involucres short-hispidulous; rays 4-6.5 mm long; plants of the Uinta Basin

E. untermannii

16(12). Caudex branches robust, 1-2.5 cm thick; plants of western Beaver and Wash-
ington counties E. wahwahensis

— Caudex branches mainly less than 1.5 cm thick, or, if broader, of different dis-
tribution 17

17(16). Stems spreading-hairy 18

— Stems strigose, or with ascending hairs 21

18(17). Leaves linear to linear-oblanceolate; plants without a prominent caudex,

mainly of lower elevations E. pumilus

Leaves oblanceolate to spatulate; plants with prominent caudex, of low to
middle or higher elevations 19

19(18). Stems glandular, with sand grains adhering; plants of lower elevations in

Emery and Wayne counties E. maguirei

— Stems lacking glands; plants of moderate and higher elevations 20

20(19). Stems commonly purplish at the base; leaves thin; plants of broad or other dis-
tribution E. eatonii

— Stems green throughout; leaves thickish; plants of the Uinta and Wasatch
mountains E. goodrichii

21(17). Caudex with spreading subrhizomatous branches, with numerous fibrous roots;

stems and lower leaf bases purplish; plants sod forming, of higher elevations

E. ursinus

— Caudex not subrhizomatous, seldom if ever with roots; stem and leaf bases not
purple; plants of various elevations 22

22(21). Stems decumbent, sharply bent from apex of caudex 23

— Stems ascending to erect, not sharply bent from caudex apex 24

23(22). Herbage glaucous, sparingly hairy; rays 15-22; basal leaf bases greatly ex-
panded, long-ciliate; plants of eastern Washington County E. canaani

— Herbage green, not especially glaucous, sparingly to moderately hairy; basal

leaf bases not greatly expanded, short-ciliate; plants of broad distribution ....£. eatonii

24(22). Cauline leaves moderately well developed, the basal ones linear-oblanceolate;

involucres 9-12 mm wide; plants of lower elevations E. engelmannii

— Cauline leaves much reduced, the basal ones spatulate; involucres less than 8

mm wide 25

25(24). Basal leaves acute or acutish; rays blue to red-purple; pappus subequal to disk

corollas; plants widely distributed E. tener

— Basal leaves obtuse to rounded; rays white to pink; pappus shorter than disk
corollas; plants of the Bear River Range, Cache County E. cronquistii

266 Great Basin Naturalist Vol. 43, No. 2

KEY IV.

Plants perennial, green; achenes mostly 2-nerved;
involucres mostly lacking; long, spreading multicellular hairs.

1. Plants substoloniferous; leaves spatulate; involucres mainly less than 8 mm

wide; plants of hanging gardens in San Juan County E. kachinensis

— Plants not substoloniferous; leaves various; involucres mainly over 8 mm wide;
plants not or seldom of hanging gardens, variously distributed 2

2(1). Cauline leaves ample, usually lanceolate or broader; plants tall and erect (more

or less asterlike) 3

— Cauline leaves definitely reduced upward, mostly linear to oblanceolate, or
broader in some low species; stems often spreading or decumbent 5

3(2). Rays mainly 2-3 mm wide E. peregrinus

— Rays 1-2 mm wide 4

4(3). Cauline leaves glabrous or minutely glandular, not ciliate, subequal to or short-
er than the internodes E. superbus

— Cauline leaves ciliate or otherwise pubescent, sometimes also glandular, usual-
ly longer than the internodes E. speciosus

5(2). Pubescence of the stem widely spreading or glandular-scabrous 6

— Pubescence of the stem appressed, ascending, or lacking 8

6(5). Involucre canescent with fine white hairs, sometimes also glandular .... E. caespitosus

— Involucre glandular and more or less spreading hairy or strigose 7

7(6). Stems hirsute with short spreading hairs, conspicuously decumbent; involucres

glandular and spreading hairy E. jonesii

— Stems glandular-scabrous, ascending or erect; involucres glandular (rarely
sparingly strigose) E. nauseosus

8(5). Basal leaves broadly oblanceolate or usually broader, the blade well-developed,

usually abruptly contracted to the petiole 9

— Basal leaves linear to oblanceolate or spatulate, tapering gradually to the pet-
iole 11

9(8). Rays purple; achenes 4- to 7-nerved; pappus simple E. peregrinus

— Rays various; achenes 2-nerved (occasionally more, but rays then pale and pap-
pus double) 10

10(9). Stems essentially scapose, the upper bracts linear; plants known from the

Wasatch Mountains E. garrettii

— Stems subscapose, the upper bracts oblong; plants rather broadly distributed ....
E. leiomeris

11(8). Peduncles and involucres densely glandular, not hairy; stems glabrous or essen-
tially SO; plants of the Wasatch Mountains E. arenarioides

— Peduncles not glandular, or, if so, the stem more or less hairy; involucres and
distribution various 12

12(11). Bases of basal leaves neither enlarged nor of different texture than the blades;
blades linear or linear-filiform; plants known from Cache and Daggett coun-
ties E. filifolius

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

267

— Bases of basal leaves somewhat enlarged, membranous or thickened, or other-
wise different from above; blades not linear 13

13(12). Leaves glabrous or nearly so, the hairs, if present, short and appressed .... E. leiomeris

— Leaves hairy, the hairs spreading or curved-ascending 14

14(13). Plants subscapose; cauline leaves reduced to acicular bracts; plants of the

Uinta Basin and west Tavaputs Plateau E. nematophyllus

— Plants caulescent; cauline leaves well developed 15

15(14). Stems decumbent-ascending, commonly curved at the base; basal leaves

sheathing basally; heads mainly solitary E. abajoensis

— Stems erect or nearly so; basal leaves not especially sheathing; heads mainly

2-4 E. awapensis

Erigeron abajoensis Cronq. Abajo Daisy.
Perennial herb, with a taproot and stout cau-
dex, the caudex branches clothed with brown
marcescent leaf bases; stems decumbent to
spreading at the base, 5-20 cm long, strigose
to strigulose, the hairs ascending; basal leaves
oblanceolate, 2-7 cm long, 2-6 mm wide,
more or less sheathing basally; cauline leaves
several to many, oblong to lance-oblong,
mostly 0.6-2.5 cm long, 1.5-4 mm wide;
heads solitary, less commonly 2-4; involucres
4-5.2 mm high, 7-12 mm wide, the bracts
subequal or slightly imbricate, somewhat
thickened dorsally, greenish brown, strigose
to strigulose, the hairs multicellular; rays
about 40-60, pink-purple to blue (or white),
3-8 mm long, 1-1.8 mm wide; pappus
double, the inner of 12-20 bristles, the outer
of setae or scales; achenes 2-nerved, hairy.
Pinyon-juniper, ponderosa pine, and spruce-
fir communities at 2135 to 3450 m in Gar-
field, Piute, San Juan, and Wayne counties;
endemic; 4 (i).

Erigeron acris L. Bitter Fleabane. Short-
lived perennial, with a slender taproot and
poorly developed caudex; stems erect or de-
cumbent at the base, 8-32 cm tall, spreading-
hairy and more or less glandular; basal leaves
spatulate-oblanceolate, 0.5-6.5 cm long, 2-10
mm wide, entire or sparingly toothed; cau-
line leaves several to many, oblong to nar-
rowly oblanceolate, lanceolate, or linear,
mostly 0.8-7 cm long, 1-8 mm wide; heads
solitary, or more commonly few to numerous,
on short to elongate peduncles; involucres
4.5-8 mm high, 9-17 mm wide, the bracts
imbricate, not especially thickened, green or
tinged pink apically in some, sparingly hairy

with spreading to ascending stiff multi-
cellular hairs and beset with short glandular
processes; rays numerous, pink or white,
erect, about 2-4.5 mm long, the inner pistil-
late flowers eligulate, with corolla tubular;
pappus of ca 25-35 slender barbellate white
to reddish bristles, surpassing the disk co-
rollas; achenes 2-nerved, sparingly hairy.
Lodgepole pine, spruce, and fir communities
at 2800 to 3500 m in Duchesne, Summit, and
Uintah counties; Alaska to Labrador, south to
California, Colorado, Michigan, and Maine;
circumboreal; 11 (ix). Varietal status of our
few specimens is unclear. One of the speci-
mens has few heads and has essentially eg-
landular bracts, one is monocephalus and has
glandular involucres, and the others are poly-
cephalus and have glandular involucres.
Names available are var. asteroides (Andrz.)
DC. and var. debilis Gray, but there appear
to be three rather poorly differentiated taxa
involved. Decisions as to proper names must
await further study.

Erigeron annuus (L.) Pers. Annual Flea-
bane. Plants annual, with slender taproots;
stems erect, 6-12 (15) dm tall, sparingly to
densely hirsute with long spreading hairs, be-
coming appressed upward; basal leaves com-
monly withered at anthesis, ovate to sub-
orbicular, petiolate; cauline leaves numerous,
lanceolate to oblong, mainly 1.5-8 cm long,
3-20 mm wide, serrate to entire; heads sever-
al to numerous, in a leafy inflorescence; in-
volucres 7.5-12 mm wide, 3-5 mm high, the
bracts subequal or the outer somewhat short-
er, greenish to brownish, acuminate-attenu-
ate, glandular and sparingly villous-hirsute
with multicellular hairs; rays ca 80-125,

268

Great Basin Naturalist

Vol. 43, No. 2

white (rarely bluish), 4-10 mm long, 0.5-1
mm wide; pappus double; achenes 2-nerved,
hairy. Roadsides, fields, and other disturbed
sites at 1370 to 1830 m in Utah and Wasatch
counties; widespread in the United States;
Europe; 8 (ii).

Erigeron aphanactis (Gray) Greene Hairy
Daisy. [E. concinnus var. aphanactis Gray].
Perennials with definite branching caudex;
stems decumbent to ascending or erect, 5-20
(30) cm tall, sparingly to copiously spreading-
hirsute with multicellular hairs; basal leaves
narrowly oblanceolate to spatulate, 0.5-8 cm

long, 1-6 mm wide, petiolate; cauline leaves
well developed or essentially lacking; heads
solitary or several; involucres 7-15 mm wide,
3.5-6 mm high, finely to coarsely spreading-
hirsute and sometimes also finely glandular;
bracts subequal or somewhat imbricate, slen-
der, acuminate, green or greenish brown, the
midrib thickened; pistillate flowers present,
tubular, eligulate, or sometimes with rays
shorter than the disk; pappus double; achenes
2-nerved, sparsely hairy. This species is rep-
resented in Utah by two rather weak
varieties.

1. Plants essentially scapose; corolla lobes sometimes becoming reddish or pur-
plish E. aphanactis var. congestus

— Plants with leafy stems; corolla lobes commonly yellowish

E. aphanactis var. aphanactis

Var. aphanactis Salt desert shrub, sage-
brush, pinyon-juniper, sagebrush, and moun-
tain brush communities at 1300 to 2700 m in
Beaver, Garfield, Juab, Piute, Sanpete, Se-
vier, Washington, and Wayne counties; Ore-
gon and Idaho south to California, Arizona,
and Colorado; 21 (iii).

Var. congestus (Greene) Cronq. [E. con-
gestus Greene]. Juniper-black sagebrush,
sagebrush, and aspen communities at 1830 to
2600 m in Garfield and Sevier counties; Cali-
fornia; 3 (0).

Erigeron arenarioides (D.C. Eaton) Gray
[E. stenophyUus D.C. Eaton, not H. & A.; As-
ter arenarioides D.C. Eaton ex Gray]. Pe-
rennial herbs, with definite branching cau-
dex, the caudex branches clothed with
brownish marcescent leaf bases; stems as-
cending to erect, 6-25 (30) cm tall, slender,
glabrous or glandular below the heads; leaves
glabrous or sparingly strigose, the basal ones
linear-filiform to linear-oblanceolate, 1.5-6
(8) cm long, 0.5-2 (4) mm wide, entire; heads
solitary or 2 or 3 (rarely more); involucres
7-9 mm wide, 3.7-5 mm high, the bracts im-
bricate in several series, greenish brown, fine-
ly glandular, the tips often purplish; rays
10-25, blue, 4-8 mm long, 0.8-1.8 mm wide;
pappus of about 10-16 bristles, and with a
few short setae; achenes 2-nerved, sparsely
strigose. Crevices in limestone and quartzite
outcrops, rarely in beach sand, at 1300 to
2440 m in Salt Lake, Tooele, Utah, and We-
ber coumties; endemic; 8 (0).

Erigeron rgentatus Gray Silver Daisy.
[Wyomingia argentata (Gray) A. Nels.]. Pe-
rennial herbs, with definite branching cau-
dex, the caudex branches more or less clothed
with brown marcescent leaf bases, the mid-
ribs not especially persistent; stems erect,
9-28 (40) cm tall, finely strigose and silvery
to gray-green; basal leaves tufted, spatulate
to oblanceolate, 1.5-7 cm long, 1-4 (6) mm
wide, petiolate, entire; cauline leaves re-
duced upward; heads solitary; involucres
10-18 mm wide, 5.5-9 mm high, the bracts
strongly imbricate, silvery strigose with ap-
pressed antrorse hairs; rays ca 20-50, blue,
lavender, or pink to white, 9-15 mm long,
1.6-2.8 mm wide; pappus double; achenes pi-
lose. Salt desert shrub, sagebrush, pinyon-
juniper, and mountain brush communities at
1600 to 2440 m in Beaver, Box Elder, Emery,
Garfield, Iron, Juab, Millard, Piute, Sanpete,
Sevier, Tooele, and Utah counties; Nevada,
California; 29 (iii).

Erigeron awapensis Welsh Awapa Daisy.
Perennial herbs from a branching caudex, the
caudex branches clothed with ragged brown
marcescent leaf bases; stems erect or nearly
so, 10-24 cm long, strigose, the hairs ascend-
ing; basal leaves 1.5-7 cm long, 2-8 mm
wide, not especially sheathing; cauline leaves
well developed, oblong to linear, mostly 1-4
cm long, 2-4 mm wide; heads 2-4, rarely
solitary; involucres 3-9 mm wide, 3.7-4.5
mm high, the bracts more or less imbricate,
thickened near the base dorsally, greenish,

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Welsh: Utah Flora: Compositae (Asteraceae)

269

strigulose, the hairs multicellular; rays 35-45,
pink-purple to pink (or white?), 5-6 mm
long, 0.9-1.8 mm wide; pappus apparently
simple, of 15-20 slender bristles, and with a
few inconspicuous shorter setae in some; ach-
enes 2-nerved, hairy. Pinyon-juniper and
sagebrush communities at 2135 to 2260 m in
Garfield and Wayne counties; endemic; 2 (1).

Erigeron bellidiastrum Nutt. Plants annual
(or biermial), the stems 3.5-32 (50) cm tall,
erect or ascending, often intricately
branched; herbage strigulose with incurved
multicellular hairs; leaves mainly cauline,
0.5-4 cm long, 1-3 (6) mm wide, linear to ob-
lanceolate, entire (or sparingly toothed to
pinnatifid), petiolate, becoming sessile up-
ward; heads solitary to numerous; involucres
5-11 mm wide, 3-5 mm high, the bracts hir-
tellous with spreading curved multicellular
hairs, thick, greenish, subequal, or the out-
ermost shorter; rays ca 30-70, pink or white,
4-6 mm long, ca 1 mm wide; pappus of ca 15
deciduous bristles; achenes 2-nerved, hairy.
Vanclevea-ephedra, blackbrush, and pinyon-
juniper communities at 1125 to 1830 m in
Garfield, Grand, Kane, San Juan, and Wash-
ington counties; Wyoming and South Dakota
to New Mexico and Texas; 23 (ii).

Erigeron caespitosus Nutt. Tufted Daisy.
[E. caespitosus var. laccoliticus Jones]. Pe-
rermial herbs with a branching caudex, the
caudex branches clothed with brown or
blackish marcescent leaf bases; stems decum-
bent at the base, 4-25 (30) cm tall, hirtellous
with short spreading hairs (especially above);
basal leaves oblanceolate to spatulate, gener-
ally rounded to obtuse apically, 1-9 cm long,
2-13 mm wide, 1- to 3-nerved; cauline leaves
reduced upward; heads solitary or few to sev-
eral; involucres 9-18 mm wide, 4-7 mm
high, the bracts subequal to imbricate, thick-
ened on the back, green, strigose to pilose
with multicellular hairs spreading laterally
from the midrib; rays ca 30-100, blue, pink,
or white, 5-15 mm long, 1-2 mm wide; pap-
pus double; achenes 2-nerved, hairy. Sage-
brush, pinyon-juniper, aspen, lodgepole pine,
spruce, and tundra communities at 2135 to
3570 m in Daggett, Duchesne, Emery, Gar-
field, Salt Lake, Sevier, Summit, Uintah,
Utah, Wasatch, and Wayne counties; Alaska
and Yukon south to Arizona, New Mexico,

and Nebraska; 40 (viii). This is a variable spe-
cies with many phases in Utah, each differing
in stature, nature of vesture, size of heads,
and other features that fail singly and in com-
bination as diagnostic criteria. The species in-
terfaces with E. nauseosus, E. abapensis, and
probably with other taxa.

Erigeron canaani Welsh Canaan Daisy.
Perennial herbs from a simple (or branched?)
caudex, this clothed with brown marcescent
leaf bases, the taproot prominent; stems 7-20
cm tall, decumbent to ascending, sometimes
purplish at the base, sparingly pubescent
with ascending hairs; leaves pubescent like
the stem, the basal ones tufted, 1-nerved,
1.4-9 cm long, 0.7-1 mm wide, linear, in-
volute, sharply acute, conspicuously ex-
panded and long-ciliate basally; cauline
leaves numerous, reduced upward; heads
1-3; involucres 9-13 mm wide, 5.3-6.5 mm
high; bracts imbricate, conspicuously glandu-
lar and sparingly to moderately villous-pilose
with multicellular hairs, green or variously
suffused with purple; rays 15-22, white or
pinkish, 3.5-5 mm long, 1.8-2.1 mm wide;
pappus single, of ca 20 slender bristles; ach-
enes 2-nerved, hairy. Ponderosa pine commu-
nity at 1585 to 2075 m in Washington Coun-
ty; endemic; 2 (i). The Canaan daisy is similar
in general aspect to E. eatonii (q.v.). The in-
volute linear glaucous leaves and few ray
flowers appear to be diagnostic.

Erigeron canus Gray Hoary Daisy. Pe-
rennial herbs, with branching caudex, the
caudex branches clothed with persistent leaf
bases, the marcescent midribs prominent;
stems erect or nearly so, 5-30 (35) cm tall,
appressed strigose; basal leaves oblanceolate,
mostly 1-6 cm long and 1-5 (7) mm wide,
hairy like the stems; cauline leaves reduced
upward; heads solitary (rarely up to 4 ); in-
volucres 9-16 mm wide, 5-7 mm high, the
bracts strigulose with ascending to spreading
multicellular hairs and more or less glandu-
lar, imbricate; rays ca 30-40, blue or white,
7-12 mm long, 0.8-1.4 mm wide; pappus
double; achenes ca 8- to 14-nerved. Gravelly
substrates of the Cedar Breaks (Wasatch)
Formation in ponderosa pine and sagebrush
communities at 2300 to 2500 m in Garfield
County; Wyoming and South Dakota to Ari-
zona and New Mexico; 3 (i).

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Vol. 43, No. 2

Erigeron carringtonae Welsh Carrington
Daisy. Pulvinate perennial herbs with a plu-
ricipital caudex, the branches clothed with
conspicuous brown to straw colored or ashy
marcescent leaf bases; leaves mainly basal,
thickish, 0.6-3.5 cm long, 1-5 mm wide,
spatulate to oblanceolate, strigose to pilosu-
lose, obtuse to rounded apically; scapes 2.5-8
cm tall; heads solitary; involucres 9.8-15 mm
wide, 5.8-7 mm high, the bracts imbricate,
suffused with purple or green, the inner
greenish with scarious margins, spreading-vil-
lous with long multicellular hairs; rays 18-30,
pink to pink-purple, 6.8-8.2 mm long,
1.4-2.3 mm wide; pappus double, the inner
of 25-35 barbellate bristles, the outer of short
setae; achenes 2-nerved, pilose. Meadows and

escarpment margins, commonly on Flagstaff
Limestone at 3050 to 3355 m in Emery and
Sanpete counties; endemic; 6 (i).

Erigeron compactus Blake [E. pulvinatus
Rydb.]. Perennial pulvinate herbs with a
branching caudex, the caudex branches
clothed with marcescent leaf bases; leaves
mainly basal, 4-20 mm long, 0.6-1.4 mm
wide, linear, finely strigose; scapes 2-10 cm
tall; heads solitary; involucres 7-17 mm
wide, 5-8.5 mm high, the bracts more or less
imbricate, straw colored or greenish brown
to green, hispidulous with short spreading
hairs; rays mainly 15-50, white or pink, 7-11
mm long, 1.4-2.5 mm wide; pappus double;
achenes 2-nerved. Two geographically segre-
gated races are recognizable as varieties.

Leaves yellowish green; involucral bracts appressed strigose; plants of the

Great Basin E. compactus var. compactus

Leaves grayish green; involucral bracts spreading-hispidulose; plants of the
Colorado Drainage system E. compactus var. consimilis

Var. compactus Pinyon-juniper community
at 1830 to 2135 m in Beaver, Box Elder, Mil-
lard, and Tooele counties; Nevada and Cali-
fornia; 4 (0).

Var. consimilis (Cronq.) Blake Salt desert
shrub and pinyon-juniper communities in
Daggett, Duchesne, Emery, and Wayne
counties; Arizona; 11 (i). The general aspect
of this variety is similar to that of E. pulcher-
rimus (q.v.), with which it is sympatric in
much of its range; the 2-nerved achenes and
low subscapose stems are diagnostic.

Erigeron compositus Pursh Fern-leaf
Daisy. Perennial caespitose cushion plants,
with a shortened pluricipital caudex, the cau-
dex branches densely clothed with brown
marcescent leaf bases; herbage glandular and
more or less spreading-hairy; leaves mainly
basal, mostly 2- or 3-temately lobed or dis-
sected, 0.5-7 cm long; cauline leaves few and
reduced upward, simple or ternate; stems
subscapose, 2-20 (25) cm tall; heads solitary;
involucres 8-20 mm wide, 5-10 mm high, the
bracts glandular and spreading-hairy, com-
monly purplish at the tips; rays lacking, or
developed and 20-60, blue, pink, or white, to
12 mm long and 2 mm wide; pappus simple;
achenes 2-nerved, villous-hirsute. Sagebrush,
rabbitbrush, aspen, aspen-fir, lodgepole pine,
spruce-fir, and alpine tundra communities at

2375 to 3965 m in Beaver, Box Elder, Dag-
gett, Duchesne, Emery, Garfield, Iron, Juab,
Millard, Piute, Salt Lake, Sanpete, Sevier,
Summit, Tooele, Uintah, Utah, Wasatch,
Wayne, and Weber counties; Alaska to
Greenland, south to California, Arizona, Col-
orado, South Dakota, and Quebec; 77 (xvi).
This is an extremely variable apomictic spe-
cies, with rare sexual individuals. Our mate-
rial has been assigned to var. glabratus Ma-
coun, which is separable from the type
variety only problematically.

Erigeron coulteri T.C. Porter in Port. &
Coult. Coulter Daisy. Perennial herbs from a
rhizome or caudex; stems more or less
spreading-hairy, mainly 1-6 dm tall; basal
and cauline leaves ample or the cauline ones
somewhat reduced, entire or toothed, the
largest 6-15 cm long, 1-2.5 cm wide, oblan-
ceolate to elliptic, lanceolate, oblong, or
ovate; heads solitary or 2 or 3; involucres
10-15 mm wide, 6-10 mm high, the bracts
densely white hirsute below with hairs hav-
ing purplish black cross-walls, at least near
the base, glandular to the tips; rays 40-80, ca
10-15 mm long, white to pink-purple; pap-
pus simple; achenes sparsely hairy. Moist
slopes in Salt Lake and Utah counties; Ore-
gon to Wyoming, south to California, Ne-
vada, and New Mexico; 8 (0).

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Welsh: Utah Flora: Compositae (Asteraceae)

271

Erigeron cronquistii Maguire Cronquist
Daisy. Perennial herb, with short caudex
branches clothed with brown leaf bases;
stems 1.5-7 cm long, sparingly strigose; basal
leaves 0.5-4 cm long, spatulate to oblanceo-
late or elliptic, petiolate, sparingly strigose;
cauline leaves few or wanting; heads solitary,
sometimes 2; involucres 5-8 mm wide, 3-5
mm high, glandular and spreading-hirsute,
the bracts imbricate, green, often suffused
with purple; rays 10-25, white or pale pink,
5-6 mm long, 1.3-2.1 mm wide; pappus
single, or with a few shorter outer ones; ach-
enes 2-nerved, sparingly hairy. Limestone
cliffs at 1750 to 2600 m in the Bear River
Range, Cache County; endemic; 2 (0). This
beautiful, tiny plant is a near congener of E.
tener (q.v.).

Erigeron divergens T. & G. Spreading
Daisy. [E. divaricatus Nutt., not Michx.]. An-
nual, biennial, or short-lived perennial herbs
from taproots; stems branched from the base
and above, pubescent with spreading hairs,
0.5-5 (7) dm tall; basal leaves oblanceolate to
spatulate, mainly 1-7 cm long, 2-10 mm
wide, spreading-hairy, petiolate, usually lack-
ing at anthesis; cauline leaves reduced up-
ward; heads several to numerous; involucres
7-11 mm wide, 4-5 mm high, finely glandu-
lar and hirsute with long, spreading hairs, the
bracts green, attenuate; rays ca 75-150, blue,
pink, or white, ca 5-10 mm long, 0.5-1.2 mm
wide, sometimes scarcely developed; pappus
double; achenes 2 (4) -nerved, sparsely hairy.
Riparian, rabbitbrush, sagebrush, pinyon-juni-
per, mountain brush, ponderosa pine, and as-
pen-spruce communities at 975 to 2900 m in
Beaver, Cache, Daggett, Davis, Duchesne,
Emery, Garfield, Grand, Iron, Kane, Millard,
Piute, Salt Lake, San Juan, Sevier, Tooele,
Uintah, Utah, Wasatch, Washington, and
Weber counties; British Columbia to South
Dakota, south to California, Arizona, and
Mexico; 109 (xiii). Our materials have been
segregated into two weak varieties differen-
tiated as follows: var. cinereus Gray, with
earliest flowers borne on long naked pe-
duncles and plants later with long leafy sto-
lons; and var. divergens, with earliest heads
on leafy peduncles and plants not developing
leafy stolons. The var. cinereus is evidently
rare in Utah; 5 (ii).

Erigeron eatonii Gray Eaton Daisy. [£. ea-
tonii f. molestus Cronq., type from the Stans-
bury Mountains]. Perennial herbs, from a
short simple or branched caudex, this clothed
with brown marcescent leaf bases, the tap-
root prominent; stems 5-38 cm tall, decum-
bent to ascending, usually purplish at the
base, strigose or rarely more or less hirsute;
leaves pubescent like the stem, the basal ones
tufted, 1- (or more commonly) 3-nerved,
acute, mainly 1.2-12 (15) cm long, 1-10 mm
wide; cauline leaves numerous, reduced up-
ward; heads 1-3 (7); involucres 8-15 mm
wide, 5-8 mm high, the bracts imbricate,
conspicuously glandular and more or less hir-
sute with spreading-ascending multicellular
hairs, green or the tips purplish; rays about
20-50, white to blue or pink, mainly 4-10
mm long, 1-2.5 mm wide; pappus single or
with a few short outer setae; achenes 2 (3)
-nerved. Sagebrush, mountain brush, pinyon-
juniper, ponderosa pine, aspen, spruce-fir,
and alpine tundra communities at 1890 to
3630 m in all Utah counties except for Box
Elder and Morgan; Oregon to Wyoming,
south to California, Arizona, and Colorado;
171 (xxv). This is a widespread and variable
species, with variants differing in size, in
head dimensions, and in nature of the pu-
bescence. The hirsute phase from the Stans-
bury Mountains has been designated as f. mo-
lestus Cronq.

Erigeron elatior (Gray) Greene Tall Daisy.
[£. grandiflorus var. elatior Gray]. Perennial
herbs, from a short caudex (seldom collected);
stems mainly 4-7 dm tall, often purplish be-
low, leafy throughout, spreading-hairy and
more or less glandular above; leaves mainly
2.2-10 cm long, 6-28 mm wide, the lower-
most smaller than the middle ones and com-
monly withered at anthesis, ovate-lanceolate
to lanceolate, entire, the lower petiolate, be-
coming sessile and somewhat clasping up-
ward; heads 1-3 (6); involucres 12-20 mm
wide, 7-11 mm high, the bracts densely
woolly-villous with long, flattened, shiny,
multicellular hairs, some of which may have
reddish purple cross-walls, subequal, long-at-
tenuate apically, the tips glandular, purple,
and reflexed; rays ca 75-150, pink or pink-
purple (white), 12-20 mm long, 0.8-1.6 mm
wide; pappus double; achenes 2-nerved,
hairy. Meadows and openings in mountain

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Vol. 43, No. 2

brush and spruce-fir communities at 2440 to
3050 m in the La Sal Mountains of Grand
and San Juan counties; Colorado and Wyom-
ing. This is a beautiful asterlike plant with
equably leafy stems and densely villous in-
volucres; 4 (0).

Erigeron engelmannii A. Nels. Engelmann
Daisy. Perennial herbs, with short branching
caudex, this clothed with straw-colored to
brown marcescent leaf bases; taproot defi-
nite; stems 3-24 (30) cm tall, decumbent to
erect, strigose or the hairs ascending, multi-
cellular; basal leaves 1-6 (10) cm long, 1.5-5
mm wide, linear-oblanceolate, the blades
hairy like the stems, the basal margins long
and coarsely ciliate; cauline leaves reduced
but well distributed upward; heads 1-4; in-
volucres 7-12 mm wide, 4-7 mm high, the
bracts hirsute and more or less glandular,
subequal, green, with brownish midrib and
scarious apices; rays ca 35-100, white (rarely
pink or blue), 5-12 mm long, 0.6-2 mm
wide; pappus double; achenes 2-nerved,
hairy. Salt desert shrub, sagebrush, rabbit-
brush, and pinyon-juniper communities at
1370 to 2200 m in Box Elder, Cache, Dag-
gett, Duchesne, Grand, Juab, Millard, Salt
Lake, Sanpete, Sevier, Summit, Tooele, Uin-
tah, and Utah counties; Oregon to Wyoming
and Colorado; 28 (iii).

Erigeron filifolius Nutt. Thread-leaf
Daisy. Perennial herbs, with branching
woody caudex, the caudex branches clothed
with brownish marcescent leaf bases; stems
10-30 (50) cm tall, more or less strigose;
leaves 1-8 cm long, 0.3-3 mm wide, linear or
filiform, strigose, the cauline ones distributed
along the stem but smaller than the basal
ones; heads 1-several; involucres 5-15 mm
wide, 4-6 mm high, the bracts villous to stri-
gose and commonly glandular as well, sub-
equal or somewhat imbricate, greenish; rays
ca 15-75, blue to pink or white, 3-12 mm
long, 1-2 mm wide; pappus single or with a
few outer setae; achenes 2-nerved, more or
less hairy. The species is reported from Utah
(Cache County, Logan, C. P. Smith 1737
RM) by Cronquist (1947), also Daggett Coun-
ty; British Columbia and Montana to Califor-
nia and Nevada; 1 (0).

Erigeron flagellaris Gray Trailing Daisy.
Biennial or short-lived perennials, with a

poorly developed caudex (if at all) and slen-
der taproot; herbage strigose or with spread-
ing hairs at stem base; stems 3-25 (40) cm
tall, the fertile ones terminated by a solitary
head, the sterile ones developed as leafy sto-
lons; basal leaves 1-5 cm long, 1.5-8 mm
wide, oblanceolate to spatulate; cauline
leaves smaller upward, linear to oblanceo-
late; heads solitary; involucres 7-13 mm
wide, 3.5-5 mm high, the bracts with ap-
pressed or spreading hairs, glandular, green
to purplish; rays mostly ca 50-100, white,
pink, or blue, 5-10 mm long, 0.8-1 mm wide;
pappus double; achenes 2-nerved, hairy to al-
most glabrous. Sagebrush, juniper, ponderosa
pine, aspen, spruce-fir, and alpine meadow
commimities at 1980 to 3180 m in Beaver,
Carbon, Daggett, Duchesne, Emery, Gar-
field, Grand, Iron, Kane, Millard, San Juan,
Sanpete, Sevier, Summit, Washington, and
Wayne counties; British Columbia to Ne-
vada, Arizona, and Texas; 63 (xi).

Erigeron formosissimus Greene Pretty
Daisy. [E. frucetorum Rydb.]. Perennial
herbs, with a simple or sparingly branched
subrhizomatous caudex; herbage variously
hirsute, glandular, or glabrous, the stems
more or less glandular above, mainly 1.5-3
(4.5) dm tall; basal leaves the largest, mainly
2-10 (15) cm long, 4-10 (15) mm wide, ob-
lanceolate to spatulate; cauline leaves com-
monly much reduced upward, lanceolate to
oblong or ovate; heads 1-6; involucres 10-20
mm wide, 5-8 mm high, the bracts subequal,
linear, acuminate, glandular, and more or less
hirsute; rays ca 75-150, 8-15 mm long, ca 1
mm wide, blue, pink, or white; pappus
double; achenes 2-nerved, hairy. Meadows in
aspen and mountain brush communities at
2440 to 1840 m in Grand, Iron, Salt Lake,
San Juan, and Sevier counties; Alberta south
to Arizona and New Mexico; 5 (i). The spe-
cies is poorly known in Utah (reports of the
species in Iron county are from Cronquist
1947).

Erigeron garrettii A. Nels. Garrett Daisy.
[£. controversus Greene]. Perennial sub-
scapose herbs, with branching caudex, the
caudex branches clothed with brown leaf
bases; stems 3-23 cm tall, sparingly strigose;
basal leaves 1.2-12 cm long, 3-13 mm wide,
oblanceolate to spatulate, glabrous, sparingly

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

273

ciliate; cauline leaves lacking or greatly re-
duced; heads solitary; involucres 8-17 mm
wide, 5-8 mm high, the bracts finely strigose
and obscurely glandular, moderately imbri-
cate; rays ca 20-35, white to pink, 7-13 mm
long, 1.4-2.7 mm wide; pappus double; ach-
enes 2-nerved, hairy. Moist cliff faces and
crevices at 2750 to 3570 m in Salt Lake,
Utah, and Wasatch coimties; endemic; 17 (0).

Erigeron glabellus Nutt. Smooth Daisy.
Perennial or biennial herbs with simple or
branched caudices, the caudex, when present,
clothed with brown to blackish leaf bases;
herbage strigose to hirsute; stems 1-6.5 dm
tall, erect or nearly so; basal and lower leaves
mainly 3-15 cm long, 3-18 mm wide, oblan-
ceolate, entire or toothed, petiolate; middle
cauline leaves lanceolate to linear, reduced
upward; heads 1-12 (15), borne on bracteate
peduncles; involucres 10-20 mm wide, 5-9
mm high, the bracts subequal to slightly im-
bricate, acuminate, strigose to strigulose; rays
ca 125-175, blue to pink, or white; pappus
double; achenes 2-nerved, hairy. Meadows
and stream sides at 1370 to 1770 m in Bea-
ver, Cache, Daggett, Davis, Duchesne, Salt
Lake, Uintah, Utah, and Wasatch counties;
Alaska and Yukon, south to Utah, Colorado,
South Dakota, and Wisconsin; 12 (0). This is
a tall handsome daisy of lower elevations in
Utah.

Erigeron goodrichii Welsh Goodrich
Daisy. Perennial herbs from a stout taproot
and caudex, the caudex branches with dark
brown marcescent leaf bases; stems 3-10 cm
tall, decumbent-ascending to erect, spread-
ing-hairy; basal leaves 0.4-6 cm long, 1.2-5
mm wide, narrowly oblanceolate, the veins
not apparent, pilosulose, obtuse apically;
cauline leaves more or less developed, but
much reduced upward; heads solitary; in-
volucres 10.5-18 mm wide, 6.4-7.8 mm high;
bracts imbricate, spreading villous-pilose
with multicellular hairs, thickened basally,
green or the apices suffused purplish, the in-
ner with scarious margins, the attenuate
apices more or less glandular and sometimes
spreading; rays 40-65, pink-purple to pink or
white, 6.8-10.4 mm long, 1.5-2 mm wide;
pappus apparently single, of 20-30 minutely
barbellate bristles; achenes 2-nerved, pilose.
Engelmann spruce krummholz and meadow
communities, often on rock outcrops or talus

at 3050 to 3400 m in Duchesne, Summit,
Uintah, and Utah counties; endemic; 8 (0).

Erigeron jonesii Cronq. Jones Daisy. Pe-
rennial herbs, from a branching or simple
caudex, the caudex branches clothed with
blackish or dark brown marcescent leaf bases;
herbage hirsute with short spreading hairs;
stems mainly 10-25 cm tall, conspicuously
decumbent and often purplish at the base;
basal leaves 3-nerved, 1.5-8 cm long, 3-12
mm wide, oblanceolate to elliptic or spatu-
late, petiolate, entire or toothed; cauline
leaves smaller than the basal; heads 1-4; in-
volucres 9-15 mm wide, 5-7 mm high, the
bracts glandular and spreading-hairy, slightly
thickened dorsally, more or less imbricate,
green, with tips often purplish; rays ca
25-50, blue, pink, or white, 4-8 mm long,
1.4-1.8 mm wide; pappus single or with a
few short outer setae; achene's 2-nerved,
hairy. Sagebrush, pinyon-juniper, mountain
brush, and alpine meadow communities at
1890 to 3350 m in Juab, Tooele, and Wash-
ington counties; Nevada; 5 (iii). The Jones
daisy simulates E. eatonii in habit and stature,
but the definite spreading hairs of the herb-
age are apparently definitive in most in-
stances. Possibly it would best be treated at
some infraspecific rank within E. eatonii.

Erigeron kachinensis Welsh & Moore
Kachina Daisy. Perennial herbs, from a short
thick branching or simple caudex, the caudex
branches clothed with brown marcescent leaf
bases; herbage glabrous throughout; stems
6-18 cm tall, decumbent to erect; basal
leaves 1.3-5 cm long, 2-13 mm wide, oblan-
ceolate to obovate or spatulate, the blade ta-
pering to the petiole, rounded or retuse api-
cally, entire; cauline leaves 5-11, reduced
upward; heads solitary or 2-4, the involucres
5-6 mm broad, 3.2-4 mm high, the bracts
distinctly imbricate, some purplish at the tip,
glabrous or minutely glandular; rays 10-15,
white or pinkish, 3.5-5.5 mm long, 0.9-1.1
mm wide; pappus double; achenes 2-nerved,
hairy. Seeps and hanging gardens at 1680 to
1890 m in White (type from Natural Bridges
National Monument) and Dark Canyons, San
Juan County, Utah, and Montrose County,
Colorado; 3 (ii). This distinctive dwarf daisy
is a Colorado Plateau endemic.

Erigeron leiomerus Gray Glaber Daisy.
Perennial herbs, from a branching caudex.

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Great Basin Naturalist

Vol. 43, No. 2

the caudex branches clothed with brown
marcescent leaf bases; herbage glabrous or
merely strigose; stems 4-12 (15) cm tall, de-
cumbent to erect; basal leaves 1.3-7 cm long,
2-11 (15) mm wide, oblanceolate to spatulate
or obovate, rounded to retuse apically, en-
larged and often purplish basally, glabrous or
strigose to glabrate; cauline leaves reduced
upward, usually several, becoming acutish;
head solitary, the involucres 7-13 mm wide,
4-6 mm high, the bracts somewhat imbricate,
purplish overall or at tips, finely glandular;
rays ca 15-60, purplish to blue or white,
6-11 mm long, 1.5-2.5 mm wide; pappus
double; achenes 2-nerved, short-hairy. Talus
slopes, boulder fields, and meadows in spruce
and lodgepole pine and alpine tundra com-
munities at 2950 to 3750 m in Beaver, Box
Elder, Cache, Daggett, Duchesne, Juab,
Piute, Salt Lake, Summit, Tooele, and Uintah
counties; Nevada and Idaho to Wyoming,
Colorado, and New Mexico; 27 (x).

Erigeron linearis (Hook.) Piper [Dauco-
pappus linearis Hook.]. Perennial herbs from
a pluricipital caudex, the branches of the
caudex clothed with broad clasping brownish
marcescent leaf bases; herbage strigose; stems
5-20 cm tall; basal leaves 1-9 cm long, 0.5-3
mm wide, linear to linear-oblanceolate,
acute, the bases enlarged, more or less
sheathing, straw colored and strongly ciliate;
cauline leaves reduced upward; heads soli-
tary or 2 or 3; involucres 8-13 mm wide, 4-7
mm high, strigose-villous with multicellular
hairs and more or less glandular; bracts sub-
equal to somewhat imbricate, green or green-
ish to straw colored, attenuate, thickened
dorsally; rays ca 20-45, yellow, 4-11 mm
long, 1.3-2.5 mm wide; pappus double, the
inner of 10-20 barbellate bristles, the outer
of scales; achenes 2-nerved, short hairy. Sage-
brush and juniper communities at 1675 to
2000 m in Box Elder County; British Colum-
bia, Washington, and Oregon, east to Idaho
and Wyoming, and south to Nevada; 1 (0).

Erigeron hnchophyllus Hook. Short-lived
perennial or biennial (?) herbs, with slender
taproots and subfibrous roots from a poorly
developed caudex; stems decumbent to erect,
5-55 (60) cm long, sparsely to densely
spreading-hairy; basal leaves oblanceolate to
spatulate, 1.2-11 (15) cm long, 2-12 mm
wide; cauline leaves several to many, mostly

0.6-8 cm long, 2-6 mm wide; heads few to
numerous, rarely solitary, borne on nearly
erect peduncles; involucres 4-9 mm high,
7-17 mm wide, the bracts evidently imbri-
cate, not especially thickened basally, green-
ish to brownish or yellowish, the tips com-
monly purplish, sparsely to moderately
strigulose with multicellular hairs; rays nu-
merous, white or pinkish, about 2-4 mm
long, lacking inner eligulate pistillate co-
rollas; pappus of ca 20-30 slender barbellate
white bristles, surpassing the disk corollas;
achenes 2-nerved, sparsely hairy. Marshes,
stream banks, seeps, and wet meadows at
1370 to 2900 m in Beaver, Daggett, Du-
chesne, Garfield, Grand, Juab, Piute, Rich,
Salt Lake, Sanpete, Sevier, Summit, Tooele,
Utah, and Washington counties; Alaska and
southern Yukon, south to California and New
Mexico, and east to Quebec and South Da-
kota; 39 (vii).

Erigeron maguirei Cronq. Maguire Daisy.
Perennial herbs, with a branching caudex, the
caudex branches clothed with brown to
straw-colored marcescent leaf bases; herbage
spreading hirsute; stems 7-18 cm high, de-
cumbent to sprawling or erect; basal leaves
2-5 cm long, 3-8 mm wide, oblanceolate to
spatulate, rounded apically; cauline leaves
well developed, but somewhat reduced up-
ward, becoming acutish; heads solitary or
2-5; involucres 5-6.5 mm high, 7-11 mm
wide, the bracts imbricate, not much thick-
ened, green or yellowish, the inner less pu-
bescent and with scarious purplish tips, all
finely glandular also; rays 12-20, white or
pinkish, ca 6-8 mm long, 1.1-2 mm wide;
pappus of 13-25 slender barbellate sordid
bristles, with a few shorter outer ones; ach-
enes 2-nerved, hairy. Canyon bottoms in
Wingate (?) and Navajo formations at 1640 to
1740 m in Emery and Wayne counties; en-
demic; 5 (ii). For the past four decades the
Maguire daisy was known officially from the
type locality in the San Rafael Swell in
Emery County. Now, other material has been
discovered at BRY and relocated in the field,
which is distinguishable only technically
from specimens at the type locality. These
latter specimens tend to have more heads per
stem, have narrower ray corollas, and shorter
disk corollas. All of these may be the result of
ecological responses, but they are recognized
as var. harrisonii Welsh.

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Welsh: Utah Flora: Compositae (Asteraceae)

275

Erigeron mancus Rydb. [E. pinnatisectus
(Gray) A. Nels. var. insolens Macbr. & Pays.].
Pulvinate caespitose subscapose perennials
from a usually branched caudex, the caudex
clothed with dark brown to straw-colored
marcescent leaf bases; herbage more or less
hirtellous and puberulent or minutely glandu-
lar; stems mainly 2-6 cm long, erect or as-
cending; basal leaves 1.2-4 cm long, 2-4 mm
wide, pinnatifid, the lobes lanceolate, some-
times again lobed; cauline leaves much re-
duced; heads solitary; involucres 5-6.5 mm
high, 7-12 mm wide, glandular, villous with
multicellular hairs, the bracts subequal, some-
what thickened basally, the acuminate tips
often purplish; ray flowers lacking; pappus
simple or nearly so, of 20-30 bristles; achenes
2-nerved, hairy. Alpine forb and grass-sedge
communities at 3050 to 3660 m in the La Sal
Mountains, astride the Grand-San Juan
County line; endemic; 3 (0).

Erigeron melanocephalus (A. Nels.) A.
Nels. [£. uniflorus var. melanocephalus A.
Nels.]. Perennial herbs, from a simple or
branched caudex, the caudex branches
clothed with dark brown marcescent leaf
bases; herbage more or less villous with mul-
ticellular hairs; stems commonly 5-12 cm
tall, erect; basal leaves 0.8-6 cm long, oblan-
ceolate to spatulate, rounded or retuse api-
cally; cauline leaves much reduced upward;
heads solitary; involucres 10-14 cm wide,
5-9 mm high, the bracts more or less densely
villous with multicellular hairs, the cross-
walls black or dark purple, equal, attenuate,
green, with purplish tips or purplish through-
out; rays 50-70, white or pink, 7-11 mm
long, 1.2-2 mm wide; pappus single, of ca
20-25 bristles; achenes 2-nerved, sparsely
hairy. Alpine meadows at 3355 to 3720 m in
Grand and San Juan counties (La Sal Moun-
tains); Wyoming, Colorado, and New Mexi-
co; 4 (0). Specimens from the Uinta Moun-
tains, which have involucral hairs with
purple cross-walls, have been assigned here
previously, but they seem to represent noth-
ing more than phases of E. simplex (q.v.).

Erigeron nanus Nutt. Dwarf Daisy. [E. in-
amoenus A. Nels.]. Perennial herbs, from a
branching caudex, the caudex branches
clothed with imbricate ashy to straw-colored
marcescent leaf bases; stems 3-8 cm high, vil-
lous with contorted multicellular hairs, sub-
scapose; basal leaves linear-oblanceolate.

1.2-4 cm long, 1-2 mm wide, hirtellous to
sparingly villous or glabrous, ciliate toward
base with spreading long hairs, the bases con-
spicuously enlarged; heads solitary; in-
volucres 7-13 mm wide, 5-8 mm high, long-
villous with multicellular hairs and more or
less finely glandular; bracts subequal, the
midstripe brown to purplish, the margins
green to scarious or purplish; rays 15-35,
purplish, 5-10 mm long, 1.3-2.4 mm wide;
pappus of 15-25 bristles and some outer
setae; achenes 2-nerved, hirsute. Sagebrush
and sagebrush-grass communities, often on
windswept ridges, at 2135 to 3270 m in Box
Elder and Daggett counties; Idaho and
Wyoming; 5 (0).

Erigeron nauseosus (Jones) A. Nels. Marys-
vale Daisy. [E. caespitosus Nutt. var. nau-
seosus Jones, type from near Marysvale]. Pe-
rennial herbs, from a stout-" branching brittle
caudex, the branches clothed with dark
brown marcescent leaf bases, the taproot sim-
ilarly colored; stems 6-25 cm tall, ascending
to erect, glandular-scabrous; basal leaves
2.3-10 cm long, 2-15 mm wide, oblanceolate
to spatulate, rounded apically, tapering to
the petiole, commonly 3-nerved; cauline
leaves well developed, only gradually re-
duced upward; heads solitary, rarely 2; in-
volucres 8-17 mm wide, 5-8 mm high, finely
glandular (rarely sparingly strigose as well);
bracts imbricate, somewhat thickened, often
purplish, attenuate; rays 30-60, white or
purplish, 6-12 mm long, 1.3-2 mm wide;
pappus double, the inner of 12-23 bristles,
the outer of inconspicuous setae; achenes 2-
nerved, hairy. Crevices in limestone, quartz-
ite, and igneous outcrops, and in talus in pin-
yon-juniper, sagebrush, mountain brush, and
Douglas fir- white fir communities at 1830 to
2900 m in Beaver, Garfield, Millard, Piute,
and Sevier counties; White Pine County, Ne-
vada; a Great Basin endemic; 24 (iii).

Erigeron nematophyllus Rydb. Needleleaf
Daisy. Perennial herbs, from a branching
caudex, the caudex branches clothed with
fibrous ashy to brown marcescent leaf bases;
herbage strigose to subglabrous; stems 4-15
cm tall; basal leaves 1-8 cm long, 1-3 mm
wide, linear to linear-oblanceolate, ciliate
near the enlarged sheathing base; cauline
leaves few and reduced, not especially ex-
ceeding the basal cluster; heads solitary; in-
volucres 6-13 mm wide, 4-6.5 mm high;

276

Great Basin Naturalist

Vol. 43, No. 2

bracts more or less imbricate, moderately
strigulose, green or brown, the inner often
with scarious margins and purplish tips; rays
15-55, white (less commonly pink), 4-8 mm
long, 1.2-2.3 mm wide; pappus of ca 15-25
bristles; achenes 2-nerved, shortly hairy.
Sagebrush, mountain brush, and pinyon-juni-
per communities, often on Green River
Shale, at 2280 to 2870 m in Carbon, Daggett,
Duchesne, and Uintah counties; Wyoming
and Colorado; 7 (i).

Erigeron peregrinus (Pursh) Greene
Strange Daisy. [E. callianthemus Greene; E.
peregrinus ssp. callianthemus (Greene)
Cronq.; £. regalis Greene; E. peregrinus var.
eucallianthemus Cronq.; E. peregrinus var.
scaposus (T. & G.) Cronq.; E. salsuginosus
var. scaposus T. & G.]. Perennial herbs, from
a rhizome, the rhizome sometimes short, dark
brown; stems 0.9-5.5 (7) dm tall, glabrous or
sparingly to moderately villous below, often
densely villous below the heads; basal leaves
2-16 (20) cm long, 0.8-3.2 (4.5) cm wide, ob-
lanceolate to spatulate or obovate, tapering
or abruptly contracted to the petiole, obtuse
or rounded to acute apically, glabrous or
rarely sparingly villous, ciliate; cauline leaves
reduced upward, becoming sessile and more
or less clasping; heads solitary, or 2-6; in-
volucres 12-22 (25) mm wide, 6-9 (11) mm
high; bracts subequal, reflexed at the attenu-
ate apices, glandular and purplish through-
out; rays ca 30-75, 8-17 (25) mm long, 1.8-4
mm wide, rose-purple to white; pappus of ca
20-30 bristles, sometimes with a few outer
setae; achenes 4- to 7-nerved, sparingly hairy.
Aspen, spruce-fir, lodgepole pine, and sedge
communities at 2280 to 3570 m in Box Elder,
Cache, Duchesne, Garfield, Grand (?), Salt
Lake, San Juan, Summit, Uintah, Wasatch,
Weber, and Washington counties; Alaska
south to California and New Mexico; 57 (x).
Our materials were segregated by Cronquist
(1947) into a dwarf alpine var. scaposus (T. &
G.) Cronq. and a taller montane var. eu-
callianthemus. On the basis of the rather
abundant materials at hand, there does not
seem to be any means of recognition of those
taxa, except arbitrarily. Thus, all our speci-
mens are herein considered as belonging to
ssp". callianthemus (Greene) Cronq. var.
callianthemus.

Erigeron proselyticus Nesom Professor
Daisy. [E. flagellaris Gray var. trilobatus Ma-
guire ex Cronq.]. Perennial herbs, from a sub-
rhizomatous or substoloniferous caudex, the
caudex branches with weakly persistent
brown marcescent leaf bases; stems 14-25 cm
tall, decumbent to ascending or erect, spar-
ingly strigose; basal leaves 0.5-6.5 (7.5) cm
long, 2-11 mm wide, oblanceolate to spatu-
late or linear, entire to pinnately few toothed
or lobed, glabrous to sparingly strigose, acute
to obtuse or rounded apically; cauline leaves
gradually to abruptly reduced upward, entire
or the lower few toothed; heads 3 to several;
involucres 3.5-7 mm wide, 2.5-4.5 mm high,
sparingly to moderately hirtellous; bracts
subequal, brown, suffused with purple, or the
inner greenish, with chartaceous margins;
rays 22-46, white to purplish, 5.4-8.5 mm
long, 1-1.4 mm wide; pappus double, the in-
ner of 10-19 bristles, the outer of short setae;
achenes 2- or 4-nerved, sparsely hairy.
Bristlecone pine, spruce-fir, and aspen com-
munities on sandstone and marly limestone
formation:; at 2440 to 3050 m in Iron and
Kane counties; endemic; 8 (i).

Erigeron pulcherrimus Heller Basin Daisy.
Perennial herbs, from a branching caudex,
the caudex branches with exfoliating brown-
ish bark, not especially clothed with per-
sistent leaf bases; herbage silvery or grayish
strigose; stems (5) 9-32 (35) cm tall, erect;
basal leaves 0.8-7 cm long, 1-3 (5) mm wide,
linear to linear-oblanceolate; cauline leaves
reduced upward, but generally developed to
stem middle or above; heads solitary; in-
volucres 10-20 mm wide, 6-9 mm high,
coarsely villous with spreading-ascending,
multicellular hairs, obscurely glandular api-
cally; bracts imbricate, greenish, the midrib
often brown, the margins chartaceous, acumi-
nate-attenuate, especially the inner; rays ca
25-60, white, pink, or violet, 8-15 mm long,
2-3.7 mm wide; pappus of ca 30-50 bristles,
the outer series more or less developed; ach-
enes (2-) 3- to 5-nerved, densely hairy. Salt
desert shrub and pinyon-juniper communities
on saline and seleniferous clays, clay-silts, and
gravelly pediments at 1310 to 2105 m in Car-
bon, Duchesne, Emery, Grand, Uintah, and
Wayne counties; Wyoming, Colorado, and
New Mexico. Our materials have been
treated as belonging to a wide-leaved (1.5-5

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

277

mm) var. wyomingia (Rydb.) Cronq. and a
narrow-leaved (1-1.5 mm) var. pulcherrima.
However, only arbitrary segregation appears
to be possible, and it seems best not to at-
tempt recognition of infraspecific taxa; 61
(xiv).

Erigeron pumilus Nutt. Vernal Daisy. Pe-
rennial herbs, arising from a caudex, the
branches clothed with ashy to brown marces-
cent leaf bases; herbage more or less hirsute
with spreading hairs; stems 4-50 cm tall,
leafy or subscapose; basal leaves 0.4-8 cm
long, mostly 2-5 mm wide, linear-oblanceo-
late to oblanceolate; cauline leaves well de-
veloped, somewhat reduced, or much re-
duced upward, or almost lacking; heads
solitary or few to numerous; involucres 7-15
mm wide, 4-7 mm high, sparingly to densely
spreading-villous with multicellular hairs;
bracts subequal, acuminate to attenuate,
green, with brownish midrib; rays mostly
50-100, white or pink to lavender, 6-15 mm
long, 0.7-1.5 mm wide (or more); pappus
double, the inner of 7-20 coarse bristles, the
outer of evident bristles or scales; achenes 2-
nerved, sparsely to moderately hairy. Black-
brush, shadscale, sagebrush, pinyon-juniper,
and mountain brush communities at 885 to
2960 m in all Utah counties; Washington to
Saskatchewan, south to California, Arizona,
New Mexico, and Kansas. Our highly variable
material was segregated on technical charac-
teristics by Cronquist (1947) into two sub-
species, each with two varieties. The bulk of
the Utah specimens belong to ssp. con-
cinnoides Cronq., segregated in large mea-
sure from the much less common and more
northern ssp. intermedius Cronq. by the few-
er (7-15, not 13-20) inner pappus bristles and
evidently puberulent (not glabrous or slightly
puberulent) corolla tubes. The varieties inter-
medius (var. euintermedius Cronq.) and grac-
ilior Cronq. of ssp. intermedius are only arbi-
trarily separable by stem thickness and head
number. The weakly segregated varieties
within ssp. concinnoides, var. concinnoides
(var. euconcinnoides Cronq.) and var. con-
densatus (D. C. Eaton) Cronq., differ in de-
gree of development of cauline leaves, with
the former having more equably leafy stems
and the latter tending to be subscapose. It
seems best to treat our material as belonging
to two variable taxa; ssp. intermedius and ssp.
concinnoides; 212 (xxvi).

Erigeron religiosus Cronq. Religious Daisy.
Short-lived perennial (or biennial?) herbs
from a slender taproot and poorly developed
caudex; herbage more or less strigose and
glandular below the heads; stems 6-35 cm
tall, decumbent-ascending to erect; basal
leaves 2-5.5 (7) cm long, 2-8 mm wide (or
more), oblanceolate to spatulate, entire or
some pinnately toothed or lobed; cauline
leaves gradually reduced upward; heads 2 to
numerous; involucres 5.5-7.5 mm wide, 2-3.5
mm high, sparingly to moderately villous and
more or less glandular; bracts with brown
midrib, somewhat thickened, scarious api-
cally; rays 35-85, white or pinkish, 3.4-6.8
mm long, 0.5-1.4 mm wide; pappus double,
the inner of 6-12 bristles, the outer of short
setae; achenes 2-nerved, sparsely hairy. Pon-
derosa pine-oak and pinyon-juniper commu-
nities at 1525 to 1830 m in Kane and Wash-
ington (type from Clear Creek Canyon)
counties; endemic; 9 (iv).

Erigeron simplex Greene Greene Daisy.
Perennial herbs, from a simple or branched
caudex, the caudex clothed with dark brown
marcescent leaf bases; herbage more or less
viscid-villous with multicellular hairs; stems
commonly 2-15 (20) cm tall; basal leaves
0.8-6 (8) cm long, 2-10 (13) mm wide, oblan-
ceolate to spatulate, obtuse to abruptly acute
or mucronate apically; cauline leaves re-
duced; heads solitary; involucres 8-22 mm
wide, 5-10 mm high, moderately to densely
villous and somewhat viscid, the hairs with
clear to reddish purple or purplish black
cross-walls; bracts equal, suffused with purple
or green, appressed or some reflexed; rays
50-125, blue-purple to pink (or white), 7-11
mm long, 1.2-2.5 mm wide; pappus double,
the inner of ca 10-15 barbellate bristles, the
outer of conspicuous setae; achenes 2-nerved,
sparsely hairy. Lodgepole pine, Engelmann
spruce, alpine fir, and alpine meadow and
tundra communities at 3355 to 3660 m (in
Deep Creek, Tushar, La Sal, and Uinta
mountains) in Beaver, Daggett, Duchesne,
Juab, Piute, San Juan, Summit, and Uintah
counties; Oregon to Montana, south to Ne-
vada, Arizona, and New Mexico; 30 (vi). Our
variable materials include specimens with
purplish-black cross-walls of the multicellular
hairs, especially on the involucres and below
the heads. These have been placed with the

278

Great Basin Naturalist

Vol. 43, No. 2

similar and related E. melanocephalus (q.v.),
but differ in shape of lower leaves and gener-
al aspect of the plants.

Erigeron sionis Cronq. Zion Daisy. Low
perennial herbs, with short stoloniferous
branches arising from a slender taproot;
stems 1.5-13.5 cm long, decumbent to erect,
glabrous or appressed pubescent; basal leaves
0.5-3.5 cm long, 2-10 mm wide, oblanceo-
late to obovate, entire or more commonly 3-
to 5-lobed, glabrous or sparsely strigose;
heads solitary or 2 to several; involucres 5-7
mm wide, 2-3 mm high, glandular and
sparsely to moderately spreading-hairy;
bracts suffused purplish or the inner green
with chartaceous margins; rays 23-38, white,
the midstripe below purplish, 3.9-6.1 mm
long, 1-1.6 mm wide; pappus double, the in-
ner of 7-13 bristles, the outer of slender
setae; achenes 2-nerved, sparsely pubescent.
Seeps and hanging gardens in ponderosa pine
and riparian communities in Navajo and
Wingate sandstones at 1350 to 1600 m in
Zion National Park, Washington and Kane (?)
counties; endemic; 3 (i).

Erigeron speciosus (Lindl.) DC. Oregon
Daisy. Rhizomatous perennial herbs with the
caudex more or less developed; stems 1.5-9
cm tall, ascending to erect, spreading-hairy
to subglabrous or glandular above; basal
leaves often lacking at anthesis, the lower-

most cauline ones oblanceolate to spatulate,
petiolate, commonly 5-15 cm long, 4-20 mm
wide; middle cauline leaves lanceolate to
oval, oblanceolate, or elliptic, 2-11 cm long,
5-28 mm wide; upper leaves gradually to
markedly reduced, lanceolate to obliquely
ovate, ciliate, the surfaces glabrous, spread-
ing-hairy, or glandular (or a combination);
heads 1-15 (or more); involucres 11-22 mm
wide, 5.5-9 mm high, glandular, with a few
long hairs, or more or less spreading-hairy;
bracts subequal, acuminate or attenuate, the
tips more or less spreading, often suffused
purplish; ray flowers ca 75-150, pink, pink-
or blue-purple, or white, 7-18 mm long,
0.7-1 mm wide; pappus double, the inner of
20-30 bristles, the outer of more of less evi-
dent setae; achenes 2- to 4-nerved, hairy. The
speciosus complex in Utah, as herein inter-
preted, consists of four variable, and more or
less intergrading, largely sympatric in-
fraspecific taxa. All have been treated pre-
viously at specific rank, or they have been
treated within E. speciosus, in part.
Cronquist (1947) discussed the problem of in-
termediacy in the complex but hesitated to
combine the taxa because "such a treatment
would distort the facts as well as being un-
wieldy." It is here contended that they are
unwieldy apart; it seems therefore best to
combine them as follows:

1. Leaves spreading-hairy on one or both surfaces; involucres spreading-hairy and

more or less glandular E. speciosus var. mollis

— Leaves glabrous on both surfaces or minutely glandular, or with minute stri-
gose hairs, rarely with a few spreading multicellular hairs 2

2(1). Leaves glandular on the surfaces (especially the upper ones), and also ciliate ....

E. speciosus var. uintahensis

— Leaves glabrous on both surfaces, ciliate 3

3(2). Involucral bracts merely glandular, rarely also somewhat spreading-hairy; up-
per leaves often ovate E. speciosus var. macranthus

— Involucral bracts glandular and commonly also spreading-hairy; upper leaves
lance-attenuate E. speciosus var. speciosus

Var. macranthus (Nutt.) Cronq. [£. grandi-
florus Nutt., not Hook.; E. macranthus
Nutt.]. Sagebrush, snowberry, aspen, spruce-
fir, and alpine meadow communities at 1760
to 3420 m in Beaver, Box Elder, Cache, Car-
bon, Duchesne, Emery, Garfield, Grand,
Juab, Kane, Millard, Piute, Rich, Salt Lake,
Sanpete, Sevier, Tooele, Utah, Wasatch,

Washington, and Weber counties; Washing-
ton and Alberta south to Nevada, Arizona,
and New Mexico; 104 (xv). This is our most
common phase, but it is only arbitrarily sepa-
rable from var. speciosus, to which it is com-
pletely transitional.

Var. mollis (Gray) Welsh comb. nov.
[based on: Erigeron glabellus var. mollis Gray

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

279

Proc. Acad. Nat. Sci. Philadelphia 1863: 64.
1864.]. Aspen, spruce-fir, and meadow com-
munities at 2070 to 3050 m in Carbon, Du-
chesne, Garfield, Grand, Juab, Salt Lake, San
Juan, Sanpete, Sevier, Uintah, Utah, and
Wasatch counties; Montana to South Dakota,
and south to New Mexico and Nebraska; 20
(i). This variety includes what has tradition-
ally been called E. subtrinervis Rydb.

Var. speciosus [Stenactis speciosa Lindl.].
Mountain brush, sagebrush, ponderosa pine,
aspen, spruce-fir, and alpine meadows at
2040 to 3300 m in Duchesne, Garfield,
Grand, Iron, Juab, Piute, San Juan, Sanpete,
Sevier, and Utah counties; British Columbia
and Montana, south to Nevada and New
Mexico; 23 (iv).

Var. uintahensis (Cronq.) Welsh comb,
nov. [based on: Erigeron uintahensis Cronq.
Bull. Torrey Bot. Club 70: 270. 1943]. Sage-
brush, mountain brush, ponderosa pine, as-
pen, lodgepole pine, spruce-fir, and alpine
meadow communities at 2070 to 3420 m in
Beaver, Carbon, Daggett, Duchesne, Juab,
Piute, Sanpete, Sevier, Summit, Uintah, Utah,
and Wasatch counties; Wyoming; 39 (vi).
The glandular condition of the leaves varies
in amount and position, and the Uinta phase
passes by degree into other taxa of the speci-
osus complex. Because of the intergradation
it seems best that this most distinctive por-
tion of the variation should be treated within
an expanded E. speciosus.

Erigeron superbus Greene ex Rydb. Splen-
did Daisy. Rhizomatous perennial herbs and
with the caudex more or less developed, the
perennating branches bearing brown marces-
cent leaf bases; herbage glabrous or glandular
above and villous in some below the heads;
stems 1-6 dm tall, erect; basal leaves smaller
than the cauline and commonly present at
anthesis, 3-15 cm long, 6-25 (33) mm wide,
oblanceolate to obovate or spatulate, petio-
late; middle cauline leaves somewhat smaller
than the lower ones, oblong to elliptic or lan-
ceolate, glandular (glabrous), the uppermost
sessile and glandular, rarely some denticulate,
not ciliate; heads 1-7; involucres 11-19 mm
wide, 7-10 mm high; bracts subequal,
glandular, sometimes with long spreading
hairs near the base, acuminate, sometimes
suffused purplish; rays 40-95, 1-2 mm wide.

12-20 mm long, rose-purple or white; pap-
pus double, the inner of 20-25 pinkish or
tawny bristles, the outer of setae; achenes 2-
nerved, hairy. Aspen, Douglas fir, lodgepole
pine, and spruce-fir communities at 2250 to
3050 m in Carbon, Garfield, Kane, Piute, San
Juan, Summit, and Uintah counties; Wyom-
ing south to Arizona and New Mexico; 18 (v).

Erigeron tener Gray Thin Daisy. Perennial
herbs, from a branching caudex, the slender
branches with ashy to brownish marcescent
leaf bases; herbage strigose; stems slender,
decumbent, ascending, or erect, 3-15 cm tall;
basal leaves 1-7.5 cm long, oblanceolate to
elliptic, rhombic, or obovate, petiolate, acute
to obtuse apically; cauline leaves much re-
duced; heads solitary or 2 or 3; involucres
6-10 (12) mm wide, 3.5-5 mm high, glandu-
lar and with spreading multicellular hairs;
bracts imbricate, somewhat thickened,
brownish, the inner membranous or some-
what scarious, sometimes suffused with
purple; rays ca 15-40, purplish or white, 4-8
mm long, 1-1.7 mm wide; pappus double,
the inner of 15-30 bristles, usually with slen-
der outer setae; achenes 2-nerved, hairy to
subglabrous. Sagebrush, mountain brush, pin-
yon-juniper, and white fir-Douglas fir com-
munities, often on limestone outcrops at 1980
to 2900 m in Beaver, Juab, Millard, Rich,
Sanpete, Tooele, and Utah counties; Oregon
to Wyoming south to California and Nevada;
10 (i)

Erigeron untermannii Welsh & Goodrich
Untermann Daisy. Perennial pulvinate herbs
with an intricately branched caudex, the cau-
dex branches mainly basal, 0.8-3.3 cm long,
1-4 mm wide, narrowly oblanceolate to
spatulate, pilosulose with ascending, often
curved, hairs; scapes 2-6 cm tall; heads soli-
tary; involucres 7-11 mm wide, 5-5.7 mm
high, the bracts more or less imbricate, green,
or the inner somewhat chartaceous, the mar-
gins hyaline, the tips suffused with purple
(sometimes throughout), densely hispidulous
with short spreading hairs; rays 14-26, white,
4-6.5 mm long, 1.5-2.1 mm wide; pappus
apparently single, of ca 20 slender fragile
bristles; achenes 2-nerved, pilose. Pinyon-
juniper community on calcareous shales and
sandstones of the Uinta and Green River for-
mations at 2135 to 2380 m in Duchesne and
Uintah counties; endemic; 4 (0).

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Great Basin Naturalist

Vol. 43, No. 2

Erigeron ursinus D.C. Eaton Bear Daisy.
Perennial rhizomatous sod-forming herbs, the
perennating organs arising from short super-
ficial branches clothed with brown marces-
cent leaf bases; herbage subglabrous to stri-
gose or variously ascending- or spreading-
hairy; stems ascending, 5-25 (30) cm tall;
basal leaves 1.2-12 cm long, 2-11 mm wide,
oblanceolate to oblong, commonly acute or
acutish apically, ciliate, the surfaces glabrous
or variously hairy; cauline leaves reduced up-
ward; heads solitary or 2 or 3; involucre 9-19
mm wide, 5-7 mm high, glandular and
spreading-hairy with multicellular heads;
bracts subequal, green or suffused purplish at
the usually reflexed tips; rays ca 30-100, pink
or blue-purple, 6-15 mm long, 1-2 mm wide;
pappus double, the inner of ca 10-20 bristles,
the outer of setae or scales; achenes 2-nerved,
hairy. Sagebrush, aspen, lodgepole pine, and
spruce-fir communities, often in forb-grass or
forb-sedge meadows at 2440 to 3660 m in
Beaver, Cache, Carbon, Daggett, Duchesne,
Emery, Garfield, Grand, Iron, Juab, Kane,

Piute, San Juan, Sanpete, Sevier, Summit,
Uintah, Utah, Wasatch, and Wayne counties;
Idaho and Montana, south to Nevada and
Arizona; 95 (x).

Erigeron utahensis Gray Utah Daisy. Pe-
rennial herbs from a branching caudex, the
branches with grayish marcescent leaf bases
and usually densely clothed with white vil-
lous-pilose hairs; stems 10-60 cm tall, erect,
appearing grayish or silvery due to strigose
hairs; basal and lowermost cauline leaves
1.5-10 cm long, 1-6 mm wide, linear-oblan-
ceolate, commonly withered or lacking at an-
thesis; cauline leaves gradually reduced up-
ward; heads solitary or few to many;
involucres 5-15 mm wide; 3-7 mm high, stri-
gose and often glandular apically; bracts im-
bricate, brownish, the inner with scarious
margins; rays ca 10-40, blue, pink, or white,
4-18 mm long, 1-2.7 mm wide; pappus
double, the inner of ca 5lO-30 bristles, the
outer of setae; achenes 4-nerved, more of less
pilose. Two rather weakly separable varieties
are present in Utah, as follows:

1. Stem bases not densely white-pilose; involucres mainly less than 8 mm wide;

plants uncommon E. utahensis var. sparsifolius

— Stem bases densely white-pilose; involucres commonly more than 10 mm wide;

plants common E. utahensis var. utahensis

Var. sparsifolius (Eastw.) Cronq. [E. spar-
sifolius Eastw. and Wyomingia vivax A. Nels,
both types from San Juan County] . Sandstone
outcrops in salt desert shrub and pinyon-juni-
per communities, often in shaded mesic
areas, at 1220 to 1900 m in Emery, Garfield,
Kane, and San Juan counties; Colorado and
Arizona; 9 (iv).

Var. utahensis [E. stenophyllus var. tetra-
pleuris Gray]. Creosote bush, blackbrush,
warm desert shrub, pinyon-juniper, and
Mountain brush communities at 900 to 2000
m in Emery, Garfield, Grand, Iron, Kane, San
Juan, Washington, and Wayne counties; Col-
orado and Arizona; 75 (vii).

Erigeron vagus Payson Payson Daisy.
Caespitose perennial herbs, from a diffuse
caudex, the branches commonly soboliferous;
herbage moderately villous and glandular;
leaves mainly basal, tufted at the apex of the
caudex branches, 0.5-2.5 cm long, palmately
3-toothed or -lobed; heads solitary, sub-
scapose; involucres 8-16 mm wide, 5-7.5 mm

high, spreading-hairy and more or less
glandular; bracts subequal, commonly suf-
fused purplish at the attenuate apices; rays ca
25-35, white or pink, 4-7 mm long, 1-2 mm
wide; pappus simple, of about 20 bristles;
achenes 2-nerved, sparingly hairy. Ponderosa
pine western bristlecone pine, and sedge-forb
communities at 2375 to 3660 m in Garfield,
Grand, Iron, and San Juan counties; Califor-
nia east to Colorado; 9 (0).

Erigeron wahwahensis Welsh Wah Wah
Daisy. Perennial herbs, from a branched cau-
dex, the caudex branches with conspicuous
fibrous brown to ash-colored marcescent leaf
bases; stems 15-40 cm long, decumbent to as-
cending; basal leaves 3-18 cm long, 4-13 mm
wide, linear-oblanceolate to oblanceolate or
elliptic, 3-nerved, petiolate, appressed to
spreading-hairy with curved hairs; cauline
leaves reduced, sessile, and bracteate above;
heads solitary or 2 or 3; involucres 13-17 mm
wide, 6-7 mm high, spreading-villous with
multicellular hairs, glandular apically; bracts

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

281

imbricate, green, the tips reddish, thickened
basally; rays 30-40, pink or white, 5.5-7 mm
long, 1.7-2.2 mm wide; disk corollas 3.5-4.2
mm long, the tube ca 2 mm long, the lobes
0.4 mm long; pappus of 15-20 bristles, with
inconspicuous outer setae; achenes 2-nerved,
short-hairy. Sagebrush, oak-maple, and pin-
yon-juniper communities at 1670 to 2440 m
in Beaver and Washington counties; endem-
ic; 7 (iii). The Wah Wah daisy stands be-
tween the distributions of E. pnesii and E.
eatonii, and it shares features of both. The
specimens examined from Washington Coun-
ty have appressed strigose stems, and are
highly variable. Those from the Wah Wah
Mountains have spreading hairy stems. Addi-
tional work is indicated.

Eriophyllum Lag.

Annual or perennial woolly herbs; leaves
alternate, entire or toothed to lobed; heads
solitary or corymbosely clustered; radiate;
rays few, pistillate and fertile, yellow or
white; involucres campanulate or hemispher-
ic; bracts 1 (apparently 2) -seriate, firm,
erect; receptacle flat to low-conic, naked;
disk flowers perfect, fertile, the tube glandu-
lar or hairy; pappus of firm nerveless chaffy
scales; style branches flattened; achenes 4-
angled.

Constance, L. 1937. A systematic study of
the genus Eriophyllum. Univ. California
Publ. Bot. 18: 69-136.

1. Plants perennial E. lanatum

— Plants annual 2

2(1). Rays white; pappus of unequal scales E. lanosum

— Rays yellow; pappus of equal scales or reduced to a short crown E. wallacei

Eriophyllum lanatum (Pursh) Forbes Pe- 2.5-4.5 mm long, slender, sparsely strigulose.

rennial herbs, the herbage tomentose; stems Creosote bush, blackbrush, and Joshua tree

erect or decumbent from a ligneus base, communities at 700 to 900 m in Washington

mainly 10-20 cm tall; leaves mainly 1-4 cm County; California, Nevada, and Arizona; 13

long, entire or 3- to 5-toothed or -lobed; (i).

heads solitary or corymbose on naked pe- Eriophyllum wallacei (Gray) Gray [Bahia
duncles 3-10 cm long; involucres cam- wallacei Gray]. Annual tomentose herb;
panulate, 6-10 mm wide, 6-8 mm high; stems mainly 1-8 cm tall, simple or branched
bracts 5-8 (10), carinate, distinct, the tips from the base; leaves 0.5-1.5 cm long, spatu-
erect; rays 5-8 (10), yellow, 6-10 mm long, late to obovate, entire or 3-lobed; heads soli-
2-5 mm wide; pappus of 8-10 variable tary, turbinate-cylindric, on short peduncles;
scales; achenes 2.5-4 mm long, 4-angled, var- involucres 4-6 mm wide, 5-7 mm high;
iously glabrous, hairy, or glandular. Sage- bracts 6-10, ovate, distinct; rays 5-10, yel-
brush community (reported from Utah in the low, 3-4 mm long, 2.5-3.5 mm wide; pappus
Pacific Northwest Flora); British Columbia to of 6-10 scales or none; achenes ca 2 mm
Montana, south to California, Nevada, and long, linear, hairy or glabrous. Larrea, black-
Wyoming; 0 (0). Our material likely belongs brush and Joshua tree communities at 700 to
to var. integrifolium (Hook.) Smiley. 900 m in Washington Co.; California, Ne-

Eriophyllum lanosum (Gray) Gray [Ac- vada, Arizona, and Mexico; 32 (iii).
Hnolepis lanosa Gray]. Annual floccose-to- „ ^

mentose herbs; stems mainly 2-10 cm tall,

simple and erect or branching from the base; Perennial herbs; leaves alternate, opposite,

leaves 0.5-1.8 cm long, 1-2 mm wide, linear or whorled, simple; heads discoid, the flowers

to linear-oblanceolate, entire or essentially all perfect and tubular; involucres cylindric

so; heads turbinate, solitary on named pe- to campanulate, the bracts striate, imbricate;

dimcles 0.5-5 cm long; involucres 5-6.5 mm receptacle naked, mainly flat; anthers obtuse

wide, 5-7 mm high; bracts 8-10, oblong, and entire basally, or minutely sagittate; style

acute, distinct or nearly so; rays 5-10, white, branches with short stigmatic lines and an

3-5 mm long, 2.5-3.5 mm wide; pappus of ca elongate papillate appendage; pappus of nu-

5 slender hyaline awn-tipped scales; achenes merous capillary bristles; achenes 10-nerved.

282

Great Basin Naturalist

Vol. 43, No. 2

1. Leaves alternate; plants of Box Elder County E. occidentale

— Leaves opposite or whorled; plants of various distribution 2

2(1). Leaves opposite; flowers white to cream E. herbaceum

— Leaves whorled; flowers purple or purplish E. maculatum

Eupatorium herbaceum (Gray) Greene [E. in Box Elder and Tooele counties; Washing-

ageratifolium var.? herbaceum Gray]. Pe- ton to Idaho, south to California and Nevada;

rennial herbs from a woody caudex; stems 2 (0).

4-7 dm tall, branched above; herbage „ ^
scabrous-puberulent; leaves mainly opposite,

the blades L5-6 cm long, 0.5-4 cm wide, White-tomentose annual herbs; leaves en-
ovate, the bases cordate or truncate, coarsely tire, alternate; heads discoid, small, in capi-
crenate-serrate, acute; heads numerous, in tate clusters; involucre reduced, the bracts
dense corymbose clusters; involucres 3.5-5 resembling those of the receptacle; outer
mm wide, 3-4 mm high; bracts green, pu- flowers pistillate, fertile, with tubular-flli-
berulent, subequal; corollas white; achenes form corolla, in several series, the outer
black, 1.5-2 mm long. Ponderosa pine and epappose and subtended by concave, partly
spruce-fir communities at 1585 to 2745 m in enclosing bracts, the inner bractless and with
Piute and Washington counties; California pappus of capillary bristles; central flowers
and Arizona; 2 (ii). 2-5, apparently perfect, but often sterile,

Eupatorium maculatum L. Joe-Pye Weed, bractless, with capillary bristles; achenes sub-

[£. bnineri Gray]. Robust perennial herbs terete, nerveless.

from short subrhizomatous caudices; stems Filago californica Nutt. Annual herbs, the
mainly 6-15 dm tall, branching in the in- stems erect, simple or branched above, 0.5-3
florescence; herbage puberulent and glandu- dm tall; leaves 0.8-2 cm long, narrowly ob-
lar-dotted; leaves in whorls of 3 or 4, mainly long to oblanceolate; heads ovoid, 3-4 mm
6-25 cm long and 1.5-7 cm wide, lanceolate high, subequal to involucrate leaves; bracts of
to lance-elliptic or lance-ovate, sharply ser- outer pistillate flowers 8-10, tomentose, boat
rate; heads numerous in corymbose clusters; shaped, the tips hyaline, the inner ones thin-
involucres 3.5-5 mm wide, 6.5-9 mm high, ner and less hairy, the inner florets ca 12-20;
the outer puberulent, the inner glabrous dor- inner achenes papillose. Warm desert shrub
sally, often ciliate, purplish to straw colored; at 915 to 1070 m in Washington County; Ari-
flowers purple; achenes ca 3 mm long, green zona and California; 3 (2).
to brown, glandular-dotted. River and canal p, ,
banks, wet meadows, bogs, and seeps at 1370 •'
to 1865 m in Box Elder, Cache, Kane, Uin- Annual herbs; leaves opposite, sessile, more
tah, and Utah counties; British Columbia to or less connate; heads several to numerous, in
Newfoundland, south to New Mexico, II- compact corymbose clusters; involucres cy-
linois, and Michigan. Our material belongs to lindric; bracts carinate, striate, 2-5, subequal;
var. bruneri (Gray) Breitimg; 15 (i). receptacles naked; ray flowers pistillate, fer-

Eupatorium occidentale Hook. Perennial tile, commonly 1 per head, yellowish, incon-

herbs from a rhizome and with a branching spicuous; disk flowers 2-5, perfect, fertile,

caudex; stems 1.5-7 dm tall, often branched yellowish; anthers not caudate at the base;

above; herbage scabrous-puberulent; leaves pappus none; achenes 8- to 10-ribbed,

alternate, the blades mainly 1.5-6 cm long, glabrous.

0.6-3 cm wide, deltoid or deltoid-ovate, ser- Flaveria campestris J.R. Johnst. Plants
rate or subentire; heads numerous, in com- 12-85 cm tall, simple or branched, glabrous
pact corymbose clusters; involucres 3-5 mm or hairy at the nodes; leaves 1-8 cm long,
wide and as high; bracts subequal, pu- 0.4-1.5 cm wide, lance-oblong to linear, ser-
berulent, green or suffused with purple; flow- rate to subentire, commonly 3-veined,
ers pink or purplish; achenes ca 3 mm long, glabrous; inflorescence leafy bracted; in-
brown, glandular-dotted. Rock crevices and volucres 5-8 mm high, the longer inner
talus (usually in quartzite) at 2135 to 2745 m bracts mostly 3, strongly keeled, glabrous;

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

283

rays ca 1-2 mm long; achenes black, ca 3 mm
long. Sand bars, stream banks, and seeps at
1220 to 1680 m in Grand and Tooele coun-
ties; Colorado to Missouri, south to New
Mexico and Texas; 7 (iii).

Gaillardia Foug.

Perennial (or biennial or annual) herbs;
leaves alternate or mainly basal, entire or

pinnatifid; heads radiate, the rays yellow, 3-
lobed, neuter or sometimes pistillate and fer-
tile; involucres 2- or 3-seriate, herbaceous,
more or less spreading, reflexed in fruit; re-
ceptacle convex, with numerous setae; disk
flowers perfect, fertile; anthers auricled at
the base; pappus of 5-10 scarious, awned
scales; achenes broadly obpyramidal, long-
hairy.

1. Disk flowers purple or purplish 2

— Disk flowers yellow 3

2(1). Base of involucral bracts densely long-villous or the corolla lobes 5-11 mm

long, or both; plants mainly montane in northeastern Utah G. aristata

— Base of involucral bracts not especially hairy, the corolla lobes mainly less than
5 mm long; plants of lower elevations in southeastern to south-
western Utah G. pinnatifida

3(1). Stems with well developed, pinnately dissected cauline leaves; plants of

canyon bottoms of the Tavaputs Plateau G. flava

— Stems subscapose, or, if the cauline leaves well developed, merely
toothed or lobed, and plants mainly of other distribution 4

4(3). Pappus scales broadly oblong or oval, awnless or abruptly short awned;

plants annual, reported for southern Utah G. arizonica Greene

— Pappus scales oblong-lanceolate, awned; plants perennial, rarely flower-
ing the first year 5

5(4). Leaves mainly basal, entire or rarely some of them toothed or lobed G. parryi

— Leaves cauline, toothed, lobed or entire G. spathulata

Gaillardia aristata Pursh Blanketflower.
Perennial herbs from a slender taproot; stems
20-80 cm tall, commonly foliose to middle or
above, less commonly with basal leaves only;
leaves 1.5-16 cm long, 3-25 mm wide, ob-
long to oblanceolate or elliptic, entire or
toothed to pinnatifid, puberulent and spar-
ingly long-villous with multicellular hairs;
heads solitary or few, long peduncled; disk
mainly 2-2.5 cm wide, purple; involucral
bracts (and/or peduncle apex) commonly
long-villous basally, green or suffused with
purple, attenuate; rays 6-16, yellow, often
purplish at the base, the lobes 5-12 mm long;
setae of receptacle well developed; disk co-
rollas densely woolly-villous, the hairs with
reddish purple cross-walls, often obscuring
the attenuate lobes; pappus of slender lance-
attenuate scales, the caudate apex entire;
achenes ca 1.5 mm long, ruffous-pilose. Pin-
yon-juniper, ponderosa pine, aspen, lodge

pole pine, and spruce-fir communities at
2135 to 2870 m in Daggett and Uintah coun-
ties; British Columbia to Saskatchewan, south
to Oregon, Colorado, and South Dakota; 9
(ii). A specimen by Neese (5711 BRY) is only
sparingly villous on the basal portion of the
bracts. The species is known from cultivation
in Utah and Emery counties; 3 (0).

Gaillardia flava Rydb. Perennial herbs
from a subrhizomatous woody caudex; stems
20-50 cm tall, foliose to the middle or above;
leaves 2-5 cm long, 4-25 mm wide, pin-
nately incised, minutely puberulent and
glandular-punctate; heads solitary, on pe-
duncles to 25 cm long; disk 17-32 mm wide,
yellow; involucral bracts sparingly to moder-
ately villous, green, caudate-attenuate; rays
8-12, yellow, the lobes 3-5 mm long; setae of
receptacle well developed, coarse and spi-
nescent; disk corollas sparingly villous, the
hairs with colorless cross-walls, the lobes

284

Great Basin Naturalist

Vol. 43, No. 2

acute; pappus scales oblong to oblanceolate,
abruptly contracted to a barbellate appen-
dage; achenes ca 1-1.5 mm long, yellowish
pilose. Stream terraces and valley bottoms,
commonly in cottonwood, willow, and tama-
rix communities at 1280 to 1650 m in Emery
(type from Lower Crossing) and Grand coun-
ties; endemic; 6 (v). The plants are extremely
resinous glandular, with a very bitter-fla-
vored exudate.

Gaillardia parryi Greene [G. acaulis
Gray]. Perennial herbs from a woody caudex;
stems 10-35 cm tall; foliose basally, less com-
monly with some leaves cauline; leaves 2.5-9
cm long, 8-25 mm wide, petiolate, the blades
ovate to elliptic, sparingly puberulent, mi-
nutely glandular-punctate, entire or irregu-
larly lobed, obtuse; heads solitary on scapose
peduncles; disks 17-32 mm wide, yellow; in-
volucral bracts sparingly villous, green, atten-
uate; rays ca 8-12, yellow, the lobes 3-5 mm
long; setae of receptacle copious, surpassing
achenes; disk corollas sparingly villous, hairs
with translucent cross-walls, the lobes acut-
ish; pappus scales lanceolate, rather abruptly
contracted to a smooth bristle; achenes ca 1.5
mm long, yellowish pilose. Pinyon-juniper
and ponderosa pine communities, often in
distiu-bed sites, at 1525 to 1830 m in Garfield,
Kane, and Washington counties; northern
Arizona; 7 (i).

Gaillardia pinnatifida Torr. [G. meamsii
Rydb.; G. crassifolia Nels. & Macbr., type
from LaVerkin; G. gracilis A. Nels., type
from Diamond Valley; and G. straminea A.
Nels., type from LaVerkin]. Perennial (less
commonly biennial or annual) herbs, the cau-
dex seldom well developed; stems 8-55 cm
tall, foliose to the middle, less commonly all
leaves basal; leaves 1-7.5 cm long, 2-15 mm
wide, petiolate; blades ellliptic to oblanceo-
late or linear-oblong, puberulent and minute-
ly glandular-punctate, pinnatifid to entire,
acute to obtuse; heads solitary, on long pe-
duncles; disks 15-35 mm wide, purple; in-
volucral bracts moderately to sparingly vil-
lous, green or suffused purplish, caudate-
attenuate; rays 7-12, yellow, the lobes 2-5
mm long; setae of receptacle spinescent; disk
corollas sparingly villous, hairs with trans-
lucent or reddish cross-walls, the lobes acute;
pappus scales oblanceolate, abruptly con-
tracted to a scabrous awn; achenes ca 2 mm

long, white-pilose. Blackbrush, shadscale,
ephedra-vanclevea, and pinyon-juniper com-
munities at 915 to 1830 m in Carbon, Emery,
Garfield, Grand, Kane, San Juan, Washing-
ton, and Wayne counties; Colorado and Ari-
zona to Texas and Mexico; 75 (ix).

Gaillardia spathulata Gray Perennial
herbs from a taproot and caudex; stems 6-35
cm tall, commonly foliose to middle or
above; leaves 1-7.5 cm long, 0.4-2.3 cm
wide, petiolate to sessile; blades oblanceolate
to elliptic or ovate to oval, sparingly villous
and glandular-punctate, entire or variously
toothed or lobed, obtuse; heads solitary or
few, on long peduncles; disks 18-33 mm
wide, yellow; involucral bracts moderately to
densely villous-pilose, green, lance-attenuate;
rays 7-10, yellow, the lobes 2-4 mm long;
setae of receptacle short, spinescent; disk co-
rollas shortly villous on the obtuse lobes, the
hairs with colorless cross-walls; pappus scales
oblong-lanceolate, abruptly contracted to a
scabrous awn; achenes ca 3.5 mm long, yel-
lowish pilose. Salt desert shrub and shrub-
grass communities at 1220 to 2320 m in Car-
bon, Emery, Garfield, Grand, and Wayne
counties; endemic; 58 (xi).

Geraea T. & G.

Annual herbs; leaves alternate; heads radi-
ate, showy, solitary or few in a corymbose
panicle; involucres hemispheric, 2- or 3-se-
riate; bracts white-ciliate; receptacle convex,
the bracts clasping the achenes; rays neuter,
yellow; pappus of two awns, connected by a
low whitish crown; disk achenes flat, cu-
neate, villous-ciliate, black.

Geraea canescens T. & G. Desert Sun-
flower. Annual herbs; stems 2-6 dm tall,
simple or branched, white-hirsute, glandular;
leaves 1-7 cm long, 0.8-4 cm wide, lanceo-
late to oblanceolate or ovate, acute to obtuse,
entire or few toothed, reduced upward; heads
showy, borne on slender, often bracteate pe-
duncles; involucres 10-25 mm wide, 7-12
mm high; bracts green, strongly ciliate,
lance-acuminate; rays 10-21, yellow, 7-20
mm long; achenes 6-7 mm long. Warm
desert shrub communities at 700 to 900 m in
Washington County; Nevada, Arizona, and
California; 3 (0).

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

285

Glyptopleura D.C. Eaton

Low annual herbs; leaves rosettiform, with
a few-toothed, white, crustaceous margin;
heads many, short peduncled, the flowers all
raylike, white or pale yellowish (or drying
pinkish); involucres of 7-12 scarious-

margined bracts subtended by a basal group
of pinnatifid or toothed bractlets; pappus of
capillary white bristles in several series, the
outer falling separately; achenes oblong, 5-
angled, each face with 2 rows of tubercles,
abruptly beaked.

1. Ray flowers showy, long exserted, 1.5-2.5 cm long; plants of Washington

County G. setulosa

— Ray flowers inconspicuous, only shortly exserted, mainly less than 10 mm long;

plants broadly distributed G. marginata

Glyptopleura marginata D.C. Eaton De-
pressed annual herbs; stems 0.5-4 cm long;
leaves crowded on the short stems, mainly
0.5-4 cm long, pinnatifid, the margins white-
crustose, extended into irregular white pro-
cesses; involucres 10-13 mm high, urceolate;
bracts green, the margins hyaline; bractlets
with white, irregular, branched processes,
crustose at the apex; rays mainly 4-7 mm
long, withered and pinkish on drying; ach-
enes 4-5 mm long, tan, sculptured. Desert
shrub communities at 1240 to 1590 m in Bea-
ver, Box Elder, Iron, Piute, San Juan, Sevier,
and Uintah counties; Oregon and Nevada; 9

(i)-

Glyptopleura setulosa Gray Low annual

herbs; stems 1.5-6 cm long; leaves crowded
on the short stems, mainly 0.3-5 cm long,
pirmately lobed, the margins white-crustose
into teeth; involucres 10-13 mm high, urceo-
late; bracts green or purplish tipped, the
bracts with expanded apices bearing simple

or coalescent processes; rays mainly 1.5-2.5
cm long, pale yellowish, showy; achenes 4-5
mm long, tan, sculptured. Larrea, blackbrush,
and Joshua tree communities at 700 to 915 m
in Washington County; Arizona, Nevada, and
California; 6 (0).

Gnaphalium L.

Annual or perennial tomentose herbs;
leaves alternate, entire; heads discoid, the
flowers white, yellowish, or suffused with
pink, borne in spikes, corymbs, or panicles;
involucres campanulate to ovoid; bracts im-
bricate, scarious apically (at least); receptacle
naked; outer flowers numerous, slender and
pistillate, the few inner ones broader and
perfect; style branches of inner flowers flat-
tened, truncate, the stigmatic portion not
sharply differentiated; anthers caudate; pap-
pus of capillary bristles; achenes small,
nerveless.

1. Heads large, mostly 4-7 mm high; clusters of heads not or rarely surpassed by

leafy bracts; plants often over 20 cm tall 2

— Heads small, the involucres 2-4 mm long; clusters of heads commonly sur-
passed or equaled by leafy bracts; plants mainly 4-20 cm tall 3

2(1). Leaves strongly decurrent; bracts of involucre yellowish or fading yellowish

G. chilense

— Leaves not strongly decurrent; bracts of involucre pearly white G. wrightii

3(1). Leaves spatulate to oblong, mainly 3-8 mm wide; plants loosely tomentose

G. palustre

— Leaves linear to narrowly oblanceolate, mainly 1-3 mm wide; plants rather
closely tomentose 4

4(3). Leafy bracts commonly less than 1.5 cm long, more loosely tomentose than the

following G. exilifolium

— Leafy bracts commonly more than 1.5 cm long, the tomentum appressed

G. uliginosum

286

Great Basin Naturalist

Vol. 43, No. 2

Gnaphalium chilense Spreng. Annual or
biennial herbs, the tomentose stems 15-40
cm tall or more; leaves 1.5-7 cm long, 2-8
mm wide, oblong to linear or the lowermost
oblanceolate, decurrent, tomentose, reduced
upward; heads numerous, in capitate clusters
at stem apices; involucres 4-7 mm high, the
bracts yellowish, tomentose only at the base.
Disturbed, often moist sites at 1370 to 1770
m in Daggett, Duchesne, Kane, Salt Lake,
and Utah counties; British Columbia to Mon-
tana, south to California, Arizona, and Texas;
5(0).

Gnaphalium exilifoUum A. Nels. [G. grayi
Nels. & Macbr.]. Annual herbs; stems 8-25
cm tall, simple or branching from the base,
tomentum appressed or somewhat loose;
leaves 0.4-4 cm long, 1-3 mm wide, linear to
linear-oblanceolate; heads clustered, in capi-
tate cymes or spicate, subtended by leafy
bracts that surpass them; involucres ca 3 mm
high; bracts with hyaline brownish tips, to-
mentose at the base. Sedge-grass commimity,
known in Utah from Wasatch (Lewis sn 1975
BRY) and Washington (Albee 2936b BRY);
Colorado, New Mexico, and Arizona; 2 (0).
This plant simulates G. uliginosum, with
which it has been synonymized by some
workers. More material is necessary to pro-
vide a definitive solution as to its proper tax-
onomic position.

Gnaphalium palustre Nutt. Annual herbs;
stems 3-20 (30) cm tall, simple or more com-
monly much branched, loosely tomentose;
leaves 1-3.5 cm long, 2-6 (10) mm wide, ob-
long to oblanceolate; heads clustered in capi-
tate terminal or axillary cymes, subtended by
leafy bracts that equal or surpass them; in-
volucres 3-4 mm high; bracts brown, usually
with whitish tips, tomentose below. Tamarix-
willow, mountain brush, ponderosa pine,
Douglas-fir, and sedge-grass communities, of-
ten on sand bars, lake shores, and pond mar-

gins, at 1370 to 2600 m in Cache, Garfield,
Iron, Juab, Millard, Piute, Salt Lake, Sanpete,
Sevier, Tooele, Uintah, Utah, Washington,
and Wayne counties; British Columbia and
Alberta, south to California and New Mexico;
31 (iii).

Gnaphalium uliginosum L. Annual herbs;
stems 3-15 (25) cm tall, simple or more com-
monly much branched, closely tomentose;
leaves 1-5 cm long, 1-3 mm wide, linear to
linear-oblanceolate; heads clustered in capi-
tate terminal or axillary cymes, subtended by
leafy bracts that much surpass them; in-
volucres 3-4 mm high; bracts brown with
pale tips, tomentose below. Lake and pond
margins and other disturbed sites at 2410 to
2830 m in Garfield and Sevier counties; 4 (0).

Gnaphalium wrightii Gray Perennial
herbs; stems 3-8 dm tall, branched in the in-
florescence; leaves 1.5-7 cm long, lance-lin-
ear, the lower ones spatulate; panicle open,
with capitate clusters of heads not subtended
or surpassed by bracteate leaves; involucres
5-6 mm high; bracts pearly white, tomentose
below. Ponderosa pine and live oak commu-
nities at 1585 to 1830 m in Washington
County; California to Texas, south to Mexico;
3(0).

Grindelia Willd.

Annual, biennial, or perennial herbs, some-
times woody at the base; leaves alternate,
simple, more or less resinous-punctate, usual-
ly sessile, often clasping; heads radiate or dis-
coid, the rays 10-45, pistillate, fertile, yel-
low; involucres imbricate, more or less
resinous; bracts thickish, with pale appressed
base and often squarrose or revolute her-
baceous tips; receptacle naked, flattish; disk
flowers fertile, yellow; style branches with
slender hispidulous appendages; pappus of
2-8 stiff, often curved, deciduous awns; ach-
enes compressed to angular, glabrous.

1. Heads discoid 2

— Heads radiate 3

2(1). Plants perennial; involucral bracts much thickened apically G. fastigiata

— Plants annual or biennial; involucral bracts only somewhat thickened

G. aphanactis

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

287

3(1). Involucral bracts, at least middle and upper ones, with appressed or erect tips,

these not revolute or thickened G. laciniata

— Involucral bracts spreading or recurved apically, often thickened apically 4

4(3). Rays mostly 12-25, rarely more; leaves entire or sharply toothed, not callous-
serrulate; achenes usually with one or more knobs on the apical margin; plants
perennial G. nana

— Rays mostly 25-40, rarely fewer; leaves regularly callous-serrulate to sharply
toothed or entire; achenes mainly lacking apical knobs; plants biennial or
perennial G. squarrosa

Grindelia aphanactis Rydb. Biennial
herbs; stems 1.5-9 dm tall, uniformly leafy,
glabrous; leaves mainly 2.5-7 cm long, 2-12
mm wide, oblong or oblanceolate, entire,
crenulate-serrate or denticulate to pinnatifid,
glabrous, the margin scabridulous; heads dis-
coid, campanulate; involucres 7-20 mm high,
10-28 mm wide, resinous, mostly in 5 or 6
series, the upper portion loosely to moder-
ately reflexed, glabrous; pappus awns 2 or 3;
achenes 2.3-3 mm long, brown, mainly trun-
cate apically. Weedy species of disturbed
sites in Kane and San Juan counties; Colo-
rado, Arizona, and Texas; 1 (0).

Grindelia fastigiata Greene Perennial
herbs; stems 5-10 dm tall or taller, glabrous;
leaves mainly 1.5-13 cm long, 10-18 mm
wide, oblanceolate to lance-oblong, entire or
denticulate to dentate or serrate, glabrous;
heads discoid; involucres campanulate, 10-14
mm high, 9-17 mm broad; bracts in ca 6
series, only the upper third or fourth spread-
ing, with revolute, thickened tips; pappus
awns 2 or 3; achenes oblong, 3.5-5 mm long.
Sandy terraces and washes at 1125 to 1375 m
in Emery, Grand, and San Juan counties; Col-
orado; 9 (iv); a Plateau endemic.

Grindelia laciniata Rydb. Perennial herbs;
stems 2.5-4.5 dm tall, glabrous; leaves mainly
2-6 cm long, 3-1.5 mm broad, pinnatifid or
the upper subentire or entire, narrowly ob-
lanceolate to oblanceolate, glabrous; heads

radiate; involucres 7-10 mm high and wide;
bracts with upper one-third to one-half
spreading, glabrous; pappus awns 3-5; ach-
enes 2.5-3.5 mm long. Sandy washes in San
Juan County (type from San Juan County);
Arizona; 1 (0).

Grindelia nana Nutt. Low Gumweed. [G.
brownii Heller; G. nana f. brownii (Heller)
Steyermark]. Perennial herbs; stems 0.8-6.5
(8) dm tall, glabrous; leaves mainly 1.5-10
cm long, 5-30 mm wide, oblanceolate,
scarcely clasping; heads radiate; involucres
campanulate; bracts in 5-7 series, reflexed or
revolute in the upper third to fifth; rays
11-28, yellow, 5-11 mm long; pappus awns
2; achenes 3.5-4 mm long. Ruderal weed at
ca 1585 to 1650 m in Cache County; Wash-
ington to Montana, south to California and
Idaho; 4 (0).

Grindelia squarrosa (Pursh) Dunal Curly
Gumweed. [Donia squarrosa Pursh; G. squar-
rosa f. depressa Steyermark, type from Salt
Lake County]. Perennial or biennial herbs;
stems 1-8 (10) dm tall, glabrous; leaves most-
ly 2-5 cm long, oblong, regularly callous
toothed, sometimes sharply toothed or entire,
the upper clasping; heads radiate, strongly
resinous; bracts with the green tips strongly
rolled back; rays 25-40, yellow, 7-15 mm
long; pappus awns 2 or 3 (to 6); achenes
2.3-3 mm long. Two more or less distinctive
varieties are present in Utah.

Main upper cauline leaves 2-4 times longer than broad, oblong-ovate to

oblong G. squarrosa var. squarrosa

Main upper cauline leaves 5-8 times longer than broad, narrowly oblong to
oblanceolate G. squarrosa var. serrulata

Var. serrulata (Rydb.) Steyermark [G. ser-
rulata Rydb.]. Salt desert shrub, sagebrush,
saline meadow, and mountain brush commu-
nities at 1310 to 1420 m in all Utah counties;

Wyoming south to New Mexico and Arizona,
and introduced widely elsewhere; 72 (vii).

Var. squarrosa [G. serrulata f. depressa
Steyermark, type from west of Salt Lake

288

Great Basin Naturalist

Vol. 43, No. 2

City]. Waste places at 1300 to 2135 m in naked or bristly, convex; disk flowers few to

Duchesne, Juab, Salt Lake, Utah, and many, yellow, perfect or sterile; pappus of

Wasatch counties; widespread mainly to the 10-12 unequal scales; achenes obovoid or ob-

east of our area; 6 (0). long, pubescent.

Lane, M. 1982. Generic limits of Xantho-

C T FRRF7IA T ap cephaluTTi, Gutierrezia, Amphiachris,

Gymnosperma, Greenella, and Thurovia
Perennial shrubs or subshrubs, glutinous, (Compositae: Asteraceae). Systematic
glabrous or hirtellous; leaves alternate, linear, Botany 7: 405-417.
often punctate; heads radiate, small, numer- Solbrig, O. T. 1960. Cytotaxonomic and
ous; rays pistillate or neutral, yellow, or lack- evolutionary studies in the North Amer-
ing; involucres cylindric to turbinate, the ican species of Gutierrezia (Compos-
bracts imbricate, chartaceous; receptacles itae). Contr. Gray Herb. 188: 1-63.

1. Heads cylindric, the ray and disk flowers 1 or 2 each G. microcephala

— Heads turbinate, with more than 4 flowers 2

2(1). Ray and disk flowers 3-8 each; involucres 2-3 mm thick; heads often clus-
tered at ends of branches; plants widespread G. sarothrae

— Ray flowers 4-10, disk flowers 5-23; involucres 2-7 (9) mm thick; heads soli-
tary or in pairs at branch ends; plants of restricted distribution 3

3(2). Disk flowers 5-12, 3.5-4.5 mm long; ray flowers 2-5 mm long; plants of

Uintah County G. pomariensis

— Disk flowers 15-23, ca 3 mm long; ray flowers 5-7 (10) mm long; plants of
eastern Millard County G. petradoria

Gutierrezia microcephala (DC.) Gray cephalum petradoria Welsh & Goodrich Brit-
Thread Snakeweed. [Brachyris microcephala tonia 33: 301. 1981]. Goldenrod Snakeweed.
DC; G. sarothrae var. microcephala (DC.) Perennial, suffrutescent; stems herbaceous ex-
Benson; Xanthocephalum microcephalum cept at the base, hirtellous, simple below the
(DC.) Shinners]. Rounded shrub, 30-100 cm inflorescence, loosely caespitose, from a stout
tall; stems slender, grayish to straw colored taproot and branching, mostly underground,
or green above, from a woody crown; leaves woody caudex; leaves arranged singly along
dimorphic, the cauline 2-5 cm long, 2-4 mm the stems, linear, 0.5-4.5 cm long, 1-3 (4)
wide, linear or linear-lanceolate, and with mm wide, reduced upward, secondary fas-
shorter, narrower fasciculate axillary ones, of- cicled leaves in some lower axils; heads soli-
ten one or both lacking at anthesis; heads tary or in pairs on bracteate peduncles, or
clustered at branch ends, sessile; involucre some almost sessile; involucres 5-9 mm high,
3-4 mm long, 1-1.5 mm wide, cylindric; 3-7 mm wide (to 9 when pressed), cam-
bracts fewer than 10, lanceolate, the tip panulate, the bracts ca 20, in 3 (4) series,
greenish, slightly thickened; ray flowers 1 or greenish, the tips thickened; ray flowers

2, 3-4 mm long; disk flowers 1-3, 2-3 mm 4-10, 5-10 mm long, 1-4 mm wide, when
long; pappus of ca 8 scales; achenes of disk fresh; disk flowers 15-23, ca 3 mm long; pap-
flowers abortive, those of ray flowers fertile, pus scales ca 10-12; achenes 3-4 mm long,
2-3 mm long, hairy. Blackbrush, vanclevea- pubescent, abortive in disk flowers. Sage-
ephedra, saltbush, purple sage, rabbitbrush, brush, oakbrush, mountain mahogany, and
and pinyon-juniper communities at 850 to white fir communities at 1920 to 2590 m in
1830 m in Emery, Garfield, Grand, Juab, eastern Millard County (Canyon and Pavant
Kane, Millard, San Juan, Utah, Washington, ranges); type from the Canyon Mountains;
and Wayne counties; Nevada and California endemic; 9 (0).

to Colorado, south to Texas and Mexico; 27 Gutierrezia pomariensis (Welsh) Welsh

(viii). comb. nov. [based on: Gutierrezia sarothrae

Gutierrezia petradoria (Welsh & Good- var. pomariensis Welsh Great Basin Nat. 30:

rich) Welsh comb. nov. [based on: Xantho- 19. 1970; Xanthocephalum sarothrae var.

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

289

pomariense (Welsh) Welsh]. Orchard Snake-
weed. Rounded subshrubs; stems 1.2-4.5 dm
tall, several to many from a persistent woody
base; leaves 1.5-5.2 cm long, 0.5-2.5 mm
wide, linear, entire, glabrous or scabrous,
glandular- punctate; heads in corymbose in-
florescences, solitary or 2 or 3 clustered at
stem ends; involucres 5-7.5 mm high, 2-5
mm broad, turbinate to cylindric; bracts
broadly obtuse, with a greenish subapical
spot, resin coated; ray flowers 5-9, the co-
rollas 2-5 mm long; disk flowers 5-12, the
corollas 3.5-4.5 mm long; pappus scales ca
5-8; achenes 1-2 mm long, hairy. Mixed
desert shrub community at 1460 to 2135 m in
Duchesne and Uintah (type from Dinosaur
National Monument) counties; endemic; 17
(iv).

Gutierrezia sarothrae (Pursh) Britt. & Rus-
hy Broom Snakeweed. [Solidago sarothrae
Pursh; Xanthocephalum sarothrae (Pursh)
Shinners]. Rounded shrubs; stems 9-90 cm
tall, profusely branched from the base, other-
wise in the inflorescence, from a woody cau-
dex and stout taproot; leaves dimorphic, the
main cauline ones 2-7 cm long, 1-3 mm
wide, linear to linear-lanceolate, the fascicled
secondary ones in lower axils, entire, glabrous
to tomentulose; heads in corymbose in-
florescences, usually in clusters of 3-10 at
branchlet ends, seldom solitary; involucres

3-4.5 mm high, 2-3.5 mm wide, turbinate;
bracts narrow, acute, with green thickened
tip; ray flowers 3-7, yellow, 2-5 mm long;
disk flowers mostly 3-8, 2-3 mm long; pap-
pus of 8-10 scales; achenes 1-2 mm long,
hairy. Warm desert shrub, sand sagebrush,
live oak, sagebrush, rabbitbrush, mountain
brush, and pinyon-juniper communities, often
in disturbed sites, at 760 to 2440 m in prob-
ably all Utah counties; British Columbia east
to Saskatchewan and south to Mexico. Our
variable material adjusts to disturbances and
increases on grazed native rangelands; it is
not considered to be palatable; 208 (xlv).

Haplopappus Cassini

Annual or perennial herbs, subshrubs, or
shrubs, usually resinous or glandular; leaves
alternate, entire or toothed to lobed; heads
discoid or radiate, usually small to large, vari-
ously clustered or solitary; involucres cylin-
dric to turbinate or campanulate, the bracts
imbricate, not aligned; receptacle flat to con-
vex, naked; rays yellow when present; disk
flowers perfect, yellow; pappus of barbellate
capillary bristles; achenes angled or striate to
smooth.
Hall, H. M. 1928. The genus Haplopappus.

Carnegie Institution of Washington.

391 pp.

1. Plants low, rounded, branched shrubs, or tall slender shrubs or subshrubs 2

— Plants aimual or perennial herbs, branched or unbranched 11

2(1). Heads borne on stems 2.5-5 dm long; plants of saline sandy drainages or sand-
stone outcrops in southern Utah 3

— Heads borne on stems less than 2.5 dm long; plants of various substrates and
distribution 4

3(2). Plants definitely shrubby, the mature branchlets ashy gray or white, of sand-
stone outcrops and canyons H. scopulorum

— Plants shrubby only at the base, the branchlets straw colored to greenish;
plants of saline drainage bottoms and terraces H. drummondii

4(2). Stems of the season white-tomentose; involucres 10-13 mm long, the bracts

only somewhat imbricate; plants commonly of high elevations H. macronema

— Stems glabrous, glandular, or hairy, not tomentose; involucres mainly less than

10 mm long, but, if longer, otherwise differing 5

5(4). Involucres campanulate, 8-12 mm long; heads showy, the rays 8-10 mm

long; plants of lower elevations in Washington County H. linearifolius

— Involucres turbinate to cylindric or campanulate, commonly less than 8 mm
long; heads not especially showy, the rays mainly 2-5 mm long, or lacking 6

290 Great Basin Naturalist Vol. 43, No. 2

6(5). Leaves densely glandular punctate, linear; ray flowers present; plants known

from Washington County H. laricifolius

— Leaves not glandular-punctate, narrowly to broadly oblanceolate or oblong;

ray flowers lacking; plants more broadly or otherwise distributed 7

7(6). Heads 12-22 mm high, the bracts subequal, the outer herbaceous and the inner
chartaceous and with broad hyaline margins; plants of limestone outcrops in
the Paunsagunt and Markagunt plateaus, or mainly of igneous outcrops in
the Pine Valley Mountains 8

— Heads 5.5-9.5 mm high, the bracts imbricate in several series, variously her-
baceous or chartaceous, but seldom any with broad hyaline margins; plants of
various substrates and distribution 9

8(7). Involucral bracts 1-nerved; achenes evenly though sparingly hairy; plants of

the Pine Valley Mountains, Washington County H. crispus

— Involucral bracts 3-nerved; achenes glabrous except for a few hairs apically;
plants of the Paunsagunt and Markagunt plateaus H. zionis

9(7). Leaves densely stipitate-glandular, oblanceolate, acute, the margins not espe-
cially repand H. watsonii

— Leaves lacking stipitate glands, narrowly oblanceolate to oblong, or, if oblan-
ceolate, the margins repand-undulate 10

10(9). Leaves oblanceolate, 2-5 mm wide H. cervinus

— Leaves narrowly oblanceolate to oblong, 0.5-2 mm wide H. nanus

11(1). Leaves strongly 3-nerved and veiny, thick and leathery; caudices thick, woody,

branched 12

— Leaves not 3-nerved and veiny; caudices simple, or, if branched, not woody 13

12(11). Bracts obtuse to rounded or less commonly acutish, strongly imbricate; plants

of the Colorado drainage system H. armerioides

— Bracts acute to attenuate, subequal to strongly imbricate; plants widespread ....
H. acaulis

13(11). Leaves with lobes or teeth spinulose tipped; involucral bracts spinulose tipped 14

— Leaves entire or toothed, but then not spinulose tipped; involucral bracts not
spinulose tipped 15

14(13). Involucral bracts glabrous or glandular dorsally; leaves pinnatifid; plants

perennial H. spinulosus

— Involucral bracts strigose dorsally, also minutely ciliate; leaves lobed or merely
toothed to entire; plants annual H. gracilis

15(13). Stems mainly 20-40 cm tall, loosely tomentose above; involucres 15-30 mm

wide; plants evidently rare H. croceus

— Stems 5-20 cm tall, or, if taller, not or seldom loosely tomentose, or the heads
smaller 16

16(15). Heads racemosely or spicately arranged; stems erect or nearly so, not strongly

bent at the base H. racemosus

— Heads solitary or corymbosely (rarely racemosely) arranged; stems strongly
bent at the base 17

17(16). Involucres 12-15 mm high, 20-30 mm wide; plants not hairy in the leaf axils ...

H. dementis

— Involucres 5-10 mm high, 10-20 mm wide, or, if larger, the plants with
hair tufts in basal leaf axils 18

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

291

18(16). Involucral bracts herbaceous throughout; achenes glabrous; plants rare, known

only from the Tushar Mountains H. apargoides

— Involucral bracts herbaceous only apically; achenes hairy; plants locally com-
mon in saline meadows H. lanceolatus

Haplopappus acaulis (Nutt.) Gray Stem-
less Goldenweed. [Chrysopsis acaulis Nutt.;
C. caespitosa Nutt.]. Perennial caespitose
herbs from a thick ligneous pluricipital cau-
dex and stout taproot, the caudex branches
clothed with brown to ashy marcescent leaf
bases and leaves; herbage resinous, scabrous
to glabrous; stems mainly 5-20 cm tall; basal
leaves 0.3-6 cm long, 1.5-10 mm wide, rigid,
narrowly to broadly oblanceolate, sharply
mucronate, 1- to 3-nerved; cauline leaves
few, developed or reduced upward; heads

solitary (rarely 2); involucres hemispheric,
6-10 mm high, 8-20 mm wide; bracts in 3
series, more or less mucronate; rays 6-15,
8-12 mm long, 2-4 mm wide; pappus white
to brownish; achenes silky-villous or
glabrous. This is a variable taxon, with sever-
al morphological phases. Despite the tenden-
cy for some of the variations to be correlated
geographically, it seems best to regard our
materials as consisting of two mainly sym-
patric varieties.

1. Cauline leaves well developed, often the main foliage leaves; herbage merely

resinous-glandular; plants of the Great Basin H. acaulis var. glabratus

— Cauline leaves usually much reduced, surpassed in size by the basal ones; her-
bage scaberulous or merely resinous-glandular; plants more widely
distributed H. acaulis var. acaulis

Var. acaulis Sagebrush-grass, pinyon-juni-
per, mountain brush, ponderosa pine, western
bristlecone, and spruce-fir communities at
1430 to 2685 m in Beaver, Box Elder, Cache,
Daggett, Emery, Garfield, Juab, Kane, Mil-
lard, Sanpete, Sevier, Summit, Tooele, Uin-
tah, and Utah counties; Oregon to Wyoming,
south to California, Nevada, and Colorado;
60 (ix). There is a narrow-leaved glabrous
phase of this taxon in the southern portion of
Duchesne County, mainly on Green River
Shale. Possibly it deserves recognition at
some taxonomic rank.

Var. glabratus D.C. Eaton [Stenotus fal-
catus Rydb., type from Iron County; S. lati-
folius A. Nels., type from Utah County].
Black sagebrush, wildrye, pinyon-juniper,
mountain brush, and grass-shrub communities
at 1525 to 2900 m in Beaver, Iron, Juab, Mil-
lard, Tooele, and Utah counties; Saskatche-
wan south and west to California and Ne-
vada; 22 (iv).

Haplopappus apargoides Gray Perennial
shortly caulescent herbs, 3-8 (15) cm tall,
from a taproot and simple or branched cau-
dex, this clothed with brown marcescent leaf
bases; basal leaves mainly 2-6 cm long, 2-6
mm wide, lanceolate to narrowly oblanceo-
late; cauline leaves reduced upward, sessile.

the margins scabrous or ciliate; herbage spar-
ingly long-villous with multicellular hairs;
heads solitary; involucres hemispheric, 8-12
mm high, 10-14 mm broad; bracts imbricate,
lanceolate to oblong, acute, cuspidate, her-
baceous almost or quite to the base, glabrous
dorsally, the margins long-ciliate; ray flowers
15-40, yellow, 8-15 mm long; pappus tawny;
achenes glabrous. Alpine tundra community
at 3355 m in Piute County (Tushar Moun-
tains); California and Nevada; 1 (i). The spec-
imen examined (Welsh and Thorne 12982
BRY) is tentatively assigned to this species,
which is known otherwise only from the east-
ern Sierra Nevada and adjacent Nevada.

Haplopappus armerioides (Nutt.) Gray
[Stenotus armerioides Nutt.]. Perennial
caespitose herbs from a thick ligneous plu-
ricipital caudex and stout taproot, the caudex
branches clothed with brown to ashy marces-
cent leaf bases and leaves; herbage resinous-
glandular, otherwise glabrous or with
scabrous leaf margins; stems 0.5-20 cm tall;
basal leaves 1.5-8 cm long, 1.5-10 mm wide,
rigid, linear to oblanceolate, sharply mu-
cronate; 1- to 3-nerved; cauline leaves few,
reduced upward; heads solitary (rarely 2); in-
volucres campanulate, 8-13 mm high, 10-18
mm wide; bracts in 3 or 4 series, imbricate.

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Great Basin Naturalist

Vol. 43, No. 2

oblong to oval or obovate, obtuse, sometimes
lobed below the apex, greenish near the apex,
glabrous; rays 8-12, 10-12 mm long, yellow,
3-5 mm wide; pappus white; achenes silky-

villous. This distinctive species is represented
in Utah by two phases, which are more or
less morphologically distinctive and geo-
graphically correlated.

Stems mainly 3-8 cm tall; leaves linear to linear-oblanceolate, mainly 1-3 mm

wide; plants of the Green River Formation, Uintah County

H. armerioides var. gramineus

Stems usually over 8 cm tall; leaves oblanceolate, mainly 3-10 mm wide;
plants widespread H. armerioides var. armerioides

Var. armerioides Blackbrush, black sage-
brush, pigmy sagebrush, salt desert shrub,
pinyon-juniper, mountain brush, and pon-
derosa pine communities at 1340 to 2120 m
in Carbon, Daggett, Duchesne, Emery, Gar-
field, Grand, Kane, San Juan, Sevier, and
Uintah counties; Montana to Arizona, east to
New Mexico and Nebraska; 85 (xii).

Var. gramineus Welsh & F. J. Smith
Desert shrub and pinyon-juniper commu-
nities at ca 1585 to 1895 m in Duchesne and
Uintah counties; endemic; 17 (0).

Haplopappus cervinus Wats. Shrubs, 1-4
dm tall, much branched; branchlets grayish
to straw colored; leaves 6-18 mm long, 2.2-6
mm wide, oblanceolate, straight or curved,
entire or repand-undulate, attenuate basally,
cuspidate apically, glabrous or resinous;
heads few, cymose; peduncles 3-10 mm long;
involucres 6.5-7.5 mm high, 5-8 mm wide;
bracts imbricate in several series, the outer
greenish ones narrowly acuminate with
straight or spreading tips, the irmer char-
taceous ones narrowly oblong, acute or cuspi-
date, all glabrous but resinous; ray flowers
5-7, yellow, 2.5-4 mm long, ca 1 mm wide;
disk flowers 5-11, glabrous or the tube spar-
ingly puberulent; pappus tawny; achenes stri-
gose. Black sagebrush, shadscale, pinyon-juni-
per, and mountain brush communities at
1670 to 2440 m in Millard and Sevier coun-
ties; Arizona; 5 (0). The type came from a
place called Antelope Canyon (possibly in
present-day western Millard County). More
collections are needed.

Haplopappus dementis (Rydb.) Blake
[Pyrrocoma dementis Rydb.; P. subcaesia
Greene, type from Panguitch Lake; P. la-
pathifolia Greene, type from "Utah"]. Pe-
rennial herbs from a simple caudex and stout
taproot, the subrhizomatous caudex clothed

with brown, often shredded marcescent leaf
bases; stems 10-30 (40) cm tall, decumbent-
ascending from an abruptly curved base, vil-
lous; basal leaves mostly 2-15 cm long, 4-17
mm wide, oblanceolate, glabrous or sparingly
puberulent, entire or dentate, tapering to a
petiole, acute; cauline leaves reduced up-
ward, sessile and somewhat clasping; heads
solitary (rarely 2 or 3); involucres broadly
hemispheric, 8-16 mm high, 18-30 mm wide;
bracts in several series, oblong to lanceolate,
green throughout or the base chartaceous,
villous; ray flowers 30-60, yellow or golden,
8-14 mm long; pappus tawny; achenes hairy.
Grass-sagebrush, spruce-fir, sedge-forb, and
meadow communities at 2590 to 3390 m in
Beaver, Daggett, Duchesne, Emery, Garfield,
Iron, Kane, Piute, Sanpete, Summit, and Uin-
tah counties; Wyoming to Colorado; 39 (vi).
Specimens from Utah that have been deter-
mined as H. integrifolius Gray apparently fall
here, including the type of Pyrrocoma la-
pathifolia, which was discussed by Hall
(1920). Involucral bracts vary from her-
baceous throughout to chartaceous at the
base. More work is indicated.

Haplopappus crispus L.C. Anderson
Shrubs, much branched from the base, 3-5
dm tall (or more); branchlets covered with
short-stalked glands; leaves 1.5-3 cm long,
3-8 mm wide, entire, green, spatulate to ob-
long-oblanceolate, acuminate, the margins
undulate-crisped, glutinous with low glands,
not crowded below the inflorescence; heads 1
or 2, more commonly more, per branch,
loosely paniculate to congested and cymose;
involucres campanulate, the heads 12.5-16
mm long, 5-9 mm wide; bracts in several
series, finely glandular, the outermost green,
leaflike; rays lacking; disk flowers 14-24,
pale yellow; pappus tawny; achenes 6.5-8.5

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

293

mm long, sparsely but evenly hairy. Ponder-
osa pine, fir, manzanita, and aspen commu-
nities at (915?) 2471 to 3050 m in Washing-
ton and Millard (?) counties; endemic; 8 (0).

Haplopappus croceus Rydb. Perennial
herbs, mainly 2-6 cm tall, from a simple cau-
dex and stout taproot, the caudex clothed
with fibrous marcescent leaf bases; basal
leaves 8-20 cm long, 6-25 (40) mm wide, el-
liptic to oblanceolate, petiolate, entire or un-
dulate, obtuse to acutish, glabrous or pu-
berulent; cauline leaves reduced upward,
sessile, more or less clasping; heads solitary
(rarely more); involucres hemispheric, 12-18
mm high, 20-30 mm wide; bracts in several
series, ovate to oblong or oblanceolate, her-
baceous apically, chartaceous to leathery ba-
sally; ray flowers 25-70, burnt orange, 10-25
mm long; pappus brownish; achenes glabrous
or pilose. Mountain brush community at ca
2470 m in San Juan (La Sal Moimtains) and
Washington (Kolob Reservoir) counties;
Wyoming south to Arizona and New Mexico;
2(0).

Haplopappus drwnTnondii (T. & G.) Blake
[Linosyris drummondii T. & G.]. Perennial
subshrub, the stems subherbaceous, arising
from a woody base, 25-75 cm tall, straw col-
ored to tan, longitudinally striate, glabrous;
leaves 1.5-7.5 cm long, 1-16 mm wide, en-
tire or irregularly lobed, linear to spatulate,
glabrous, resinous; heads few to numerous,
borne in corymbose cymes, peduncled; in-
volucres turbinate, 6-8 mm high, 4-7.2 mm
wide; bracts in 4 or 5 series, lance-oblong, co-
riaceous, with a thick green or brownish sub-
apical spot, acute, resinous; ray flowers lack-
ing; pappus tawny; achenes silky. Saline
riparian areas in greasewood, saltgrass, rab-
bitbrush, saltbush, and tamarix communities
at 1050 to 1800 m in Emery, Garfield,
Grand, Kane, and San Juan counties; Colo-
rado, Arizona, New Mexico, and Texas; 25

Haplopappus gracilis (Nutt.) Gray [Die-
teria gracilis Nutt.]. Annual herbs, 3-25 (30)
cm tall, commonly branched from near the
base; leaves 4-25 mm long, 1-3 mm wide,
linear to narrowly spatulate, spinulose-den-
tate to piimatifid, white-strigose, progressive-
ly/reduced and entire upward; heads solitary
or few to several and corymbosely arranged;
involucres 6-8.5 mm high, 8-12 mm wide;

bracts in 5 or 6 series, linear-lanceolate, awn
tipped, herbaceous medially, strigulose, not
glandular; rays 15-30, yellow, 6-9 mm long;
strigulose, not glandular; rays 15-30, yellow,
6-9 mm long; pappus tawny to white; ach-
enes pilose. Larrea-gutierrezia, ponderosa
pine, and spruce-fir communities at 850 to
960 m in Iron, Kane, and Washington coun-
ties; California to Colorado, south to Mexico;
10 (0).

Haplopappus lanceolatus (Hook.) T. & G.
[Donia lanceolata Hook.; H. tenuicaulis D.C.
Eaton; H. lanceolatus var. tenuicaulis (D.C.
Eaton) Gray; Pyrrocoma subviscosa Greene;
H. lanceolatus ssp. subviscosus (Greene) Hall;
Donia uniflora Hook.; H. uniflorus (Hook.)
T. & G.]. Perennial herbs from a simple cau-
dex and stout taproot, the caudex clothed
with brown to ashy marcescent, often fibrous,
leaf bases; stems decumbent-ascending,
abruptly bent at the base, 5-68 cm long; bas-
al leaves 3-16 cm long, 3-35 mm wide, ellip-
tic-oblong or lanceolate, glabrous or tomen-
tose, petiolate, entire or dentate to lobed,
often densely tomentose in the axils; cauline
leaves reduced upward, finally sessile and
clasping; heads solitary or few to several, and
subcorymbose or less commonly racemose;
involucres hemispheric, 5-12 mm high,
10-18 mm wide; bracts imbricate in 3 or 4
series, with green tips, glabrous or tomentu-
lose; ray flowers 10-45, yellow, 5-10 mm
long; pappus tawny; achenes densely hairy.
Saline meadows at 1300 to 2500 m in Beaver,
Cache, Carbon, Duchesne, Emery, Garfield,
Iron, Juab, Millard, Piute, Rich, Salt Lake,
Sevier, Tooele, and Utah counties; Oregon to
Saskatchewan, south to California, Nevada,
Colorado, and Nebraska; 42 (xi). This is a
highly variable taxon of saline meadows
through much of our area. Heads vary from
solitary to numerous, from solitary to cor-
ymbosely or racemosely arranged. Vesture is
lacking or tomentose, or rarely glandular.
Recognition of taxonomic categories within
the variation appears to be only arbitrarily
possible, and it seems best to treat our speci-
mens conservatively.

Haplopappus laricifolius Gray Rounded
shrubs 3-8 dm tall; branchlets resinous, yel-
lowish, becoming gray in age; leaves 5-18
mm long, 1-1.5 mm wide, thick, linear, res-
inous-punctate; heads few to several in com-
pact cymes, shortly pedunculate; involucres

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Great Basin Naturalist

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campanulate, 3-5 mm high, 3-6 mm wide;
bracts imbricate in ca 3 series, narrowly ob-
long, acute, yellowish or hyaline, glabrous or
puberulent-ciliate; ray flowers 3-6, yellow,
4-5 mm long; disk flowers 9-16, glabrous or
minutely pubescent; pappus tawny; achenes
white hairy. Saltgrass seep margin in warm
desert shrub at 1220 m in Washington Coun-
ty; Arizona to Texas and Mexico; 2 (ii).

Haplopappus linearifolius Gray [H. interi-
or Gov.; H. linearifolius ssp. interior (Gov.)
Hall]. Shrubs, mainly 4-10 (12) dm tall;
branchlets yellowish, resinous, becoming gray
in age; leaves 6-28 mm long, 1-2.5 mm wide,
thickish, linear to narrowly oblanceolate, res-
inous-punctate; heads few to many, solitary
on naked peduncles mainly 2-7 cm long; in-
volucres hemispheric, 8-10 mm high, 10-18
mm wide; bracts biseriate, lance-linear, acute
or acuminate, herbaceous medially, some-
times minutely glandular; rays 12-18, yellow,
9-15 mm long, 4-5 mm wide; disk flowers
numerous; pappus white; achenes densely
hairy. Joshua tree, creosote bush, blackbrush,
juniper, live oak, and sagebrush communities
at 700 to 1375 m in Washington Gounty;
Galifomia, Nevada, Arizona, and Baja Gali-
fomia; 30 (ii). Our material is assignable to
var. interior (Gov.) Jones.

Haplopappus macronema Gray [Mac-
ronema discoideum Nutt.]. Shrubs, mainly
1-5 dm tall; branchlets white-tomentose;
leaves 8-32 mm long, 2-7 mm wide, oblan-
ceolate to oblong, entire or more commonly
undulate-crisped, acute to obtuse, mucronate,
glandular-scabrous; heads solitary or 2 to sev-
eral; involucres campanulate, 9-13 mm high,
6-12 mm wide; bracts subequal, the outer
few herbaceous, oblong, the inner lance-
acuminate, chartaceous, glandular-scabrous;
ray flowers lacking; disk flowers 10-25; pap-
pus tawny; achenes villous. Douglas fir,
lodgepole pine, spruce-fir, and alpine tundra
communities at 2135 to 3420 m in Beaver,
Box Elder, Duchesne, Garfield, Iron, Juab,
Piute, Salt Lake, Sanpete, Sevier, Tooele,
Salt Lake, and Utah counties; Oregon to
Wyoming, south to Galifornia, Nevada, and
Colorado; 38 (vi).

Haplopappus nanus (Nutt.) D.G. Eaton
[Ericameria nana Nutt.]. Compact shrubs,
mainly 1-3 (5) dm tall; branchlets yellowish,
resinous, becoming gray in age; leaves 3-18

mm long, 0.5-2 mm broad, narrowly oblan-
ceolate to linear, entire, acute, resinous but
not punctate; heads solitary or few to several
in compact cymes, sessile or shortly peduncu-
late; involucres narrowly turbinate, 5.5-8.5
mm high, 3-7 mm wide; bracts imbricate in
4 or 5 series, the outer often greenish me-
dially, the inner chartaceous, with hyaline
margins, glabrous; rays 1-7, yellow, 2-3 mm
long; disk flowers 4-10; pappus tawny; ach-
enes villous or glabrous. Desert shrub, shrub-
grass, and juniper or pinyon-juniper commu-
nities at 1310 to 2820 m in Beaver, Juab, Mil-
lard, Piute, Sevier, Tooele, and Washington
counties; Oregon, Galifomia, Nevada, and
Idaho; 18 (iii).

Haplopappus racemosus (Nutt.) Torr.
[Homopappus racemosus Nutt.]. Perennial
herbs, from a simple caudex and stout tap-
root, the caudex clothed with fibrous marces-
cent leaf bases; stems 20-60 (100) cm tall,
erect, not abruptly bent at the base (in ours);
basal leaves mainly 6-25 cm long, 5-30 mm
wide, the blades elliptic to oblong or oblan-
ceolate, petiolate, rigidly erect, entire or
toothed, glabrous or puberulent; cauline
leaves reduced, sessile, clasping; heads ra-
cemose, in panicles or spikes, shortly pe-
dunculate; involucres 8-12 mm high, 4-18
mm wide; bracts in 3 or 4 series, with green
tips and coriaceous bases, abruptly pointed
apically; rays 10-35, yellow, 5-12 mm long;
pappus tawny; achenes hairy or glabrous. Sa-
line meadows at 1370 to 1470 m in Millard
and Utah counties; Oregon to Idaho, south to
Galifornia and Nevada; 2 (i). Utah lies at the
eastern margin of the range of this species
complex, in which Hall (1928) recognized
nine subspecies. Our material is hardly repre-
sentative of the variation within the assem-
blage of forms that lie to the west of this re-
gion. One of our specimens (Welsh et al.
14514 BRY) belongs to the spiciform narrow-
headed var. sessiliflorus (Greene) Welsh stat.
nov. (based on: Pyrrocoma sessiliflora Greene
Leafl. Bot. Obs. & Grit. 2: 12. 1909), and the
other is a paniculiform large-headed phase
apparently nearest to var. prionophyllus
(Greene) Welsh stat. nov. (based on Pyrro-
coma prionophylla Greene Leafl. Bot. Obs. &
Grit. 2: 12. 1909). Much more material is re-
quired to evaluate the nature of the speci-
mens in Utah. Racemose phases of the closely

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

295

related H. lanceolatus (q.v.) have been mis-
taken for H. racemosus. The erect or suberect
stems and stiffly erect leaves appear to be di-
agnostic for our specimens of H. racemosus.

Haplopappus scopulorum (Jones) Blake
[Bigelovia nienziesii var. scopulorum Jones,
type from Zion Canyon; H. scopulorum var.
hirtellus Blake, type from Cedar Canyon].
Shrubs, mainly 3-10 dm tall; branchlets
green to straw colored or white, glabrous;
leaves 0.7-7.8 cm long, 1-8 mm wide, nar-
rowly lanceolate to oblong, entire, 3-nerved,
glabrous, the margins scabrous, attenuate to a
spinulose apex; heads few to many, borne in
loose to subcompact cymes, peduncled; in-
volucre narrowly campanulate, 6.5-9.5 mm
high, 3-5.5 mm wide; bracts in 5 or 6 series,
oblong, chartaceous and pale, or the tips
greenish or often brownish, rounded-obtuse,
glabrous, not resin coated; ray flowers lack-
ing; disk flowers 10-20; pappus white; ach-
enes white-pilose. Pinyon-juniper, mountain
brush, and ponderosa pine commimities at
1370 to 1830 m in Iron, Kane, San Juan, and
Washington counties; Arizona; 10 (vii).

Haplopappus spinulosus (Pursh) DC.
[Amellus spinulosus Pursh]. Perennial herbs
from a ligneus caudex; stems mainly 12-50
(60) cm tall, branching above the base; leaves
0.5-6 cm long, 1-10 mm wide, pinnatifid to
bipinnatifid or the upper ones entire, or
merely toothed, spinulose; heads solitary, or
few in corymbose clusters; involucres 5-8
mm high, 8-12 mm wide; bracts in 4-6
series, linear, awn-tipped, herbaceous me-
dially, glandular, not strigulose; rays 15-50,
yellow, 8-10 mm long; pappus brownish;
achenes pilose. Desert shrub commimity at ca
1300 m in San Juan County (Atwood 7175
BRY); Alberta to Minnesota, south to Califor-
nia, Arizona, New Mexico, Texas, and Mexi-
co; 1 (0).

Haplopappus watsonii Gray Shrubs, 1-4
dm tall; herbage stipitate-glandular; branch-
lets yellowish, becoming whitish to straw col-
ored or grayish in age; leaves 4-28 mm long,
3-10 mm wide, oblanceolate to obovate or
spatulate, entire or undulate, abruptly cuspi-
date-acuminate apically; heads several to nu-
merous (rarely some solitary) in loose cymes,
the peduncles 1-7 mm long; involucres sub-
cylindric to narrowly campanulate, 5.5-8
mm high, 3-76 mm wide; bracts in ca 5

series, the outer ones greenish, the inner
chartaceous or greenish at the tips; rays 5-10,
yellow, 4-6 mm long; disk flowers 5-15; pap-
pus brownish; achenes hairy. Rock outcrops
(limestone, sandstone, or quartzite) in desert
shrub, pinyon-juniper, mountain brush, and
ponderosa pine commimities at 1310 to 3440
m in Beaver, Box Elder, Cache, Davis, Juab,
Millard, Salt Lake, Summit, Tooele, and We-
ber counties; Nevada and Utah; 33 (ii). Our
material belongs to one of a vicarious pair of
infraspecific taxa within the Great Basin
known as var. rydbergii (Blake) Welsh comb,
nov. (based on: H. rydbergii Blake Contr. U.S.
Natl. Herb. 25: 545. 1925, nom. nov. pro
Macronema obovatum Rydb. Bull. Torrey
Bot. Club. 27: 68. 1900, type from City
Creek Canyon). The var. rydbergii differs in
having fewer disk flowers (5-15 not 15-25).
Other supposedly diagnostic features (i.e., the
green outer involucral bracts) fail, being
present to a greater or lesser degree in both
phases. The type variety has not been discov-
ered in Utah, but should be expected in the
western border region.

Haplopappus zionis L.C. Anderson Shrubs,
mainly 1-3 dm tall; herbage minutely and
shortly stipitate-glandular; leaves 0.8-3.5 (4)
cm long, 2-4.5 (7) mm wide, oblong to nar-
rowly oblanceolate, 1 -nerved, entire, abrupt-
ly mucronate; heads solitary or 2 or 3, in cy-
mose clusters, peduncled; involucres
cylindric-campanulate, 12-15 mm high, 6-12
mm wide; bracts subequal, herbaceous (outer)
and greenish, the inner chartaceous or with a
subapical green spot and broadly hyaline
margins; rays lacking; disk flowers 8-21; pap-
pus tawny; achenes glabrous below, strigose
apically. Ponderosa pine and spruce-fir com-
munities, commonly on limestone members
of the Cedar Breaks (Wasatch) Formation, at
2440 to 3050 m in Garfield, Iron, and Kane
counties; endemic; 5 (i).

Helenium L.

Annual or perennial herbs; leaves alter-
nate, glandular-pimctate, decurrent or clasp-
ing; heads solitary or few to numerous in co-
rymbose clusters, radiate, yellow; involucral
bracts in 2 or 3 series, the bracts subequal or
the inner shorter and narrower, herbaceous

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Great Basin Naturalist

Vol. 43, No. 2

or essentially so, soon deflexed; receptacle
naked, convex or conic; rays pistillate or neu-
ter; disk flowers numerous, perfect; pappus

of 5-10 scarious or hyaline scales; achenes
truncately obpyramidal, 4- or 5-angled, with
as many intermediate ribs.

1. Leaves sessile, clasping; stems not winged; plants of aspen communities and

upward H. hoopesii

— Leaves decurrent; stems winged below the leaf bases; plants of riparian com-
munities at lower elevations H. autumnale

Helenium autumnale L. Common Sneeze-
weed. Perennial herbs; stems mainly L5-10
(12) dm tall, puberulent and glandular, cor-
ymbosely branched above; leaves 1.5-15 cm
long, 3-35 (40) mm wide, serrate to entire,
glandular-punctate; heads 3 to many, the disk
hemispheric to subglobose, yellow, 1-2 cm
wide; rays 10-20, yellow, mainly 8-12 mm
long, soon reflexed; pappus scales lance-
ovate, with slender awn-tip as long as the
body; achenes ca 1.5 mm long, hirsute and
glandular. Cattail-willow, tamarix-grease-
wood, and sedge-rush communities at 1220 to
1830 m in Box Elder, Daggett, Emery, Rich,
Uintah, and Utah counties; British Columbia
to Quebec, south to Arizona, and Florida; 18
(iii). This species is poisonous to livestock.

Helenium hoopesii Gray Orange Sneeze-
weed. [Heleniastrum hoopesii (Gray) Kuntze;
Dugaldia hoopesii (Gray) Rydb.]. Perennial
herbs, mainly 2-8 (10) dm tall, with a sub-
rhizomatous caudex and fibrous roots; herb-
age more or less villous-tomentose to gla-
brate; basal leaves 2-30 cm long, 0.5-5 cm
wide, oblanceolate, tapering to a clasping
base; cauline leaves reduced upward, oblan-
ceolate to elliptic or lanceolate, entire; heads
2-11, in loose corymbs; disks hemispheric,
2-3.5 cm wide; involucres 5-8 mm high, the

bracts lanceolate to elliptic; rays 13-21, yel-
low or yellow-orange, 15-35 mm long, finally
reflexed; pappus scales hyaline, lanceolate,
attenuate; achenes 3-4 mm long, hairy. Sage-
brush, mountain brush, aspen, and spruce-fir
communities, often in openings or riparian
zones, at 1830 to 3200 m in Beaver, Box El-
der, Carbon, Duchesne, Garfield, Iron, Juab,
Piute, San Juan, Sevier, Summit, Utah,
Wasatch, and Washington counties; Oregon
to Wyoming, south to California, Arizona,
and New Mexico; 69 (xiv). This is a poisonous
plant, causing spewing sickness in sheep.

Helianthella T. & G.

Perennial herbs; leaves simple, opposite or
alternate, entire; heads radiate, solitary or
few to several in loose subcorymbose clusters;
bracts imbricate to subequal, more or less
herbaceous; receptacle plano-convex, chaffy
throughout, the persistent bracts clasping the
achenes; disk flowers numerous, fertile, yel-
low, or purple; rays yellow; pappus of 2 slen-
der awns and short scales; achenes strongly
compressed at right-angles to involucral
bracts.
Weber, W. A. 1952. The genus Helianthella

(Compositae). Amer. Midi. Naturalist

48: 1-35.

2(1).

Heads 3-12 or more, mainly less than 20 mm broad; rays 7-13 mm long, in-
conspicuous; disk flowers normally purple H. microcephala

Heads solitary or 2 or 3, mainly over 20 mm broad; rays 15-30 mm long,
showy; disk flowers yellow 2

Heads erect; involucral bracts lance-oblong, short-ciliate H. uniflora

Heads nodding; involucral bracts oblong-ovate, long-ciliate with multicellular
hairs H. quinquenervis

Helianthella microcephala (Gray) Gray

[Encelia microcephala Gray]. Perennial
herbs; stems 20-65 cm tall; herbage ap-
pressed hispidulous; basal leaves mainly 4-30
cm long, 0.5-3 cm wide, petiolate, the blades

elliptic to lanceolate, scabrous and harshly
ciliate, acute to obtuse; cauline leaves re-
duced upward; heads 3-12 or more; bracts
imbricate in ca 3 series, oblong to lanceolate
or oblanceolate, strigose and roughly ciliate

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

297

and glandular; rays 8-10, yellow, 7-13 mm
long; disk flowers commonly purple; achenes
7-8 mm long, long-pilose. Desert shrub, pin-
yon-juniper, ponderosa pine, mountain brush,
and Douglas fir-limber pine communities at
1220 to 2745 m in Carbon, Duchesne,
Emery, Garfield, Grand, Kane, San Juan, Se-
vier, and Uintah counties; Colorado and Ari-
zona; 34 (x). In one plant from Navajo Moun-
tain the disk flowers are apparently yellow. A
single collection from west of Richfield
(Welsh et al. 17487 BRY) is the only record
examined for the Great Basin.

Helianthella quinquenervis (Hook.) Gray

[Helianthus quinquenervis Hook.]. Perennial
herbs; stems 5-15 dm tall, glabrous or villous
above; basal leaves 0.3-40 cm long, 0.8-4 cm
wide, petiolate, the blades elliptic to oblong
or oblanceolate, entire, obtuse to acute; cau-
line leaves often enlarged to near stem
middle then reduced, becoming subsessile or
sessile, the largest (at least) prominently 5-
nerved; heads nodding, solitary or 2 or 3; disk
2.5-4 cm wide; bracts ovate-lanceolate,
acuminate, long-ciliate; rays 12-21, yellow,
15-35 mm long; achenes 8-10 mm long, pi-
lose. Sagebrush, aspen, ponderosa pine, and
spruce-fir communities at 2115 to 3175 m in
Carbon, Daggett, Duchesne, Emery, Gar-
field, Grand, San Juan, Sevier, Summit, Uin-
tah, and Wasatch counties; Oregon to South
Dakota, south to Nevada, Arizona, and New
Mexico; 13 (0).

Helianthella uniflora (Nutt.) T. & G. [He-
lianthus uniflorus Nutt.]. Perennial herbs
from a branching caudex; stems mainly 3-10
dm tall, glabrous below or more or less
spreading-hairy throughout; basal leaves
3-15 cm long, 0.6-5.5 cm wide, petiolate, the

blades oblanceolate to elliptic or lanceolate,
entire, obtuse to acute; cauline leaves often
enlarged to near stem middle, then reduced,
becoming sessile or subsessile, the largest
prominently 3-nerved; heads erect, solitary
or 2 or 3; disk 1.5-3 cm wide; bracts lance-
linear, acuminate or obtuse, scabrous-
puberulent, shortly ciliate; rays 13-17, yel-
low, 2-4.5 cm long; achenes 6-7 mm long,
pilose. Sagebrush, pinyon-juniper, mountain
brush, ponderosa pine, aspen, and spruce-fir
communities at 1525 to 3175 m in Beaver,
Box Elder, Cache, Carbon, Davis, Duchesne,
Garfield, Grand, Iron, Millard, Morgan,
Piute, Rich, Salt Lake, Sanpete, Sevier, Sum-
mit, Tooele, Uintah, Utah, Wasatch, and We-
ber counties; Alberta to Montana, south to
Nevada and Colorado; 64 (v).

Helianthus L.

Annual or perennial herbs; leaves simple,
opposite below, usually alternate above;
heads radiate, showy, solitary or few in cor-
ymbs; involucral bracts imbricate or sub-
equal, herbaceous; receptacle flat to convex,
chaffy throughout, its bracts clasping the
achenes; ray flowers conspicuous, yellow,
neuter; disk flowers yellow or reddish, fertile;
pappus usually of 2 main awns, scalelike at
base, sometimes with additional scales pres-
ent; achenes narrowly obovate in outline, 4-
angled or obcompressed.
Blauer, a. C. 1965. Helianthus (Compositae)
in Utah. Proc. Utah Academy 42:
240-251.
Heiser, C. B. Jr. 1947. Hybridization be-
tween sunflower species Helianthus an-
nuus and H. petiolaris. Evolution 1:
249-262.

1. Plants perennial; disk flowers yellow; leaves mainly opposite, lanceolate to

linear-lanceolate H. nuttallii

— Plants annual; disk flowers reddish brown to purplish; leaves mainly alternate;
leaves lanceolate to ovate 2

2(1). Involucral bracts linear to narrowly lanceolate; pappus of numerous unequal

scales 3

— Involucral bracts lanceolate to ovate; pappus commonly of 2 distinct awns 4

3(2). Involucral bracts surpassing the disk; pappus scales linear; stems moderately

hirsute to glabrous; plants widespread H. anomalus

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Great Basin Naturalist

Vol. 43, No. 2

— Involucral bracts subequal to the disk; pappus scales ovate; stems markedly
hirsute; plants of Washington County H. deserticola

4(3). Leaves canescent on both sides; involucral bracts narrowly lanceolate; plants

of San Juan County H. niveus

— Leaves hispid to strigose; involucral bracts lanceolate to ovate; plants wide-
spread 5

5(4). Involucral bracts ovate, rather abruptly narrowed to an acuminate tip, the cen-
tral ones inconspicuously hairy; leaves often cordate and with serrate margins ..
H. annuus

— Involucral bracts lanceolate, tapering to the tip, the central ones often white
bearded apically; leaves seldom cordate, usually entire H. petiolaris

Helianthus annuus L. Common Sun-
flower. Annual herbs; stems commonly hispid
and rough, 3-40 dm tall, simple or branched;
leaves alternate above (3) 5-40 cm long, 2-40
cm wide, lance-ovate to broadly ovate, acute
to obtuse, serrate, truncate or cordate (alter-
nate) basally, hispid to hispidulous on both
sides, petiolate; heads solitary or few; disks
mainly 2-5 cm wide; involucral bracts lance-
ovate to ovate, attenuate to caudate, hispid
to hispidulous, ciliate; disk corolla lobes
purplish red (rarely yellow); pappus of 2
awnlike ovate-lanceolate scales; achenes
glabrous to strigose. Saltgrass-muhly grass,
desert shrub, pinyon-jimiper, and mountain
brush communities, commonly where dis-
turbed, at 1200 to 2440 m, probably in all
Utah counties; widespread in the United
States, Canada, Mexico, and elsewhere; 70
(xv). Our common weedy sunflower is assign-
able to ssp. lenticularis (Dougl.) Cockerell;
the cultivated large-headed phase to var.
macrocarpus (DC.) Cockerell.

Helianthus anomalus Blake Sand Sun-
flower. Annual herbs; stems sparingly hispid
to glabrate, 5-70 cm tall; leaves mainly alter-
nate, petiolate, the blades L2-10 cm long,
0.4-4 cm wide, narrowly lanceolate to lance-
ovate, yellowish green, acute, cuneate to ob-
tuse basally, hispidulous to hispid on both
sides; heads solitary or few, showy; disks
mainly 12-24 mm wide; involucral bracts lin-
ear, commonly 10-25 mm long and 2-3 mm
wide, hispid above, definitely hispid-ciliate,
at least below, often some much surpassing
the disk; disk corolla lobes purple; pappus of
2 large linear scales and numerous similar
subequal scales; achenes 3.5-5.5 mm long,
appressed pilose. Blackbrush, ephedra,
purple-sage, vanclevea, psorothamnus, and

pinyon-juniper communities, commonly in
dunes or other sandy sites, at 1150 to 1830 m
in Emery, Garfield, Grand, Juab, Kane, Mil-
lard, San Juan, Tooele, and Wayne (type
from near Hanksville) counties; Arizona; 33
(xi). This is a Colorado Plateau endemic, with
an extension onto dunes of the eastern Great
Basin.

Helianthus deserticola Heiser Desert Sun-
flower. Annual herbs; stems strongly hispid
(at least below), 2-12 dm tall; leaves mainly
alternate, petiolate, the blades 2-6 cm long,
0.5-2 cm wide, lanceolate to lance-ovate,
green, acute, cuneate to obtuse basally, his-
pid to hispidulous on both sides; heads few to
several (solitary), showy; disks 1.3-2.5 cm
wide; involucral bracts linear, 8-14 mm long,
1.5-2.5 mm wide, hispid dorsally and ciliate,
subequal to the disk; disk corolla lobes
purple; pappus of 2 large lanceolate to lance-
ovate scales and ca 10 smaller ones; achenes
4-5 mm long, pilose. Blackbrush, creosote
bush, matchweed, and live oak communities
at 850 to 1070 m in Washington County; Ari-
zona and Nevada; 2 (0). This is an obscure
taxon, despite the passage of two decades
since its description; more specimens are re-
quired. Perhaps it is too nearly allied to H.
anomalus, with which it shares some mor-
phological features.

Helianthus niveus (Benth.) Brandegee
Snowy Sunflower. [Encelia nivea Benth.].
Annual herbs; stems mainly 5-15 dm tall, ca-
nescent and hispid; leaves mainly alternate,
petiolate, the blades 3-12 cm long, 0.9-6 cm
wide or more, lanceolate to ovate, entire or
serrate, mostly acute apically, canescent on
both sides; heads solitary or few to many;
disk 1-2.5 cm wide; bracts narrowly lanceo-
late, 1.5-2.5 mm wide, 8-12 mm long, sub-
equal to the disk; disk corolla lobes purplish;

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

299

pappus of 2 lanceolate scales and several
shorter scales. Sandy sites at ca 1375 m in
San Juan Counties; Texas to California and
Mexico; 1 (0). Our limited material is assign-
able to ssp. canescens (Gray) Heiser [H. pet-
iolaris var. canus Britt.].

Helianthus nuttallii T. & G. Nuttall Sun-
flower. [H. bracteatus E.E. Watson, type
from Logan; H. giganteus var. utahensis D.C.
Eaton, type from Wasatch Mountains; H.
utahensis (D.C. Eaton) A. Nels.]. Perennial
rhizomatous herbs with tuberous roots; stems
3-20 dm tall or more, glabrous or sparingly
scabrous or hispid; leaves mainly opposite,
shortly petiolate, the blades 4-16 cm long,
0.8-3 cm wide, narrowly lanceolate, acute to
attenuate, entire or denticulate, cuneate ba-
sally, scabrous on both sides; heads solitary or
few to many; disks mainly 12-28 mm wide;
bracts lance-linear, 1.5-3 mm wide, subequal
to, or surpassing, the disk, attenuate, ap-
pressed pubescent and more or less ciliate;
disk corolla lobes yellow; pappus of 2 narrow
awnlike scales; achenes 3-4 mm long,
glabrous. Seeps, springs, wet meadows, and
canal banks at 1280 to 2200 m in Cache, Car-
bon, Duchesne, Garfield, Juab, Rich, Salt
Lake, Summit, Tooele, Uintah, Utah,
Wasatch, Washington, and Weber counties;
British Columbia to Saskatchewan, south to
Nevada, Arizona, and New Mexico; 18 (0).
Note: The perennial sunflower, H. tuherosus
L., is grown for its edible roots in our area. It
persists following cultivation and is difficult
to eradicate. The leaves are broadly lanceo-
late to ovate.

Helianthus petiolaris Nutt. Prairie Sun-
flower. Annual herbs; stems 0.5-12 dm tall,
strigose to hispid or glabrous; leaves mainly
alternate, petiolate, the blades 1-8 cm long,
4-25 (30) mm wide, lanceolate to ovate,
acute to obtuse, entire or rarely serrate, cu-
neate to truncate basally, hispidulous to stri-
gose; heads solitary or few; disk 10-25 mm
wide; involucral bracts 2-5 mm wide, 7-15
mm long, lanceolate, acuminate or attenuate,
hispidulous, usually short-ciliate; disk corolla
lobes purplish; pappus of 2 lanceolate awn-
like scales; achenes 3-4.5 mm long, hairy.
Salt desert shrub, desert shrub, pinyon-juni-
per, and riparian communities, often where
disturbed, at 1220 to 1920 m in Beaver,
Duchesne, Emery, Garfield, Grand, Kane,
Millard, San Juan, Sevier, Uintah, Wasatch,
Washington, and Wayne counties; Alberta to
Maine, south to California, Arizona, New
Mexico, Texas, Louisiana, and South Caro-
lina; 62 (xxii). Our material has been assigned
to ssp. fallax Heiser. The material appears to
be indigenous in the Colorado drainage sys-
tem, but the rare specimens in the Great Ba-
sin seem to be adventive.

Heliomeris Nutt.

Annual or perennial herbs; leaves opposite
(at least below), simple; heads radiate, soli-
tary or cymose; involucres 2- or 3-seriate;
rays yellow, neuter, pubescent dorsally; re-
ceptacles chaffy, the chaffy bracts clasping
the achenes; disk flowers fertile; pappus
none; achenes laterally compressed, 4-angled.

1. Plants perennial, widespread in montane habitats, less commonly in saline low

elevation sites H. multiflora

— Plants annual, restricted in low elevation saline habitats 2

2(1). Plants subscapose, with long naked peduncles; leaves ovate to lanceolate . H. soliceps

— Plants caulescent, the peduncles bracteate or leafy; leaves linear 3

3(2). Leaves canescent with appressed hairs; plants of southern Utah H. longifolia

— Leaves hispidulous; plants of central and western Utah H. hispida

Heliomeris hispida (Gray) Blake Hairy
Goldeneye. [H. multiflora var. hispida Gray;
Gymnolomia hispida var. ciliata Robins. &
Greenm., type from Utah; Viguiera ciliata
(Robins. & Greenm.) Blake]. Annual herbs;

stems simple or variously branched, 10-70
cm tall, hispidulous; leaves 0.6-9 cm long,
1-3 mm wide, linear, hispid and hispid-
ciliate, acute; heads solitary or 2-5 or more;
disks 7-15 mm wide, the corollas yellow; rays

300

Great Basin Naturalist

Vol. 43, No. 2

ca 9-15, yellow, 6-13 mm long; involucral
bracts 5.5-10 mm long, lance-attenuate, his-
pid and coarsely ciliate; pappus lacking; ach-
enes ca 2.5 mm long, glabrous. Saline mar-
shes and meadows at ca 1300 to 1470 m in
Millard, Salt Lake, and Utah counties; Ari-
zona, New Mexico, and Mexico; 9 (ii).

Heliomeris longifolia (Robins. & Greenm.)
Cockerel! [Gymnolomia longifolia Robins. &
Greenm.; Viguiera longifolia (Robins. &
Greenm.) Blake]. Annual herbs; stems simple
or variously branched, 14-60 cm tall, stri-
gose; leaves 1-6 cm long, 1.2-7.5 mm wide,
linear to oblong, strigose, rarely hispid-ciliate
near the bases, acute; heads solitary, or 2 to
numerous; disks 7-10 mm wide, the corollas
yellow; rays ca 8-10, yellow, 6-12 mm long;
involucral bracts lance-acuminate to -attenu-
ate, strigose, not especially ciliate; pappus
lacking; achenes 2-2.5 mm long, brown,
glabrous. Salt desert shrub and pinyon juni-

per communities at 1150 to 1525 m in Kane
and Washington counties; Arizona to Texas
and Mexico; 7 (iii). Our material is assignable
to var. annua (Jones) Yates [Gymnolomia
multiflora var. annua Jones, type from
Utah?].

Heliomeris multiflora Nutt. Showy Gold-
eneye. [Viguiera multiflora (Nutt.) Blake].
Perennial herbs, from a woody taproot and
pluricipital caudex; stems 2-10 (13) dm tall,
strigose to scabrous-puberulent; leaves lan-
ceolate to linear, mainly opposite, entire or
serrate, 1-8 (10) cm long, 2-20 (25) mm
wide, short-petiolate, plane or revolute, acute
to obtuse; heads commonly 2 to several; disk
6-14 mm wide; involucral bracts linear or
narrowly lanceolate, strigose; rays 10-14, yel-
low, 7-18 mm long; pappus lacking; achenes
1.2-1.8 mm long, brown, glabrous. Two
weakly discernible varieties are included in
our material.

Leaves commonly over 5 mm wide, plane; plants of mesic montane sites

H. multiflora var. multiflora

Leaves commonly less than 5 mm wide, the margins revolute; plants of arid
plains and mountains H. multiflora var. nevadensis

Var. multiflora Sagebrush, juniper, cotton-
wood, pinyon-juniper, aspen, and spruce-fir
communities, often in riparian sites, at 1340
to 2870 m in all Utah counties; Montana
south to California, Arizona, and New Mexi-
co; 137 (xvi).

Var. nevadensis (A. Nels.) Yates [Gymo-
nolmia nevadensis A. Nels.]. Shadscale, mat-
atriplex, pinyon-juniper, and mountain brush
communities at 1370 to 2135 m in Grand,
Juab, Uintah, and Washington counties; Ne-
vada; 13 (i).

Heliomeris soliceps (Bameby) Yates Tropic
Goldeneye. [Viguiera soliceps Bameby]. An-
nual herbs, 10-41 cm tall; stems branched be-
low, terminating in subscapose, merely brac-
teate peduncles that overtop the foliage;
leaves opposite below, the blades 15-38 mm
long, 6-20 mm wide, ovate to lanceolate,
strigose, 3-nerved, petiolate, obtuse to cu-
neate, becoming smaller upwards; peduncles
7-28 cm long; involucres biseriate, the bracts
lance-acuminate, acute, 5-6 mm long, stri-

gose; rays 10-12, yellow, 10-15 mm long;
pappus lacking; achenes 2.8-3.3 mm long,
blackish. Mat-saltbush community on Tropic
Shale Formation at 1400 to 1470 m in Kane
County; endemic. This is a striking species,
forming masses of yellow blossoms in years of
adequate rainfall; 5 (ii).

Heterotheca Cass.

Annual, biennial, or perennial herbs; leaves
alternate, simple, entire; heads radiate; in-
volucres campanulate to hemispheric; bracts
numerous, narrow, imbricated in several
series; receptacle convex, naked; rays yellow,
pistillate and fertile; pappus of capillary
bristles; disk flowers numerous, the pappus
present and usually double, the inner of ca-
pillary bristles, the outer (when present) of
short scales or bristles; achenes hairy.
Wagenknecht, B. L. 1960. Revision of Het-
erotheca, Section Heterotheca (Com-
positae). Rhodora 62:61-76, 97-109.

1. Plants low, creeping, arising from subrhizomatous caudex branches; heads nod-
ding, solitary or 2 or 3, known from sandy sites in Garfield, Kane, and Wash-
ington (?) counties H. pnesii

April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 301

— Plants various in habit, the caudex branches, if present, not rhizomatous; heads

few to numerous, seldom nodding, distribution various 2

2(1). Plants perennial, from a woody root crown, the stems numerous, forming

rounded clumps, common H. villosa

— Plants annual or biennial, the root crown herbaceous; stems solitary or few, not
forming rounded clumps, rare 3

3(2). Upper leaves cordate-clasping basally; involucres glandular-puberulent and

canescent H. psammophila

— Upper leaves tapering to a sessile base; involucres glandular, pubescent, not
canescent H. grandiflora

Heterotheca grandiflora Nutt. Telegraph herbs; stems stout, 5-12 (20) dm tall, branch-
Weed. Annual or biennial herbs; stems stout, ing above, hispid hirsute and glandular up-
branched above, 5-12 (20) dm tall, hirsute, ward; leaves mainly 1-7 cm long, ovate to
glandular-pubescent; leaves 2-6 cm long, lance-oblong, serrate to subentire, the lower
0.8-2.5 cm wide (or more), ovate to elliptic, petiolate, the upper cordate-clasping; heads
oblong, or oblanceolate, serrate, the lower numerous; involucres 8-12 mm high, glandu-
petiolate and lobed at base; heads numerous; lar and canescent; rays 20-30, yellow, mainly
involucres 7-9 mm high; rays 25-35, 6-8 mm 3-7 mm long; pappus tawny; achenes 2.4-3.8
long, ca 1 mm wide, the tube hairy; disk mm long. Sandy roadside at ca 970 to 1350 m
flowers numerous, slender; pappus tawny, in Grand and Washington counties; Arizona
Sandy roadside at ca 915 m in Washington to Texas, south to Mexico; 1 (i). The speci-
County; California and Arizona; 1 (i). Our men from Grand County (Welsh & Moore
material has the pappus merely tawny, not 2745) is missing from BRY, the Washington
brick red as reported elsewhere for the County locality is reported by Meyer,
species. Heterotheca villosa (Pursh) Shinners Hairy

Heterotheca jonesii (Blake) Welsh & At- Goldenaster. [Amellus villosus Pursh; Chry-
wood Jones Goldenaster. [Chrysopsis caespi- sepsis villosa (Pursh) Nutt. ex DC.]. Perennial
tosa Jones, not Nutt.; Chrysopsis jonesii herbs, from a ligneous root-crown and tap-
Blake]. Perennial caespitose herbs from a root; stems several to numerous, forming
creeping subrhizomatous caudex; stems 4-8 rounded clumps, mainly 1.5-5 dm tall; herb-
cm tall, loosely villous; leaves 5-11 mm long, age hirsute to strigose and more or less
1.5-4 mm wide, petiolate, the blades obovate glandular; leaves 0.5-5 cm ong, 2-10 mm
to spatulate, pilose; heads solitary or 2 or 3; ^i^^' oblanceolate to spatulate or elliptic,
., c % c uucin A green or silvery to gray-green, petiolate or
involucres 5-7.5 mm high, 6-10 mm wide, => , ... / . & -^ & ' r
^, , ^ Till ^ • subsessiie; heads few to numerous, mainly co-

the bracts narrowly lance-oblong, stngose-pi- , . , -7 m u- u -7 io

, , , 1. -^ . 11.1 n rymbose; involucres 7-10 mm high, 7-12 mm
lose, the hyaline margins reddish; ray flowers ^.^^_ ^^^^^^ lance-linear, green or char-
5-13, yellow, 4-6 mm long, 1.5-2.5 mm ^^^^^^^ ^^^ .^^ ^^^^.^^^ sometimes red-
wide; pappus tawny; achenes 2-3 mm long, ^^^. ^^^^ ^q_25 yellow, 6-10 mm long; pap-
hairy. Ponderosa pine, manzanita, and ^^^ ^^wny; achenes 2-3 mm long, hairy. Our
Douglas fir communities, on sandstone or in materials represent only a small portion of
sand, at 1580 to 2745 m in Garfield, Kane, the vast array of variation within the villosa
and Washington (?) counties (the type pre- complex. Three infraspecific taxa are appar-
sumably came from Springdale); endemic; 7 ent among our specimens, but application of
(ii). names is difficult. The following treatment is

Heterotheca psammophila Wagenkn. [H. therefore tentative, with a definitive treat-

subaxillaris, authors]. Annual or biennial ment awaiting monographic study.

1. Leaves green or gray-green, the surface apparent through the spreading to sub-

appressed hairs; plants widespread H. villosa var. hispida

— Leaves silvery or grayish, the surface seldom apparent through the usually ap-

pressed hairs; plants restricted 2

302

Great Basin Naturalist

Vol. 43, No. 2

2(1). Stems mainly appressed hairy, or with some hairs ascending to spreading

H. villosa var. villosa

— Stems with appressed and spreading contorted long hairs H. villosa var. foliosa

Var. foliosa (Nutt.) V.L. Harms [Chry- Var. villosa Blackbrush, pinyon-jimiper,

sopsis foliosa Nutt.]. Mountain brush and and ponderosa pine communities at 1090 to

bunchgrass (Agropyron et al.) communities at 2785 m in Garfield, Kane, Washington, and

1280 to 2135 m in Davis, Duchesne, Salt Wayne counties; widespread in western U.S.;

Lake, San Juan (?), Utah, and Weber coun- 8 (iii).
ties; widespread in western U.S.; 9 (0).

Var. hispida (Hook.) V.L. Harms [Dip- Hieracium L.
lopappus hispidus Hook.; Chry sopsis hispida

(Hook.) Nutt.; C. villosa var. scabra Eastw. Perennial rhizomatous herbs with milky

(?), type from San Juan County; C. viscida juice; leaves alternate or basal and still alter-

var. cinerascens Blake, type from Beaver nate, entire or toothed, simple; heads few to

Canyon]. Shadscale-rabbitbrush, ephedra-ly- numerous, in corymbose clusters; flowers all

cium-dropseed, sagebrush-grass, pinyon-juni- raylike, yellow to orange or white; involucres

per, mountain brush, ponderosa pine-manza- cylindric to hemispheric; bracts more or less

nita, and aspen commimities at 1150 to 2745 imbricate; receptacle naked; pappus usually

m in all Utah counties; widely distributed in of brownish capillary bristles; achenes terete

the western U.S.; 105 (xii). or prismatic, more or less strongly ribbed.

1. Flowers white; stems long-setose basally if at all, the petioles and leaves long-
setose on the lower midrib H. albiflorum

— Flowers yellow; stems long-setose throughout or only above; leaves variously
setose or glabrous 2

2(1). Leaves glabrous or short-hairy, 2-10 cm long, mainly basal, the stems merely

bracteate H. gracile

— Leaves long-hairy, mainly 10-20 cm long, the cauline ones well developed, re-
duced above H. cynoglossoides

Hieracium albiflorum Hook. White stems erect, 20-75 cm tall, pubescent with
Hawkweed. Perennial herbs; stems erect, long, coarse, loose or spreading white setae
15-75 cm tall, long-hairy at the base, becom- that dry yellowish; basal leaves commonly
ing glabrous upward; basal leaves 2-12 (17) withered at anthesis; lower cauline leaves
cm long, 0.8-4.5 cm wide, oblanceolate, pet- 10-25 cm long, 1-3 cm wide, petiolate, the
iolate, entire or remotely toothed, long-hairy blades oblanceolate to elliptic, long-hairy;
on petioles and midvein, commonly long- middle and upper leaves reduced, sessile;
ciliate; cauline leaves sessile, reduced up- heads yellow, few to many, 15- to 40-flow-
ward; stellate hairs lacking; heads few to ered, corymbose, the inflorescence more or
many, on slender peduncles, the inflorescence less open; involucres 7-12 mm high; bracts
open; involucres 6-11 mm high, blackish greenish, the margins chartaceous to hyaline,
green, glandular or sparingly long-hairy, or minutely stellate and stipitate-glandular;
glabrous; flowers 12-35, white; pappus taw- pappus tawny. Grass-forb, aspen, and spruce-
ny. Lodgepole pine, spruce, and spruce-fir fir communities at 200 to 2990 m in Box El-
communities at 1980 to 3420 m in Daggett, der. Salt Lake, Summit, Utah, Wasatch, and
Duchesne, Sanpete, Uintah, and Utah coun- Weber counties; British Columbia and Al-
lies; Alaska and Yukon to Saskatchewan, berta, south to Oregon and Wyoming; 18 (0).
south to California, Nevada, and Colorado; Hieracium gracile Hook. Slender Hawk-
12 (ii). weed. Perennial herbs; stems erect, 8-40 cm

Hieracium cynoglossoides Arv.-Touv. tall, tomentulous to glabrous; basal leaves

Houndstongue Hawkweed. [H. griseum 2-10 cm long, 0.4-2 cm wide, petiolate,

Rydb.; H. scouleri, authors]. Perennial herbs; blades oblanceolate, entire or denticulate.

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

303

stipitate-glandular to glabrous or less com-
monly with a few long blackish setae; cauline
leaves much reduced; heads solitary or more
commonly greenish black, stellate hairy, and
with long black setae; pappus tawny. Lodge-
pole pine, spruce-fir, and grass-forb commu-
nities at 3050 to 3390 m in Duchesne, Sum-
mit, and Uintah counties; Alaska and Yukon
to Mackenzie, south to California and New
Mexico; South America; 13 (v).

wide, sessile, and more or less clasping; heads
conspicuous, in racemose or corymbose clus-
ters; bracts with long-attenuate, often reddish
tips, subequal to the disk; rays reddish to
purple or yellowish, hairy and glandular;
pappus scales unequal, lacerate; achenes 6-7
mm long. Pinyon-juniper community at 2135
m in the Beaverdam Mountains, Washington
County (Higgins 1410 BRY); Nevada and
California; 1 (0).

HOFMEISTERIA Walp.

Shrubs; leaves opposite below, alternate
above, simple, petiolate; heads discoid, few
to several in terminal corymbose clusters; in-
volucre campanulate; bracts striate, narrow,
imbricated; receptacle naked; disk flowers
whitish; pappus of 10-12 scabrous bristles
and other short scales; achenes 5-angled, cal-
lous-thickened.

Hofmeisteria pluriseta Gray Arrowleaf.
Shrubs, low, rounded, and intricately
branched, mostly 3-8 dm tall; branchlets
green, glandular-puberulent, becoming white
barked in age; leaves long petioled, the pet-
ioles 0.8-4 cm long, the blades hastately
lobed to entire, 4-10 mm long, 2-4 mm
wide; heads small; involucres 4-9 mm high;
bracts 3-lined, acuminate; disk flowers whit-
ish. Reported for Utah in Munz (Flora of Cal-
ifornia, p. 267); to be sought on rock out-
crops at lower elevations in Washington
County; Nevada, California, and Arizona; 0
(0).

HULSEA T. & G.

Perennial viscid-pubescent aromatic herbs;
leaves alternate, simple; heads radiate; in-
volucres hemispheric, the bracts subequal in
2 or 3 series, herbaceous, finally reflexed; re-
ceptacle convex, naked; ray flowers yellowish
to purplish, pistillate, fertile; disk flowers
perfect, fertile; pappus of 4 hyaline scales
united at the base; achenes compressed, an-
gled, villous.

Hulsea heterochroma Gray Perennial
herbs from a stout taproot; herbage viscid-
villous, scented, 3-10 (12) dm tall; basal
leaves oblanceolate or spatulate, tapering to
a broadly petioled base, dentate; cauline
leaves mainly 3-10 cm long, 1.5-3.5 cm

Hymenoclea T. & G.

Xerophytic shrubs; leaves alternate, linear,
usually entire; heads discoid, small, numer-
ous, mostly glomerate-paniculate, with both
sexes in each leaf axil, the staminate above
the pistillate; staminate heads several-flow-
ered; pistillate heads 1-flowered; involucre
becoming indurated and beaked in fruit, the
bracts persistent as scarious wings; pappus
none.

Hymenoclea salsola T. & G. Burrobrush.
Shrubs, 6-12 (15) dm tall; branchlets green,
becoming straw colored to gray in age; herb-
age yellow green, resinous, glabrous or
scabrous; leaves 2-5 cm long, linear, entire;
staminate heads 2-3 mm high, 2.5-5 mm
wide, the bracts obtuse to rounded, ciliate on
the hyaline margin; pistillate heads mainly
6-9 mm high at maturity, the middle and up-
per bracts with white, chartaceous, broadly
rounded, erose margins, longitudinally
veined. Blackbrush, creosote bush, and Joshua
tree communities at 670 to 900 m in Wash-
ington County; Nevada, Arizona, and Cali-
fornia; 19 (ii).

Hymenopappus L'Her

Perennial herbs; leaves alternate or mainly
basal (and still alternate), mainly pinnatifid;
heads discoid, the flowers perfect; involucral
bracts in 2 or 3 series, subequal, at least the
inner with broad rounded scarious or hyaline
margins; receptacle flat, naked or rarely
chaffy; corollas yellow or white; anthers sag-
ittate; pappus of several membranous scales;
achenes 15- to 20-nerved, 4- or 5-angled.
Turner, B. L. 1956. A cytotaxonomic study
of the genus Hymenopappus (Compos-
itae). Rhodora 58:163-308.

Hymenopappus filifolius Hook. Perennial
subscapose herbs; stems 5-60 (100) cm tall.

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Great Basin Naturalist

Vol. 43, No. 2

tomentose to glabrate; basal leaves 3-20 cm
long, twice pinnately dissected, the ultimate
divisions mainly 2-25 mm long, minutely
punctate; cauline leaves lacking or several,
much reduced upward; heads solitary or
more commonly few to numerous, turbinate
to campanulate, with 10-59 flowers or more,
on peduncles 0.5-10 cm long or more; in-
volucral bracts mainly 3-14 mm long; co-
rollas yellow or white, 2-7 mm long; pappus

of narrowly oblong scales; achenes 3-7 mm
long, densely hairy. This is a polymorphic
species, which consists of a series of geo-
graphic and/or edaphically correlated in-
fraspecific taxa. Those taxa peripheral to the
main body of the species in the Colorado
Plateau province are the most distinctive.
The following treatment differs from that of
Turner (1956) and represents a more conserv-
ative approach.

1. Basal leaf axils sparingly tomentose or glabrous; stems scapose, or with 1 or 2

leaves; plants of high elevations H. filifolius var. alpestris

— Basal leaf axils prominently white-tomentose; stem leaves often more than 2;
plants of middle and lower elevations 2

2(1). Corollas 2-3 mm long; flowers fewer than 30; plants of Daggett and Uintah

counties H. filifolius var. luteus

— Corollas 3-7 mm long, or, if shorter, not of Daggett or Uintah counties; flow-
ers in main heads often more than 30; leaves more coarsely dissected; plants of
various distribution 3

3(2). Flowers white; achene hairs 0.5-1 mm long; plants of Washington County

H. filifolius var. eriopodus

— Flowers yellow; achene hairs 1-2 mm long; plants more widely distributed 4

4(3). Leaves mainly basal; plants of the Great Basin H. filifolius var. nanus

— Leaves cauline and basal; plants of the Colorado drainage system

H. filifolius var. cinereus

Var. alpestris (Maguire) Shinners [H. nu-
dipes var. alpestris Maguire; H. nudipes Ma-
guire; H. filifolius var. nudipes (Maguire)
Turner]. Ponderosa pine, western bristlecone
pine, sagebrush-grass, limber pine, aspen, and
alpine tundra communities, commonly on
limestone or thermally modified igneous out-
crops, at 2445 to 3450 m, in Beaver, Carbon,
Duchesne, Emery, Iron, Garfield, Kane, Mil-
lard, Piute, Sanpete, Sevier, Summit, Utah,
and Washington counties; Wyoming; 46 (ix).
This is the most distinctive of the varieties
within H. filifolius in Utah.

Var. cinereus (Rydb.) Johnst. [H. cinereus
Rydb.; H. lugens Greene; H. filifolius var. lu-
gens (Greene) Jepson; H. filifolius var. me-
gacephalus Turner, as to Utah materials; H.
pauciflorus Johnst., type from near Bluff; H.
filifolius var. pauciflorus (Johnst.) Turner; H.
tomentosus Rydb., type from St. George; H.
filifolius var. tomentosus (Rydb.) Turner; H.
niveus Rydb., type from Springdale]. Black-
brush, warm desert shrub, salt desert shrub,
sand sagebrush-ephedra, pinyon-juniper, pon-
derosa pine, and sagebrush communities at

1065 to 2685 m in Carbon, Daggett, Du-
chesne, Emery, Garfield, Grand, Kane, San
Juan, Uintah, Washington, and Wayne coun-
ties; Colorado, California, Arizona, New
Mexico, and Texas; 144 (xxix). The variety
cinereus, as interpreted herein, includes three
largely sympatric phases that were treated by
Turner (1956) at varietal rank. Although
there is a tendency for these phases to be
geographically correlated, they are con-
nected completely by series of intermediates,
and they can be segregated only arbitrarily.
Turner noted that herbarium specimens of
var. megacephalus from eastern Utah, inter
alia, carried a "hodge-podge of annotations:
H. lugens, cinereus, pauciflorus, tomentosus,
eriopodus, etc.," and further that "the possi-
bility exists that the variety [megacephalus]
here typified includes only the individuals
from Clark County, Nevada, and vicinity,
and that most of the remaining material to
the east represents either a weakly defined
separate variety or a common area of exten-
sive hybridization and introgression among
the several peripheral taxa mentioned

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

305

above, ..." A phase of the cinereus complex
from Washington County has cauline leaves
well developed and plant bases appearing
bulbous due to a copious tomentum. These
plants are apparently intermediate between
var. tomentosus (Rydb.) Turner, in a narrow
sense, and var. eriopodus (A. Nels.) Turner,
which share the feature of the "bulbous"
bases.

Var. eriopodus (A. Nels.) Turner [H. eriop-
odus A. Nels., type from Diamond Valley].
Pinyon-juniper community at 1675 to 2135 m
in Washington County; California to Nevada;
3 (0). This variety is evidently rare in Utah,
and might best be treated within an ex-
panded var. cinereus (q.v.).

Var. luteus (Nutt.) Turner [H. luteus
Nutt.]. Salt desert shrub, mixed cool desert
shrub, and pinyon-juniper communities at
1525 to 1830 m in Daggett, Summit, and
Uintah counties; Wyoming and Colorado; 7
(ii). The small flowers and finely divided
leaves appear to be diagnostic for this
variety.

Var. nanus (Rydb.) Turner [H. nanus
Rydb.]. Black sagebrush-rabbitbrush, pinyon-
juniper, and ponderosa pine communities at
1490 to 2300 m in Beaver, Garfield, Iron,
Juab, Millard, Piute, Sevier, Tooele, and
Washington counties; Nevada, California,
and Arizona; 26 (v).

Hymenoxys Cass.

Perennial or biennial herbs from a taproot
and commonly with a pluricipital caudex;
stems simple or branched; leaves basal or bas-
al and cauline, simple and entire or pinnately
to ternately divided; heads radiate, peduncu-
late; involucres hemispheric; bracts in 2 or 3
series, the outer distinct or connate basally,
subequal or imbricate, herbaceous or carti-
laginous; receptacle naked, hemispheric; ray
flowers yellow, pistillate, fertile, prominently
veined, 3-toothed; disk flowers perfect, fer-
tile; pappus scales usually 5, hyaline, nerved
or nerveless, the nerve often produced into
an awn; achenes obpyramidal, more or less 5-
angled, appressed hairy.

1. Leaves entire, essentially all basal 2

— Leaves pinnatifid or palmatifid, or some entire, the cauline ones well
developed 5

2(1). Involucral bracts sparsely pubescent or glabrous apically, the margins thin and

scarious or hyaline 3

— Involucral bracts moderately to densely villous-pilose or some rarely glabrous,

the margins not at all or only narrowly scarious 4

3(2). Plants depressed pulvinate-caespitose, acaulescent; outer involucral bracts re-
curved, thickened and reddish apically; disks less than 10 mm wide H. lapidicola

— Plants merely caespitose, scapose; outer involucral bracts erect, not thickened

and seldom reddish apically; disks over 10 mm wide H. torreyana

4(2). Plants pulvinate-caespitose; caudex branches clothed with a marcescent thatch

of erect-ascending leaf bases; leaves mainly linear, cuspidate apically H. depressa

— Plants seldom pulvinate-caespitose; caudex branches without a definite thatch
of ascending or erect leaf bases; leaves various, sometimes cuspidate apically ...
H. acaulis

5(1). Disks 18-30 mm wide or more; involucral bracts similar, distinct, in 2 or 3 in-
definite subequal series; herbage villous-tomentose; plants of high elevations ....
H. grandiflora

— Disks 7-22 mm wide; involucral bracts in 2 dissimilar series, the outer thick-
ened and united at the base 6

6(5). Plants silvery-canescent; leaves entire or 3-cleft, the blades or segments 1.5-4

mm wide H. subintegra

— Plants green, or, if silvery-canescent, the leaves commonly 3- to 5-cleft 7

7(6). Plants apparently biennial, with an evident basal rosette and taproot; cauline

leaves numerous, gradually reduced upward, temate or palmatifid H. cooperi

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Great Basin Naturalist

Vol. 43, No. 2

— Plants perennial, from a taproot and caudex; cauline leaves rather well devel-
oped, palmatifid to entire 8

8(7). Stems merely glandular or glandular-scabrous; plants of low elevation saline

meadows in western Utah H. lemmonii

— Stems more or less villous; plants of various habitats, but seldom of saline
meadows and not of western Utah 9

9(8). Stems few to several from a pluricipital caudex; leaf bases conspicuously long-

villous below; leaf segments mainly 1-2.5 mm wide H. richardsonii

— Stems solitary or few from a simple or branched caudex; leaf bases glabrous or
only somewhat hairy; leaf segments 2-6 mm wide H. helenoides

Hymenoxys acaulis (Pursh) Parker [Gail-
lardia acaulis Pursh]. Perennial caespitose
herbs from a short multicipital caudex, the
caudex branches clothed with short brownish
or blackish marcescent leaf bases, 2-50 cm
tall, villous to glabrous; leaves 1-6 cm long,
2-8 mm wide, all basal or some cauline,
glandular-punctate or epunctate, linear to

disk 7-20 mm broad; bracts distinct, in 2 or 3
subequal series, 4-9 mm high; rays 5-9, yel-
low, 6-15 mm long; pappus scales 2.5-4.5
mm long, acute or shortly awned; achenes
2.5-4.5 mm long. This is a complex entity,
consisting of a series of morphological phases,
which are more or less geographically or
edaphically correlated.

oblanceolate, entire; heads solitary (rarely 2);

1. Plants with 1-4 (or more) cauline leaves; stems simple or branched

//. acaulis var. ivesiana

— Plants scapose, with cauline leaves lacking or rarely with 1; scapes un-
branched 2

2(1). Leaves linear to linear-oblanceolate, conspicuously glandular-punctate, spar-
ingly long-hairy to glabrous; plants of the Colorado drainage system

H. acaulis var. arizonica

— Leaves narrowly to broadly oblanceolate, inconspicuously glandular-punctate,
merely punctate, or epunctate, or plants of the Great Basin, densely pilose to
villous or glabrous 3

3(2). Leaves epunctate or nearly so, glabrous or less commonly silky-hairy; plants of

the Colorado drainage system H. acaulis var. caespitosa

— Leaves punctate, silky-hairy, or less commonly glabrous; plants of the Great
Basin H. acaulis var. acaulis

Var. acaulis Sagebrush, mixed desert
shrub, pinyon-juniper, and bunchgrass com-
munities, often on windswept ridges, at 1525
to 2990 m in Beaver, Box Elder, Juab, Mil-
lard, Sanpete, Sevier, Tooele, and Washing-
ton counties; Idaho east to Saskatchewan,
south to Nevada, Colorado, and Texas; 42
(viii). Specimens from the Great Basin might
not belong to var. acaulis in a strict sense,
and perhaps should be regarded as a separate
variety. The problem cannot be solved on the
basis of Utah specimens alone.

Var. arizonica (Greene) Parker [Tetra-
neuris arizonica Greene]. Salt and sandy
desert shrub, pinyon-juniper, sagebrush, blue
grama, aspen, Douglas fir, white fir, and pon-
derosa pine communities at 1220 to 3175 m

in Carbon, Daggett, Duchesne, Emery,
Grand, Kane, San Juan, and Uintah counties;
Colorado and Arizona; 108 (xvi).

Var. caespitosa (A. Nels.) Parker [Tetra-
neuris acaulis var. caespitosa A. Nels.; Tetra-
neuris epunctata A. Nels., type from Dyer
Mine]. Shadscale-eriogonum, black sage-
brush, sagebrush, pinyon-juniper, mountain
brush, and alpine tundra, often on plateau
margins and windswept ridges, at 1585 to
3510 m in Carbon, Daggett, Duchesne,
Emery, Grand, Sanpete, Sevier, Summit, and
Uintah counties; Wyoming south to New
Mexico; 87 (viii).

Var. ivesiana (Greene) Parker [Tetraneuris
ivesiana Greene; H. ivesiana (Greene) Park-
er]. Sand sagebrush, ephedra, pinyon-juniper,

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

307

and ponderosa pine communities at 1150 to
2505 m in Garfield, Grand, Kane, San Juan,
and Wayne counties; Colorado, New Mexico,
and Arizona; 66 (xii). This variety approaches
phases of the partially sympatric var. arizo-
nica in stature, and it is possible to confuse
some specimens when cauline leaves are
lacking and the stems are unbranched. The
varieties acaulis, arizonica, and caespitosa are
tetraploids, i.e., 2n = 60, whereas var. ive-
siana is diploid, i.e., 2n = 30. Because of this
difference, Parker (1960. Leafl. W. Bot. 9:
93) elevated this taxon to specific rank.

Hymenoxys cooperi (Gray) Cockerell [Ac-
tinella cooperi Gray; A. biennis Gray, type
from Washington County?]. Biennial or
short-lived perennial herbs; stems 16-60 (80)
cm tall, leafy, simple below, branched in a
corymbose inflorescence above, often red-
dish, scurfy villous, canescent; basal rosette
leaves mainly 2-10 cm long, pinnately di-
vided, the linear lobes often again divided,
mainly 1-1.5 mm wide; stem leaves longer
than the internodes; heads (1) 3-50; in-
volucres 5-6 mm high, 10-24 mm wide,
hemispheric; bracts thickened and united ba-
sally, more or less pubescent and glandular;
rays 7-13, yellow, 6-15 mm long; pappus
scales acuminate; achenes 2-3 mm long,
densely pilose. Sagebrush and pinyon-juniper
communities at 975 to 2380 m in Garfield,
Juab, Kane, and Washington counties; Ne-
vada, California, and Arizona; 18 (i).

Hymenoxys depressa (T. & G.) Welsh &
Reveal [Actinella depressa Gray]. Pulvinate-
caespitose scapose perennial herbs from a
multicipital caudex, the caudex branches
clothed with conspicuous, commonly erect
marcescent leaf bases (often forming a
thatch), 1-4 cm tall; scapes villous; leaves
0.4-3 (4) cm long, 1-2 (4) mm wide, linear to
oblanceolate, the outer sparingly if at all
glandular-punctate, the inner definitely so,
sparingly villous to glabrous, cuspidate; heads
solitary; disk 6-10 mm wide; involucres 4-6
mm high; bracts in 2 or 3 subequal series,
long villous, the margins nonscarious, the
apices erect; rays 5-7, yellow, 3-6 mm long;
pappus scales 2-3 mm long, long-acuminate;
achenes 2-3 mm long. Ephedra, sagebrush,
shadscale, and pinyon-juniper woodland at
1340 to 2170 m in Duchesne, Emery, and
eastern Sevier counties; endemic? There is

justification for inclusion of H. depressa with-
in the H. acaulis complex, at some in-
fraspecific rank. And the plants have been
suggested as merely depauperate phases of
that group. However, if they are ecologically
controlled variations, they should be ex-
pected through much of the range of H.
acaulis; but they are not. Dwarf forms of H.
acaulis, especially of the var. caespitosa, have
been mistaken for this species, but they are
more hairy, have usually broader leaves, and
lack glandular punctate. There is also a ques-
tion of typification; the type of H. depressus
was taken by Fremont, on his second expedi-
tion in the Rocky Mountains. Fremont evi-
dently traversed the area occupied by H.
depressa in 1845, and the material could have
come from western Emery County; 24 (iv).

Hymenoxys grandiflora (T. & G.) Parker
[Actinella grandiflora T. & G.]. Perennial
herbs from a taproot and usually simple cau-
dex, this clothed with brown marcescent leaf
bases; stems mainly 5-25 cm tall, 1 to sever-
al, simple or branched basally, densely vil-
lous; leaves basal and cauline, 2-10 cm long,
2- or 3-times ternately or palmately divided,
the lobes linear, villous to glabrate; heads
solitary; disk 1.5-3 cm wide or more; in-
volucral bracts subequal, in 2 or 3 series,
8-14 (16) mm high, densely villous-tomen-
tose; rays 15-50, yellow, 25-35 mm long;
pappus scales 3.5-7 mm long, attenuate;
achenes 3-5 mm long. Sedge-forb commu-
nities at or above timberline, often in talus or
rockstripes, at 3050 to 3660 m in Duchesne,
Grand, Salt Lake, San Juan, Summit, and
Utah counties (Uinta, Wasatch, and La Sal
mountains); Idaho to Montana, south to Colo-
rado; 14 (ii). This is a strikingly beautiful yel-
low sunflower of alpine tundra in our
mountains.

Hymenoxys helenioides (Rydb.) Cockerell
[Picradenia helenioides Rydb.]. Perennial
herbs from a simple or branched caudex, this
clothed with broad brown marcescent leaf
bases; stems mainly 25-45 cm tall, simple be-
low, branched above, scurfy and more or less
villous; leaves basal and cauline, mainly 5-15
cm long, entire or 2- to 5-lobed, the lobes
mainly 3-8 mm wide, finely glandular-punc-
tate, glabrous or puberulent; heads 3-13, in
corymbose clusters; disks 10-21 mm wide; in-
volucres 6.5-8 mm high, the outer bracts

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Great Basin Naturalist

Vol. 43, No. 2

green, connate in the lower portion, more or
less villous and glandular; rays 5-11, yellow,
8-19 mm long; pappus scales 2.5-3.5 mm
long, acuminate; achenes 2.5-3 mm long.
Mountain brush, sagebrush, and aspen com-
munities, often in meadows, at 2440 to 2990
m in Emery, Garfield, Sanpete, and Sevier
counties; Colorado and Arizona; 10 (i). This
handsome plant has long remained obscure in
Utah, partially due, no doubt, to its resem-
blance to Helenium hoopesii (q.v), with
which it occurs in the aspen communities of
central and southern Utah.

Hymenoxys lapidicola Welsh & Neese Pul-
vinate caespitose herbs from a multicipital
caudex, this densely clothed with brown mar-
cescent leaf bases, acaulescent; leaves all bas-
al, 0.3-1.2 cm long, 0.8-2 mm wide, narrow-
ly oblanceolate, the inner conspicuously
glandular-punctate, the blades glabrous, the
axils long-villous; heads solitary, immersed in
the leaves; disks 5.5-9 mm wide; involucres
5-8 mm high; bracts distinct, in 2 or 3 sub-
equal series, sparingly villous and suffused
reddish, the margins scarious, the tips more
or less squarrose-spreading and somewhat
thickened; rays 5 or 6, yellow, 5-6 mm long;
pappus scales lance-acuminate, 2.3-3 mm
long; achenes 2-2.5 mm long, pilose. Pinyon-
juniper and ponderosa pine-manzanita com-
munities, often in rock crevices, at 1830 to
2476 m in Uintah County; endemic; 4 (0).

Hymenoxys lemmonii (Greene) Cockerel!
[Picradenia lemmonii Greene; H. lemmonii
ssp. greenei Cockerell, type from Washington
County (?)]. Perennial herbs from a taproot
and short ligneus caudex, the caudex clothed
with brown to straw-colored or purplish
clasping leaf bases; stems 20-60 cm tall, 1 to
few, glabrous; leaves cauline and basal, 2-15
cm long, pinnately parted, the lobes linear,
2-3 mm wide, glabrous, glandular-punctate;
cauline leaves longer than the internodes, the
uppermost often unlobed; heads 5-12; in-
volucres 4.5-7 mm high, hemispheric, 8-14
mm wide; outer bracts green, sparsely scurfy
and glandular, thickened dorsally and con-
nate below; rays 6-10, yellow, 6-13 mm
long; pappus lance-attenuate, 1.6-2 mm
long; achenes 2.5-3 mm long, pilose. Saline
rabbitbrush-alkali sacaton meadows at 1660
m in Millard, Tooele, and Washington (?)
counties; Nevada and California; 3 (i).

Hymenoxys richardsonii (Hook.) Cock-
erell Colorado Rubberweed. [Picradenia rich-
ardsonii Hook.]. Perennial caespitose herbs
from a pluricipital ligneus caudex, the caudex
branches clothed with a thatch of marcescent
brown leaf bases, usually with villous leaf
axils; stems few to numerous, 6-40 (50) cm
tall, simple below, branched; leaves basal and
cauline 2-12 (15) cm long, ternate or with
5-7 linear segments, or some entire, pu-
bescent to glabrous; involucres hemispheric,
5-8 mm high, the outer bracts connate ba-
sally, thickened dorsally, green or char-
taceous, more or less villous; rays 9-14, yel-
low, 8-20 mm long; pappus 2-4.5 mm long,
acuminate; achenes 2.5-4 mm long, pilose.
Salt desert shrub, cool desert shrub, pinyon-
juniper, sagebrush, mountain brush, pon-
derosa pine, aspen, fir, and western bristle-
cone pine communities at 1460 to 2870 m.
Our material falls into two varieties, a low
plant with 1-5 large heads of Daggett and
Uintah counties, belonging to var. richard-
sonii, and a taller plant with 3-20 smaller
heads of Beaver, Carbon, Duchesne, Emery,
Garfield, Kane, Millard, Piute, Sanpete, Se-
vier, Uintah, and Wayne counties, belonging
to var. florabunda (Gray) Parker; Alberta and
Saskatchewan to Arizona and Texas; 129
(xvii). The plants are considered poisonous to
sheep, cattle, and goats.

Hymenoxys subintegra Cockerell Per-
ennial (or biennial) herbs from a taproot;
stems solitary or few, 10-60 cm tall,
branched above; herbage silvery canescent;
basal leaves often withered at flowering;
cauline leaves numerous, 1.5-8 cm long, 2-4
mm wide, entire or 2- or 3-lobed; heads few
to several; disks 9-12 mm wide; involucres
5-7 mm high; outer bracts connate basally,
villous; rays ca 12-20, yellow, 5-10 mm long;
pappus scales lance-acuminate, 2.8-3.2 mm
long; achenes ca 3 mm long. Ponderosa pine,
aspen, and spruce-fir communities in Sanpete
(Maguire 20049 BRY) and Washington (re-
ported by Meyer) counties; Arizona; 1 (0).

Hymenoxys torreyana (Nutt.) Parker [Ac-
tinella torreyana Nutt.]. Perennial caespitose
scapose herbs from a stout pluricipital cau-
dex, the caudex branches densely clothed
with brown to straw-colored or ashy leaf
bases, 3-10 cm tall, villous; leaves 1-7.5 (9)

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

309

cm long, 2-6 mm wide, all basal, glandular-
punctate, narrowly oblanceolate, entire;
heads solitary; disk 12-20 mm wide; in-
volucres hemispheric, 5-10 mm high; bracts
distinct, in 2 or 3 subequal series, less pu-
bescent to glabrous apically, the margins
scarious, not thickened apically, the tips
erect, sometimes reddish; rays 10-16, yellow,
8-20 mm long; pappus scales ovate-acumi-
nate, 2.8-3.5 mm long; achenes 2-3 mm long.
Pinyon-juniper, sagebrush, and mountain
brush communities at 1830 to 2200 m in
Daggett and Uintah counties; Wyoming; 6
(0).

Hypochaeris L.

Perennial subscapose herbs from taproots,
the juice milky; leaves primarily basal,
simple, pirmately lobed to pinnatifid, the
cauline leaves small and bractlike; heads soli-
tary or few in a branching inflorescence; in-
volucral bracts in several series, greenish
black, the irmer ones with hyaline margins;
receptacle chaffy; corollas of ray flowers
only, perfect, yellow or purplish on the dor-
sal surface; pappus of plumose capillary
bristles; style branches semicylindrical;
achenes several-nerved, subterete, minutely
roughened, long beaked.

Hypochaeris radicata L. Cat's-ears. Plants
1.5-5 dm tall, the stems simple or branched
above, glabrous or spreading-hairy below;
basal leaves 3-16 (25) cm long, 0.5-3.5 (5) cm
broad, oblanceolate, pinnately toothed or
piimatifid, sparsely to moderately spreading-
hairy above and below, rounded to obtuse
apically, tapering to a broad petiole basally;
cauline leaves alternate, minute or lacking;
heads solitary, or more commonly 2-5; pe-
duncles glabrous; involucres 5-15 mm high,
7-20 mm wide; bracts glabrous or stiffly
hairy along the midribs; corollas numerous,
longer than the bracts; achenes 4-7 mm long,

the beak mostly 2-3 mm long. Weedy species
of disturbed soils in Davis and Salt Lake
counties; widespread in North America; ad-
vent! ve from Europe; 2 (0).

Inula L.

Perennial tomentose herbs; leaves basal
and cauline, alternate; heads radiate, large,
hemispheric, few to numerous in cymose
clusters; involucral bracts imbricate in sever-
al series; receptacle naked; ray flowers pistil-
late, yellow, 3-toothed; disk flowers perfect,
fertile; anthers sagittate at the base; style
branches of disk flowers linear; pappus of
capillary bristles; achenes 4- or 5-ribbed.

Inula helenium L. Elecampane. Perennial
herbs, mainly 6-20 dm tall, from thick roots;
stems simple below; basal leaves 2-5 dm
long, petiolate, the blades ovate to oblong,
denticulate, rough-hairy above, velvety be-
neath; cauline leaves reduced upward, cor-
date-clasping, acute; heads large and showy;
involucres 15-23 mm high, 30-50 mm wide;
outer bracts foliaceous, ovate; ray flowers nu-
merous, 18-30 mm long, narrow; achenes
glabrous. Canal banks and moist meadows at
1370 to 1830 m in Sanpete and Utah coun-
ties; widespread in North America; adventive
from Eurasia; 2 (i).

IvaL.

Annual or perennial herbs; leaves opposite,
at least below; heads discoid, the pistillate
flowers few, with corolla tubular or lacking;
involucres campanulate; bracts subequal or
imbricate in 1-3 series, sometimes with a
short inner series subtending the achenes; re-
ceptacle chaffy, the receptacular bracts lin-
ear to spatulate; staminate flowers with abor-
tive pistils, the styles undivided, the filaments
monadelphous; anthers obtuse basally, almost
distinct; pappus none; achenes compressed.

Leaves sessile or shortly petiolate, entire; plants rhizomatous, mainly less than

40 cm tall, of saline low-elevation sites 7. axillaris

Leaves petiolate, serrate; plants taprooted annuals, mainly much over 40 cm
tall, ruderal weeds /. xanthifolia

Iva axillaris L. Poverty Weed. Perennial
herbs from elongate rhizomes; stems 15-50
(60) cm tall, branched from the base; herbage

strigose to strigulose and more or less glandu-
lar; leaves opposite below, alternate above,
0.8-4.5 cm long, 4-15 mm wide, elliptic to

310

Great Basin Naturalist

Vol. 43, No. 2

obovate or lanceolate; heads numerous in ter-
minal bracteate spicate clusters, nodding, 3-7
mm wide; bracts connate, shallowly 4- or 5-
lobed; pistillate flowers 4-8, perfect; achenes
2-3 mm long, glandular. Commonly in saline
riparian sites in the warm desert shrub, salt
desert shrub, pinyon-juniper, and aspen com-
munities at 760 to 2440 m in all Utah coun-
ties; British Columbia to Manitoba, south to
California, New Mexico, and Oklahoma; 60
(viii).

Iva xanthifoUa Nutt. Marsh-elder. Coarse
perennial herbs, mainly 4-25 dm tall, simple
or branched, essentially glabrous below,
glandular above; leaves opposite below, pet-
iolate, the blades 4-20 cm long and about as
wide, broadly ovate to lance-ovate, serrate
and sometimes lobed, green above, canescent
beneath; heads 2-4 mm thick, numerous,
borne ebracteate in paniculate clusters; in-
volucral bracts distinct, ovate; pistillate flow-
ers 5; achenes sparsely pilose apically, ca 2
mm long. Ruderal weeds of disturbed soils at
1370 to 2290 m in Beaver, Duchesne, Emery,
Iron, Kane, Millard, Salt Lake, Sevier, Sum-
mit, Uintah, Utah, and Wayne counties; Al-
berta to Saskatchewan, south to Washington,
Arizona, and New Mexico; widely distributed
elsewhere; 22 (iii).

KUHNIA L.

Perennial herbs from a woody caudex and
taproot; stems branched, erect or ascending;

leaves alternate or some lower ones opposite;
entire or lobed; heads discoid, several to nu-
merous in paniculate clusters; involucres
campanulate; involucral bracts in 4-7 series,
the outer ones only graduated; receptacle
naked; disk flowers perfect, fertile, whitish;
style tips flattened, clavate; pappus of plu-
mose bristles; achenes 10-ribbed.

Kuhnia chlorolepis Woot. & Standi. Per-
ennial clump-forming herbs; stems 30-75 cm
tall, much branched, minutely hairy; leaves
8-50 mm long (or more), 1-3 mm wide, en-
tire or with a pair of basal lobes, linear; in-
volucres 8-12 mm high; bracts linear to nar-
rowly oblong, striate; corollas 6-7.5 mm
long; achenes 4.8-5.2 mm long, dark brown,
short-hairy. Rabbitbrush community in inter-
mittent stream courses at 1890 to 2045 m in
Uintah County; Colorado to Arizona, New
Mexico, Texas, and Mexico; 2 (i).

Lactuca L.

Annual, biennial, or perennial herbs; leaves
alternate, entire or pinnatifid; flowers all
raylike, yellow, blue, or white; heads panic-
ulately arranged; involucres cylindrical;
bracts imbricate in several series; receptacle
flat, naked; pappus copious, of white or
brownish capillary bristles; achenes oval, ob-
long, or linear in outline, compressed, ribbed
on each face, short- to long-beaked.

1. Plants perennial, rhizomatous; rays long-exserted, blue L. tatarica

— Plants annual or biennial; rays not long-exserted, yellow (often fading blue) or
blue to white (in L. biennis) 2

2(1). Achenes prominently 1-nerved on each side 3

— Achenes prominently several nerved on each side 4

3(2). Involucres 10-15 mm high in fruit; pappus 5-7 mm long; achenes 4.5-6.5

mm long, including the beak L. canadensis

— Involucres 15-22 mm high in fruit; pappus 7-12 mm long; achenes 7-10 mm
long L. ludoviciana

4(2). Involucres cylindrical at anthesis; flowers not fading blue; plants cultivated

and occasionally escaping L. sativa

— Involucres tapering to the apex at anthesis; flowers fading blue 5

5(4). Achenes with a long filiform beak as long as or longer than the body of the

achene; pappus white L. serriola

— Achenes with a short beak much shorter than the body of the achene; pappus
brownish L. biennis

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

311

Lactuca biennis (Moench) Fern. [Sonchus
biennis Moench]. Annual or biennial,
glabrous or hairy (on midvein of leaves)
herbs; stems erect, mainly 6-20 (35) dm tall;
leaves mainly 10-40 cm long, 4-20 mm wide,
pinnatifid or merely toothed; heads 13- to 50-
flowered, nimierous, arranged in a narrow
paniculate inflorescence; rays bluish to
white; pappus brownish; achenes 4-5.5 mm
long, prominently several nerved on each
face, beakless or short beaked. Moist sites at
ca 1800 m in Salt Lake County (Arnow 2561
BRY, UT); Alaska to Newfoundland, south to
California, Colorado, and North Carolina; 1
(0).

Lactuca canadensis L. Annual or biennial,
glabrous or hirsute herbs; stems erect, 3-25
dm tall; leaves mainly 10-35 cm long, 1-12
cm wide, entire to pinnatifid; heads mostly
13- to 22-flowered, arranged in open pan-
icles; rays yellow (fading blue); pappus
white; achenes black, obovate, transversely
rugose and with 1 prominent longitudinal
vein on each face, 4.5-6.5 mm long, includ-
ing the beak from half as long to as long as
the body. Weedy species of moist sites at ca
1155 m in Kane County (Atwood 4118 BRY);
widespread in U.S.; 1 (0).

Lactuca ludoviciana (Nutt.) Riddell [Son-
chus ludovicianus Nutt.]. Biennial or short-
lived perennial herbs; stems 6-15 dm tall or
more; leaves 10-35 cm long or more, mainly
1-10 (20) cm wide, commonly pinnatifid and
weakly spinose-toothed, setose-hispid on the
lower midrib, the uppermost auriculate-
clasping; heads numerous in an open pan-
iculate cluster, the peduncles bracteate; in-
volucres 15-22 mm high in fruit; heads most-
ly 20- to 50-flowered, the flowers yellow or
sometimes blue, fading blue; pappus white,
7-10 mm long at maturity; achenes flattened,
blackish, with a longitudinal median nerve on
each face, transversely rugulose, 4-5 mm
long. Collected once in Salt Lake County
(without collector, UT); widespread in the
northwestern U.S.; 1 (0).

Lactuca sativa L. Lettuce. Annual herbs;
stems erect, mostly 5-12 dm tall, glabrous;
leaves mainly 10-30 cm long and as broad,
undulate-crisped and serrate, glabrous; in-
volucres 7-10 mm high; heads ca 15-flow-
ered, the flowers yellow, not fading blue,

numerous in a paniculate cluster; pappus
white; achenes brownish, oblanceolate in out-
line, flattened, hispid apically, 3.5-4.5 mm
long, with 5-7 longitudinal nerves on each
face, the beak 2.5-3.5 mm long. Cultivated
food plant in much of Utah; introduced from
Europe; 2 (0).

Lactuca serriola L. Prickly Lettuce. [L.
scariola, scarriola, orthographic variants].
Biennial or winter annual herbs; stems erect,
3-18 dm tall, hispid below or glabrous over-
all; leaves mainly 3-30 cm long, 1-10 cm
wide, pinnatifid or pinnately lobed, or
merely spinose-toothed, the blades vertically
oriented (twisted at the base), setose-hispid
on main veins beneath; involucres 7-15 mm
high at maturity; heads mostly 6- to 12-flow-
ered, the flowers yellow, fading blue, several
to numerous in a paniculate cluster; pappus
white; achenes brown, the body obovate to
oblong in outline, flattened, hispid along
margin apically, 3-4.5 mm long, with 5-8
longitudinal nerves on each face, the beak
3-7 mm long. Ruderal weeds at 850 to 2440
m, probably in all Utah counties, widely dis-
tributed in the U.S.; adventive from Europe;
37 (v). This species invades lower elevation
range lands, where it is eaten by wildlife and
livestock. It is reported to produce fertile
hybrids with L. sativa (q.v.).

Lactuca tatarica (L.) CA. Mey. Blue Let-
tuce. Perennial rhizomatous herbs; stems
2-12 dm tall, glabrous; leaves 4-20 cm long,
5-35 mm wide, linear to lanceolate or ob-
long, entire, toothed, lobed, or pinnatifid,
short-petiolate below, sessile above; in-
volucres 10-20 mm high; heads cylindric, 15-
to 50-flowered, the flowers blue, numerous in
an elongate paniculate cluster; pappus white;
achenes black to pale, oblong-lanceolate in
outline, flattened, 4-7 mm long, with several
longitudinal nerves on each face, the beak ca
2 mm long. Marshes, canal and stream banks,
and roadsides at 1370 to 2440 m in Cache,
Daggett, Duchesne, Garfield, Grand, Iron,
Juab, Kane, Millard, Piute, Salt Lake, Sevier,
Tooele, Uintah, Utah, and Weber counties;
Alaska to Minnesota, south to California and
Missouri; 39 (vii). Our material belongs to
ssp. puhhella (Pursh) Stebbins [Sonchus pul-
chellus Pursh], the North American phase of
a circumboreal species.

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Great Basin Naturalist

Vol. 43, No. 2

Lapsana L.

Annual herbs from taproots, the juice
milky; leaves alternate, simple, subentire to
toothed or lyrate-pinnatifid; heads numerous;
involucral bracts in 2 series, the inner ones
large and keeled, the outer minute, greenish;
receptacle naked; corollas of ray flowers
only, perfect, yellow; pappus none; style
branches semicylindrical; achenes subterete,
several-nerved, tapering at both ends,
beakless.

Lapsana communis L. Nipplewort. Plants
mostly 2.5-10 dm tall, the stems erect, simple
or branched, pubescent with stipitate glands
or glabrous; leaves mostly 3-10 cm long and
1.4-5 (7) cm wide, the blades subentire to
toothed, or the lower ones lyrate-pinnatifid,
sparsely hairy to glabrous above and below;
heads numerous, the peduncles glabrous or
nearly so; involucres 5-8 mm high, 3-9 mm

broad, the bracts glabrous; flowers mostly
10-14; achenes 3-5 mm long. Weedy species
of disturbed sites in Salt Lake County (Arnow
4747, BRY; UT); widely established in North
America; adventive from Eurasia; 1 (0).

Layia H. & A.

Annual herbs from taproots; leaves mainly
alternate, subentire to toothed or pinnatifid;
heads radiate, solitary or few to several, sub-
corymbose; ray and disk flowers both fertile;
involucres campanulate to broadly hemi-
spheric; bracts with thin margins abruptly di-
lated below, enclosing the ray achenes; re-
ceptacle plano-convex, chaffy marginally; ray
flowers 8-24, yellow or with the tips white;
pappus of numerous bristles, awns, or scales,
the bristles often plumose below; ray achenes
obcompressed, commonly glabrous and epap-
pose; disk achenes pubescent and pappose.

Ray flowers yellow with a white tip; pappus setae merely scabrous; anthers

black; plants rare in San Juan County L. platyglossa

Ray flowers white; pappus setae plumose; anthers yellow; plants locally com-
mon, widespread L. glandulosa

Layia glandulosa (Hook.) H. & A. Tidy-
tips. [Blepharipappus glanduhsus Hook.].
Plants slender, the stems simple or branched,
0.8-3 dm tall or more, often reddish, with
long spreading-ascending multicellular setae;
leaves 0.8-6 cm long, 1.5-16 mm wide, often
mainly basal, hispid, toothed to lobed, the
cauline ones reduced upward and finally en-
tire; heads solitary or 2 to numerous; in-
volucres 6-9 mm high, 10-18 mm wide;
bracts hispid and with some tacklike purplish
black stipitate glands; rays white, 6-15 mm
long; disk flowers numerous; ray achenes 3-4
mm long; disk achenes 3-6 mm long; pappus
of 10-12 white flattened setose scales plu-
mose to above the middle with straight capil-
lary and tangled woolly hairs. Sagebrush-
grass, grassland, and pinyon-juniper commu-
nities at 1370 to 1865 m in Daggett, Garfield,
Juab, Kane, Millard, Salt Lake, Sanpete,
Tooele, Utah, and Washington counties; Brit-
ish Columbia, south to Baja California and
Arizona; 24 (i).

Layia platyglossa (Fisch. & Mey.) Gray
[Callichroa platyglossa Fisch. & Mey.]. Plants
slender, the stems erect, simple or branched.

setose with long, multicellular hairs, often
reddish; leaves mainly 1-6 cm long, 2-7 mm
wide, with long, slender, spreading multi-
cellular hairs, the cauline leaves reduced up-
ward and finally entire; heads solitary or few;
involucres 6-12 mm high, 12-20 mm wide;
bracts hairy like the leaves and with some
tacklike purplish black stipitate glands; rays
yellow with white tips, 6-18 mm long; disk
flowers numerous; ray achenes 3-4 mm long;
disk achenes 3-5 mm long; pappus of
scabrous setae. Dunes at ca 1375 m in San
Juan County (Harrison 2545 BRY); Califor-
nia. Our material apparently belongs to var.
breviseta Gray [ssp. campestris Keck], and
this is apparently the only known station for
the species east of California. The collection
was taken in 1927. The plants resemble those
of Gaillardia, in a general way, and our ma-
terial has been filed for more than four de-
cades in a folder of that genus.

Lepidospartum Gray

Shrubs; leaves alternate, linear, entire;
heads several to numerous, in corymbose or

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

313

racemose clusters; heads discoid, the flowers
perfect, yellow; involucres subcylindric;
bracts chartaceous, inbricate in several series,
rounded apically (at least the inner); recep-
tacle flat, naked; anthers sagittate; style
branches flattened; pappus of copious capil-
lary bristles; achenes oblanceolate in outline,
long-pilose.

Lepidospartum latisquamum Wats. Ne-
vada Broomshrub. Shrubs mainly 6-15 dm
tall or more; branchlets with prominent lon-
gitudinal striae, the striae glandular, the in-
tervening areas tomentose; leaves 0.5-3 cm
long, linear, 0.5-1 mm wide, apiculate; heads
4- to 7-flowered; involucres 8-10 mm high,
3.5-6 mm wide; bracts chartaceous, tomen-
tose, the outer apiculate, very short, the inner
broadly rounded and more or less hyaline
margined; achenes 4-5 mm long, long-pilose
with copious white hairs 3-4 mm long. Rab-
bitbrush community along a wash at 1705 to
1740 m in Millard County (Pine Valley); Ne-
vada and California; 7 (iii).

Leucelene Greene

Perennial rhizomatous herbs; leaves alter-
nate, simple, entire, linear or subulate; heads
radiate, solitary or few to many; involucres
turbinate; bracts imbricate in several series,
green, the margins scarious; ray flowers
white or tinged pink, pistillate; disk flowers
perfect, fertile, yellow; pappus of capillary
bristles; achenes subcylindric or somewhat
compressed.

Shinners, L. H. 1946. Revision of the genus
Leucelene Greene. Wrightia 1:82-89.

Leucelene ericoides (Torr.) Greene Rose-
heath. [Inula ? ericoides Torr.; L. arenosa
Heller; Aster bellus Blake; A. leucelene
Blake; A. hirtif alius Blake]. Perennial herbs
from a branching caudex and rhizome, simple
or more commonly branched, 3-17 cm tall,
strigose and more or less glandular; leaves
2-10 mm long, 1-2 (3) mm wide, linear to
spatulate, becoming subulate upward; heads
solitary or few to many; involucres 5-7 mm
high, 5-12 mm wide; bracts in 3-5 series;
rays 12-25, white to pink, 3-6 mm long;
achenes appressed-hairy. Blackbrush, desert
shrub, salt desert shrub, pinyon-juniper, and
ponderosa pine communities at 1370 to 2595
m in Beaver, Carbon, Daggett, Duchesne,
Emery, Garfield, Grand, Iron, Juab, Kane,
Millard, Piute, Salt Lake, San Juan, Sanpete,
Sevier, Tooele, Uintah, Utah, Washington,
and Wayne counties; Nevada and California,
east to Kansas, south to Arizona and Mexico;
145 (xvii).

Lygodesmia D. Don

Perennial or annual herbs with milky juice;
leaves alternate or mainly basal and still al-
ternate, entire or pinnatifid; heads solitary or
few to many in corymbose or paniculate clus-
ters; flowers all raylike, pink to pink-purple
or white; involucres cylindric; bracts mostly
5-9, with some more or less reduced outer
ones; receptacle naked; pappus of numerous
capillary bristles; achenes linear, subterete,
prominently several nerved.

1. Rays 10-12 mm long, ca 4 mm wide; pappus 6-9 mm long L. juncea

— Rays 15-25 mm long, 6-10 mm wide; pappus 12-17 mm long 2

2(1). Flowers white (or pink?, and drying pinkish); stems ligneous, branching from
the base, forming rounded clumps; leaves stiff, spreading; plants of Emery and
Grand counties L. entrada

— Flowers pink or pink-purple; stems various, but, if branched from the base, the
leaves eitiier lax or the plants of different distribution L. grandiflora

Lygodesmia entrada Welsh & Goodrich

Entrada Rushpink. Perennial herbs from a
subterranean caudex, branching from the
base, the branches ligneus and wiry, mainly
25-45 cm tall; leaves entire, linear or acicu-
lar, 5-30 (70) mm long; peduncles with nu-
merous bracts, 12-20 cm long; involucral

bracts hyaline-margined, the outer 5-10 mm
long, fimbrillate, the inner ca 6, 16-18 mm
long, puberulent at the apex; rays white, ca 3
cm long; pappus barbellate, sordid, 10-15
mm long; achene ribs glabrous. Juniper and
mixed desert shrub communities at 1340 to
1465 m in Emery and Grand (type from near

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Great Basin Naturalist

Vol. 43, No. 2

Courthouse Wash) counties; endemic; 3 (i).
The status of this entity is unclear; certainly
it is a portion of the grandiflora complex.
Further work is indicated.

Lygodesmia grandiflora (Nutt.) T. & G.
Showy Rushpink. [Erythremia grandiflora
Nutt.]. Perennial herbs from deeply placed
elongate rhizomes; stems 0.6-5 dm tall,
simple or branched from the base or above;
leaves alternate, 0.5-10 cm long or more, 1-5
mm wide, attenuate, gradually to abruptly

reduced upward; involucres cylindric, 18-25
mm high, densely hairy to glabrous (?), the
outer mostly short and ovate to lance-ovate,
the inner 5-9 equal, narrowly oblong; rays
5-10, pink, pink-purple, or rarely white,
mostly 2-4 cm long; pappus of numerous
barbellate tawny bristles; achenes 12-18 mm
long, ribbed, glabrous. Our material consists
of three taxa, which have been regarded at
specific rank. Intermediates between the taxa
suggest a more conservative approach.

2(1).

Main involucral bracts 8 or 9; flowers 8-12; plants of east central and north-
eastern Utah L. grandiflora var. grandiflora

Main involucral bracts 5 or 6; flowers 5-7 (10); plants of southeastern and
western Utah 2

Uppermost leaves reduced to linear scales mainly 3-10 mm long; achenes

13-19 mm long, smooth on the lower surface L. grandiflora var. dianthopsis

Uppermost leaves not reduced to scales, mainly 20-40 mm long; achenes
10-13 mm long, rugose on the lower surface L. grandiflora var. arizonica

Var. arizonica (Tomb) Welsh comb. nov.
[based on: Lygodesmia arizonica Tomb Sida
7:530. 1970]. Blackbrush-ephedra and Indian
ricegrass-dropseed communities at 1125 to
1590 m in Kane and Wayne counties; Ari-
zona; 7 (ii).

Var. dianthopsis (D.C. Eaton) Welsh
comb. nov. [based on: Lygodesmia juncea
var. dianthopsis D.C. Eaton in Watson Rep.
U.S. Geol. Explor. 40th Parallel, Bot. 5:200.
1871; L. diantliopsis (D.C. Eaton) Tomb].
Sagebrush-grass, pinyon-juniper, and moun-
tain brush communities at 1370 to 2440 m in
Beaver, Cache, Kane, Millard, Salt Lake, Se-
vier, and Utah counties; Nevada; 22 (iii). In-
termediate specimens transitional to var.
arizonica occur in south central Utah.

Var. grandiflora [L. grandiflora var. stricta
Maguire, type from south of Price]. Shad-
scale, sagebrush, pinyon-juniper, mountain
brush, ponderosa pine, and aspen-sagebrush
communities at 1460 to 2750 m in Carbon,
Daggett, Duchesne, Emery, Garfield, Grand,
and Uintah counties; Wyoming south to New
Mexico; 37 (vii). The var. stricta is a phase
with stiffly erect leaves, but seems to repre-
sent only an ecological variant. Specimens of
intermediate nature occur southward with
both varieties arizonica and dianthopsis.

Lygodesmia juncea (Pursh) D. Don [Pre-
nanthes juncea Pursh]. Perennial glabrous
herbs from a deeply placed elongate root
(rhizome?); stems mainly 1.5-6 dm tall, much

branched; leaves stiff, entire, mainly 1-4 cm
long, 1-4 mm wide, the upper ones reduced
to subulate scales; heads few to several,
mainly 5 (4-10) -flowered; flowers pink or
less commonly white; involucres 9-16 mm
high, with 4-8 main bracts and several short-
er outer ones; pappus tawny; achenes ca 5-7
mm long, several nerved. Our few specimens
from sandy sites in mixed desert shrub and
juniper communities at ca 1400 to 1590 m in
Emery and Juab counties; British Columbia
to Minnesota, south to Arizona and Arkansas;
3 (0). This is mainly a Great Plains species,
with disjunct populations westward, often in
sandy habitats.

Machaeranthera Nees

Annual, biennial, or perennial herbs from
taproots; leaves alternate, entire or pinnatifid
to toothed or lobed, spinulose apically and
the teeth, when present, spinulose; heads soli-
tary or 2 to numerous; involucral bracts in
several series, herbaceous apically, char-
taceous or coriaceous basally, mainly squar-
rose; rays pistillate and fertile, pink, laven-
der, pink-purple, or white, or lacking;
receptacle naked; anthers not caudate; pap-
pus of capillary bristles; achenes narrowly
oblong in outline.

Cronquist, a. and D. D. Keck. 1957. A re-
constitution of the genus Machaeran-
thera. Brittonia 9:231-239.

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

315

1. Heads discoid; leaves spinose-toothed M. grindelioides

— Heads radiate; leaves various, but not conspicuously spinose-toothed 2

2(1). Plants perennial, from a definite caudex, of montane sites, commonly on gran-
ite, limestone, or quartzite Aster kingii D.C. Eaton (q.v.)

— Plants biennial, winter annual, or annual (rarely short-lived perennial), the
caudex not well developed; plants of various habitats and substrates 3

3(2). Leaves pinnately dissected; plants annual M. tanacetifolia

— Leaves merely toothed to entire; plants mainly biennial or short-lived per-
ennial 4

4(3). Involucral bracts with green tip commonly equaling or longer than the char-

taceous base, the long-tapering apices often reflexed M. bigelovii

— Involucral bracts with green tip much shorter than the chartaceous base, the
reflexed to erect tips shortly attenuate to acute M. canescens

Machaeranthera bigelovii (Gray) Greene

[Aster bigelovii Gray; M. mucronata Greene,
sensu Utah materials]. Short-lived perennial
(biennial in some?) herbs from a taproot, a
caudex not or only poorly developed; stems
11-35 cm long, puberulent below, becoming
glandular to stipitate-glandular above; basal
leaves often withered at anthesis; cauline
leaves oblanceolate to linear or oblong,
mainly 1-7.5 cm long, 1.5-8 mm wide, the
surfaces glabrous and more or less glandular
or stipitate-glandular, ciliate, entire to spi-
nose-toothed; heads few to many in corym-
bose inflorescences; involucres 9-12 mm
high, 12-23 mm wide; bracts lance-linear, at-
tenuate apically, the green apex subequal to
the coriaceous base, especially in the outer
bracts, commonly spreading-reflexed, glandu-
lar and glandular-ciliate; rays 21-31, violet or
pink-purple, 10-15 mm long, 2-4.2 mm
wide; pappus off-white; achenes glabrous or
sparingly strigose, 2.5-4.2 mm long. Moun-
tain brush, aspen, spruce-fir, and alpine
meadow communities at 2440 to 3355 m in
Garfield, Iron, Kane, and Washington coun-
ties (Henry Mountains, Markagunt Plateau,
and Kolob Terrace); Colorado, New Mexico,
and Arizona; 18 (iii).

Machaeranthera canescens (Pursh) Gray
[Aster canescens Pursh]. Biennial (winter an-
nual) or short-lived perennial herbs from a

taproot, a caudex seldom developed; stems
8-60 cm tall or more, variously glabrous,
glandular, or puberulent; basal leaves with-
ered or persistent at anthesis; cauline leaves
linear to oblong or oblanceolate, 1-10 cm
long, 1-22 mm wide, the surfaces glabrous,
puberulent, or glandular, commonly ciliate,
entire or toothed; heads few to many in pa-
niculate to corymbose clusters; involucres
6-10 (12) mm high, 6-18 mm wide; bracts
linear to oblong, attenuate to abruptly atten-
uate, the green apex commonly much shorter
than the coriaceous base, sometimes spread-
ing-reflexed, glandular and or puberulent;
rays 15-25, pink to pink-purple or white,
5-12 mm long, 1.5-2.5 mm wide; pappus off-
white; achenes pilose, ca 2.5 mm long. The
canescens complex consists of a series of in-
tergrading taxa, which, in the extreme, are
distinctive and geographically or edaphically
correlated. Many names have been applied to
members of the complex, and specimens of-
ten bear annotations of several of the names
involved. This is partially in recognition of
the intermediate nature of the specimens and
partially due to the quality of diagnostic cri-
teria. It seems best to treat the materials
from Utah as belonging to a single poly-
morphic species, consisting of four inter-

grading varieties.

Leaf surfaces glabrous, the upper leaves commonly glandular to stipitate-
glandular; upper branches usually with numerous bracteate leaves; plants of

southeastern Utah, rarely elsewhere M. canescens var. vacans

Leaf surfaces puberulent, the upper leaves sometimes also glandular; upper
branches lacking bracteate leaves or variously so; plants of broad distribution,
but not of southeastern Utah 2

316

2(1).

Great Basin Naturalist

3(2).

Vol. 43, No. 2

Upper branches with numerous bracteate leaves; rosette leaves abruptly and
angularly lobed or toothed; plants biennial, of central and southwestern Utah ...

M. canescens var. leucanthemifolia

Upper branches seldom especially bracteate; rosette leaves various; plants
biennial or short-lived perennial, of various distribution 3

Involucral bracts 1-1.5 (2) mm broad, abruptly attenuate apically; plants often

perennial, mainly of higher middle elevations M. canescens var. commixta

Involucral bracts 0.5-1 mm wide, rather gradually attenuate apically; plants
often biennial, mainly of lower to middle elevations M. canescens var. canescens

Var. canescens [M. pulverulenta (Nutt.)
Greene]. Salt desert shrub, mixed desert
shrub, pinyon-juniper, mountain brush, as-
pen-sagebrush, Douglas fir, and lodgepole
pine communities at 1250 to 2900 m in Bea-
ver, Carbon, Daggett, Duchesne, Emery,
Garfield, Grand, Iron, Juab, Kane, Millard,
Piute, Salt Lake, Sanpete, Sevier, Summit,
Uintah, and Washington counties; British Co-
lumbia to Saskatchewan, south to California,
Arizona, and Colorado; 102 (xiii). This is a
variable complex of forms that differ in sev-
eral morphological features, but further seg-
regation seems unwarranted. I have been
unable to distinguish M. tephrodes (Gray)
Greene from among our rather large
collection.

Var. commixta (Greene) Welsh comb. nov.
[based on: Machaeranthera commixta Greene
Pittonia 4:71. 1899, type from the Henry
Mountains; M. latifolia A. Nels., type from
Big Cottonwood Canyon, M. leptophylla
Rydb., type from Logan; M. paniculata A.
Nels., type from Parleys Canyon; M. rubri-
caulis Rydb.; Aster rubrotinctus Blake].
Mountain brush, aspen, Douglas fir, sage-
brush, spruce-fir, and alpine meadow com-
munities at 1705 to 3420 m in Beaver, Cache,
Carbon, Duchesne, Emery, Garfield, Iron,
Juab, Millard, Salt Lake, Sanpete, Sevier,
Summit, Tooele, Uintah, Utah, and Wayne
counties; Wyoming and Colorado; 82 (v).

Var. leucanthemifolia (Greene) Welsh
comb. nov. [based on: Aster leucanthemi-
folius Greene Erythaea 3:119. 1895; M. leu-
canthemifolia (Greene) Greene]. Blackbrush,
mixed desert shrub, pinyon-juniper, mountain
brush, and ponderosa pine communities at
915 to 2135 m in Beaver, Iron, Juab, Sanpete,
Sevier, Utah, and Washington counties; Ne-
vada and Arizona; 43 (xiii). This plant is
mainly a xerophyte of sandy and silty habi-

tats at lower elevations in the Great Basin
and lower Virgin River drainage systems; it is
transitional at higher elevations with the pre-
ceding varieties. Phases of var. canescens
from northeastern Utah have been regarded
as portions of this variety, but they seem not
to fit the concept of var. leucanthemifolia,
whose type is from Mineral County, Nevada.

Var. vacans (A. Nels.) Welsh comb. nov.
[based on: Machaeranthera pulverulenta var.
vacans A. Nels. Bot. Gazette 56:70. 1913,
type from San Juan County, Utah]. Salt
desert shrub, mixed desert shrub, pinyon-juni-
per, and ponderosa pine communities at 1155
to 2380 m in Carbon, Emery, Garfield,
Grand, Kane, Juab, San Juan, and Washing-
ton counties; Colorado, Arizona, and New
Mexico; 61 (xix). This material has been
treated as M. linearis Rydb., a glabrous-
leaved phase of M. canescens whose type
came from Yellowstone Park, Wyoming.
Work of a monographic nature is necessary
for the entire canescens complex. Additional
research might indicate an older name at va-
rietal rank for this taxon.

Machaeranthera grindelioides (Nutt.)
Shinners [Eriocarpum grindelioides Nutt.;
Haplopappus nuttallii T. & G.]. Perennial
herbs from a woody caudex and stout tap-
root, the caudex branches more or less
clothed with marcescent leaf bases; stems
2-30 cm tall, pilosulose or spreading-hairy
below, stipitate-glandular and/or hairy
above; basal leaves withered or persistent at
anthesis; cauline leaves oblanceolate to
spatulate or oblong, mainly 0.5-4.5 cm long,
2-12 mm wide, serrate, the teeth with spi-
nulose tips 1-3 mm long, the surfaces pilosu-
lose and/ or stipitate-glandular; heads solitary
or few to many in corymbose clusters; in-
volucres 6.5-9.5 mm high, 8-15 mm wide;

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

317

bracts narrowly oblong, attenuate to an acute
apex, the apical portion green or brown, the
base chartaceous, erect, glandular; rays lack

ing; pappus off-white to brownish; achenes
densely pilose, ca 3 mm long. Two distinctive
varieties are present in Utah.

1. Plants dwarf, often monocephalous; leaves commonly clustered at stem bases;

plants of semibarren habitats in the Great Basin M. grindelioides var. depressa

— Plants seldom dwarf, often with more than 1 head; leaves mainly cauline;

plants seldom of the Great Basin M. grindelioides var. grindelioides

Var. depressa (Maguire) Cronq. & Keck

[Haplopappus nuttallii var. depressa Ma-
guire, type from Millard County]. Mixed
desert shrub, pinyon-juniper, and mountain
brush communities at 1465 to 2320 m in Bea-
ver, Juab, and Millard counties; Nevada, a
Great Basin endemic; 24 (xiv).

Var. grindelioides Blackbrush, mixed
desert shrub, sagebrush, pinyon-juniper, and
mountain brush communities at 1340 to 3175
m in Carbon, Daggett, Duchesne, Emery,
Garfield, Grand, Juab, Kane, Millard, Rich,
San Juan, Sanpete, Sevier, Summit, Uintah,
and Utah counties; Montana to Saskatche-
wan, south to Nevada, Arizona, and New
Mexico; 98 (xiii). There is a tendency for
leaves of plants from the Great Basin to be
more glandular than for those in the main
body of distribution in the Colorado drainage
system.

Machaeranthera tanacetifolia (H.B.K.)
Nees [Aster tanacetifolius H.B.K.]. Annual
(winter annual) herbs; stems 8-50 cm tall,
glandular-puberulent and more or less vil-
lous; leaves 1-6 cm long, 1- or 2-pinnatifid,
the segments ending in spinulose bristles;
heads 1 to many, in corymbose clusters; in-
volucres 8-12 mm high, 10-18 mm wide,
hemispheric; bracts lance-linear, attenuate.

chartaceous basally, green apically, spread-
ing, the reflexed tips glandular; rays 11-23
(36), pink- or blue-purple, 11-14 mm long;
pappus off-white; achenes ca 2.5 mm long,
pilose. Mixed desert shrub, salt desert shrub,
and pinyon-juniper communities at 1125 to
1830 m in Emery, Garfield, Grand, Juab,
Kane, San Juan, Sevier, Utah, Wasatch,
Washington, and Wayne counties; 31 (vi).
The plants are somewhat weedy, colonizing
disturbed sandy and silty soils. The similar M.
parviflora Gray [Aster parvulus Blake] is re-
ported for Utah by various authors. It differs
in having once-pinnatifid leaves, involucres
4-6 mm long, and rays 5-7 mm long. No ma-
terial has been seen from Utah by me.

Madia Molina

Annual or biennial tar-scented herbs from
taproots; leaves opposite below, alternate
above, simple, entire; heads radiate, the rays
pistillate, fertile, yellow, or inconspicuous;
involucral bracts uniseriate, equal, enfolding
the ray achenes; receptacle flat or convex,
with a single series of bracts between the ray
and disk flowers; disk flowers perfect; pappus
none, a short crown, or a few scales; achenes
finely striate, commonly incurved,
compressed.

1. Heads turbinate-ovoid, 6-12 mm wide (when pressed); rays 4-7 mm long,

showy M. gracilis

— Heads ellipsoid, 2-5 mm wide (when pressed); rays to 2.5 mm long, or lacking .

M. glomerata

Madia glomerata Hook. Tarweed. Annual
herbs; stems mainly 8-40 (60) cm tall; herb-
age strigose and with long setiform multi-
cellular hairs on leaf bases and on stems
above, and stipitate-glandular upward, mal-
odorous; leaves 1.2-9 cm long, 1.5-7 mm
wide, linear; heads in dense terminal clusters
or sometimes open; involucres 5.5-9 mm
high, 2-5 mm wide; rays inconspicuous.

mostly 1.5-2.5 mm long, yellow or purplish;
disk flowers 1-10; achenes 5-nerved,
glabrous. Sagebrush, mountain brush, aspen,
spruce-fir, grass-forb, and alpine meadow
communities at 1830 to 3175 m in Cache,
Carbon, Davis, Duchesne, Emery, Iron, Juab,
Piute, Salt Lake, Sanpete, Sevier, Summit,
Tooele, Utah, Wasatch, Washington, and
Weber counties; Alaska to Saskatchewan,

318

Great Basin Naturalist

Vol. 43, No. 2

south to California, Arizona, and Colorado;
38 (iv).

Madia gracilis (Smith) Keck [Sclerocarpus
gracilis Smith in Rees]. Annual herbs; stems
mainly 10-60 (100) cm tall; herbage pilosu-
lose, becoming hirsute with long multi-
cellular hairs upward, stipitate-glandular
with dark capitate glands on the peduncles
and sepals; leaves 1-10 cm long, 2-7 (10) mm
wide, linear to elliptic or oblong; heads sev-
eral to many in an open corymbose cluster
involucres 6-11 mm high, 6-12 mm wide
rays conspicuous 5-13, yellow, 4-7 mm long
disk flowers 15-35; achenes often mottled.
Opening in mountain brush community at ca
1925 m in Salt Lake County; British Colum-
bia to Montana, south to California; 1 (0).

Malacothrix DC.

Annual (winter annual) or perennial herbs
from taproots with milky juice; leaves alter-
nate or mainly basal, mostly pinnatifid; heads
of ray flowers only, long-peduncled, solitary
or few to several and more or less corymbose;
flowers yellow; involucres campanulate;
bracts subequal in 2-4 series, with a few
short outer ones; receptacle flat, setose or
naked; rays 5-lobed; pappus of capillary
bristles, these more or less united at the base
and falling together or with some persistent;
achenes columnar, glabrous, 10- to 15-ribbed,
crowned or denticulate at the summit.
Williams, E. W. 1957. The genus Malacoth-
rix (Compositae). Amer. Midi. Natural-
ist 58:494-512.

1. Leaves merely dentate, elliptic to oblong or lanceolate, the cauline ones clasp-
ing basally; involucral bracts orbicular to ovate and with broad scarious mar-
gins M. coulteri

— Leaves pinnatifid or incised to pinnately lobed, the cauline ones not especially
clasping; involucral bracts linear to narrowly lanceolate 2

2(1). Leaves linear-filiform or pinnately dissected into linear segments M. glahrata

— Leaves with triangular to oblong lobes or teeth, these sometimes attenuate but

not linear 3

3(2). Involucres longer than broad (when pressed); persistent pappus setae 1 or 2;

stems decidedly tapering upward; plants rare, in Washington County . M. clevelandii

— Involucres broader than long (when pressed); persistent pappus setae 1-5 or
lacking; stems various 4

4(3). Leaves with lateral lobes regularly toothed; involucres mainly less than 10 mm

long; achenes 2-2.8 mm long; pappus bristles all deciduous M. sonchoides

— Leaves with lateral lobes irregularly toothed or lobed; involucres more than 10
mm long; achenes 3-4 mm long; pappus often with 1 or few persistent
bristles M. torreyi

Malacothrix clevelandii Gray Cleveland
Malacothrix. Annual herbs; stems mainly
10-40 cm tall, often branched, glabrous,
commonly reddish; leaves basal and cauline,
1-10 cm long, 5-15 mm wide, oblanceolate
to elliptic, pinnately lobed or merely
toothed; heads few to many in a sub-
corymbose cluster; involucres campanulate,
6-7 mm high; main involucral bracts linear,
glabrous, green, the tips often purple, the
margins narrowly scarious; rays yellow, ca
2-3 mm long; pappus deciduous or with 1 or
2 persistent bristles; achenes ca 2 mm long,
slender, striate. Pinyon-juniper and live oak
communities at ca 1375 m in Washington

County; California, Nevada, and Arizona;
2(0).

Malacothrix coulteri Harv. & Gray in
Gray Snakeshead Malacothrix. Annual (win-
ter annual) herbs; stems mainly 10-50 cm
tall, often branched, glabrous and straw col-
ored to whitish tan; leaves basal and cauline
1.2-10 cm long, oblong to oblanceolate or
lanceolate, the cauline ones clasping basally;
heads few to several, corymbose; involucres
hemispherical, 10-15 mm high; bracts imbri-
cate, suborbicular to ovate, with broad
scarious margins, the midline broad, purplish;
rays yellow to off-white, 5-18 mm long; pap-
pus with 1-4 persistent bristles; achenes

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

319

2-2.8 mm long, striate. Warm desert shrub
community at ca 950 m in Washington
County (Galway sn BRY); Arizona and Cali-
fornia; 2 (0).

Malacothrix glabrata (D.C. Eaton) Gray
[M. califomica var. glabrata D.C. Eaton]. An-
nual (winter-armual) or biennial herbs; stems
mainly 10-60 cm tall, often branched from
the base and above, glabrous; leaves basal
and cauline, 0.5-15 cm long, pinnately lobed,
glabrous or more or less villous, with rachis
and lobes linear to linear-filiform, the cauline
ones similar to the basal except reduced up-
ward; head solitary or more commonly few
to many and subcorymbosely arranged; in-
volucres broadly campanulate 10-14 mm
high, the main bracts linear to narrowly ob-
long, with narrow hyaline margins, glabrous,
the outer bracts commonly more or less vil-
lous; rays yellow, 10-20 mm long; pappus
with usually 2 persistent bristles; achenes 2-3
mm long, striate. Joshua tree, blackbrush,
Vanclevea-ephedra, and pinyon-juniper com-
munities at 700 to 1525 m in Kane, Millard,
San Juan, and Washington counties; Oregon
to Idaho, south to California and Arizona; 22
(0).

Malacothrix sonchoides (Nutt.) T. & G.
[Leptoseris sonchoides Nutt.]. Annual or win-
ter aimual herbs; stems mainly 6-37 cm tall,
often branched from the base and above,
glabrous or with short yellowish glandular
hairs in the inflorescence; leaves basal and
cauline, 0.7-12 cm long, 1-28 mm wide, the
basal ones at least pinnatifid and the lobes
regularly toothed; heads solitary or more
commonly few to many and subcorymbosely
arranged; involucres campanulate 7.5-10.2
mm high, 6.5-12 (14) mm wide, the main
bracts lance-oblong to linear, with narrowly
hyaline margins, glabrous, the outer some-
times with yellowish stipitate glands; rays
yellow, 7-12 mm long; pappus bristles all de-
ciduous; achenes 2-2.8 mm long, striate.
Blackbrush, krameria-psorothamnus, mixed
desert shrub, sagebrush, and pinyon-juniper
communities at 915 to 1856 m in Beaver,
Duchesne, Emery, Garfield, Grand, Juab,
Kane, Millard, San Juan, Tooele, Uintah,
Washington, and Wayne counties; California
-and Nevada, east to Nebraska and New Mexi-
co; 72 (vi).

Malacothrix torreyi Gray [M. sonchoides
var. torreyi (Gray) Williams]. Annual or win-
ter annual herbs; stems mainly 8-29 cm tall,
often branched from the base and above,
glabrous, or with yellowish stipitate glands in
the inflorescence; leaves basal and cauline,
1.7-9.5 cm long, 5-27 mm wide, the basal
ones at least pinnatifid, and the lobes irregu-
larly toothed or lobed, often more or less
white villous; heads solitary, or more com-
monly few to several or many and sub-
corymbosely arranged; involucres broadly
campanulate, 10.5-15 mm high, 12-21 mm
wide, the main bracts lance-linear, with hya-
line margins, glabrous or some with stipitate
yellowish glands, the outer bracts often stipi-
tate-glandular; rays yellow, 10-14 mm long;
pappus all deciduous or with 1-5 persistent
setae; achenes 3-4 mm long, striate. Shad-
scale, greasewood, other salt desert shrub,
and mixed desert shrub communities at 1460
to 1925 m in Beaver, Box Elder, Carbon, Du-
chesne, Emery, Garfield, Grand, Juab, Mil-
lard, Piute, Salt Lake, Sevier, Tooele (type
from Great Salt Lake), and Uintah counties;
Oregon to Wyoming, south to California and
Arizona; 28 (i).

Matricaria L.

Biennial or perennial herbs; leaves alter-
nate, 2- to 3- pinnatisect, with ultimate seg-
ments linear- filiform; heads radiate, few to
many in corymbose clusters; involucres
broadly campanulate, the bracts in several
series, the margins scarious; receptacle hemi-
spheric, solid, naked; rays pistillate, white;
disk flowers 5-lobed, perfect, yellow; pappus
a small crown; achenes laterally compressed,
with 3 smooth ribs on the ventral surface and
1 or 2 (rarely more) resin glands at the apex
of the dorsal face. Note: Tentatively I have
chosen to follow authors of Flora Europaea
(Vol. 4) in segregating Chamomilla (q.v.)
from Matricaria. The genera are much alike
and are separated mainly on technical char-
acteristics that are discernible only when
fruit is mature.

Matricaria maritima L. Biennial or, less
commonly, perennial, essentially unscented
herbs; stems 1-6 dm tall, glabrous or nearly
so; leaves 1-8 cm long, the ultimate segments
long and slender; heads several to many, the

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Great Basin Naturalist

Vol. 43, No. 2

disk 8-15 mm wide; rays 10-25, white, 6-13 ternate or all basal, entire or pinnatifid;
mm long. Ruderal weed of moist sites at 1830 heads many flowered, erect or nodding in
to 2135 m in Salt Lake, Sanpete, and Sevier bud; involucres cylindric to campanulate, the
counties; widespread in North America; ad- innermost bracts lance-attenuate, subequal,
ventive from Europe; 2 (i). the outer ones shorter and imbricate; recep-

tacle naked; corollas all raylike, showy, yel-
MiCROSERis D. Don 1«7 t° yellow-orange (fading bluish); pappus

or awn-tipped scales or or plumose capillary
Annual or perennial, scapose or caulescent bristles; achenes columnar to fusiform, not or
taprooted herbs with milky juice; leaves al- only short beaked, ca 10-ribbed.

1. Plants annual; pappus of 5 scales, entended into scabrous bristles apically

M. lindleyi

— Plants perennial; pappus of numerous plumose capillary bristles arising from

short scales M. nutans

Microseris lindleyi (DC.) Gray [Calais
lindleyi DC.; Microseris linearifolia (Nutt.)
Schultz-Bip; Uropappus linearifolius Nutt.].
Armual herbs from slender taproots; herbage
puberulent or glabrate; stems lacking or
more or less developed, the scapose pe-
duncles 10-25 cm high; leaves 6-15 (30) cm
long, pinnately lobed to entire, linear to nar-
rowly elliptic; heads many flowered, erect,
the main bracts lance-attenuate, 15-30 mm
long, subequal, the outer ones shorter and un-
equal; rays yellow (drying blue); pappus
10-20 mm long, silvery, deciduous, of 5
lance-linear scales, each terminating in a
scabrous awn from a bifid apex; achenes dark
brown, 9-13 mm long, tapering apically,
scabrous on the ribs. Blackbrush, creosote
bush, and pinyon-juniper communities at 915
to 1375 m in Washington County; Washing-
ton to Idaho, south to Baja California and
Arizona; 5 (i).

Microseris nutans (Geyer) Schultz-Bip
[Scorzonella nutans Geyer]. Perennial herbs
from tuberous roots; herbage glabrous or
sparsely scurfy; stems more or less developed,
the scapose peduncles mainly 12-40 (60) cm
high, pinnately lobed to entire, linear to el-
liptic, lanceolate, or oblanceolate; heads soli-

tary or 2-5, many flowered, nodding in bud,
the main bracts 10-20 mm long, lance-atten-
uate, subequal, the outer bracts shorter and
unequal; rays yellow (drying lavender or
blue); pappus of numerous narrow scales,
each with a plumose terminal bristle. Sage-
brush, pinyon-juniper, mountain brush,
Douglas fir, and aspen communities at 1675
to 2745 m in Box Elder, Cache, Daggett,
Davis, Juab, Millard, Rich, Salt Lake, San-
pete, Sevier, Summit, Uintah, Utah, and We-
ber counties; British Columbia to Montana,
south to California, Nevada, and Colorado;
33 (ii).

MoNOPTiLON T. & G. ex Gray

Annual herbs, branched from base, the
herbage hispid; leaves alternate, spatulate,
entire; heads radiate, solitary on branch tips,
closely subtended by upper leaves; involucre
campanulate, the bracts subequal, linear, her-
baceous; receptacle flat, naked; ray flowers
pistillate, white to pink; disk flowers perfect,
fertile, yellow (purplish); pappus of a short
scarious cup and 1 apically plumose bristle,
or of numerous bristles alternating with short
scales; achenes compressed, marginally
nerved, pubescent.

1. Pappus of usually several nonplumose bristles alternating with scales; disk

corollas sparsely if at all pilose; reported for Utah by Abrams and Ferris
(Illustrated Flora of the Pacific States), but not seen by me . M. bellioides (Gray) Hall

— Pappus consisting of minute scales and a single apically plumose bristle; disk

corollas densely pilose below M. bellidiforme

Monoptilon bellidiforme T. & G. in Gray

Depressed annual branching herbs; stems 1-5

cm high; leaves 4-10 mm long, 0.5-2.5 mm
wide, narrowly oblanceolate; heads showy;

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

321

involucres 4-5 mm high; bracts linear, atten-
uate or acuminate, hirsute, and minutely
glandular; rays 12-20, ca 4-5 mm long, the
tube pilose; pappus of 1 apically plumose
bristle and several shorter scales, or the pap-
pus rarely lacking; achenes ca 2 mm long.
Warm desert shrub at 700 to 900 m in Wash-
ington County; California, Nevada, and Ari-
zona; 2 (0).

Onopabdum L.

Biennial caulescent spiny herbs from tapr-
oots, the juice watery; leaves basal and caul-
ine, alternate, winged-decurrent; heads soli-
tary or few to several; involucral bracts in
several series, imbricate, spine tipped; recep-
tacle flat, fleshy, honeycombed, often with
short bristles on the partitions, not densely
bristly; corollas all discoid, reddish purple or
pink, perfect; pappus bristles barbellate; ach-
enes glabrous, subquadrangular, 4- or 5-
ribbed.

Onopardum acanthium L. Biennial herbs;
stems mainly 5-15 (30) dm tall; leaves of bas-
al rosettes 5-50 cm long or more and 2-15
cm wide, pinnately lobed and serrate-den-
tate, tomentose on both surfaces, but less so
above, spinose; cauline leaves pinnatifid, to-
mentose to glabrate, strongly winged-decur-
rent along the stem length; involucres 25-35
mm high, 30-65 mm wide, the bracts lance-
attenuate, with spreading spine tips, tomen-
tose to glabrate marginally, the inner erect;
spines 3-5 mm long, yellowish; corollas red-
dish purple to pink. Ruderal weeds at low
elevations in Millard, Tooele, Utah, Wasatch,
and Washington counties; adventive from
Europe; 6 (ii). This handsome but trouble-
some thistle is spreading through the state,
but less vigorously than the musk thistle, Car-
duus nutans (q.v.).

tips; pappus lacking; achenes obovoid, dense-
ly villous, 1 -ridged on each face.

Oxytenia acerosa Nutt. Copperweed. Pe-
rennial herbs; stems erect, mainly 5-12 dm
tall, broomlike in the inflorescence, striate;
leaves 3-15 cm long, pinnately 3- to 7-lobed,
or the upper ones simple; herbage strigulose;
heads 3-4 mm wide, erect or ascending; in-
volucral bracts herbaceous, strigulose; ach-
enes 1.5-2 mm long, black, long villous-pi-
lose. Saline riparian areas and near seeps and
springs at 1220 to 2135 m in Carbon, Emery,
Garfield, Grand, Kane, San Juan, and Wash-
ington counties; Colorado, New Mexico, Ari-
zona, Nevada, and California; 25 (vii). Cop-
perweed is poisonous to livestock.

Palafoxia Lag.

Annual herbs; leaves alternate, entire;
heads discoid, few to several, corymbose or
paniculate; involucres cylindric; bracts in 1
series, herbaceous; receptacle flat, naked;
flowers white, all alike or the outer with im-
equal lobes; pappus scales 4-8, slender, un-
equal, with a strong nerve; achenes linear,
quadrangular.

Palafoxia linearis (Cav.) Lag. Spanish
Needle. [Ageratum lineare Cav.]. Annual
herbs; herbage hispid with slender multi-
cellular hairs, glandular upward; stems com-
monly branched above the base, 1-7 dm tall;
leaves petiolate, the blades 1-7.5 cm long,
2-8 mm wide, linear-lanceolate, long-attenu-
ate; involucres 12-18 mm high, glandular,
and more or less hispid, 10- to 20-flowered,
the corollas white with pink exserted styles;
pappus scales usually 4; achenes strigose.
Warm desert shrub community at 700 to
1000 m in Washington County; California to
Arizona and Mexico; 5 (0).

Oxytenia Nutt.

Perennial riparian herbs from a ligneus
caudex; leaves alternate, pinnately divided or
some entire, the segments linear-filiform, in-
volute; heads discoid, numerous, in elongate
paniculate inflorescences; marginal flowers 5,
pistillate, inner flowers 10-30, staminate;
flowers yellowish white; involucral bracts 5,
orbicular, mucronate; receptacle chaffy, the
chaffy bracts slender, with dilated villous

Parthenium L.

Herbs or shrubs; leaves alternate, entire or
lobed; heads solitary or few and more or less
clustered, inconspicuously radiate; ray flow-
ers 5, white, pistillate, fertile, persistent; disk
flowers staminate; receptacle plano-convex,
chaffy throughout; pappus of 2 or 3 awns or
scales; ray achenes dorsiventrally com-
pressed, rotund in outline, the margins thick-
ened into riblike structures attached to the

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Great Basin Naturalist

Vol. 43, No. 2

contiguous pair of infertile disk flowers and
the subtending bract, the achene, the 2 at-
tached flowers, and the bract falling as a unit.

Rollins, R. C. 1950. The guayule rubber
plant and its relatives. Contr. Gray.
Herb. 179: 1-73.

Plants shrubs, the internodes apparent; heads seldom solitary; known from

Washington County P. incanum

Plants pulvinate-caespitose herbs, the internodes not apparent; heads solitary;
plants of eastern Utah P. ligulatum

Parthenium incanum H.B.K. Aromatic
shrub, 4-10 dm tall, much branched, the
branchlets loosely tomentose, becoming gla-
brate; leaves short-petioled, the blades 0.5-5
cm long, 0.4-1.5 cm wide, lobed, white-to-
mentose below, less so above; heads several
to many, corymbose, 3-5 mm wide, outer in-
volucral bracts oblong, acute, villous, the in-
ner ones suborbicular, membranous; rays
white, emarginate to incised, 1-2 mm long;
pappus of 2 or more pubescent awns; achenes
black, oblanceolate, 1.5-2 mm long, pu-
bescent on the ventral surface. Limestone
cliffs in creosote bush-blackbrush community
at ca 1220 m in Washington County (Higgins
4102 BRY); Arizona to Texas, south to Mexi-
co; 1 (0).

Parthenium ligulatum (Jones) Bameby [P.
alpinum var. ligulatum Jones, type from The-
odore (Duchesne); Bolophyta ligulata (Jones)
W.A. Weber]. Pulvinate caespitose to merely
caespitose acaulescent mound- forming herbs
to ca 3 cm high, from a taproot and branched
caudex, the caudex branches densely clothed
with brownish marcescent leaf bases and of-
ten with ashy leaves of the previous year;
leaves 3-20 mm long, 1.5-4 mm wide, spatu-
late to oblanceolate, strigose; heads solitary
at branch ends, sessile, 5-7 mm high, 4.5-6
mm wide; outer bracts oblong, densely pu-
bescent apically; pappus scales distinct or ad-
nate to the corolla tube; rays white, 1-2 mm
long, emarginate; achenes oblanceolate,
densely hairy, 4-5 mm long, 2-3 mm wide.
Barren or semibarren calciferous or gypsi-
ferous outcrops of the Green River, Uinta,
Ferron, and Carmel formations in salt desert
shrub and pinyon-juniper communities at
1705 to 2135 m in Daggett, Duchesne,
Emery, and Uintah counties; Colorado (a
Colorado Plateau endemic); 42 (iv). This
amazing plant is one of a series of edaphi-
cally restricted mound-formers in semibarren
habits on shales and clays of arid sites in
Utah. It belongs to a closely related assem-

blage of two or three taxa within section Bo-
lophytum, and has been regarded at specific
status within the segregate genus Bolophyta.
Its phylogenetic position was reviewed by
Rollins (1950), and its status within Parthe-
nium, seems to represent best its generic
affinities.

Pectis L.

Annual herbs; leaves opposite, entire,
glandular-dotted; heads radiate, few to sever-
al in cymose clusters; involucres turbinate to
subcylindric; bracts 3-12 in one series, ex-
panded basally, enclosing the ray flowers, of-
ten with translucent glands; receptacle
naked; ray flowers perfect, yellow; disk flow-
ers few; anthers entire, obtuse at base; style
branches short, hispidulous; pappus of short-
plumose bristles on disk flowers, that of ray
flowers a short crown of united scales; ach-
enes terete.

Pectis papposa Harv. & Gray in Gray
Chinch-weed. Annual herbs; stems dichoto-
mously branched, often decumbent-spread-
ing, 5-20 (25) cm long, the herbage yellowish
green; leaves 6-40 (60) mm long, 0.5-2 mm
wide, with a few setae at the base, glabrous,
bearing oval to elliptic large yellowish
glands; heads on peduncles mainly 0.3-1 (2)
cm long; involucres gibbous at the base,
rounded dorsally, sparingly glandular like the
leaves; ray flowers yellow, 7-9, ca 4-6 mm
long; achenes 4-5 mm long, stipitate-glandu-
lar. Sandy soil in warm and sandy desert
shrub communities at 850 to 1650 m in Kane,
San Juan, and Washington counties; Califor-
nia to New Mexico, and south to Mexico; 9
(i)-

Perezia Lag.

Perennial herbs from a caudex, this clothed
with rusty woolly hairs; leaves alternate,
simple, clasping; heads numerous in corym-
bose cymes, apparently discoid; involucres

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

323

campanulate, strongly imbricate; flowers per-
fect, pink to pink-purple, the corollas bila-
biate, the outer lip 3-toothed, the inner lip
recurved, 2-toothed; anthers appendaged;
style branches flattened, truncate apically;
pappus of white capillary bristles; achenes
subterete, minutely glandular.

Perezia wrightii Gray [Acourtia wrightii
(Gray) Reveal & King]. Perennial herbs;
stems 4-6 (10) dm tall, often purplish at the
base, the rusty hairs at stem base copious;
leaves lance-oblong to ovate or lanceolate,
spinulose-dentate, glandular-puberulent on
both sides, the lower ones petiolate, becom-
ing sessile and clasping upward; involucres
5-8 mm high and about as broad, the bracts
graduated, the outer ones ovate, the inner
lance-oblong, obtuse, green, the margins of-
ten purplish, ciliate; corollas pink to pink-
purple; achenes 4.8-5.2 mm long. Warm
desert shrub and juniper communities at 915
to 1525 m in Kane, San Juan, and Washing-

ton counties; Arizona to Texas, south to Mex-
ico; 6 (i).

Perityle Benth.

Annual herbs or perennial subshrubs;
leaves mostly opposite below, alternate
above, simple, petiolate; heads few to numer-
ous, corymbose, radiate or discoid; involucres
hemispheric or turbinate, the bracts some-
what keeled, in 1 or 2 subequal series; recep-
tacle flat, naked; ray flowers (when present)
pistillate, white or yellow; disk flowers per-
fect; anthers subentire to auriculate at the
base; style branches linear or subulate; ach-
enes flattened; pappus of scales, or of 1 or 2
awnlike bristles, or lacking.
Powell, A. M. 1973. Taxonomy of Perityle

section Laphamia (Compositae-Helen-

ieae-Peritylinae). Sida 5: 61-128.
1974. Taxonomy of Perityle section

Perityle (Compositae-Peritylinae). Rho-

dora 76: 229-305.

1. Plants annual P. emoryi

— Plants subshrubs 2

2(1). Heads radiate; plants glandular-hispidulous, of the Great Basin P. stansburyi

— Heads discoid; plants villous or glandular-hispidulous, of the Colorado or Vir-
gin drainages 3

3(2). Herbage short-villous and more or less glandular; pappus bristles 1; plants of

Washington County P. tenella

— Herbage hispidulous; pappus of 3 (4) unequal bristles; plants of Grand County
P. specuicola

Perityle emoryi Torr. in Emory Emory
Rock-daisy. Annual herbs, mainly 2-5 dm
tall; stems erect or spreading, commonly
branched above, puberulent; leaves mostly
alternate, petiolate, the bases 0.5-4 cm long,
0.6-3 (5) cm wide, ovate, cordate, or sub-
orbicular, toothed, lobed, cleft, or divided,
the lobes again toothed or lobed, hirsute to
glandular-pubescent; heads radiate; in-
volucres 5-6 mm high and usually broader;
rays 8-12, white, 1.5-5 mm long; disk flowers
numerous; pappus vestigial or a crown of
scales and 1 slender bristle; achenes 2-3 mm
long, the flattened faces nearly glabrous, the
margin thickened and bearing short stiff
hairs. Sand sagebrush community at lower
elevations in Washington County (Tanner sn
1941 BRY); Nevada, California, Arizona, and
Mexico; 1 (0).

Perityle specuicola Welsh & Neese Alcove
Rock-daisy. Perennial suffruticose herbs,
mainly 50-75 cm tall; stems sprawling or
pendulous, much branched; herbage glandu-
lar-hispidulose; leaves mostly alternate, short-
petiolate, the blades 3-6 mm long, 1.5-3 mm
wide, ovate-elliptic, entire, hispidulous; heads
few to many in a branching corymbose in-
florescence; involucres 3.5-5 mm high, 5-6
mm wide; bracts 11-16, oblong to elliptic,
keeled; ray flowers lacking; disk flowers nu-
merous, ca 2.5 mm long, whitish (?); pappus
of 3 unequal scabrous bristles and often with
1 apically flattened and sigmoid scale; ach-
enes 3-3.8 mm long, the faces flattened,
glabrous, the margin thickened and with
short ascending hairs. Hanging garden com-
munities at ca 1220 m in Grand County; en-
demic; 2 (ii).

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Great Basin Naturalist

Vol. 43, No. 2

Perityle stansburyi (Gray) Macbride
Stansbury Rock-daisy. [Laphamia stansburyi
Gray, type from Stansbury Island]. Suffruti-
cose perennials, clump-forming, 7-30 cm tall
and as broad or more; herbage glandular-hir-
tellous; leaves mainly alternate, the blades
3-14 mm long, 1.5-12 mm wide, broadly
ovate to deltoid or orbicular, typically few to
several lobed; petioles 1-14 mm long; heads
few to many in a branching corymbose in-
florescence; involucres 5-6.5 mm high, 5-10
mm wide; bracts 16-21, lanceolate to oblan-
ceolate, strongly keeled; ray flowers 10-14,
yellow, 3-5.5 mm long; disk flowers numer-
ous, yellow, 4-5 mm long; pappus of 1 stout
bristle and a very short crown of hyaline
scales; achenes 2.3-3.5 mm long, with thin
callous margins, short-pubescent on margins
and on faces. Limestone, dolomite, and ig-
neous ignimbrite (ashflow tuff) outcrops, in
mixed desert shrub, pinyon-juniper, and
mountain brush communities, at 1280 to
1895 m in Beaver, Juab, Millard, Salt Lake,
Sanpete, Sevier, and Juab counties; Nevada (a
Great Basin endemic); 39 (v).

Perityle tenella (Jones) Macbride Jones
Rock-daisy. [Laphamia palmeri Gray, type
from Beaverdam, Arizona?, not P. palmeri
Wats.; L. palmeri var. tenella Jones, type
from Springdale]. Suffruticose perennials,
clump-forming, 9-25 cm tall and as broad or
more; herbage villous and glandular; leaves
mainly alternate, the blades 4-13 mm long,
3-15 mm wide, deltoid-ovate, the base obtuse
to truncate or cordate; petioles 1-8 mm long;
heads solitary or few to many, corymbose; in-
volucres 4-6.5 mm long, 5-10 mm wide;
bracts 11-18, lance-elliptic, keeled; ray flow-
ers absent; disk flowers numerous, yellow,
3-4 mm long; pappus of a single bristle; ach-
enes 2.5-3 mm long, with thin callous mar-
gins, short-pubescent on margins and on
faces. Joshua tree, creosote bush, blackbrush,
warm desert shrub, pinyon-juniper, and pon-
derosa pine communities at 915 to 2135 m in
Washington County; Arizona; a Mohave en-
demic; 7 (0). Plants from the Beaverdam
Moimtains have heads that average larger,
but they seem not to differ otherwise from
the typical materials taken near Zion Nation-
al Park.

Petradoria Greene

Suffrutescent perennials from a taproot
and woody caudex; stems herbaceous, leafy;
leaves basal and cauline, alternate, entire, 3-
to 5-veined, coriaceous; heads radiate (in
ours), congested at branch ends in an open
corymbose inflorescence; involucres cylin-
dric; bracts in several series, in more or less
vertical ranks; flowers 4-7, yellow, the co-
rollas glabrous; pappus of brownish capillary
bristles; achenes somewhat compressed,
glabrous.

Anderson, L. C. 1964. Studies on Petradoria
(Compositae); anatomy, cytology, tax-
onomy. Trans. Kansas Acad. Sci. 66:
632-684.
Petradoria pumila (Nutt.) Greene Rock
Goldenrod. [Chrysoma pumila Nutt.]. Plants
from a well-developed caudex, the caudex
branches clothed with dark to ashy or tan
marcescent leaf bases; leaves 1.5-12 cm long,
1-11 mm wide, oblanceolate to lanceolate,
elliptic, or linear; cauline leaves reduced up-
ward; heads numerous; involucres 5-9.5 mm
high, 1.3-2.8 mm wide; involucral bracts
10-21, in 3-6 series, more or less keeled;
flowers 2-8, the rays 1-3, yellow, 4-9 mm
long; achenes 4-5 mm long, glabrous, 6- to 9-
nerved. Shadscale, mixed desert shrub, pin-
yon-juniper, sagebrush, and ponderosa pine
communities at 1525 to 3050 m in all (?)
Utah counties; Idaho and Wyoming, south to
Nevada, California, Arizona, and New Mexi-
co; 100 (xv). Most of our specimens belong to
the broad-leaved var. pumila, but one speci-
men from Emery County (Harris 546 BRY)
seems to be clearly allied to var. graminea
(Woot. & Standi.) Welsh comb nov. [based
on: Petradoria graminea Woot. & Standi.
Contr. U.S. Natl. Herb. 16: 183. 1913; ssp.
graminea (Woot. & Standi.) L.C. Anderson].
That taxon has been known previously only
from Arizona.

Platyschkuhria (Gray) Rydb.

Perennial herbs from a woody caudex and
rootstock; leaves alternate, simple, co-
riaceous, often impressed-punctate; heads
few to many in a cymose paniculate cluster,
radiate, campanulate to hemispheric; in-
volucral bracts subequal in 2 series; recep-
tacle essentially flat, naked; rays pistillate.

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

325

fertile, yellow; disk flowers numerous, per-
fect; anthers more or less sagittate basally;
pappus of 8-16 scales with midribs some-
times produced apically; achenes narrowly
obpyramidal and 4-sided, hairy or glabrous
on the sides.
Ellison, W. L. 1971. Taxonomy of Platysch-

kuhria (Compositae). Brittonia 23:

269-279.
Platyschkuhria integrifolia (Gray) Rydb.
[Schkuhria integrifolia Gray; Bahia nudi-
caulis Gray; B. integrifolia (Gray) Macbride].

Perennial herbs; stems solitary or few to sev-
eral, mainly 12-55 cm tall; herbage white-
strigulose or stipitate-glandular, especially
above; main leaves near the stem base, pe-
tiolate, the blades 1.5-9.5 cm long, 0.5-4 cm
wide, ovate to lanceolate, elliptic, or oblan-
ceolate; cauline leaves reduced upward, fi-
nally merely bracteate; heads (1) 2-10; rays
7-11, yellow, 6-14 mm long; achenes 5-8
mm long. Three rather distinctive varieties
are present in eastern Utah, as indicated
below.

1. Stems leafy almost or quite to the apex; plants of San Juan County

P. integrifolia var. oblongifolia

— Stems leafy mainly below the middle; plants not known from San Juan County 2

2(1). Pubescence of upper stems merely white-strigulose; involucral bracts caudate-
attenuate P. integrifolia var. ourolepis

— Pubescence of upper stems stipitate-glandular; involucral bracts mainly obtuse

to acute P. integrifolia var. desertorum

Var. desertorum (Jones) Ellison [Bahia
desertorum Jones, type from Cisco]. Salt
desert shrub, pinyon-juniper, and mountain
brush communities, mainly in saline sub-
strates, at 1280 to 2565 m in Carbon, Du-
chesne, Emery, Garfield, Grand, Sevier, Uin-
tah, and Wayne counties; Colorado; a
Colorado Plateau endemic. A report by Elli-
son (1971) of var. integrifolia (a Wyoming-
Montana endemic) belongs here; 39 (xi). The
var. desertorum is closely allied with var. in-
tegrifolia, as indicated by pubescence and
bract shape similarities. This variety is transi-
tional with var. ourolepis.

Var. oblongifolia (Gray) Ellison
[Schkuhria integrifolia var. oblongifolia Gray;
Bahia oblongifolia (Gray) Gray; Platysch-
kuhria oblongifolia (Gray) Rydb.]. Desert
shrub communities in San Juan County; Ari-
zona, Colorado, New Mexico; 0 (0). The vari-
ety is reported from San Juan County, but no
specimens have been seen by me.

Var. ourolepis (Blake) Ellison [Bahia ouro-
lepis Blake, type from Green River]. Salt
desert shrub and pinyon-juniper communities
at 1280 to 1830 m in Duchesne, Emery,
Grand, and Uintah counties; endemic; 24 (iii).
The main body of this variety lies in Uintah
County.

Pluchea Cass.

Shrubs; leaves alternate, simple, entire, se-
riceus; heads discoid, few to several, aggre-
gated in terminal cymose clusters; involucres
campanulate; bracts imbricate in several
series, scarious, the outer ones sericeus; re-
ceptacle flat or concave, naked; outer flowers
pistillate, numerous, their filiform corollas 3-
or 4-toothed; central flowers perfect but the
innermost sterile, their tubular corollas 5-
toothed; anthers sagittate basally; pappus of
outer flowers merely capillary bristles, those
of inner flowers clavate apically.

Pluchea sericea (Nutt.) Gov. Arrowweed.
[Polypappus sericeus Nutt.; Tessaria sericea
(Nutt.) Shinners]. Shrubs with slender, erect,
willowlike branches, mainly 0.8-3 m tall, se-
riceus throughout, longitudinally striate;
leaves 0.8-4.5 cm long, 2-9 mm wide, ellip-
tic to narrowly lanceolate or lanceolate, en-
tire, sessile; heads more or less conspicuous;
involucres 3.5-5 mm high, 4-7 mm wide;
outer bracts ovate to ovate-lanceolate,
abruptly acute, deciduous, often purplish;
pistillate flowers purplish, numerous; perfect
flowers purplish, fewer; achenes glabrous;
pappus bristles of perfect flowers dilated api-
cally. Riparian areas at 460 to 1220 m in
Garfield, Kane, San Juan (?), and Washington

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Great Basin Naturalist

Vol. 43, No. 2

counties; California to Texas, south to Mexi-
co; 22 (iii). The genus Pluchea, in a broad
sense, includes annual and perennial herbs
and shrubs. Tessaria, when segregated from
Pluchea, consists of the shrubby species that
have dimorphic corollas and the inner per-
fect flowers with apically flared pappus
bristles. The residue within Pluchea contains
only herbs with uniformly 4-lobed corollas
and pappus of uniform barbellate capillary
bristles. I follow tradition in maintaining our
species in Pluchea.

PoROPHYLLUM (VaiU.) Adans.

Suffruticose perennial; leaves alternate or
opposite, simple, with at least some elliptic
to oval oil glands; heads discoid, solitary, or
few to several in corymbose clusters; in-
volucres cylindric, the bracts usually 5, in
subequal series, glandular like the leaves; re-
ceptacle naked; flowers perfect, fertile,
purplish; anthers rounded basally; style
branches slender, hirtellous, subulate; pappus
of scabrous bristles; achenes slender, striate.

Porophyllum gracile Benth. Odora.
Rounded bushy perennials from a woody
base; stems much branched, 1.5-4 dm tall;
herbage dark green or often purplish,
glaucous, odoriferous; leaves 1-4 cm long,
linear-filiform, entire; involucre subcylindric,
10-15 mm long; bracts 5, dark green, tinged
purplish, oblong, the hyaline margin pink,
gibbous basally, bearing conspicuous glands,
especially apically; corollas purplish, white;
pappus bristles pinkish; achenes 8-9 mm
long, hispidulous. Desert shrub communities
in Washington County (Cottam 5522 UT);
California to Arizona and Mexico; 1 (0).

Prenanthella Rydb.

Annual herbs; leaves basal and alternate,
simple, pinnately lobed, toothed, or pinnati-

fid; heads small, few to numerous; involucres
campanulate; bracts in 2 series, the inner sub-
equal, 3 or 5, the outer much reduced, 1 or 2,
herbaceous; flowers all raylike, 4 or 5; ach-
enes 5-ribbed; pappus of white capillary
bristles.

Prenanthella exigua (Gray) Rydb. [Pre-
nanthes exigua Gray; Lygodesmia exigua
(Gray) Gray]. Annual; stems branched from
the base, forming rounded clumps, 7-24 (30)
cm tall; inflorescence paniculate, comprising
more than half the plant height; lower leaves
1-4 (6.5) cm long, 3-12 (20) mm wide, spatu-
late to oblanceolate, the rosette often with-
ered at anthesis; cauline leaves reduced up-
ward, finally bracteate scales; herbage
sparingly stipitate-glandular; involucres 3-5.5
mm long, 1.2-3.5 mm wide; inner bracts ob-
long, herbaceous, apically constricted in bud;
rays pink or white, 1.5-2 mm long; achenes
3-3.5 mm long, 5-ridged, scabrous; pappus of
white capillary bristles. Blackbrush, creosote
bush, other warm desert shrub, salt desert
shrub, and pinyon-juniper communities at
850 to 1925 m in Beaver, Carbon, Emery,
Garfield, Grand, Juab, Kane, Millard, San
Juan, Tooele, Uintah, and Washington coun-
ties; California, Nevada, Colorado, Arizona,
and New Mexico; 20 (ii).

Psathyrotes Gray

Annual or perennial (?) herbs; leaves alter-
nate, petiolate, simple, entire or lobed to
toothed; heads discoid, the flowers yellow or
purplish in age; involucres campanulate;
bracts biseriate, the outer often shorter or
otherwise different; receptacle flat, naked;
anthers minutely sagittate; style branches
flattened; achenes hairy; pappus of capillary
bristles.

2(1).

Plants lanate-tomentose as well as scurfy; outer involucral bracts expanded api-
cally, oblong-obovate; reported for Utah by Munz (A California Flora), but not

seen by me P. ramosissima (Torr.) Gray

Plants scurfy and less commonly somewhat tomentose; outer involucral bracts
tapering apically, lanceolate 2

Leaves entire; herbage scurfy and with long-piliferous multicellular hairs

P. pilifera

Leaves toothed; herbage scurfy but not long-piliferous P. annua

Psathyrotes annua (Nutt.) Gray Mealy Ro-
settes [Bulbostylis annua Nutt.]. Annual or

winter annual herbs forming low rounded
cushions, mainly 2-18 cm tall; leaves petio-

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

327

late, the blades 5-17 mm long, 5-20 mm
wide, orbicular to fan shaped, dentate; her-
bage scurfy; heads few to numerous, corym-
bose; involucres 5.5-7.5 mm high, 5-8 mm
wide; outer bracts lanceolate to oblong, more
or less constricted above the middle, scurfy
and ciliate; disk corollas 3.5-4.2 mm long,
yellow, becoming purplish; achenes 2-2.5
mm long, pilose. Warm desert shrub, salt
desert shrub, and pinyon-juniper commu-
nities, commonly on limestone and dolomitic
gravels, at 790 to 1740 m in Juab, Millard,
Tooele, and Washington counties; Idaho
south to California, Nevada, and Arizona; 17

Psathyrotes pilifera Gray Annual or win-
ter annual herbs forming hemispheric cush-
ions, mainly 5-15 cm tall; leaves petiolate,
the blades 5-15 mm long, 4-16 mm wide,
obovate, ovate, or oval-elliptic, entire; herb-
age scurfy and piliferous with long multi-
cellular hairs; heads few to many, corymbose;
involucres 8.5-10 mm high, 4-5.5 mm wide;
outer bracts lanceolate, seldom constricted
above the middle, scurfy and with long pili-
ferous setae marginally; disk corollas 6-6.5
mm long, yellow, becoming purplish; achenes
3.8-4.8 mm long. Warm desert shrub and salt
desert shrub, commonly on gypsiferous sub-
strates of the Moenkopi and Chinle forma-
tions, at 760 to 2260 m in Grand, Kane, and
Washington counties; Arizona; endemic; 9

PSILOCARPUS Nutt.

Low floccose-wooUy armual herbs; leaves
opposite, simple, entire; heads discoid, sub-

globose; involucre per se essentially lacking;
receptacle chaffy, subglobose; pistillate flow-
ers numerous, imbricate, each enclosed by
and deciduous with its subtending bract,
woolly, with sides meeting in the center,
bearing below the rounded tip on inner side a
scarious appendage; corollas filiform; pappus
lacking; perfect flowers few, central, ebrac-
teate, the corollas 4- or 5-toothed, epappose;
anthers sagittate.

Psilocarphus brevissimus Nutt. Low
white-woolly annuals; stems simple or with
decumbent-prostrate branches mainly 1.5-20
cm long; leaves 5-15 mm long, 1-3 mm
wide, spatulate to lanceolate, apiculate;
heads solitary or clustered, long-woolly, ca
5-7 mm thick, subtending leaves about as
long as the head or longer; pistillate flowers
20-34 or more, the enclosing bracts 2.5-3.2
mm long, woolly, the appendage horizontally
produced to erect, ca 0.5 mm long; perfect
flowers ca 6-10; achenes subcylindric, terete,
1.3-2 mm long. Lake and reservoir beds at ca
1710 m in Cache and Salt Lake counties;
Washington to Montana, south to California,
Mexico, and South America; 6 (0).

PSILOSTROPHE DC.

Perennial herbs or shrubs; leaves alternate,
simple, entire or merely lobed; heads few to
many, corymbose; involucres campanulate;
bracts in 1 series, subequal; receptacle naked;
ray flowers yellow, pistillate, becoming pa-
pery and persistent; disk flowers perfect, 5-
lobed; anthers obtuse basally; style branches
truncate; pappus of 4-6 hyaline scales;
achenes obtusely angled, glabrous or hairy.

1. Plants shrubby; stems closely white-tomentose; of Washington County

P. cooperi

— Plants herbaceous, from a definite caudex; stems glabrous or loosely tomentose;

not of Washington County 2

2(1). Stems loosely tomentose; involucres densely white villous-tomentose; plants of

Grand County P. bakeri

— Stems glabrous, or tomentose only at the base; involucres sparingly tomentose,
greenish; plants of Wayne, Garfield, and Kane counties P. sparsiflora

Psilostrophe bakeri Greene Perennial
herbs from a caudex; stems 10-35 cm tall,
densely white-woolly below, loosely tomen-
tose upward; leaves 0.8-10 cm long, 2-15

mm wide, spatulate to oblanceolate, entire or
lobed, pubescent like the stems; cauline
leaves reduced upward; involucres loosely
villous-tomentose, 5-9 mm high, ca 3-4 mm

328

Great Basin Naturalist

Vol. 43, No. 2

wide; rays 3-6, yellow, 6-12 (15) mm long;
pappus scales ca 2 mm long, rounded;
achenes whitish, ca 2-2.5 mm long. Sandy
warm desert shrub community at ca 1285 m
in Grand County (Trotter 101 BRY); Colo-
rado (a Plateau endemic?); 1 (0).

Psilostrophe cooperi (Gray) Greene Paper-
flower. [Riddellia cooperi Gray]. Shrubs;
stems closely white-tomentose, mainly 30-60
cm tall; leaves 1.2-7 cm long, linear, entire,
sparingly tomentose, finally glabrate; in-
volucres tomentose, 6-8 mm high, 5-8 mm
wide; rays 4-8, yellow, 8-20 mm long; pap-
pus scales ca 2 mm long, acute; achenes whit-
ish, 4.5-7 mm long. Joshua tree, creosote
bush, blackbrush, and pinyon-juniper com-
munities at 915 to 2135 m in Washington
County; Nevada, California, Arizona, and
New Mexico; 11 (ii).

Psilostrophe sparsiflora (Gray) A. Nels.
[Riddellia tagetina var. sparsiflora Gray].
Perennial herbs from a caudex; stems 14-60
cm tall, densely to moderately pilose basally.

sparingly villous-tomentose upward; leaves
0.9-11.5 (14.5) cm long, 1-11 mm wide,
spatulate to oblanceolate or linear, pubescent
like the stems or glabrate; involucres 4.5-6
mm high, 4-6 mm wide; rays usually 3, yel-
low, 6-12 mm long; pappus scales 1.5-2.5
mm long, acutish; achenes yellowish, 2.5-3
mm long. Salt desert shrub, pinyon-juniper,
and sagebrush communities at 1430 to 2045
m in Garfield, Kane, and Wayne counties;
Arizona, New Mexico, and Mexico; 42 (vi).

Rafinesquia Nutt.

Annual herbs; stems fistulous; leaves alter-
nate, pinnatifid; heads 2 to several, large,
showy, with white or rose-tinged flowers; in-
volucres essentially cylindric; bracts 7-15, in
2 series, the inner subequal, the outer ones
much shorter, obtuse or subcordate basally;
flowers all raylike; pappus white or tawny, of
8-15 slender long-plumose bristles; achenes
terete, obscurely few ribbed, attenuate into a
beak.

Rays mainly 5-8 mm long; achene beak as long as the body; plumose hairs of
pappus straight; plants reported for Utah by Munz (A California Flora), but

not seen by me R. californica Nutt.

Rays mainly 12-18 mm long; achene beak shorter than the body; plumose
hairs of pappus crinkled; plants of Washington County R. neomexicana

Rafinesquia neomexicana Gray Desert
Chicory. Annual (winter annual) herbs; stems
mainly 15-40 (50) cm tall, simple or
branched, often growing up through shrubs;
basal leaves 1.2-9 cm long, pinnatifid, often
withered at anthesis; cauline leaves sessile
and auriculate-clasping, reduced upward; in-
volucre 15-25 mm high, 5-9 mm wide; main
bracts lance-attenuate, the margins scarious,
the outer ones more or less cordate basally;
rays 12-18 mm long, white or suffused with
pink, 5-toothed or -lobed apically; pappus
bristles white, the bases flattened, plumose to
near the apex; achenes 12-15 mm long, pa-
pillate-puberulent. Joshua tree, creosote
bush, blackbrush, and desert almond commu-
nities at 700 to 1070 m in Washington Coun-
ty; California to Texas and Mexico; 6 (i).

Ratibida Raf.

Perennial herbs from a caudex and stout
taproot; leaves alternate, pinnatifid; heads

radiate, solitary or few and corymbose; rays
neuter, commonly yellow (sometimes purple
in part or throughout); involucre in 1 series,
green; receptacle columnar, chaffy through-
out, the bracts more or less clasping the
achenes; disk flowers perfect, fertile; anthers
sagittate; style branches flattened; achenes
compressed at right angles to the involucral
bracts, glabrous, the margins sometimes cil-
iate; pappus of an evident tooth and some-
times with a second one.

Ratibida columnifera (Nutt.) Woot. &
Standi. Prairie Coneflower. [Rudbeckia col-
umnifera Nutt.; R. columnaris Pursh]. Pe-
rennial herbs; stems mainly 3-6 (12) dm tall,
several, often branched above, strigose;
leaves 2-9 cm long, pinnatifid, with the ter-
minal division often the largest; heads borne
on slender peduncles 6-18 cm long, the disk
grayish in bud, purplish brown in flower,
1.5-3 cm long; rays 3-7, yellow (or purple),
1-3 (4.5) cm long, spreading or reflexed; pap-
pus an evident awn tooth on the inner angle

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

329

of the achene, often also a shorter one on the
outer edge; achenes ciliate and more or less
winged on the inner edge. Salt desert shrub
and sagebrush communities at 1585 to 2565
m in Garfield, Millard, and Washington
counties; British Columbia to Minnesota,
south to Arizona, Texas, and Mexico; 4 (i).
Our material appears to be adventive from
the main body of the species in the prairies
and plains provinces to the east of Utah.

RiGiOPAPPUs Gray

Annual herbs; leaves alternate, linear, en-
tire; heads radiate, solitary or few to several,
cymose; rays pistillate, fertile, yellow, incon-
spicuous; involucres campanulate; bracts ob-
long-alternate, subequal, partly clasping out-
er achenes; receptacle flat, with a row of
bristles between ray and disk flowers; disk
flowers perfect, fertile, yellow; anthers not
toothed; style branches flattened; pappus
usually of 3-5 awnlike scales (or lacking);
achenes linear, transversely rugulose.

Rigiopappus leptocladus Gray Wireweed.
Slender annual herbs; herbage puberulent to

glabrate; stems 2-20 (30) cm tall; lateral
branches, when present, very slender, over-
topping the early flowers; leaves 0.3-2 (3) cm
long, linear, the lower often withered at an-
thesis; heads small; involucres 4-7 mm high
and about as broad; bracts herbaceous,
glabrous, thickened dorsally; rays few, yel-
lowish, 1.5-2 mm long; pappus scales linear-
subulate, ca 3 mm long; achenes 5-6 mm
long. Reported for Utah by Cronquist (Flora
of the Pacific Northwest), but not seen by
me; 0 (0).

RUDBECKIA L.

Perennial caulescent herbs; leaves alter-
nate, serrate or pinnately to palmately lobed;
heads radiate or discoid, the rays (when pres-
ent) neuter, commonly yellow; involucral
bracts in 2 or 3 series, mainly unequal, her-
baceous, spreading or reflexed; receptacle
conic or columnar, chaffy throughout, the
bracts clasping the achenes; disk flowers fer-
tile; anthers obtuse or sagittate basally; style
branches flattened; pappus a crown or none;
achenes quadrangular or flattened at right
angles to the involucral bracts.

1. Heads radiate; disks 1-2 cm wide and about as long, little elongate in fruit;

plants of San Juan County H. laciniata

— Heads discoid; disks 1.5-2.5 cm wide, mostly 2-5 cm long, elongating in fruit;
plants not of San Juan County 2

2(1). Leaves laciniately lobed; plants glabrous or merely scabrous-ciliate on leaf

margins; known from Iron and Washington counties R. montana

— Leaves crenate-serrate, dentate, undulate, or entire, not lobed; plants evidently
short-hairy to almost glabrous; known from mountains of central northern to
south central Utah R. occidentalis

Rudbeckia laciniata L. Cutleaf Cone-
flower. Perennial herbs; stems erect from a
coarse ligneus base, mainly 5-10 (20) dm tall,
glabrous or scabrous-ciliate; leaves petiolate,
the blades laciniate-pirmatifid to palmatifid,
mainly 4-15 cm long and as broad; heads
showy, the disk 1-2 cm wide and about as
high; rays yellow, 6-16, deflexed-spreading,
2-5 cm long; pappus a short crown. Moist
meadows at 1890 to 2200 m in San Juan
County; Montana to Quebec, south to Ari-
zona and Florida; 2 (0).

Rudbeckia montana Gray? Perennial
herbs; stems erect, from a short sub-
rhizomatous caudex, 6-12 dm tall, glabrous;

leaves petiolate, the blades laciniate-pinnati-
fid, mainly 4-20 cm long and about as broad;
heads discoid, the disk 1.5-2.5 cm wide, 3-5
cm high; rays lacking; pappus an irregularly
margined, almost toothed crown. Iron and
Washington counties; Colorado; 2 (0).

Rudbeckia occidentalis Nutt. Western
Coneflower. Perennial herbs; stems erect
from a coarse ligneus rhizome, mainly 5-20
dm tall, glabrous or strigulose; leaves petiol-
ate, the blades 5-20 cm long, 2.5-10 cm
wide, ovate to ovate-lanceolate or lanceolate,
attenuate to acuminate, entire, crenate-ser-
rate, or dentate; heads discoid, the disks
1.5-2.5 cm wide, 3-6 cm long; rays lacking;

330 Great Basin Naturalist Vol. 43, No. 2

pappus a short crown. Mountain brush, as- margins scarious or hyaUne, or variously col-
pen, grass-tall forb, and spruce-fir commu- ored; receptacle flat or convex, naked; ray
nities at 2135 to 3175 m in Cache, Carbon, flowers yellow or orange, or sometimes lack-
Duchesne, Emery, Piute, Salt Lake, Sanpete, ing; pappus or capillary bristles; style
Sevier, Summit, Tooele, Utah, Wasatch, and branches flattened; achenes subterete, 5- to
Weber counties; Washington to Montana, 10-nerved, glabrous or pubescent. Note: This
south to California and Wyoming; 42 (iii). genus consists of a series of species that inter-
grade freely when they are in contact with
bENECio L. others of the group. Because of hybridization
Annual, biennial, or perennial herbs with the species lines tend to be blurred, and it is
rhizomes, caudices, or taproots, the juice wa- not possible to place all specimens with con-
tery; stems erect, ascending, or decumbent at fidence. Keys are, and have been, based on
the base; leaves alternate, simple, entire, features that are subject to interpretation; the
toothed, or lobed to pinnatifid; heads soli- present one is not an exception, being tenta-
tary, or few to many in corymbose cymes; in- tive at best.

volucral bracts in 1 series, often with smaller Barkley, T. M. 1978. Senecio. N. Amer.

bractlets at the base, green throughout or the Flora II. 10:50-139.

1. Leaves pinnatilobate with linear-filiform divisions or entire and linear-filiform;
stems with leaves only gradually reduced upward, often more or less woody
below 2

— Leaves variously lobed, toothed, or entire, but the segments and leaves not
linear- filiform; stems with leaves various, seldom, if at all, woody at the base 3

2(1). Heads cylindric, subcylindric, or narrowly campanulate; main involucral bracts
8-13, the outer ones much reduced and inconspicuous; plants glabrous ..S. spartioides

— Heads campanulate to broadly campanulate; main involucral bracts 13-21, the
outer ones conspicuous, or, if inconspicuous, the plants tomentose S. douglasii

3(2). Heads nodding, especially in bud, or, if erect, with both distinctly black

triangular tips on involucral bracts and cauline leaves prominently clasping 4

— Heads erect, even in bud; plants various but not as above 7

4(3). Heads discoid 5

— Heads radiate 6

5(4). Heads 8-12 mm high, 6-9 mm wide, narrowly campanulate, conspicuously

pedunculate S. pudicus

— Heads 12-20 mm high, 14-20 mm wide, broadly campanulate, short-
pedunculate S. bigelovii

6(4). Heads erect, the involucral bracts black-tipped S. crassulus

— Heads nodding, the involucral bracts often suffused with purple throughout,

but not especially black tipped S. amplectens

7(3). Plants annual or winter annual, introduced weedy species S. vulgaris

-^ Plants perennial, indigenous species 8

8(7). Stems uniformly leafy to the inflorescence, or the leaves concentrated upward 9

— Stems few leaved, or the upper leaves definitely reduced in size and distribu-
tion 12

9(8). Stems 1-3 dm tall, more or less sprawling, arising from a subrhizomatous cau-

dex S. fremontii

— Stems mostly 2-15 dm tall, erect or ascending, arising from a rhizome or a
caudex 10

April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 331

10(9). Plants mainly 2-4 dm tall; leaves pinnatifid to lobed or laciniate; involucral

bracts with dark tips S. eremophilus

— Plants mainly 5-10 dm tall; leaves dentate to serrate; involucral bracts uni-
formly greenish or brownish 11

11(10). Leaf blades acute to obtuse basally, the teeth all about alike S. serra

— Leaf blades truncate to obtuse basally, or more or less hastately lobed, the
lowermost teeth often the largest S. triangularis

12(8). Plants glaucous tall herbs, semiaquatic; leaves entire or denticulate, thick and

leathery S. hydrophilus

— Plants not or seldom glaucous, terestrial; leaves entire, toothed, or pinnatifid,

not thick and leathery 13

13(12). Rays orange or orange-red (see also S. pauperculus) S. crocatus

— Rays yellow or lacking 14

14(13). Heads discoid; plants tomentose, soboliferous S. fendleri

— Heads radiate (or rarely discoid in some individuals); plants tomentose, gla-
brate, or glabrous, not soboliferous (except in S. werneriifolius) 15

15(14). Leaves pinnatifid, at least the cauline ones, or the basal ones commonly

roimded apically or oblanceolate to ovate or oval in outline 16

— Leaves serrate to entire, the basal ones variously shaped, but mainly acute to
attenuate apically 20

16(15). Basal leaves distinctly pinnately divided, the lobes often again toothed

S. multilobatus

— Basal leaves merely toothed to subentire 17

17(16). Basal and lower cauline leaves entire to dentate, but not pinnatifid S. hartianus

— Basal and lower cauline leaves toothed to pinnatifid 18

18(17). Middle and upper cauline leaves clasping with large auriculate bases

S. dimorphophyllus

— Middle and upper cauline leaves without a prominent clasping or auriculate
base 19

19(18). Basal leaves obovate to oblanceolate or ovate, rounded apically, thickish;

plants of dryish habitats S. streptanthifolius

— Basal leaves oblanceolate to elliptic, obtuse, but usually pointed apically, thin;
plants of meadows S. pauperculus

20(15). Cauline leaves rounded and more or less clasping basally, long-attenuate api-
cally, entire or denticulate S. integerrimus

— Cauline leaves tapering to the base or with a few basal clasping lobes in some,
usually not attenuate apically 21

21(20). Stems subscapose, the cauline leaves none or few and bractlike; plants often so-
boliferous or with a branching rhizomatous caudex S. werneriifolius

— Stems more or less leafy, the cauline leaves gradually reduced upward, but
hardly bractlike; plants seldom as above 22

22(21). Involucral bracts ca 8 or fewer; heads 5-6 mm wide, mainly 20-60 per in-
florescence S. atratus

— Involucral bracts mostly 13-21; heads 8-12 mm wide or more, fewer or
larger than above 23

332

Great Basin Naturalist

Vol. 43, No. 2

23(22). Achenes glabrous; plants often less than 20 cm tall 24

— Achenes hirtellous or hispidulous; plants often over 20 cm tall 25

24(23). Main leaves regularly and evenly pinnatifid or pinnatisect; plants often with

slender rhizomes S. fendleri

— Main leaves entire to dentate, not as above; plants shortly rhizomatous S. canus

25(23). Main leaves 10-15 cm long or more, entire or denticulate; plants 50-70 cm

tall, of northern Utah S. sphaerocephahis

— Main leaves 2-8 cm long, dentate, serrate, or subentire; plants mainly 15-35

cm tall, of southern Utah S. neomexicanus

Senecio amplectens Gray Alpine Ground-
sel. Perennial short-rhizomatous herbs; stems
ascending to erect, mainly 8-30 cm tall;
herbage glabrous or sparingly tomentose;
main leaves middle and lower cauline, the
lower ones broadly petiolate, more or less
clasping the stem, the blades 3-10 cm long,
0.8-3 cm wide, dentate to shallowly lobed;
cauline leaves becoming short-petiolate or
sessile upward, finally bractlike; heads 1-5
(rarely more), conspicuously nodding, corym-
bose; involucres broadly hemispheric, 10-15
mm long and about as wide or wider; bracts
mainly ca 21, usually brown, with scarious
margins, glabrous; outer bracts to about half
as long as the inner; rays 7-17, yellow, 10-25
mm long; pappus white; achenes glabrous.
Spruce-fir and alpine tundra communities, of-
ten in talus or on ridge margins, at 3050 to
3570 m in Beaver, Grand, Piute, San Juan,
Sanpete, and Utah counties (Wasatch, Tush-
ar, and La Sal mountains, and Wasatch
Plateau); Montana, Wyoming, Colorado, and
Nevada; 24 (v). Our material belongs to var.
holmii (Greene) Harrington [S. holmii
Greene; Ligularia holmii (Greene) W.A.
Weber].

Senecio atratus Greene Perennial sub-
rhizomatous herbs from a branching caudex;
stems erect or ascending, 2-8 dm tall; herb-
age floccose-tomentose; basal and lower cau-
line leaves petiolate, mainly 8-30 cm long,
1-4 cm wide, the blade oblanceolate or ob-
long, conspicuously dentate to subentire;
cauline leaves gradually reduced upward, be-
coming sessile or subsessile and finally brac-
teate; heads ca 15-60, in more or less com-
pact corymbose clusters; involucres 6-8 mm
high, 3-6 mm wide; main bracts 8 or fewer,
greenish to brownish, the margins scarious,
the tips black, tufted-hairy apically; rays 3-5,
yellow, 4-8 mm long; pappus white; achenes

glabrous. Aspen, spruce-fir, mixed conifer,
and tall forb communities at 2440 to 3335 m
in Duchesne, Garfield, Iron, San Juan, Salt
Lake, Sanpete, and Uintah counties; Colo-
rado and New Mexico; 22 (iv).

Senecio bigelovii Gray in Torr. Bigelow
Groundsel. Perennial subrhizomatous herbs;
stems erect, mainly 3-8 (10) dm tall; herbage
floccose-tomentose to glabrate or glabrous;
main leaves cauline, largest below, reduced
gradually upward, petiolate below, sessile
and clasping to auriculate above, mostly 7-15
cm long, 0.6-3 (5) cm wide, the blades oblan-
ceolate to oblong or elliptic, subentire to ser-
rate; heads 3-8, nodding, racemosely ar-
ranged; involucres 8-12 mm long, 12-25 mm
wide; bracts mainly ca 21, usually brown,
with scarious margins, the outer to half as
long as the inner, all sparingly tomentose; ray
flowers lacking; achenes glabrous. Mountain
brush, ponderosa pine, aspen, and spruce-fir
communities at 2745 to 3175 m in San Juan
County; Wyoming south to New Mexico and
Arizona; 2 (0). Our material has been as-
signed to var. hallii Gray; the type variety is
more southern.

Senecio canus Hook. Gray Groundsel. [S.
piirshianus Nutt.; S. convalliiim Greenm.,
type from Rabbit Valley]. Perennial short-
rhizomatous herbs, often with a caudex;
stems 8-30 cm tall (rarely more), erect or as-
cending; herbage wooUy-tomentose; basal
leaves petiolate, the blades 1-5 cm long,
3-30 mm wide, lanceolate to oblanceolate,
elliptic or ovate, entire or denticulate, obtuse
to rounded apically; cauline leaves reduced
upward, the upper ones often clasping, final-
ly bracteate, occasionally lobed in some in-
trogressant forms; heads mainly 2-10, sub-
umljellate or corymbose; involucres 3-8 mm
long, 4-10 mm wide; main bracts 13-21,
lance-attenuate, greenish or with brownish

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

333

midstripe, glabrous or tomentose; outer
bracts very short; rays 8-13, yellow, 5-10
mm long; achenes glabrous. Pinyon-juniper,
sagebrush, Douglas fir, aspen, spruce-fir, and
alpine timdra commvmities, often in talus or
on windswept ridges, at 2105 to 3815 m in
Beaver, Box Elder, Cache, Carbon, Daggett,
Duchesne, Garfield, Iron, Juab, Millard,
Piute, San Juan, Sanpete, Sevier, Summit,
Uintah, and Utah counties; British Columbia
to Manitoba, south to California, Nevada,
Colorado, and Kansas; 56 (xiii). This attrac-
tive grayish white species forms inter-
mediates with S. multilobatus, S. streptanthi-
folius, and S. werneriifolius.

Senecio crassulus Gray Perennial short-
rhizomatous herbs, often with a caudex;
stems 15-50 cm tall or more, erect; herbage
glabrous; lower leaves broadly petiolate, the
main ones 3-15 cm long, 0.6-3 (5) cm wide,
lanceolate to elliptic or oblanceolate, dentate
to entire; cauline leaves reduced upward, be-
coming sessile and clasping; heads solitary or
2-12, corymbose; involucres 8-13 mm high,
12-21 mm wide; main bracts 8-21, oblong to
lance-oblong, greenish to brown, with
scarious margins, the tips black and tufted-
hairy; outer bracts to half as long as the inner
or more; rays 8-13, yellow, 5-12 mm long;
achenes glabrous. Aspen, lodgepole pine, and
spruce-fir communities, often in forb-grass
meadows, at 1830 to 3355 m in Box Elder,
Cache, Carbon, Duchesne, San Juan, Salt
Lake, Sanpete, Sevier, Summit, and Utah
counties; Oregon to Montana, south to New
Mexico; 34 (iv).

Senecio crocatus Rydb. Perennial sub-
rhizomatous herbs, the caudex more or less
developed; stems erect, mainly 2-8 dm tall;

herbage glabrous or with minute hairs in the
inflorescence; basal leaves with long slender
petioles, the blades 1-8 cm long, 1-4 cm
wide, ovate to oblong, lanceolate, or elliptic,
subcordate to acute basally, often rounded
apically, entire to crenate-dentate; cauline
leaves reduced upward, becoming lobed or
sublyrate, sessile and sometimes auriculate
and/or clasping; heads mainly 3-30; in-
volucres 4-8 mm long, 5-8 mm wide; main
bracts 13-21, lance-oblong, green or suffused
with red or purple; outer bracts very short;
rays 6-13, orange or orange-red; pappus
white; achenes glabrous. Rush-grass, willow,
aspen-forb, and lodgepole pine communities
at 2195 to 2990 m in Cache, Duchesne, Rich,
Summit, and Utah counties; Colorado; 23 (i).
One specimen from Rich County (Thome
1465 BRY) is apparently intermediate with S.
eremophilus. The species is remarkably like
the next.

Senecio dimorphophyllus Greene Per-
ennial subrhizomatous herbs; stems erect,
mainly 30-70 cm tall; herbage glabrous or es-
sentially so; basal leaves with long slender
petioles, the blades 1-7 cm long, 1-5 cm
wide, oval to oblong or elliptic, subcordate to
acute basally, commonly rounded apically;
cauline leaves becoming sessile, lyrate-pin-
natifid, and auriculate-clasping, the auricles
often lobed; heads mainly 2-25, sub-
umbellately to corymbosely arranged; in-
volucres 5-8 mm high, 6-10 mm wide; main
bracts 13-21, lance-attenuate, green, some-
times suffused reddish, the tips not black,
tufted-hairy; outer bracts very short; rays
8-13, yellow, 5-8 mm long; pappus white;
achenes glabrous. Two weakly discernible va-
rieties are present in Utah.

Cauline leaves merely lobed to subentire; plants of the La Sal Mountains

S. dimorphophyllus var. intermedius

Cauline leaves sharply lobed; plants of Uinta Mountains and Wasatch Plateau
S. dimorphophyllus var. dimorphophyllus

Var. dimorphophyllus Aspen-tall forb and
spruce-fir communities at 1860 to 3265 m in
Duchesne, Emery, Sanpete, and Utah coun-
ties; Wyoming and Colorado; 9 (0). Utah ma-
terials approach S. crocatus in most morpho-
logical features, including the tall stature. If
the flower color is discounted and the larger
heads are not definitive, then the specimens

could be considered as a portion of S. cro-
catus. Some specimens from Duchesne Coun-
ty appear to be transitional to S.
sphaerocephalus.

Var. intermedius T.M. Barkley Wet mead-
ows at 3050 to 3115 m in the La Sal Moun-
tains, San Juan County (type from Geyser
Pass); endemic; 2 (0).

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Great Basin Naturalist

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Senecio douglasii DC. Suffrutescent pe-
rennials; stems erect or ascending, mainly
3-8 (10) dm tall; herbage glabrous or tomen-
tose; leaves simple and linear-filiform or pin-
natifid into linear-filiform segments, 2-11 cm
long, 0.8-3 mm wide; heads few to numer-
ous, in paniculately branched subcorymbose
cymes; involucres campanulate, mainly 5-10
mm long, 6-14 mm wide; main bracts 13-21,

lance-oblong, green, with scarious margins,
minutely tufted-hairy apically; the outer
bracts short and inconspicuous or to half as
long as the inner ones; rays 8-17, yellow,
10-18 mm long; pappus white; achenes
hairy. Two infraspecific taxa, previously
treated at specific rank with some justifica-
tion, are present in Utah.

1. Herbage grayish or whitish tomentose; outer involucral bracts short and incon-
spicuous; plants rather broadly distributed S. douglasii var. longilobus

— Herbage green, glabrous or essentially so; outer involucral bracts to about half

as long as the inner ones; plants of Washington County ... S. douglasii var. monoensis

Var. longilobus (Benth.) L. Benson [S.

longilobus Benth.; S. filifolius var. iamesii T.
& C, nom. illeg.]. Warm desert shrub, salt
desert shrub, sagebrush-rabbitbrush, saltgrass,
and pinyon-jimiper communities at 1095 to
2200 m in Beaver, Duchesne, Garfield, Iron,
Kane, Millard, Piute, San Juan, Sevier, Wash-
ington, and Wayne counties; Arizona to
Texas; 52 (x).

Var. monoensis (Greene) Jepson [S. mon-
oensis Greene]. Creosote bush, blackbrush,
other warm desert shrub, and pinyon-juniper
communities at 760 to 1465 m in Washington
Coimty; California to Texas; 23 (vi).

Senecio eremophilus Richards. Perennial
subrhizomatous herbs; stems rather equably
leafy, erect or ascending, mainly 2.5-9 dm
tall; herbage glabrous or essentially so; lower
leaves often deciduous or withered at an-
thesis; cauline leaves 2-15 cm long (or more),
0.4-5 (7) cm wide, oblanceolate to elliptic, or
lanceolate in outline, pinnatifid or pinnately
lobed or toothed, the lower ones petiolate,
becoming sessile upward; heads several to
numerous, corymbose; involucres 5-8 mm
high, 6-10 mm wide; main bracts 8-17,
lance-oblong, brownish or greenish, with
scarious margins, blackish tips, and hair-
tufted apices; outer bracts very short; rays
7-10, yellow, 5-10 mm long; pappus white;
achenes glabrous or puberulent along the
ribs. Grass-forb,- ponderosa pine, aspen,
lodgepole pine, spruce-fir, and alpine timdra
communities, at 1615 to 3450 m in Beaver,
Carbon, Duchesne, Emery, Garfield, Grand,
Iron, Juab, Piute, San Juan, Salt Lake, San-
pete, Sevier, Summit, Tooele, Uintah, Utah,

Wasatch, Washington, and Wayne counties;
British Columbia and Mackenzie south to
Arizona and New Mexico; 104 (xiv). Our ma-
terial belongs to var. kingii (Rydb.) Greenm.
[S. kingii Rydb., type from Cottonwood Can-
yon]. This plant forms intermediates with S.
spartioides.

Senecio fendleri Gray Perennial rhizo-
matous herbs, with a caudex more or less de-
veloped; stems mainly 5-30 cm tall, erect or
ascending; herbage floccose-tomentose; basal
leaves petiolate, the blades 1-6 cm long,
4-20 mm wide, pinnatifid or pinnately lobed;
cauline leaves reduced upward, becoming
sessile, finally bracteate; heads 3 to many, co-
rymbose; involucres 4-6 mm high, 5-8 mm
wide; main bracts ca 13, lance-attenuate,
greenish, the margins scarious or hyaline,
minutely hairy apically, more or less tomen-
tose below; outer bracts very short; ray flow-
ers lacking; pappus white; achenes glabrous.
Ridge tops on limestone barrens near Musi-
nea Peak, at ca 2960 to 3295 m in Sanpete
County (Lewis 4274, 5516 BRY); Wyoming
south to New Mexico; 2 (0). Our specimens
approach S. canus, more or less.

Senecio fremontii T. & G. Perennial herbs,
subrhizomatous or from a caudex and tap-
root; stems 0.6-4 dm tall; herbage glabrous;
leaves cauline, 1-6 cm long, 0.5-2 cm wide,
oblanceolate to obovate, shortly petiolate or
sessile and somewhat clasping, dentate to
subentire; heads 1-5; involucres 6-12 mm
high, 7-12 mm wide; main bracts 8-17,
lance-oblong or lance-attenuate, green or
brown, the margins scarious, tufted hairy api-
cally; outer bracts short and inconspicuous or

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

335

to half as long as the inner ones; rays 7-10,
yellow, 5-12 mm long; pappus white;

achenes glabrous or hairy. Two rather weak
varieties are present.

1. Involucres mostly 8-10 mm high; stems mostly less than 20 cm high

S. fremontii yar. fremontii

— Involucres 10-12 mm high; stems often over 30 cm tall S. fremontii var. blitoides

Var. blitoides (Greene) Cronq. [S. blitoides
Greene]. Alpine communities, often in talus
or on rock outcrops, at 2745 to 3355 m in
Salt Lake (?), Tooele, and Utah counties;
Wyoming to Colorado; 9 (0).

Var. fremontii Spruce-lodgepole pine and
alpine timdra communities at 3050 to 3965 m
in Duchesne, San Juan, Salt Lake, Summit,
Uintah, and Utah counties; British Columbia
and Alberta, south to Oregon and Wyoming;
16 (v).

Senecio hartianus Heller Perennial herbs
from a subrhizomatous or stoloniferous cau-
dex; stems erect, 2-5 dm tall; herbage floc-
cose-tomentose, sometimes glabrate; basal
leaves petiolate, the blades 1-5 cm long,
0.5-3 cm wide, oval to obovate or elliptic,
serrate or crenate, rounded apically; cauline
leaves reduced upward, subpinnatisect to en-
tire; heads 3-12, corymbose; involucres 4-7
mm high and as broad; main bracts 13-21,
lance-attenuate, greenish, the tips glabrous;
rays ca 10-13, yellow, 5-8 mm long; achenes
glabrous. Ponderosa pine community at ca
2290 m in Kane County (Atwood 7425 BRY);
Arizona; 1 (0).

Senecio hydrophilus Nutt. Water Ground-
sel. Perermial subaquatic herbs from a caudex
and fibrous roots; stems erect, mainly 4-10
dm tall; herbage glacuous, blue-green; basal
and lower cauline leaves petiolate, the broad
petioles with clasping bases, the blades 5-35
cm long or more, 1-10 cm wide, elliptic to
oblanceolate, entire or denticulate, thick and
leathery; cauline leaves reduced upward, be-
coming sessile, finally bracteate; heads nu-
merous in a branching corymbose cluster; in-
volucres 5-8 mm long, 4-76 mm wide; main
bracts 8-13, oblong or lance-attenuate, yel-
lowish, the tips often black, tufted-hairy; rays
3-5 or lacking, yellow, 3-8 mm long; pappus
white; achenes glabrous. Stream banks, pond
margins, and wet meadows at 1375 to 2745
m in Carbon, Garfield, Juab, Kane, Piute,
Salt Lake, Sanpete, Sevier, Sujnmit, and Utah

counties; British Columbia, south to Califor-
nia and Colorado; 25 (iv).

Senecio integerrimus Nutt. Wet-the-bed.
Perennial herbs with a short subrhizomatous
caudex; stems mainly 1-6 (7) dm tall, erect;
herbage arachnoid- villous or glabrate; basal
and lower cauline leaves broadly petiolate,
3-20 cm long, 0.8-4 cm wide, lanceolate to
elliptic or oblanceolate to oblong, entire or
serrate to dentate, rounded to obtuse api-
cally; cauline leaves reduced upward; heads
few to many, in a corymbose to subumbellate
cyme; involucres 6-12 mm high, 8-18 mm
wide; main bracts 13-21, lance-attenuate,
green, with scarious margins and black tips,
the tips tufted-hairy; outer bracts very short;
rays 8-13 (or lacking), yellow, 4-15 mm long;
pappus white; achenes glabrous. Sagebrush,
pinyon-juniper, forb-grass, mountain brush,
ponderosa pine, aspen, and spruce-fir com-
munities at 1460 to 3660 m in probably all
Utah counties; British Columbia to Montana,
south to California; 132 (xiv). Presumed hy-
brids with S. dimorphophyllus are known
(Hansen sn 1976 BRY).

Senecio multilobatus T. & G. Uinta
Groundsel. [S. lapidum Greenm., type from
Silver Reef]. Perennial (or biennial?) herbs
from a taproot; stems mainly 1-6 dm tall;
herbage glabrous, glabrate, or tomentose
throughout or only in axils of basal leaves;
basal leaves 2-12 cm long, 0.3-3.5 cm wide,
spatulate to obovate in outline, pinnatifid to
lyrate-pinnatifid, the segments variously
again toothed, petiolate; cauline leaves re-
duced upward, finally bracteate; heads few
to many, corymbose or subumbellate; in-
volucres 4-9 cm high, 4-10 mm wide; main
bracts 13-21, lance-attenuate or oblong-at-
tenuate, the margins scarious, the apices hair
tufted; rays 7-13, yellow, 4-10 mm long, or
lacking; pappus white; achenes glabrous.
Blackbrush, sandy desert shrub, pinyon-juni-
per, sagebrush, mountain brush, ponderosa
pine, aspen, lodgepole pine, and spruce-fir

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Great Basin Naturalist

Vol. 43, No. 2

communities at 915 to 3420 m in all Utah
counties (type from the Uinta River); Idaho
and Wyoming to California, Arizona, and
New Mexico; 312 (xliv). This widespread and
common species forms presumed hybrids
with S. streptanthifolius and S. neo-
mexicanus.

Senecio neomexicanus Gray Perennial (or
biennial?) herbs from a taproot; stems 14-40
cm tall, erect; herbage tomentose; basal and
lower cauline leaves petiolate, the blades 1-5
cm long, 0.6-2 cm wide, oblanceolate to obo-
vate or oval, dentate, serrate or subentire,
toothed to obtuse apically; cauline leaves re-
duced upward, toothed to lobed or entire,
bracteate in inflorescence; heads few to
many, corymbose or subumbellate; involucres
4-7 mm high, 5-12 mm wide; main bracts
13-21, lance-attenuate, green or brown, with
scarious margins, not especially hairy api-
cally; rays 8-13, yellow, 4-10 mm long; pap-
pus white; achenes pubescent. Sagebrush,
mountain brush, ponderosa pine, and aspen
communities at 2105 to 3050 m in Garfield,
Kane, San Juan, and Wayne counties; Colo-
rado, New Mexico, and Arizona; 10 (0). Our
materials are assigned to var. mutahilis
(Greene) Barkley [S. mutabilis Greene].
Through this variety there is virtually a com-
plete intergrading series into S. wernerii-
folius, S. streptanthifolius, and S. multi-
lobatus (Barkley 1978).

Senecio pauperculus Michx. Perennial
herbs from a subrhizomatous caudex; stems
erect, mainly 2-4 dm tall; herbage glabrous
or somewhat tomentose in axils of basal
leaves; basal leaves petiolate, the blades
mainly 2-6 cm long, 0.5-3 cm wide, oblan-
ceolate to elliptic, obovate or ovate, crenate,
dentate, or subentire; cuneate basally,
toothed to obtuse apically; cauline leaves re-
duced upward, becoming sessile, pinnatifid,
not especially auriculate, finally bracteate;
heads few to many, corymbose or sub-
umbellate; involucres 4-8 mm long, 5-9 mm

wide; main bracts 13-21, lance-attenuate, of-
ten with scarious margins, the tips not espe-
cially tufted-hairy; outer bracts very short;
rays 8-13, yellow or yellow-orange, 4-10 mm
long; pappus white; achenes glabrous or pu-
berulent along the angles. Lodgepole pine
and spruce-fir communities, usually in moist
meadows, at 2345 to 2745 m in Daggett,
Garfield, and Rich counties; Alaska to Lab-
rador, south to Oregon and Georgia; 4 (0).
Our material is intermediate to both S. strep-
tanthifolius and S. crocatus.

Senecio pudicus Greene [S. cernuus Gray,
not L.f.; Ligularia pudica (Greene) W.A.
Weber]. Perennial herbs from a sub-
rhizomatous caudex; stems 20-50 cm tall,
erect; herbage glabrous; basal and lower
cauline leaves petiolate, the blades 3-15 cm
long, 0.5-3 cm wide, lanceolate to oblanceol-
ate or narrowly elliptic, tapering basally,
acute apically, entire or shallowly dentate;
cauline leaves reduced upward, finally brac-
teate; heads few to many, nodding; in-
volucres 5-9 mm long and as broad; main
bracts 8-13, lance-oblong, green to brown,
the margins scarious, tufted-hairy apically;
outer bracts very short; ray flowers lacking;
pappus white; achenes glabrous. Aspen,
spruce-fir, and alpine tundra communities at
2650 to 3480 m in Carbon and Garfield coun-
ties; Colorado; 11 (i).

Senecio serra Hook. Perennial herbs from a
caudex, with coarse, felt-covered roots; stems
equably leafy, erect, 4-15 dm tall (or more),
glabrous or sparingly tomentose; leaves 3-15
cm long, 0.4-4 cm wide, short-petiolate, the
blades lanceolate to narrowly lanceolate or
linear, dentate to subentire; heads several to
numerous, corymbose; involucres 4-11 mm
high, 2-10 mm wide; main bracts 8-13,
lance-oblong, greenish to brownish, the mar-
gins scarious, black-tipped, hair tufted; outer
bracts very short; rays 5-8, yellow, 3-10 mm
long; pappus white; achenes glabrous or es-
sentially so. Two rather distinctive varieties
are present.

Involucral bracts 4-6 mm long, 2-6 mm wide; disk flowers ca 12; plants of

central and northern Utah S. serra var. serra

Involucral bracts 6-8 mm long, 6-10 mm thick; disk flowers ca 20; plants of
San Juan County S. serra var. admirabilis

Var. admirabilis (Greene) A. Nels. [S. ad-
mirabilis Greene]. Ponderosa pine commu-

nity at ca 1830 m in San Juan County;
Wyoming and Colorado; 1 (0).

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

337

Var. serra Sagebrush, mountain brush, as-
pen, forb-grass, lodgepole pine, and spruce-
fir communities at 1830 to 3035 m in Box El-
der, Cache, Davis, Duchesne, Juab, Rich, Salt
Lake, Summit, Utah, and Weber counties;
Washington to Montana, south to California
and Nevada; 42 (vi).

Senecio spartioides T. & G. Broom
Groundsel. Perennial herbs from a taproot;
stems equably leafy, erect or ascending, 2-10
dm tall or more, often in clumps; herbage

glabrous; leaves 2-10 cm long or more, lin-
ear, simple and entire or with linear lobes,
mainly 1-3 mm wide (wider in some hybrid
derivatives); heads several to many in branch-
ing corymbose cymes; involucres subcylindric
to narrowly campanulate, 5-10 mm high, 4-8
mm wide; main bracts 8-13, lance-linear,
green, the margins scarious, not tufted-hairy;
outer bracts very short; rays 4-8, yellow,
7-12 mm long; pappus white; achenes white-
hairy. Two intergrading varieties are present.

1. Leaves simple and unlobed, or, if lobed, lower cauline leaves often over 4 mm

wide; plants widespread S. spartioides var. spartioides

— Leaves commonly with 4-6 lateral lobes, seldom if ever more than 2.5 mm

wide; plants of southeastern Utah S. spartioides var. multicapitatus

Var. multicapitatus (Greenm. in Rydb.)
Welsh comb. nov. [based on: Senecio multi-
capitatus Greenm. in Rydb. Bull Torrey Bot.
Club 33: 160. 1906]. Warm desert shrub and
pinyon-juniper communities, often in saline
riparian sites, at 1220 to 1895 m in Garfield,
Grand, San Juan, and Wayne counties; Colo-
rado, Arizona, New Mexico, and Texas; 11
(vii). Barkley (1978) hesitated to combine S.
multicapitatus with S. spartioides, because of
field distinctions. They are, however, much
alike and evidently lack diagnostic criteria
that will allow segregation of all specimens.
Further, specimens intermediate between S.
spartioides and S. eremophilus bear "multi-
capitatus" leaves. I follow a moderate course
in maintaining this taxon at varietal level.

Var. spartioides [S. incurvus A. Nels., type
from Zion National Park]. Warm desert
shrub, pinyon-juniper, sagebrush, mountain
brush, and aspen communities, often in sand,
at 1155 to 2870 m, in Beaver, Duchesne,
Emery, Garfield, Grand, Iron, Kane, Piute,
San Juan, Sanpete, Sevier, Uintah, Washing-
ton, and Wayne counties; Wyoming to South
Dakota, south to California and New Mexico;
75 (xxv). Intermediates are formed with S.
eremophilus.

Senecio sphaerocephalus Greene [S. lugens
var. hookeri D.C. Eaton, type from Summit
(?) County]. Perennial herbs from a short
stout rhizome; stems erect or ascending, 3-8
dm tall; herbage tomentose; basal leaves pet-
iolate, the blades 4-15 cm long, 1-3.5 cm
wide, oblanceolate to elliptic, entire or den-
ticulate, obtuse apically; cauline leaves re-

duced upward, becoming sessile, finally brac-
teate; heads few to many, corymbose; in-
volucres 3-7 mm long, 6-12 mm wide; main
bracts 13-21, oblong- to ovate-lanceolate,
greenish or brownish, with scarious margins,
the tips black, hair-tufted apically; outer
bracts very short; rays 8-13, yellow, 4-10
mm long; pappus white; achenes hairy.
Lodgepole pine and spruce-fir communities,
in meadows, at 2315 to 3205 m in Daggett,
Duchesne, Summit, and Wasatch counties;
Oregon and Montana, south to Nevada and
Wyoming; 10 (i).

Senecio streptanthifolius Greene [S. aqua-
riensis Greenm., type from Aquarius Plateau;
S. jonesii Rydb., type from Alta; S. leonardii
Rydb., type from American Fork Canyon; S.
malmstenii Blake in Tidestr., type from
Wasatch Mountains; S. ruhricaulis var. aph-
anactis Greenm., type from Logan; S. wardii
Greene, type from Fish Lake Mountain]. Pe-
rennial herbs from a taproot and simple or
branched and infrequently subrhizomatous
caudex; stems erect, mainly 8-47 cm tall;
herbage glabrous or rarely sparingly tomen-
tose; leaves thickish; basal leaves petiolate,
the blades 1-5 cm long, 0.3-3 cm wide, ob-
lanceolate to obovate, suborbicular, elliptic,
or ovate, crenate, dentate, or subentire, less
commonly lobed; cauline leaves reduced up-
ward, commonly some of them pinnatifid, fi-
nally bracteate; heads few to many, corym-
bose to subumbellate; involucres 4-8 mm
high, 5-12 mm wide; main bracts 8-21,
lance-oblong, green or brownish, the margins
scarious, sparingly hair-tufted apically; outer

338

Great Basin Naturalist

Vol. 43, No. 2

bracts very short; rays 8-13, yellow, 5-8 mm
long; pappus white; achenes glabrous. Sage-
brush, mountain brush, ponderosa pine, as-
pen, lodgepole pine, spruce-fir, and alpine
tundra communities, often in meadows, at
1370 to 3415 m in Box Elder, Cache, Carbon,
Daggett, Duchesne, Emery, Garfield, Grand,
Juab, Kane, Millard, Salt Lake, Sanpete, Se-
vier, Summit, Tooele, Uintah, Utah, Wasatch,
Washington, Wayne, and Weber counties;
Yukon to Northwest Territories, south to Cal-
ifornia and New Mexico; 107 (vii). This spe-
cies forms a plexus around which revolves
such species as S. pauperculus, S. multi-
lobatus, S. neoniexicanus, and S. canus, as
judged from morphological intermediates,
which are presumed to be hybrids.

Senecio triangularis Hook. Arrowleaf
Groundsel. Perennial herbs from a caudex
and more or less well-developed rhizome;
stems equably leafy, erect, 2.5-12 dm tall or
more; herbage glabrous or sparingly tomen-
tose; leaves petiolate, the blades mainly 3-15
cm long, 0.5-6 cm wide, lance-oblong to
triangular, abruptly contracted or subhastate
at the base, dentate to sinuate dentate or sub-
entire, finally bracteate in the inflorescence;
heads few to many, subcorymbose; involucres
6-12 mm high, 8-17 mm wide; main bracts
8-12, lance-attenuate, the margins often
scarious, tufted-hairy apically; outer bracts
very short; rays 5-9, yellow, 6-15 mm long;
pappus white; achenes glabrous. Aspen-
mountain brush, Douglas fir-white fir, lodge-
pole pine, and spruce-fir communities at
1765 to 3265 m in Cache, Duchesne, Grand,
Salt Lake, Summit, Utah, and Weber coun-
ties; Alaska and Yukon, south to California
and New Mexico; 25 (viii).

Senecio vulgaris L. Common Groundsel.
Plants annual or biennial, with fibrous roots,
1-5.5 dm tall, the stems glabrous or sparingly
villous; basal leaves smaller than the main
cauline ones, often withered by anthesis;
cauline leaves not much reduced upward,
2-10 cm long, 0.5-4.5 cm wide, irregularly
piimatifid, the lobes again toothed, glabrous

or more or less villous, especially along the
veins beneath, the lower ones petiolate, the
upper ones becoming sessile and auriculate-
clasping; heads few to many; involucres 5-8
mm high, 4-10 mm wide; the outer bracts
short and black tipped, the inner lance-linear,
green, with scarious margins, black tipped;
ray flowers lacking; pappus white; achenes
hairy. Weedy species of disturbed sites in Salt
Lake and Utah counties; adventive from Eu-
rope; 6 (0).

Senecio werneriifolius (Gray) Gray [S.
aureus var. werneriifolius Gray]. Plants com-
monly rhizomatous or soboliferous herbs;
stems erect or ascending, 3-18 cm tall; herb-
age tomentose, often glabrate or glabrous in
age; basal leaves petiolate, the blades 0.6-3
cm long, 0.4-2 cm wide, oval to elliptic, obo-
vate, or oblanceolate, thickish, sometimes
revolute; cauline leaves few, commonly in-
conspicuous and bracteate; heads 1-4; in-
volucres 4-10 mm long, 7-15 mm wide; main
bracts 13-21, lance-oblong, green or suffused
with purple, the margins scarious, hair tufted
apically; outer bracts to half as long as the in-
ner; rays 8-13, yellow, 4-10 mm long; pap-
pus white; achenes glabrous. Ponderosa pine,
western bristlecone pine, aspen-conifer, and
spruce-fir communities, often in semibarrens,
at 2375 to 3600 m in Beaver, Duchesne, Gar-
field, Iron, Juab, Piute, Salt Lake, Summit,
and Utah counties; Idaho and Montana, south
to California, Nevada, and Arizona; 28 (ii).

SOLIDAGO L.

Perennial herbs from a caudex or rhizome;
leaves alternate, simple; heads numerous,
radiate, yellow, borne in paniculate, race-
mose, or cymose clusters; involucres imbri-
cate in several series or subequal, commonly
chartaceous or with the tips green; recep-
tacle flat, naked; ray flowers fertile; disk
flowers perfect, fertile; anthers subentire ba-
sally; style branches with lanceolate ap-
pendages; pappus of capillary bristles;
achenes few nerved, pubescent.

2(1).

Heads in corymbs or flat-topped cymes; leaves punctate; plants of lower eleva-
tions riparian habitats [Euthamia] S. occidentalis

Heads racemose or panicled; leaves not punctate; plants of various habitats 2

Stems glabrous 3

Stems puberulent with short incurved hairs or villous with multicellular hairs 4

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

339

3(2). Plants definitely rhizomatous; involucres 2.5-4 mm long; plants of lower ele-
vations S. missouriensis

— Plants subrhizomatous; involucres 4-6 mm long; plants of higher elevations

S. spathulata

4(2). Stems villous with multicellular hairs; petioles long-ciliate S. multiradiata

— Stems puberulent with short incurved hairs; petioles scabrous or strigose mar-
ginally 5

5(4). Involucres 6-11 mm high, the outer bracts subfoliaceus S. parryi

— Involucres 2-5 mm high, the bracts not subfoliaceus 6

6(5). Leaves very numerous and much longer than the internodes, gradually attenu-
ate or acuminate, not dimorphic, strongly 3-nerved S. canadensis

— Leaves not very numerous, often less than twice as long as the internodes,
acute or rounded apically, often dimorphic, with lateral nerves obscure or
moderately apparent 7

7(6). Leaves sparingly hairy to glabrous, the margins rough-hairy; plants widespread

and common S. sparsiflora

— Leaves cinereus-puberulent with disoriented hairs, the margins hairy like the
surfaces; plants more restricted and less common S. nana

Solidago canadensis L. Goldenrod. [S. al-
tissima L.; S. lepida DC.]. Perennial herbs
from creeping rhizomes; stems 3-12 dm tall
or more; herbage puberulent with short in-
curved hairs, or the stems glabrous below;
basal leaves often deciduous or withered at
anthesis; cauline leaves numerous and
crowded, 2-10 cm long or more, 3-20 mm
wide, lanceolate to lance-linear, or narrowly
elliptic, tapering to a sessile base, 3-nerved,
serrate to entire, attenuate to acuminate api-
cally; inflorescence commonly (but not al-
ways) of recurved branches with secund
heads; involucres 2-5 mm high and about as
broad, the bracts lance-attenuate, scarious or
greenish; rays 10-17, yellow, 1-3 mm long.
Riparian and other mesic sites at 350 to 2290
m in all Utah counties; widespread in North
America; 87 (xvi). This plant serves as host
for a peculiar red and black leaf beetle. A
phase of the species is cultivated as an orna-
mental in Utah. Designation of varietal level
in Utah seems academic. The species is tran-
sitional to S. sparsiflora.

Solidago missouriensis Nutt. Missouri
Goldenrod. Perennial herbs from creeping
rhizomes; stems* 2-5 (9) dm tall; herbage
glabrous or sparingly puberulent in in-
florescence only; basal leaves oblanceolate,
often withered at anthesis; main cauline
leaves 2-13 cm long, 0.4-1.5 cm wide, oblan-
ceolate to elliptic or linear, tapering to a ses-

sile base, mainly 3-nerved, entire or essen-
tially so, acute to obtuse apically; in-
florescence compact, with ascending
branches, somewhat or not at all secund; in-
volucres mostly 3-5 mm high and as broad,
the bracts lance-attenuate, greenish to
scarious; rays 7-13, yellow, 2-3 mm long. Ri-
parian communities at 1525 to 2475 m in Box
Elder, Carbon, Daggett, Duchesne, Emery,
Salt Lake, Sanpete, Summit, Uintah, and
Wasatch counties; British Columbia to On-
tario, south to Arizona, Texas, and Tennessee;
13 (ii).

Solidago multiradiata Ait. Low Gold-
enrod. [S. ciliosa Greene]. Perennial herbs
from a rhizome or rhizomatous caudex; stems
5-45 cm tall; herbage villous with multi-
cellular hairs, at least on upper stem and pet-
iole bases; basal and lower cauline leaves
1.5-14 cm long, 5-24 mm wide, oblanceolate
to spatulate or elliptic, tapering to a con-
spicuously ciliate petiole, obscurely 3-nerved,
entire or serrate, rounded to obtuse apically;
inflorescence loosely to densely corymbose;
involucres 4-6 mm high, 5-7 mm wide, the
bracts lance-oblong, green apically, with
prominent midvein; rays ca 13, yellow, 4-5
mm long. Aspen, lodgepole pine, spruce-fir,
and alpine tundra communities at 2745 to
3660 m in Beaver, Cache, Carbon, Duchesne,
Garfield, Grand, Iron, Juab, Kane, Piute, San
Juan, Salt Lake, Sanpete, Sevier, Summit,

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Great Basin Naturalist

Vol. 43, No. 2

Uintah, and Utah counties; Alaska to Quebec,
south to California and New Mexico; 84
(xvii). Our specimens belong to var. scopulo-
rum Gray.

Solidagu nana Nutt. Dwarf Goldenrod. [S.
radulina Rydb., type from Cottonwood Can-
yon]. Perennial herbs from a rhizome or sub-
rhizomatous caudex; stems 13-48 cm tall;
herbage densely canescent with fine hairs of
mixed orientation; basal and lower cauline
leaves petiolate, 1.5-9 cm long, 0.7-2.3 cm
wide, oblanceolate to spatulate, tapering to a
petiole, weakly 3-nerved, entire or slightly
toothed, rounded to obtuse apically; cauline
leaves definitely reduced upward; in-
florescence corymbose, seldom if at all se-
cund; involucres 4-6 mm high and about as
broad; rays 5-8, yellow, 3-4 mm long. Desert
shrub upward to spruce-fir communities,
mainly in riparian or wet meadow sites, at
1460 to 2745 m in Duchesne, Kane, Salt
Lake, Sevier, Summit, Uintah, Utah, and
Wasatch counties; Idaho to Montana, south
to Arizona and Colorado; 13 (i).

Solidago occidentalis (Nutt.) T. & G.
Western Goldenrod. [Euthamia occidentalis
Nutt.]. Perermial herbs from elongate rhi-
zomes; stems erect, branched above, mainly
4-12 (20) dm tall; herbage essentially
glabrous; leaves numerous, sessile, linear to
lance-linear, 2-10 cm long, 1-10 mm wide;
inflorescence usually large, leafy-bracted,
broadly roimded; involucres 3.5-4.5 mm high
and about as broad, the bracts narrowly ob-
long, greenish apically, the midnerve con-
spicuous; rays 15-30, yellow, 1.5-2.5 mm
long. Riparian habitats at 850 to 1650 m in
Box Elder, Cache (?), Carbon, Duchesne,
Emery, Garfield, Grand, Juab, Kane, San
Juan, Salt Lake, Uintah, Utah, Washington,
and Weber counties; British Columbia and
Alberta, south to California, New Mexico,
and Nebraska; 42 (x). I follow tradition by in-
cluding this taxon in Solidago; it might best
be treated in Euthamia.

Solidago parryi (Gray) Greene Parry
Goldenrod. [Haplopappus parryi Gray; H.
parryi var. minor Gray, type from Alta]. Per-
ennial rhizomatous herbs; stems erect or as-
cending, 8-50 cm tall; herbage scabrous to
hispidulose; basal and cauline leaves petio-
late, mainly 3-20 cm long, 0.9-3.8 cm wide,
oblanceolate to elliptic, entire, obtuse to

rounded apically; cauline leaves becoming
sessile and smaller upward, more or less
clasping; heads few to many in compact
branched cymes; involucres 8-11 mm high,
7-14 mm wide; outer bracts ovate to ovate-
lanceolate, green, ciliate, the bases often
scarious; inner bracts narrower and with
scarious or hyaline margins; rays 12-20, yel-
low, 5-8 mm long. Aspen, tall forb, lodge-
pole pine, spruce-fir, and alpine tundra com-
munities at 2285 to 3570 m in Beaver,
Carbon, Duchesne, Emery, Garfield, Juab,
Kane, Millard, Piute, San Juan, Salt Lake,
Sanpete, Sevier, Summit, Tooele, Uintah,
Utah, and Wasatch counties; Wyoming, New
Mexico, Arizona; 50 (ix).

Solidago sparsiflora Gray [S. garrettii
Rydb., type from Big Cottonwood Canyon].
Perennial rhizomatous herbs; stems erect or
ascending, mainly 15-50 dm tall; herbage
puberulent (often sparingly so on leaf sur-
faces); leaves cauline or basal, oblanceolate
to elliptic or spatulate, mainly 1-10 cm long,
2-25 mm wide, entire or less commonly some
of them serrate, acute to attenuate or obtuse
to rounded apically, often dimorphic, with
the upper ones reduced in size; inflorescence
a pyramidial to conic or cylindric cluster,
compact or with branches curved and heads
secund; involucres 4-6 mm high and about as
broad; bracts oblong to subulate, chartaceous
basally, green apically, the midvein con-
spicuous; rays 5-10 or more, yellow, 3-4 mm
long. Pinyon-juniper, mountain brush, sage-
brush, aspen, ponderosa pine, and spruce-fir
communities at 1125 to 3050 m in all Utah
counties (except Box Elder and Morgan);
Wyoming and South Dakota, south to Ari-
zona and Nevada; 141 (xix). Our materials are
far from uniform; in the hanging gardens of
southeastern Utah they are transitional to S.
canadensis (having more ray flowers), and at
high elevations they are more or less inter-
mediate with S. spathulata. Possible addition-
al influence of S. mollis Bartl. and/or S.
nemoralis Ait. is indicated, although neither
of them is known from the state currently.

Solidago spathulata DC. Perennial herbs
from a subrhizomatous caudex; stems 5-30
cm tall (rarely more), erect or ascending;
herbage glabrous or somewhat scabrous and
often glutinous above; basal leaves oblanceo-
late to spatulate, 2-15 cm long, 8-30 mm

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

341

wide, serrate to entire, obtuse to rounded
apically; cauline leaves reduced upward, fi-
nally sessile and more or less clasping; in-
florescence compact to elongate, narrow, the
heads not secund; involucres 4-6 mm high
and as broad or more; bracts oblong, scarious
or greenish along the prominent midvein;
rays 5-10, yellow, 2.5-4 mm long. Aspen,
spruce-fir, and alpine tundra communities at
2440 to 3510 m in Beaver (?), Daggett, Du-
chesne, Emery, Garfield, Grand, San Juan,
Sevier, Summit, and Uintah counties; Alaska
to Quebec, south to California, Arizona, and
New Mexico; 21 (v). Two completely inter-
grading phases, regarded as varieties, are
present in Utah; a tall montane phase known
as var. neomexicana (Gray) Cronq. [S. multi-
radiata var. neomexicana Gray], and a dwarf

alpine phase known as var. nana (Gray)
Cronq. [S. humilis var. nana Gray; S. decum-
bens Greene].

SONCHUS L.

Annual or perennial herbs from taproots or
deep-seated, rhizome-like roots, the juice
milky; leaves chiefly cauline, alternate,
simple, entire to lobed or pinnatifid; heads
few to several; involucral bracts imbricate in
several series, green or greenish (drying
brownish), the inner ones with hyaline mar-
gins; receptacle naked; corollas of ray-flow-
ers only, yellow, perfect; pappus of capillary
bristles; style branches semicylindrical;
achenes compressed, several to many nerved,
beakless, glabrous.

1. Plants perennial, spreading from rhizomelike roots; involucres more than 14

mm long in fruit 2

— Plants armual from taproots; involucres less than 14 mm long in fruit 3

2(1). Involucres and peduncles bearing coarse stipitate glands S. arvensis

— Involucres and peduncles glabrous or tomentose, not stipitate-glandular

S. uliginosus

3(1). Leaves sharply and narrowly toothed, and sometimes lobed; achenes not trans-
versely wrinkled, merely longitudinally nerved S. asper

— Leaves sharply and broadly toothed, or merely toothed and lyrate pinnatifid;
achenes transversely wrinkled and longitudinally nerved S. oleraceus

Sonchus arvensis L. Field Sow-thistle.
Plants perennial with deep-seated rhizome-
like roots; stems 4-10 dm tall or more, pu-
bescent with coarse stipitate glands, at least
above, and often glabrous below; leaves 5-40
cm long, 0.8-10 cm broad, more or less pin-
natifid, auriculate-clasping basally, acute to
obtuse apically, prickly margined; heads few
to several, the peduncles stipitate-glandular;
involucres 14-20 mm high and 10-30 mm
broad in fruit, the bracts lance-oblong to
lance-linear, glandular like the peduncles;
rays yellow, mostly 10-20 mm long; achenes
transversely wrinkled. Weedy species of dis-
turbed soils at 1370 to 2135 m in Cache,
Duchesne, Salt Lake, and Utah counties;
widely distributed and considered as a
"noxious" weed in North America; adventive
from Europe; 10 (0).

Sonchus asper (L.) Hill Spiny Sow-thistle.
Plants annual from taproots; stems 3-10 dm
tall, pubescent with coarse stipitate glands, at

least above, often glabrous below (less com-
monly throughout); leaves 3-15 cm long, 1-5
cm broad, merely lobed or lobeless, auricu-
late-clasping basally, acute to acuminate or
less commonly obtuse apically, the margins
armed with slender sharp prickles; heads few
to several, the peduncles stipitate-glandular
or glabrous; involucres 9-14 mm long and
10-16 mm wide in fruit, the bracts lance-ob-
long to lance-linear, glabrous or with few
stipitate glands; rays yellow, mostly 5-10 mm
long; achenes 2-3 mm long, several nerved,
not transversely wrinkled. Weed of disturbed
sites at 760 to 2135 m in Box Elder, Du-
chesne, Garfield, Grand, Kane, Millard,
Piute, San Juan, Salt Lake, Sevier, Tooele,
Uintah, Utah, and Washington counties;
widespread in North America; adventive
from Europe; 27 (ii).

Sonchus oleraceus L. Common Sow-thistle.
Plants annual from taproots, the stems 2-10
dm tall or more, glabrous throughout or

342

Great Basin Naturalist

Vol. 43, No. 2

sometimes with stipitate glands above; leaves
4-20 cm long, 0.6-10 cm broad, more or less
lyrate-pinnatifid, auriculate-clasping basally,
acute to obtuse apically, irregularly and
broadly toothed, the teeth weakly prickly;
heads few to several, the peduncles glabrous
or stipitate-glandular; involucres 10-13 mm
high and 8-20 mm broad in fruit, the bracts
lance-linear to lance-oblong, glabrous or with
a few stipitate glands; rays yellow, mostly
8-12 mm long; achenes 2-3 mm long, several
nerved and transversely wrinkled. Weeds of
disturbed sites at 850 to 2135 m in Duchesne,
Garfield, Salt Lake, Utah, and Washington
counties; widely distributed in North Ameri-
ca; adventive from Europe; 9 (i).

Sonchus uUginousus Bieb. Meadow Sow-
thistle. Plants perermial from deeply seated
rhizomelike roots; stems 4-10 dm tall or
more; herbage glabrous or obscurely tomen-
tose; leaves 5-40 cm long, 0.8-10 cm wide,
pinnatifid, auriculate-clasping basally, acute
to obtuse apically, prickly margined; heads
few to several, the peduncles glabrous; in-
volucres mainly 14-16 mm high and 10-20
mm broad in fruit; bracts lance-linear to ob-
long, glabrous or tomentose; rays yellow,
mostly 10-20 mm long; achenes 2-3.5 mm
long, several nerved, transversely wrinkled.
Weeds of disturbed sites at 1220 to 2260 m in
Daggett, Duchesne, Garfield, Grand, Juab,

Salt Lake, Uintah, and Utah counties; wide-
spread in North America; adventive from Eu-
rope; 22 (ii). Authors of Flora Europaea (Tut-
in et al. 1976) treat this entity as S. arvensis
ssp. uliginosus (Bieb.) Nyman. Amow et al.
(Flora of the Central Wasatch Front, Utah)
discount the usefulness of stipitate glands as
diagnostic features, noting that glandular and
eglandular plants occur together in the same
populations, and that glands are not corre-
lated with other features. On a statewide
basis the plants act like legitimate taxa, and
the eglandular plants do seem to have some-
what smaller heads.

Sphaeromeria Nutt.

Perennial herbs or subshrubs; leaves alter-
nate or mainly basal, simple and entire or
pinnatifid to palmatifid; heads discoid, few to
several, corymbose to subcapitate; involucres
hemispheric to campanulate; bracts in 2 or 3
series, imbricate to subequal; receptacle con-
ic or concave, naked; outer flowers pistillate,
fertile; disk flowers perfect, fertile; pappus
lacking or a short crown; achenes usually 5-
to 10-ribbed, glabrous or glandular.
Holmgren, A. H., L. M. Shultz, and T. K.
Lowrey. 1976. Sphaeromeria, a genus
closer to Artemisia than to Tanacetum
(Asteraceae: Anthemidae). Brittonia 28:
255-262.

2(1).

Plants pulvinate-caespitose; heads capitately arranged on subscapose branches;

known from Garfield County S. capitata

Plants caulescent subshrubs; heads in paniculate or corymbose clusters on leafy
branches, not of Garfield County 2

Leaves pinnatifid, at least some, tomentose; heads paniculate; plants of Wash-
ington County S. ruthiae

Leaves entire or pinnatifid, glabrous; heads corymbose; plants not of Washing-
ton County S. diversifolia

Sphaeromeria capitata Nutt. [Tanacetum
capitatum (Nutt.) T. & G.]. Pulvinate-caespi-
tose herbs; herbage canescent with malpi-
ghian hairs; stems subscapose, 2-12 (20) cm
tall; leaves mainly basal, 4-10 mm long, 1- or
2-palmately lobed, the cauline entire and re-
duced upward; heads few to numerous in a
compact headlike cluster; involucres 3-5 mm
high, the broad bracts with hyaline margins;
corollas 2.5-3 mm long. With western
bristlecone pine on Cedar Breaks limestone,

at ca 2380 m in Garfield County; Montana
and Wyoming; 1 (0).

Sphaeromeria diversifolia (D.C. Eaton)
Rydb. [Tanacetum diversifolium D.C. Eaton].
Subshrubs, mainly 1-4 dm tall; herbage
glabrous; leaves simple, entire, or some of
them pinnately lobed, 8-55 mm long, 0.5-5
mm wide, linear; heads several to many in
compact to open corymbose clusters; in-
volucres 3-4 mm high, the broad bracts with
hyaline margins; corollas 2-2.5 mm long.

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Welsh: Utah Flora: Compositae (Asteraceae)

343

Juniper, mountain brush, mixed conifer, and
aspen communities upward to alpine tundra,
often in rock crevices, at 1370 to 3205 m in
Davis, Juab, Millard, Salt Lake, Tooele, and
Utah counties; Nevada; 33 (i). This is a Great
Basin endemic.

Sphaeromeria ruthiae Holmgren, Shultz,
and Lowrey Subshrubs, mainly 3-7 dm tall;
herbage tomentose-canescent with malpi-
ghian hairs; leaves pinnately lobed or the up-
per ones entire, 1-9 cm long, 2-4 mm wide
or more; heads several to many, paniculate;
involucres 3-5 mm high, the broad bracts

with hyaline margins; corollas 1.8-2 mm
long, yellow. Crevices in Navajo Sandstone,
ponderosa pine community, in Washington
County; endemic; 3 (0).

Stephanomeria Nutt. Nom. Cons.

Annual, biennial, or perennial herbs with
milky juice; leaves alternate, often pinnatifid;
flowers all raylike, perfect, pink or white; in-
volucres cylindric; main bracts few, subequal;
outer bracts much shorter; receptacle naked;
pappus of plumose bristles (barbellate in S.
spinosa); achenes 5-angled or -ribbed.

1. Plants annual, from slender taproots S. exigua

— Plants perennial, a caudex often more or less developed 2

2(1). Plants spinescent; pappus barbellate S. spinosa

— Plants unarmed; pappus plumose 3

3(2). Involucres 12-15 mm high; heads with 10 or more flowers S. parryi

— Involucres 5-12 mm high (rarely higher); heads with 3-9 flowers 4

4(3). Main leaves runcinate-pinnatifid; plants commonly 1-2 dm tall S. runcinata

— Main leaves entire or pinnatifid, often deciduous at anthesis; plants commonly

2-8 dm tall 5

5(4). Stems very slender; leaves filiform to linear, entire or dentate; pappus bristles

white (rarely brownish), plumose to the base S. tenuifolia

— Stems not very slender; leaves linear-subulate, often pinnatifid or lobed; pap-
pus brownish, scabrous toward the base S. pauciflora

Stephanomeria exigua Nutt. Annual or
biennial (winter annual) herbs from slender
taproots; herbage glabrous or puberulent;
stems 5-60 cm tall, erect and commonly
branched from the base upward, often fis-
tulous; main leaves 1-6 cm long, pinnatifid to
bipinnatifid, deciduous or withered by an-
thesis; cauline leaves soon reduced and brac-
teate upward; heads more or less corymbose,
terminating bracteate branchlets; involucres
5-10 mm high, 3-4.5 mm wide; main bracts
usually 3-5; rays pink or white, 3-5 mm
long; pappus of white to off-white bristles
plumose in the upper half; achenes 3-4 mm
long, tuberculate. Warm, mixed cool, and salt
desert shrub, and pinyon juniper commu-
nities, often in sand, at 850 to 2230 m in Bea-
ver, Emery, Garfield, Grand, Kane, Millard,
San Juan, Sevier, Tooele, Uintah, Utah,
Washington, and Wayne counties; Oregon to
Wyoming, south to California and New Mex-
ico; 84 (x).

Stephanomeria parryi Gray Perennial
herbs; stems 1 to few, weak, branching, 8-25
cm tall; herbage glabrous; leaves 2-8 cm tall,
runcinate-pinnatifid, thickish, the lobes
weakly spinulose-tipped; heads terminating
very short bracteate branches, 10- to 14-flow-
ered; involucres 12-15 mm high; rays whit-
ish, 15-20 mm long; pappus bristles tawny,
scabrous at the base only; achenes 3-4 mm
long, not rugose. Blackbrush community at
ca 1460 m in Kane County (Atwood & Allen
2822a BRY); California to Arizona; 1 (0).

Stephanomeria pauciflora (Torr.) A. Nels.
in Coult. & Nels. [Prenanthes? pauciflora
Torr.]. Perennial herbs (or somewhat woody
below) from a caudex, branched from the
base, mostly 30-60 cm tall; herbage glabrous;
main leaves 2-7 cm long, runcinate-pinnati-
fid, the lobes weakly spinulose-toothed; heads
terminating short to elongate branchlets, 3-to
5-flowered; involucres 8-10 high, 3-5 mm
wide; main bracts 5; rays pink or white,

344

Great Basin Naturalist

Vol. 43, No. 2

mainly 4-7 mm long; pappus bristles brown-
ish, plumose except at the base; achenes
3.5-7 mm long, striate, more or less wrinkled.
Warm, salt, and mixed desert shrub, and juni-
per communities, often in sandy soil, at 760
to 1525 m in Beaver, Garfield, Grand, Juab,
Kane, Millard, San Juan (?), Tooele, and
Washington counties; California to Kansas,
south to Texas and Mexico; 26 (v).

Stephanomeria runcinata Nutt. Perennial
herbs from a caudex; stems branched from
the base, mostly 8-25 (30) cm tall; herbage
glabrous, scabrous, or sparingly villous; main
leaves 2-7 cm long, runcinate-pinnatifid, the
lobes merely cuspidate; heads terminating
naked or sparingly bracteate branchlets, com-
monly 5-flowered; involucres 9-12 mm high,
3.5-7 mm wide; rays pink, mainly 8-12 mm
long; pappus bristles white, plumose almost
to the base; achenes 4-5 mm long, tubercu-
late. Salt Desert shrub and pinyon-juniper
communities at 1250 to 2535 m in Daggett,
Duchesne, Emery, Grand, Uintah, and
Wayne counties; Montana to Nebraska and
Colorado; 17 (i).

Stephanomeria spinosa (Nutt.) Tomb
[Lygodesmia spinosa Nutt.]. Perennial herbs
from a woody caudex, the caudex branches
clothed with brownish marcescent leaf bases,
the axils copiously villous-hairy; stems 11-52
cm tall, thorny; herbage glabrous upward or
the branches puberulent; leaves linear 0.5-7
cm long, 1-3 mm wide, reduced to bracteate
scales upward, often lacking at anthesis;

heads terminal on short lateral branches or
sessile, 3- to 5-flowered; involucres 5.7-10
mm high, 3-5 mm wide; main bracts oblong
to lance-oblong, green or often suffused with
purple; outer bracts proportionately broader;
rays pink, 3-5 mm long; pappus bristles off-
white, scabrous throughout; achenes 4-6.5
mm long, smooth. Desert shrub, sagebrush-
grass, pinyon-juniper, mixed conifer, and as-
pen communities, often in moist sites, at 1675
to 3050 m in Beaver, Emery, Garfield, Juab,
Kane, Millard, Piute, Sevier, Tooele, Wash-
ington, and Wayne counties; British Colum-
bia to Montana, south to California and Ari-
zona; 41 (ii).

Stephanomeria tenuifolia (Torr.) Hall
Slender Wirelettuce. [Prenanthes? tenuifolia
Torr.]. Perennial herbs from a woody caudex;
caudex branches lacking or with few marces-
cent leaf bases, not hairy; stems 25-100 cm
tall or more; herbage glabrous or puberulent;
leaves filiform to linear, 1-8 (11) cm long,
1-3 (8) mm wide, entire or dentate, much re-
duced upward; heads terminating elongate or
short lateral bracteate branchlets, 5-flowered;
involucres 8-11.2 (16) mm high, 3-5 mm
wide; main bracts lance-oblong, green, pu-
berulent or glabrous; outer bracts very short;
rays 4-8 (10) mm long, pink; pappus bristles
white, dull white, or less commonly brown-
ish, plumose to the base; achenes 4-6 mm
long, longitudinally ribbed, smooth. Two
more or less distinctive phases are present,
recognizable as varieties.

Involucre 10-16 mm high, the bracts attenuate; basal leaves bipinnatifid, at

least some; plants of Uintah County S. tenuifolia var. uintaensis

Involucres mainly 8-11.2 mm high, the bracts not especially attenuate; basal

leaves seldom if ever bipinnatifid; plants of rather broad distribution

S. tenuifolia var. tenuifolia

Var. tenuifolia Desert shrub, hanging gar-
den, pinyon-juniper, mountain brush, pon-
derosa pine, and white-fir communities, at
1155 to 2746 m in Beaver, Duchesne, Emery,
Garfield, Grand, Iron, Kane, Piute, San Juan,
Sevier, Uintah, Washington, and Wayne
counties; British Columbia to Montana, south
to California, Arizona, and Texas; 46 (xvi).
The great sprawling plants of the canyon-
lands portion of Utah might be worthy of tax-
onomic consideration; sometimes they ap-
proach S. pauciflora in having tawny pappus
bristles.

Var. uintahensis Goodrich & Welsh Pon-
derosa pine community at ca 2490 m in Uin-
tah County; endemic; 2 (0).

Stylocline Nutt.

Woolly annual herbs; stems commonly
branched; leaves alternate, simple, entire;
heads discoid, leafy bracted; involucre per se
lacking; outer receptacular bracts subtending
and enclosing pistillate flowers; receptacle
cylindric; pistillate flowers many, deciduous
with the enclosing bract, the bract apex hya-
line; corollas filiform; pappus none; perfect

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

345

flowers (functionally staminate) few, sur-
rounded by linear hyaline bracts; corollas
tubular, the ovaries vestigial; pappus of 3-5
deciduous bristles; anthers sagittate basally;
achenes ellipsoid, few nerved.

Stylocline micropoides Gray Desert Nest-
straw. Annual woolly herbs; stems usually
branched, 4-12 cm tall; leaves 4-12 mm
long, 0.5-1.5 mm wide, acute; bracteate
leaves 6-10 mm long, 1.5-2.5 mm wide, lan-
ceolate; heads clustered at branch tips, dense-
ly woolly; pistillate flowers with bracts boat
shaped, densely long-woolly, hyaline mar-
gined; staminate flowers with pappus of 3-5
deciduous bristles; achenes ellipsoid, ca 1.5
mm long. Blackbnish, bursage, and indigo
bush communities at 915 to 1160 m in San
Juan and Washington counties; California to
New Mexico, south to Mexico; 3 (i).

Syntrichopappus Gray in Torr.

Aimual herbs; stems simple or branched;
leaves alternate (or some opposite below),
simple, entire or lobed; heads radiate, many,
terminating branchlets; involucres sub-
cylindric; bracts few, in 1 series, partly en-
closing ray achenes; receptacle flat, naked;
ray flowers pistillate, fertile, yellow; disk
flowers perfect, fertile, yellow; anthers ob-
tuse at base; style branches flattened; pappus
of barbellate bristles; achenes 5-angled.

Syntrichopappus fremontii Gray Annual
herbs, 2-14 cm tall; herbage floccose-tomen-
tose; leaves 5-22 mm long, narrowly spatu-
late to spatulate, rounded to 3-lobed apically,
cuneate basally; heads few to many; in-
volucres 5-6 mm high, 3-4 mm wide; bracts
5, oblong, greenish, with scarious margins,
abruptly acute apically; rays 5, yellow, 2-5
mm long; disk corollas numerous, yellow;
pappus of white barbellate bristles falling to-

gether. Joshua tree, creosote bush, black-
brush, sagebrush, and juniper communities at
760 to 1375 m in Washington County; Cali-
fornia, Nevada, Arizona; 6 (i).

Tanacetum L.

Perennial herbs from a rhizome; leaves al-
ternate, 2- to 3-pinnatifid; heads discoid, nu-
merous, corymbose; flowers perfect; in-
volucres hemispheric; bracts in 2 or 3 series,
more or less imbricate, the margins scarious;
receptacle low-convex, naked; anthers entire
at the base; pappus a minute crown; achenes
5-angled, truncate.

Tanacetum vulgare L. Tansy. Aromatic,
glabrous or sparingly tomentose perennials,
3-10 (15) dm tall; leaves 6-15 cm long, ses-
sile or subsessile, the blades 2- to 3-pinnatifid;
heads many, discoid, yellow; involucres ca
4-5 mm high and 6-10 mm broad; bracts lan-
ceolate; marginal flowers 3-lobed; inner flow-
ers 5-lobed; achenes glandular, 5-angled, ca 1
mm long. Weedy species of disturbed soils at
1370 to 1985 m in Emery, Uintah, and Utah
counties; widespread in the U.S.; adventive
from Europe; 3 (0).

Taraxacum Hall.

Perennial scapose herbs with milk juice,
from taproots; leaves all basal, pinnatifid to
subentire; heads solitary on a scape; in-
volucral bracts in 2 series, herbaceous, the
outer shorter, the inner often dilated or ap-
pendaged apically, usually with broad hya-
line or scarious margins, at least basally; re-
ceptacle naked; corollas of ray flowers only,
perfect, yellow; pappus of capillary bristles;
style branches semicylindric; achenes angular
or terete, prominently nerved or ribbed, usu-
ally spinulose or with ridges near the body
apex, glabrous, beaked.

2(1).

Irmer involucral bracts commonly dilated or bearing appendages apically, over

10 cm long; plants indigenous, of high elevations T. ceratophorum

Inner invofucral bracts usually not dilated or with appendages apically; plants
various 2

Outer bracts reflexed or spreading, the inner ones 12-18 mm long; achenes

straw colored to olive drab or brownish; plants adventive T. officinale

Outer bracts erect, the inner ones 6-10 mm long; achenes black to grayish;
plants indigenous at high elevations T. lyratum

346

Great Basin Naturalist

Vol. 43, No. 2

Taraxacum ceratophorum (Ledeb.) DC.

Rough Dandelion. [Leontodon ceratophorus
Ledeb.]. Plants mostly 4-10 cm tall, from a
simple or branched caudex; leaves 4-8 cm
long, 0.7-2 cm broad, subentire to toothed;
scapes sparingly villous, moderately so below
the head; involucres 12-17 mm high in flow-
er, the outer bracts ovate to lanceolate, ap-
pressed or ascending, the inner ones lance-
oblong, attenuate, the apex dilated or
appendaged; rays yellow; achene bodies 3-7
mm long, straw colored to olive-drab or
brownish, the beak usually 2-4 times longer
than the body; pappus white. Spruce krumm-
holz and sedge-forb meadows at 3230 to 3660
m in Daggett, Duchesne, and Uintah counties
(Leidy Peak); Alaska to Yukon, east to the
Atlantic, south to California, New Mexico,
and Massachusetts; circumboreal; 2 (0).

Taraxacum lyratum (Ledeb.) DC. Alpine
Dandelion. [Leontodon lyratus Ledeb.].
Plants mostly 2-8 cm tall, from a simple or
branched caudex; leaves 1-6 cm long, 0.3-1
cm wide, pinnately lobed to pinnatifid or
subentire; scapes glabrous or nearly so; in-
volucres 6-10 mm high, the outer bracts lan-
ceolate-ovate, appressed or ascending-spread-
ing, the inner ones lance-oblong to oblong,
scarcely or slightly dilated; rays yellow (fad-
ing bluish); achene bodies 3-6 mm long,
black or grayish, the beak subequal to the
body; pappus white. Alpine tundra and
meadows in spruce-fir communities at 3325
to 3965 m in Duchesne and Summit counties;
Alaska and Yukon, south to Nevada, Arizona,
and Colorado; Asia; 5 (i).

Taraxacum officinale Weber ex Wiggars

Common Dandehon. Plants mostly 3-60 cm
tall, from a simple or branched caudex;
leaves 5-40 cm long, 1-10 cm wide, pin-
nately lobed to pinnatifid, the terminal lobe
broader than the lateral ones; scapes villous
to subglabrous, often moderately to densely
villous below the head; involucres 15-25 mm
high in flower, the outer bracts lance-acumi-
nate, reflexed, the inner ones lance-attenuate,
not or scarcely dilated apically, rarely appen-
daged; rays yellow, or bluish externally; ach-
ene bodies 3-4 mm long, straw colored to
olive drab, the beak usually 2-4 times longer
than the body; pappus white. Ubiquitous
brightly flowered weedy species at 885 to
3205 m throughout Utah; widespread in
North America; adventive from Eurasia; 65
(xiii). This handsome plant is among the
earliest of our spring flowers, and among the
last to bloom in autumn.

Tetradymia DC.

Armed or unarmed shrubs; stems pannose-
tomentose; leaves alternate, entire, foliaceous
or modified as spines, with secondary leaves
fasciculate in the axils; heads discoid, corym-
bose or racemose; involucres cylindric to tur-
binate or hemispheric; receptacle naked;
bracts 4-6, equal or nearly so; flowers 4-8,
yellow or cream; style branches truncate to
rounded or conic apically; anthers sagittate
basally; pappus of capillary bristles or barbel-
late scales; achenes striate.
Strother, J. L. 1974. Taxonomy of Tetra-
dymia (Compositae: Secenioneae). Brit-
tonia 26:177-202.

1. Heads solitary or 2 or 3, axillary; primary leaves modified as spines 2

— Heads several to many in terminal corymbose clusters; primary leaves fo-
liaceous or modified as spines 3

2(1). Spines commonly recurved, mainly 5-20 mm long, pannose-tomentose;

achenes 6-8 mm long; plants widespread, not of Washington County T. spinosa

— Spines straiglit, mainly 20-40 mm long, glabrescent; achenes 4-5 mm long;
plants of Washington County T. axillaris

3(2). Primary leaves modified as persistent spreading, straight or recurved spines

5-25 mm long T. nuttallii

— Primary leaves not modified as persistent spines, if at all spinescent then ap-
pressed-ascending 4

April 1983

4(3).

Welsh: Utah Flora: Compositae (Asteraceae)

347

Primary leaves linear-subulate, spinescent apically, appressed-ascending, to-

mentose; secondary leaves obtuse apically, glabrous or essentially so T. glabrata

Primary leaves various but not spinescent, not contrasting in shape and
pubescence with the secondary ones T. canescens

Tetradymia axillaris A. Nels. Longspine
Horsebrush. Spiny shrubs, mainly 4-12 dm
tall; branchlets evenly white-pannose; pri-
mary leaves modified as persistent spines 1-5
cm long, straight or becoming curved, to-
mentose at first, becoming glabrate; second-
ary leaves linear to spatulate, 2-12 mm long,
essentially glabrous; heads solitary or 2 or 3,
from nodes of the previous year; involucres
8-11 mm high; bracts 5, subequal, tomentose;
flowers 5-7, pale yellow, the corollas 7.5-9
mm long; pappus of slender bristles; achenes
4.5-5.5 mm long; achenes pilose, the hairs
9-11 mm long. Salt and warm desert shrub
communities at 850 to 1375 m in Washington
County; Nevada and California; 18 (ii). Our
material belongs to var. longispina (Jones)
Strother [T. spinosa var. longispina Jones,
type from St. George].

Tetradymia canescens DC. Gray Horse-
brush. [T. linearis Rydb., type from Iron
County]. Unarmed shrubs, mainly 1-9 dm
tall; branchlets white-pannose except for gla-
brate streaks below the primary leaves; pri-
mary leaves 0.5-4 cm long, 1-6 mm wide,
lanceolate to oblanceolate or spatulate, to-
mentose; secondary leaves similar to the pri-
mary ones but shorter and narrower; heads
few to several at branch tips; involucres 6-8
mm high or more; bracts 4, subequal, tomen-
tose; flowers 4, yellow to cream, the corollas
7-11 mm long; pappus of white or tawny
bristles; achenes 2.5-5 mm long, glabrous or
hairy. Sagebrush-grass, mountain brush, pon-
derosa pine, mixed conifer, and aspen com-
munities at 1525 to 3150 m throughout Utah;
British Columbia to Montana, south to Cali-
fornia, Arizona, and New Mexico; 75 (viii).

Tetradymia glabrata T. & G. Shrubs,
mainly 3-12 dm tall; branchlets pannose ex-
cept for glabrate or glabrous streaks below
the primary leaves; primary leaves mainly
5-15 mm long, 0.8-1.4 mm wide, linear-sub-
ulate, spinose tipped, soon deciduous; second-
ary leaves linear to narrowly spatulate,
glabrous or thinly tomentose; heads few to

many on branch tips; involucres 7-10 mm
high; bracts 4, subequal, tomentose to
glabrous; flowers 4, yellow to cream, the co-
rollas 9-10 mm long; pappus of white
bristles; achenes 3-5 mm long, hirsute. Shad-
scale, greasewood, sagebrush, rabbitbrush,
and juniper communities at 1370 to 2370 m
in Emery, Juab, Millard, Sanpete, Sevier,
Tooele, and Wayne counties; Oregon and
Idaho, south to California and Nevada; 44

(V).

Tetradymia nuttallii T. & G. Nuttall
Horsebrush. Spinescent shrubs, 3-12 dm tall;
branchlets white-pannose except for glabres-
cent streaks below the primary leaf bases;
primary leaves modified as persistent straight
or recurved spines 5-25 mm long, tomentose
to glabrous; heads in terminal clusters of (2)
3-6; involucres 6-9 mm high; bracts 4, equal;
flowers 4, yellow, the corollas 8-10 mm long;
pappus of white or tawny bristles; achenes
4-6 mm long, hirsute. Shadscale, greasewood,
sagebrush-rabbitbrush and pinyon-juniper
communities at 1370 to 1830 m in Box Elder,
Daggett, Duchesne, Juab, Millard, Tooele,
and Uintah counties; Wyoming and Nevada;
25 (i).

Tetradymia spinosa H. & A. Spinescent
shrubs, 3-12 dm tall; branchlets evenly pan-
nose; primary leaves modified as spines, 5-20
mm long, tomentose, finally glabrate; second-
ary leaves linear to spatulate, glabrous or gla-
brescent; heads borne singly or in pairs, later-
ally, on stems of the previous season;
involucres 8-12 mm high; bracts 4-6, sub-
equal, tomentose; flowers 5-8, yellow, the
corollas 6-10 mm long; pappus of slender
bristles, white; achenes 6-8 mm long, hairy,
the trichomes 9-12 mm long. Mixed desert
shrub, shrub-grass, and pinyon-juniper com-
munities at 1250 to 1925 m in Carbon, Dag-
gett, Duchesne, Emery, Garfield, Grand,
Juab, Millard, Salt Lake, Uintah, and Utah
counties; Oregon to Montana and Wyoming,
south to California, Nevada, and New Mexi-
co; 15 (vi).

348

Great Basin Naturalist

Vol. 43, No. 2

Thelesperma Less.

Perennial glabrous or sparingly puberulent
herbs; leaves opposite, pinnately to pal-
mately parted, or the upper ones entire;
heads pedunculate, solitary or few per stem;
involucres hemispheric to campanulate;
bracts in 2 unlike series, the outer ones

spreading and distinct, the inner ones con-
nate to the middle and calyxlike; receptacle
flat, chaffy with broad scarious scales; rays
present (or lacking), neuter, yellow; disk
flowers perfect, fertile; anthers not caudate
basally; pappus of 2 retrorsely hispid awns, a
crown, or lacking; achenes oblong to linear.

Plants 30-80 cm tall; rays normally lacking; pappus of 2 awns; known from

San Juan and Washington counties T. megapotamicum

Plants 3-35 cm tall; rays normally present; pappus a crown or none T. suhnudum

Thelesperma megapotamicum (Spreng.)
Kuntze Greenthread. [Bidens megapotamica
Spreng]. Perennial herbs from a caudex and
stout root; stems 30-80 cm tall; leaves mainly
2-7 cm long, once or twice pinnatifid, the
lobes linear, or the uppermost simple; outer
bracts 4-6, oblong to ovate, obtuse, much
shorter than the inner; inner bracts 6-12 mm
high, connate to above the middle, the lobes
with narrow scarious margins; rays lacking;
disk flowers yellow (or brownish); pappus of
2 or 3 retrorsely hispid awns; outer achenes
somewhat papillose dorsally. Desert shrub
commimity at ca 915 to 1375 m in San Juan
and Washington counties; Wyoming to Ne-
braska, south to Arizona, Texas, and Mexico;
1(0).

Thelesperma subnudum Gray Perennial
herbs from a taproot and less commonly with
a caudex and creeping rootstock; stems 3-35
cm tall, subscapose; leaves mainly at base of

stem, 1.5-9 cm long, pinnately to sub-
palmately lobed or some or all of them en-
tire; petioles often ciliate and blades more or
less puberulent; involucres 6.3-14 mm high,
9-22 mm wide; outer bracts oblong to lan-
ceolate, with narrow scarious margins, to half
as long as the inner ones; inner bracts united
to below the middle, conspicuously scarious-
margined; rays present and bright yellow,
10-28 mm long and 6-18 mm wide, or lack-
ing; disk flowers yellow; pappus a toothed
crown or lacking; achenes glabrous or hairy
apically, 3.5-4.5 mm long. This taxon is vari-
able being radiate or discoid, in division of
leaves, and in position of leaves along the
stem. They occur mainly at elevations below
2135 m elevation. A dwarf alpine phase oc-
curs above that elevation, and because of its
small size, lack of rays, and apparent eco-
typical differences these plants are herein
designated at varietal level.

1. Plants mainly 3-7 cm tall; involucres 6.3-9 mm high, 9-14 mm wide; heads

discoid T. subnudum var. alpinum

— Plants mainly 9-35 cm tall; involucres 8-14 mm high, 12-22 mm wide;

heads commonly radiate T. subnudum var. subnudum,

Var. alpinum Welsh Pinyon-juniper,
mountain brush, and western bristlecone pine
communities at ca 2745 m in Wayne County;
endemic; 2 (0).

Var. subnudum Mixed desert shrub, salt
desert shrub, and pinyon-juniper commu-
nities at 1065 to 2135 m in Carbon, Du-
chesne, Garfield, Grand, Iron (type from Red
Creek), Kane, San Juan, Uintah, Washington,
and Wayne counties; Colorado, Arizona, and
New Mexico; 109 (xiv).

TOWNSENDIA Hook.

Annual, biennial, or perennial herbs, cau-
lescent or acaulescent; leaves alternate, en-
tire or rarely lobed or toothed; heads radiate,
solitary or few, terminating branches, or ses-
sile; receptacle convex, naked; involucres
campanulate to hemispheric; bracts in 2-7
series; rays pistillate, fertile, the corollas
white, pink, or yellow; disk flowers perfect,
yellow; disk pappus of barbellate capillary
bristles; ray pappus similar to that of the disk

April 1983 Welsh: Utah Flora: Compositae (Asteraceae) 349

or shortened; achenes 2- or 3-ribbed, com- Contr. Gray Herb. 183:1-151.

pressed, usually hairy. Reveal, J. L. 1970. A revision of the Utah

Beaman, J. H. 1957. The systematics and species of Townsendia (Compositae).

evolution of Townsendia (Compositae). Great Basin Nat. 30:23-52.

1. Plants caulescent, the internodes apparent, annual or biennial (short-lived

perennial) 2

— Plants acaulescent, the internodes not elongating, perennial 5

2(1). Plants annual or winter annual; disk pappus shorter than disk-corollas; plants

of southeastern Utah (Navajo Basin) T. annua

— Plants biennial or short-lived perennials; disk pappus subequal to or longer
than the disk corollas 3

3(2). Achenial hairs unevenly branched; ray flowers usually dark pink-purple dor-
sally; plants biennials of western Utah T. florifer

— Achenial hairs glochidiate; ray flowers variously colored, but if dark pink-
purple dorsally then the plants perennial and of different distribution 4

4(3). Stems gray-white, the pubescence dense; plants of broad distribution,

perennial T. incana

— Stems thinly strigose, evident beneath the hairs; plants of the Uinta Basin, bien-
nial T. strigosa

5(1). Involucral bracts linear to narrowly lanceolate, in 5-7 series 6

— Involucral bracts lanceolate to ovate or elliptic, in 2-5 series 9

6(5). Involucral bracts hair tufted apically, linear, acuminate; plants of Carbon,

Duchesne, and Daggett counties T. hookeri

— Involucral bracts not hair tufted apically, narrowly lanceolate, acute; plants
variously distributed 7

7(6). Rays glandular dorsally; leaves canescent; plants of Duchesne and Uintah

counties T. mensana

— Plants glabrous or sparingly pubescent dorsally; leaves greenish or grayish-
canescent; plants not or seldom of Duchesne and Uintah counties 8

8(7). Disk pappus 3-6 mm long; leaves green, the midveins not conspicuous; plants

of the Wasatch Plateau and Uinta Mountains T. leptotes

— Disk pappus 6-11 mm long; leaves grayish canescent, the midveins con-
spicuous; plants of Sevier, Iron, Wayne, and Garfield counties T. exscapa

9(5). Rays yellow ventrally, densely glandular and often purplish dorsally; ray pap-
pus 1-2 mm long; plants of Emery and eastern Sevier counties T. aprica

— Rays white or pink or bluish, or rarely yellow ventrally, but, if yellow, the ray
pappus 2-4.5 mm long and plants of other distribution 10

10(9). Plants green or greenish; flowers often bluish or purplish to pink, mainly of

higher elevations in mountains and plateaus T. montana

— Plants grayish canescent or whitish; flowers seldom bluish or purplish, usually
white to pink or yellowish ventrally; mainly of low elevations 11

11(10). Involucral bracts sparingly strigose; ray pappus 2-4.5 mm long; plants mainly

of western Utah T. jonesii

— Involucral bracts moderately strigose; ray pappus 0.3-0.6 mm long; plants
mainly of eastern Utah T. incana

350

Great Basin Naturalist

Vol. 43, No. 2

Townsendia annua Beaman Caulescent
annual or winter annual herbs, 2-18 cm tall;
herbage strigose; leaves of basal rosettes soon
withered or poorly developed; cauline leaves
5-28 mm long, 1-5 mm wide, oblanceolate
to spatulate or linear, sparingly to moder-
ately strigose, green or greenish; heads soli-
tary or few; involucres 4.5-7 mm long, 6-14
mm wide; bracts in 2-4 series, green or suf-
fused with purple, scarious, ciliate; rays
13-34, the corollas white or pink to lavender,
4-8 mm long, 1-2.3 mm wide, glabrous; disk
corollas yellow, 2.2-3.5 mm long; achenes
1.9-2.6 mm long, pubescent with glochidiate
hairs; ray pappus 0.2-0.8 mm long, that of
disk flowers 1.8-3 mm long. Sandy desert
shrub and blackbrush communities at 1125 to
1590 m in Carbon, Emery, Garfield, Grand,
Kane, and San Juan counties; Colorado, Ari-
zona, New Mexico, and Texas; 23 (v).

Townsendia aprica Welsh & Reveal Pulvi-
nate-caespitose acaulescent perennial herbs
from a caudex, 1.5-2.5 cm tall; leaves 7-13
(16) mm long, 1-3.5 mm wide, spatulate to
oblanceolate, strigose; heads sessile, sub-
mersed in the leaves; involucres 4-8 mm
high, 7-13 mm wide; bracts in 3-4 series,
lanceolate, fimbriate, red-scarious, hyaline-
ciliate, the outermost sparsely strigose; rays
13-21, the corollas yellow to golden ven-
trally, purplish dorsally and glandular, 4-7
mm long; disk corollas yellow, 3.7-4.5 mm
long; achenes 2-2.5 mm long, 2-ribbed, the
hairs glochidiate; ray pappus 0.7-1 mm long;
pappus of disk flowers 4-5 mm long. Salt
desert shrub and pinyon-juniper commu-
nities, commonly on clay or clay-silt expo-
sures of the Mancos Shale (Blue Gate Mem-
ber), at 1860 to 2440 m in Emery and
adjacent Sevier (type from south of Fremont
Junction) counties; endemic; 10 (ii). The yel-
low flowers and short pappus of ray flowers
are diagnostic.

Toumsendia exscapa (Richards.) T.C. Por-
ter [Aster? exscapa Richards.]. Caespitose
acaulescent perennial herbs from a simple or
branched caudex, 2-3.5 cm high; leaves 0.6-5
cm long, 1-3.5 mm wide, oblanceolate to lin-
ear, acute and mucronate apically, strigose,
with midvein apparent; involucres 10-18 mm
high, 15-30 mm wide; bracts in 4-7 series,
linear to narrowly lanceolate, ciliate on
scarious margins, sparingly strigose to

glabrous; ray flowers 21-40, the corollas
white or pinkish, 8-15 mm long, 1.2-3 mm
wide; disk corollas yellow; achenes 2- or 3-
ribbed, pubescent with glochidiate hairs; ray
pappus 4-8 mm long; disk pappus 6-12 mm
long. Ponderosa pine, mountain sagebrush,
and spRice-fir communities, often in mead-
ows, at 2135 to 3295 m in Garfield, Iron, Se-
vier, and Wayne counties; British Columbia
to Manitoba, south to Nevada, Arizona, Mex-
ico, and Texas; 8 (ii).

Townsendia florifer (Hook.) Gray [Erig-
eron? florifer Hook.; T. watsonii Gray, type
from Stansbury Island; T. scapigera var. am-
higua Gray, type from Rabbit Valley; T.
florifer var. communis Jones, type from
Marysvale]. Caulescent winter annual or
biennial herbs 3-20 cm tall; basal leaves 6-50
mm long, 3-12 mm wide, spatulate; cauline
leaves narrowly oblanceolate to linear, 10-40
mm long, 1-5 mm wide, strigose, petiolate,
grayish; heads solitary or few; involucres
6.5-13 mm high, 15-30 mm wide; bracts in 3
or 4 series, green or suffused with purple,
scarious, ciliate; rays 13-34, the corollas
white or pink ventrally, dark pink or laven-
der dorsally, 7-12 mm long, 1.2-3 mm wide,
often glandular; disk corollas yellow, 3.3-6
mm long; achenes 3.3-4.5 mm long, pu-
bescent with unequally forked hairs; ray pap-
pus 1-6 mm long; disk pappus 3.5-7.5 mm
long. Mixed desert shrub communities at
1280 to 1985 m in Beaver, Box Elder, Gar-
field, Juab, Millard, Sanpete, Sevier, Tooele,
Utah, and Wayne counties; Washington to
Idaho, Oregon, and Nevada; 56 (vii).

Townsendia hookeri Beaman Caespitose
acaulescent perennial herbs from a simple or
branched caudex, 2.5-3.5 cm high; leaves
10-40 mm long, 1-2.5 mm wide, linear to
linear-oblanceolate, strigose; involucres 9-13
mm high, 9-14 mm wide; bracts in 5-7
series, linear to lance-linear, tufted-hairy api-
cally, green or suffused with purple, strigose;
rays 13-34, the corollas 6-9 mm long, 1-1.9
mm wide, white or pink ventrally, pinkish
dorsally, glabrous; disk corollas yellow, 4.5-6
mm long; achenes 3.5-4.5 mm long, pu-
bescent with glochidiate hairs; ray pappus
1-1.5 mm long; disk pappus 5.5-8.5 mm
long. Sagebrush, sagebrush-grass, and mixed
conifer communities at 2165 to 2716 m in
Carbon, Daggett, Duchesne, and Uintah

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

351

counties; Yukon to Saskatchewan, south to
South Dakota and Colorado; 5 (0).

Townsendia incana Nutt. [T. incana var.
amhigua Jones, type from Thompson]. Sub-
caulescent to acaulescent caespitose herbs,
the caudex often branched; stems con-
spicuously white strigose, mainly 2-6 cm
high, forming clumps to 2 dm wide; leaves
5-40 mm long, 1-5 mm wide, spatulate to
oblanceolate, strigose; heads solitary or few;
involucres 7-11 mm high, 8-20 mm wide;
bracts in 3 or 4 series, lanceolate, green, the
margins scarious and ciliate, strigose; rays
13-34, the corollas white ventrally, pink to
lavender dorsally, 6-10 mm long, 1.5-3 mm
wide; achenes 2.5-4.5 mm long, pubescent
with glochidiate hairs; ray pappus 0.3-0.6
mm long; disk pappus 4-7.5 mm long. Black-
brush, salt desert shrub, mixed desert shrub,
pinyon-juniper, and sagebrush commimities
at 1310 to 2290 m in Beaver, Carbon, Dag-
gett, Duchesne, Emery, Garfield, Grand,
Iron, Kane, Piute, San Juan, Sevier, Uintah,
and Wayne counties; Wyoming to Nevada,
Arizona, and New Mexico; 183 (xxiii). This is

the common townsendia of the Colorado
drainage system in Utah; its Great Basin
counterpart is T. pnesii, from which it can be
distinguished by the white strigose stems and
shorter ray pappus.

Townsendia jonesii (Beaman) Reveal [T.
mensana var. jonesii Beaman, type from
Mammoth]. Subcaulescent to acaulescent
caespitose herbs, the caudex commonly
branched; stems not conspicuously white stri-
gose, mainly 2-4 cm tall, forming clumps to
1 dm wide; leaves 10-40 mm long, 1-4 mm
wide, oblanceolate to spatulate or almost lin-
ear, strigose; heads mostly solitary; involucres
9-12.5 mm high, 8-14 mm wide; bracts in 4
or 5 series, lanceolate, green or suffused
purple, sparsely strigose; rays 13-21, the co-
rollas white to pink, cream, or yellow ven-
trally, pink to red-purple dorsally, glandular,
4-7 mm long; disk corollas yellow, ca 3.5 mm
long; achenes 3-5.5 mm long; pubescent with
glochidiate hairs; ray pappus 2-4.5 mm long;
disk pappus 5-8 mm long. Two weak, but
geographically and edaphically correlated,
varieties are present.

1. Ray flowers yellow to lemon-yellow ventrally; plants of gypsiferous substrates

in Sevier and Piute counties T. pnesii var. lutea

— Ray flowers pink to white or cream ventrally; plants of various substrates,

rather broadly distributed T. pnesii var. pnesii

Var. jonesii Sagebrush, shadscale, rabbit-
brush, pinyon-juniper, mountain brush com-
munities at 1525 to 2745 m in Beaver, Juab,
Millard, Sanpete, and Sevier counties; Ne-
vada; 13 (ii). The type of T. mensana var.
pnesii consists of strictly acaulescent plants
with very slender leaves and smallish heads;
it is unmatched in the specimens examined,
and it is understandable why the taxon was
placed initially with T. mensana.

Var. lutea Welsh Salt desert shrub and
juniper communities at ca 1675 to 1830 m in
Sevier and Piute counties (on Arapien shale
and clays in volcanic rubble); endemic; 6 (i).

Toumsendia leptotes (Gray) Osterh. [T. se-
ricea var. leptotes Gray]. Perennial acaules-
cent herbs from a simple or more commonly
branched caudex, 1-3 cm tall; herbage spar-
ingly strigose, greene; leaves 0.6-4 cm long,
1.3-2.6 mm wide, linear to narrowly oblan-
ceolate; involucres 5-10 mm high, 9-14 mm
wide; bracts in 4-7 series, lanceolate to lin-

ear, the margins scarious, ciliate, often suf-
fused purple; rays 13-34, the corollas white,
cream, or pink ventrally, sometimes lavender
dorsally, 6-10 mm long, 1.2-2 mm wide; disk
corollas yellow, 3-5 mm long; achenes pu-
bescent with glochidiate hairs; ray pappus
0.8-6.5 mm long; disk pappus like the ray
pappus. Montane sagebrush and grass-forb
communities, often on ridge crests and
plateau margins at 2680 to 3145 m in Du-
chesne, Sanpete, and Summit counties (Uinta
Mountains and Wasatch Plateau); Idaho and
Montana, south to California, Nevada, and
New Mexico; 7 (0).

Townsendia mensana Jones Perennial
acaulescent herbs from a simple or more
commonly branched caudex 1-2.5 cm high;
herbage strigose; leaves 3-17 mm long,
0.6-1.3 mm wide, narrowly oblanceolate to
linear; involucres 5-9 mm high, 76-10 mm
wide; bracts in 4 or 5 series, lanceolate, the
margin scarious and ciliate; rays 13-21, the
corollas whitish, cream, or pinkish, glandular

352

Great Basin Naturalist

Vol. 43, No. 2

dorsally, 5-7.5 mm long, 0.9-1.4 mm wide;
disk corollas yellow, 3.5-4.8 mm long;
achenes pubescent with glochidiate hairs; ray
pappus 2.5-4 mm long; disk pappus 5-6.5
mm long. Salt desert shrub, pinyon-juniper,
and sagebrush commimities, especially on
barren and semibarren sites, at 1705 to 2715
m in Duchesne (type from near Duchesne,
then Theodore) and Uintah counties; Colo-
rado (?); a Uinta Basin endemic; 38 (v).

Townsendia montana Jones Perennial
acaulescent or rarely subcaulescent herbs
from a simple or branched caudex, sometimes
with soboliferous rhizomatous branches, from

a taproot, 2-6 cm high; herbage glabrate to
strigose; leaves 5-40 mm long, 2-8 mm wide,
spatulate, thickish; involucres 6-12 mm high,
8-20 mm wide; bracts in 3-6 series, oblong,
obovate, oblanceolate or lanceolate, glabrous
or sparingly strigose, the margins scarious;
ciliate, often suffused with purple; rays
12-30, the corollas blue, pink, lavender, or
white, 6-12 mm long, 1-3.5 mm wide; ach-
enes 3.7-5.2 mm long, glabrous or sparingly
pubescent with bifurcate or glochidiate hairs;
ray and disk pappus alike, 3-5.5 mm long.
Three more or less distinctive varieties are
present.

1. Heads usually sessile; leaves mainly 1-3.5 mm wide, rather abruptly obtuse

apically; plants of Garfield and Kane counties T. montana var. minima

— Heads usually at least shortly pedunclulate; leaves mainly broader (at least
some), rounded to obtuse; plants not of Garfield or Kane counties 2

2(1). Leaves rounded apically, broadly spatulate; plants of calciferous outcrops in

southern Duchesne, Wasatch, and Sanpete counties T. montana var. caelilinensis

— Leaves obtuse to subacute apically; plants of various substrates in the Uinta

and Wasatch mountains , T. montana var. montana

Var. caelilinensis Welsh Pinyon-juniper,
spruce-fir, and limber pine communities on
Flagstaff Limestone and Green River forma-
tions at 2135 to 3735 m in southern Du-
chesne, Wasatch, and Sanpete counties; en-
demic; 13 (i).

Var. minima (Eastw.) Beaman [T. minima
Eastw., type from Bryce Canyon]. Ponderosa
pine, western bristlecone, limber pine, and
Douglas fir-white fir communities, on white
and pink members of the Cedar Breaks For-
mation, at 2375 to 3115 m in Garfield and
Kane counties; endemic; 14 (i).

Var. montana [T. dejecta A. Nels., type
from Dyer Mine]. Spruce-fir and lodgepole
pine communities at 3050 to 3510 m in
Cache, Juab, Salt Lake (type from Alta),
Summit, and Uintah counties; Idaho, Mon-
tana, and Wyoming; 2 (0).

Townsendia strigosa Nutt. Caulescent
biennial herbs; stem-s branched from the base
and above, 3-15 cm long; herbage strigose to
strigulose; basal leaves 1.2-4.5 cm long, 1.2-7
mm wide, oblanceolate to spatulate, more or
less persistent; cauline leaves mostly smaller
and narrower, often clustered below and
overtopping the heads; involucres 5-10 mm

high, 7-20 mm wide; bracts in 3 or 4 series,
lance-ovate to lanceolate, the margins
scarious, ciliate, strigose; rays 12-30, the co-
rollas white to pink, sometimes darker dor-
sally, 5-14 mm long, 1.5-3 mm wide; disk co-
rollas 3.3-5 mm long; achenes pubescent
with glochidiate hairs; ray pappus 0.5-1.6
mm long; disk pappus 3.3-5 mm long. Salt
desert shrub, mixed desert shrub, and pinyon-
juniper communities at 1460 to 1895 m in
Daggett, Duchesne, and Uintah counties;
Wyoming; 14 (ii).

Tragopogon L.

Biennial (annual or perennial) herbs from
thickened taproots, the juice milky; leaves al-
ternate, entire, clasping basally; heads soli-
tary or few and corymbose; flowers all ray-
like, perfect, yellow or purple; involucres
cylindric or campanulate; bracts uniseriate,
equal; receptacle naked; pappus of plumose
bristles united at the base; achenes 5- to 10-
nerved, slender-beaked or the outer beakless.
OwNBEY, M. 1950. Natural hybridization and

amphiploidy in the genus Tragopogon.

Amer. J. Bot. 37:487-499.

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

353

1. Peduncles scarcely if at all inflated, even in fruit; achenes 15-25 mm long (in-
cluding the beak); bracts subequal to the rays; plants rare in Utah T. pratensis

— Peduncles strongly inflated apically; achenes 25-36 mm long (including the
beak); bracts usually longer than the rays; plants locally common 2

2(1). Rays purple; involucral bracts mainly 8 or 9 T. porrifolius

— Rays yellow; involucral bracts usually 13 T. dubius

Tragopogon dubius Scop. Biennial herbs;
stems erect, 3-10 dm tall, simple or branched;
leaves mainly 5-25 cm long, linear-subulate
from an expanded base, floccose, becoming
glabrate; peduncles enlarged and fistulous be-
low the heads; involucres cylindric to cam-
panulate; bracts commonly 13 (8 on later
heads), 2.5-4 cm long in flower, 4-7 cm long
in fruit; rays pale lemon yellow, shorter than
the bracts; achenes 25-36 mm long; pappus
whitish to tawny. Disturbed soils and in low
quality range sites at 1370 to 3205 m in all
Utah counties; widely distributed in the U.S.;
adventive from Europe; 58 (vi).

Tragopogon porrifolius L. Oyster-plant;
Salsify. Biennial herbs; stems erect, 3-10 dm
tall, simple or branched above; leaves mainly
5-30 cm long, linear-subulate, the apex not
recurved; peduncles enlarged and fistulose
below the heads; involucres cylindric to
campanulate; bracts commonly 8 (5-11),
2.5-4 cm long in flower, 4-7 cm long in fruit;
rays purple, subequal to or shorter than the
bracts; achenes 25-35 mm long; pappus
brownish. Cultivated plants, escaping and
persisting on canal banks, in moist meadows,
and along roadsides at 1370 to 2595 m in
Carbon, Millard, Salt Lake, Sanpete, Summit,
and Weber counties; widespread in much of
the U.S.; introduced from Europe; 10 (0).

Tragopogon pratensis L. Biennial herbs;
stems erect, 1.5-8 dm tall, simple or
branched; leaves mainly 5-30 cm broad, ta-
pering from a broadly expanded base to 2 cm
wide, recurved apically; peduncles not espe-
cially enlarged in flower or in fruit; in-
volucres campanulate; bracts commonly 8,
12-24 mm long in flower, 18-38 mm long in
fruit; rays chrome-yellow, equaling or sur-
passing the bracts; achenes 15-25 mm long;
pappus off-white. Disturbed sites in Rich,
Salt Lake, and Summit counties; widespread
in the U.S.; adventive from Europe; 2 (0).

Vanclevea Greene

Shrubs; branchlets glutinous-resinous,
green to tan, finally white to gray barked;

leaves alternate, sessile, entire or serrate, fal-
cately curved; heads discoid, yellow, solitary
or cymose; involucres campanulate; bracts in
4 or 5 series, imbricate, glutinous; receptacle
naked, resinous; styles long-exserted, the
branches flattened, papillose; achenes cla-
vate, 5-angled; pappus of 12-16 linear per-
sistent slender scales.

Vanclevea stylosa (Eastw.) Greene [Grin-
delia stylosa Eastw., type from San Juan
County]. Shrubs, mainly 5-12 dm tall;
branchlets glutinous-resinous; bark tan to
white or grayish black in age; leaves 0.6-3.5
cm long, 1-9 mm wide, narrowly lanceolate
to oblong or elliptic, commonly entire, atten-
uate to a spinulose tip; heads solitary or more
commonly few to many in corymbose or cy-
mose clusters; involucres 8-10 mm high,
9-15 mm wide; bracts lanceolate to lance-at-
tenuate, sometimes abruptly acuminate and
recurved apically, resin coated; corollas yel-
low to cream, 6-7 mm long; achenes 4-5 mm
long, compressed, glutinous and spreading
hairy. Four-wing saltbush, ephedra, sand
dropseed, Indian ricegrass, blackbrush, and
juniper communities, in sand, at 1125 to 1620
m in Emery, Garfield, Grand, Kane, San
Juan, and Wayne counties; Arizona (a Colo-
rado Plateau endemic); 32 (viii). The genus is
monotypic.

Verbesina L.

Annual (biennial or perennial?) herbs;
leaves opposite, at least below, simple,
toothed; heads radiate, showy; involucres
biseriate, about equal, herbaceous; receptacle
convex, chaffy, the bracts enfolding the ach-
enes; rays yellow or yellow-orange, pistillate;
disk flowers perfect, fertile; anthers subentire
basally; style branches with hispidulous ap-
pendages; pappus of 2 slender awns; achenes
flattened, 2-winged.

Verbesina encelioides (Cav.) Benth. [Xime-
nesia encelioides Cav.]. Annual herbs; stems
4-10 dm tall, cinereous-strigose, often

354

Great Basin Naturalist

Vol. 43, No. 2

branched above; lowest leaves opposite, al-
ternate upward, petiolate, often with stipule-
like appendages at base; blades 1.2-10 cm
long, 0.7-6 cm wide, ovate to lanceolate,
acute to attenuate, irregularly toothed, stri-
gose beneath, green and sparingly strigose
above; involucres 7-12 mm high, 15-25 mm
wide; bracts lance-ovate to lance-linear, her-
baceous, strigose; rays 10-15, yellow or yel-
low-orange, 8-20 mm long; pappus of 2 short
slender awns; achenes thickly 2-winged, pu-
bescent. Sagebrush, rabbitbrush, saltgrass,
pinyon-juniper, and ponderosa pine commu-
nities, often in disturbed sites, at 1280 to
2260 m in Beaver, Garfield, Juab, Kane, San
Juan, and Washington counties; Montana to
California and Texas; 20 (v). Most of our ma-
terial belongs to var. exariculata Robins. &
Greenm. The bright flowers contrast sharply

with the grayish-strigose pubescence, result-
ing in a strikingly beautiful plant.

Wyethia Nutt.

Perennial herbs from thick taproots; stems
erect or ascending; leaves alternate, simple;
heads large, solitary or several, radiate; in-
volucral bracts in 2-4 series, herbaceous or
coriaceous; receptacle convex, chaffy, the
bracts folded, persistent; rays yellow, pistil-
late, fertile; disk flowers perfect, yellow;
pappus a crown of scales or lacking; achenes
trigonal or 4-angled, glabrous or pubescent.
Weber, W. A. 1946. A taxonomic and cy-
tological study of the genus Wyethia,
family Compositae, with notes on the
related genus BalsaTuorhiza. Amer.
Midi. Nat. 35:400-452.

1. Leaves mainly cauline, the basal reduced or lacking, scabrous-roughened;

plants of sandy desert shrublands W. scahra

— Leaves basal and cauline, the basal often larger than the cauline ones, smooth

or, if rough-hairy, not of lower elevations 2

2(1). Herbage glabrous, resinous; upper leaves rounded and clasping basally

W. amplexicaulis

— Herbage hirsute to glabrate; upper leaves petiolate W. arizonica

Wyethia amplexicaulis (Nutt.) Nutt.
Mulesears. [Espeletia amplexicaulis Nutt.].
Perennial herbs; stems mostly 2.5-9 dm tall,
glabrous; basal leaves 12-40 cm long, 2-15
cm wide, entire or dentate, petiolate, res-
inous; cauline leaves smaller, sessile, rounded
and clasping basally; heads large, solitary or
several; involucres hemispheric, 25-35 mm
high, 25-50 mm wide; outer bracts fo-
liaceous, subequal; rays 6-16, yellow, 2.5-4.5
cm long; pappus a crown, sometimes pro-
longed into filiform awns; achenes 8-10 mm
long, glabrous. Sagebrush, oak,, pinyon-juni-
per, aspen-fir, and forb-grass communities at
1525 to 2745 m in Box Elder, Cache, Juab,
Millard, Morgan, Salt Lake, Sanpete, Sevier,
Summit, Tooele, Utah, Weber, and Washing-
ton counties; Washington to Montana, south
to Nevada and Colorado; 38 (ii).

Wyethia arizonica Gray Perennial herbs;
stems mainly 30-80 cm tall, spreading hairy,
especially upward; basal leaves 15-40 cm
long or more, 3-15 cm wide, petiolate, the
blades oblanceolate to elliptic or lanceolate;

cauline leaves smaller, attenuate basally to a
short petiole; heads large, solitary or several;
involucres hemispheric or campanulate,
20-30 mm high, 15-40 mm wide; outer
bracts foliaceous, subequal; rays 6-16, yel-
low, 2.5-4 cm long; pappus a crown, some-
times prolonged into filiform awns; achenes
8-10 mm long, glabrous. Pinyon-juniper, oak,
and ponderosa pine communities at 1430 to
2440 m in Grand, Kane, San Juan, and Wash-
ington counties; Colorado, New Mexico, and
Arizona; 9 (0).

Wyethia scabra Hook. Robust, clump-
forming perennial herbs; stems several to
many, 1.5-6 dm high or more, scabrous and
hispidulose; leaves mainly cauline, the lower
ones rudimentary, 3-15 cm long, 3-17 mm
wide, elliptic to oblong or linear, scabrous;
heads solitary or few, terminating stems and
branches; involucres hemispheric, 20-40 mm
high, 20-55 mm wide; bracts lanceolate to
linear, attenuate to caudate-attenuate; rays
10-23, yellow, 18-40 mm long; pappus a
crown; achenes 6-8 mm long, glabrous.

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

355

Three more or less distinctive varieties are
present. Diagnostic features are based on the
nature of surface and habit of the involucral

bracts, which in the typical, common phase is
almost sufficiently variable as to include the
others.

1. Involucral bracts long-attenuate from short dilated bases, ciliate with multi-
cellular hairs, glabrous but with shiny resin droplets dorsally; plants of Kane
County W. scabra var. attenuata

— Involucral bracts variable in shape, ciliate or not, scabrous to pubescent and
more or less glandular dorsally, but seldom if ever with resin droplets 2

2(1). Involucral bracts closely imbricate, the outer recurved-spreading, pubescent

with short fine hairs; plants of San Juan, Grand, and eastern Kane counties

W. scabra var. canescens

— Involucral bracts various, scabrous to long-hairy dorsally; plants rather widely
distributed W. scabra var. scabra

Var. attenuata W. A. Weber Ponderosa
pine, oak, and pinyon-juniper (less commonly
in desert shrub) communities, in sand, at 1370
to 1985 m in Kane County (type from north
of Kanab); Arizona; 13 (iii). This handsome
plant is a botanical motif of the Coral Pink
dunes area, and is also present on East Clark
Bench.

Var. canescens W. A. Weber. Warm desert
shrub and mixed desert shrub communities at
1125 to 1680 m in Grand, Kane, and San
Juan counties; Colorado, Arizona, and New
Mexico; 4 (i). This is a variable entity transi-
tional to the typical variety, especially in
Grand and eastern Kane counties.

Var. scabra Bl^ckbrush, vanclevea-
ephedra, other mixed desert shrub, pinyon-
juniper, and ponderosa pine communities at
1220 to 2625 m in Carbon, Daggett, Du-
chesne, Emery, Garfield, Grand, Kane, and
Uintah counties; Wyoming; 48 (vii).

Xanthium L.

Aimual herbs with fleshy large cotyledons
and a taproot; leaves alternate, petiolate, the
blades broad, rough-hairy; heads unisexual,
discoid, or the corolla lacking; staminate
heads uppermost, many flowered; involucral
bracts in 1-3 series, separate; receptacle cy-
lindric, chaffy; fikments monadelphous, the
anthers separate; pistil vestigial, the styles
unbranched; involucre of pistillate heads en-
closing the 2 flowers, forming a 2-chambered
bur armed with hooked prickles, the corolla
lacking; achenes large, solitary in each cham-
ber; pappus none.

Xanthium strumarium L. Cocklebur. [X.
italicum Moretti; X. pensylvanicum Wallr.].
Annual monoecious herbs; stem 1.5-10 dm
tall or more, simple or branched, scabrous,
often purple mottled; leaves petiolate, the
blades mainly 2-12 cm long and about as
broad, ovate to oval or orbicular, obtuse to
cuneate or cordate basally, scabrous, dentate
and often lobed; heads in few to many short
axillary clusters; burs broadly cylindric to
ovoid, 1-3.5 cm long, with 2 more or less in-
curved beaks apically, covered with stout
hooked prickles. Weedy species of cultivated
and other disturbed lands, at 850 to 1925 m
in much of Utah; adventive (?) from the east-
ern U.S. or possibly from Europe; 33 (iii).
The seedlings are poisonous to livestock, and
they produce dermatitis in some people.

Xylorhiza Nutt.

Subshrubs or suffrutescent perennial herbs;
branchlets green to straw colored or whitish;
leaves alternate, simple; heads solitary at
branch ends; involucres campanulate to
hemispheric; bracts imbricate in several
series, herbaceous to largely scarious, erect;
ray flowers pistillate, fertile, yellow; achenes
somewhat compressed, hairy; pappus of taw-
ny to whitish capillary bristles. Note: Mem-
bers of this genus are all primary or second-
ary selenium indicators.

Cronquist, a. and D. D. Keck. 1957. A re-
constitution of the genus Machaeran-
thera. Brittonia 9:231-239.
Watson, T. J. 1977. The taxonomy of Xylor-
hiza (Asteraceae-Astereae). Brittonia
29:199-216.

356
1.

2(1).
3(2).

Great Basin Naturalist

Vol. 43, No. 2

4(2).

Leaves linear to linear-filiform, the margins entire and more or less involute;

plants of Kane and Garfield counties X. confertifolia

Leaves serrate to serrate-dentate, or, if entire, of other distribution (except X.
cronquistii) 2

Leaves serrate to serrate-dentate (at least some); plants of south central and
southwestern Utah, and of canyons of the Colorado 3

Leaves entire; plants of eastern Utah 4

Leaves only sparingly serrate, linear-oblanceolate to elliptic; involucral bracts
shortly attenuate, short-villous dorsally; plants of north central Kane County ....
X. cronquistii

Leaves sharply serrate-dentate, narrowly oblanceolate, elliptic, oblong, or lan-
ceolate; involucral bracts long-attenuate, glandular or villous-pilose dorsally;
plants of canyons of the Colorado and southwestern Utah X. tortifolia

Peduncles mainly less than 5 cm long; stems usually leafy to much above the

middle X. glabriuscula

Peduncles mainly more than 5 cm long; stems usually to the middle or below ...
X. venusta

Xyhrhiza confertifolia (Cronq.) T.J. Wat-
son [Macliaeranthera glabriuscula var. con-
fertifolia Cronq., type from NE of Henrie-
ville]. Perennial herbs from a woody caudex
and taproot, with rootstocks sometimes de-
veloped; stems 9-23 cm high, sparingly pi-
lose to glabrate and sparingly to densely
stipitate-glandular; leaves 1-4.5 cm long,
1-2.5 mm wide, linear, pilose to glabrate,
commonly involute; peduncles 1.8-14 cm
long; involucres 9-12 mm high, 12-18 mm
wide; bracts lanceolate to lance-acuminate,
pilose to glabrate and glandular; rays 4-12,
white, 9-18 mm long, 2-4 mm wide; disk
flowers yellow, the corollas 6-9 mm long;
pappus of capillary bristles to 6.5 mm long;
achenes 3.5-6 mm long, pubescent. Salt
desert shrub and pinyon-juniper communities
at 1675 to 1985 m in Garfield and Kane
counties; endemic; 6 (i).

Xylorhiza cronquistii Welsh & Atwood in
Welsh Cronquist Woody-aster. Subshrubs,
forming rounded clumps, from a stout tap-
root; stems numerous, whitish, ca 30 cm tall,
villous at the nodes, almost glabrous other-
wise; leaves 2.5-5 cm long, 2.5-5 mm wide,
linear-lanceolate, sparingly serrate-dentate to

entire, sparsely villous, ciliate, the midrib
prominent; heads solitary on branches; in-
volucre 10-12.5 mm high, 13-19 mm wide;
bracts oblanceolate to lance-attenuate, acute
to acuminate, herbaceous above the middle,
chartaceous below, short-villous and glandu-
lar dorsally; rays white, 14-16, 20-25 mm
long; achenes compressed, villous; pappus of
capillary bristles to 7.2 mm long. Pinyon-
juniper community, on the Kaiparowits For-
mation, at 1890 to 2075 m in Kane County;
endemic; 1 (0).

Xylorhiza glabriuscula Nutt. Subshrubs or
suffrutescent perennial herbs from a woody
caudex and taproot; stems 7-37 cm tall, vil-
lous to glabrous; leaves 1-7.5 cm long, 1-9
mm wide, villous to glabrate, lanceolate to
narrowly lanceolate or oblanceolate; heads
solitary at branch ends; involucres 9-13 mm
high, 15-27 mm wide; bracts lanceolate, at-
tenuate to acute or acuminate, herbaceous
above the middle, scarious below, villous to
glabrous; rays 10-22, white to bluish or
purplish, 11-20 mm long; achenes com-
pressed, villous; pappus of capillary bristles
to 5 mm long. Two allopatric varieties are
present.

Leaves with attenuate bases; rays white; plants of Daggett County

X. glabriuscula var. glabriuscula

Leaves with truncate or rounded bases; rays bluish, purplish, or white; plants

of San Juan Coimty X. glabriuscula var. linearifolia

April 1983

Welsh: Utah Flora: Compositae (Asteraceae)

357

Var. glabriuscula [Aster glabriuscula
(Nutt.) T. & G.; Machaeranthera glabriuscula
(Nutt.) Cronq. & Keck]. Salt and mixed desert
shrub communities at ca 1525 to 2135 m in
Daggett County; Colorado, Montana, South
Dakota, and Wyoming; 0 (0).

Var. linearifolia T.J. Watson Salt desert
shrub community, mainly on Chinle and
Moenkopi formations, in Grand and San Juan
counties; endemic; 3 (iii).

Xyhrhiza tortifolia (T. & G.) Greene Sub-
shrubs; stems 15-50 cm tall or more, villous
or tomentose and more or less stipitate-

glandular; leaves 1-10 cm long, 4-20 mm
wide, lanceolate to elliptic or oblanceolate,
villous to tomentose and glandular, spinulose-
dentate; heads terminating branches; in-
volucres mainly 12-20 mm high and 15-30
mm wide; bracts narrowly lance-attenuate to
-acuminate, herbaceous above, scarious be-
low; rays 17-60 or more, bluish or purplish
to white, 10-33 mm long, 1.8-5.5 mm wide;
pappus of capillary bristles to 9 mm long;
achenes compressed, pilose. Two varieties are
present.

1. Involucres merely glandular dorsally; plants of canyons of the Colorado

X. tortifolia var. imberbis

— Involucres villous-pilose as well as glandular; plants of Washington County

X. tortifolia var. tortifolia

Var. imberbis (Cronq.) T.J. Watson [Mach-
aeranthera tortifolia var. imberbis Cronq.].
Blackbrush, pinyon-juniper and sagebrush
communities at 1220 to 2290 m in Garfield,
Grand, Kane, San Juan, and Wayne counties;
Arizona (Colorado canyons endemic); 32
(viii).

Var. tortifolia [Haplopappus tortifolius T.
& G.; Aster abatus Blake; Machaeranthera
tortifolia (T. & G.) Cronq. & Keck]. Black-
brush and other warm desert shrub commu-
nities at 760 to 1010 m in Washington Coun-
ty; Arizona, Nevada, and California; 10 (i).

Xylorhiza venusta (Jones) Heller [Aster
venustus Jones, type from Cisco]. Suffrutes-
cent to herbaceous perennial herbs from a

woody caudex and taproot; stems mainly
10-40 cm tall, glabrous to densely pilose;
leaves 2.4-9 cm long, 2-17 mm wide, oblan-
ceolate to spatulate, villous to glabrate, at-
tenuate basally; heads terminating branches;
peduncles 5-20 cm long; involucres 10-18
mm high, 18-50 mm wide; bracts lance-at-
tenuate to caudate-acuminate, herbaceous
above, scarious below; rays 12-36, white or
bluish to purplish, 12-27 mm long; pappus
bristles to 10 mm long; achenes sericeus. Salt
desert shrub communities at 1250 to 1985 m
in Carbon, Daggett, Emery, Garfield, Grand,
San Juan, Uintah, and Wayne counties; Colo-
rado (a Colorado Plateau endemic); 99 (xv).

HAPLOPAPPUS CRISPUS AND H. ZIONIS (ASTERACEAE):
NEW SPECIES FROM UTAH

Loran C. Anderson'

Abstract. — The new species, Haplopappus crispus and H. zionis of section Macronema, are formally described
and illustrated. They are endemic to southern Utah. Also, H. bloomeri ssp. compactus is raised to species. Chromo-
.some numbers of all three are n = 9. Aspects of anatomy are detailed. Comparisons are made to H. bloomeri and H.
suffntticosus. Relationships are discussed, and a key to the species is given.

The only comprehensive monograph of
Haplopappus is that of H. M. Hall (1928). In
recent years the generic integrity of Hap-
lopappus has been questioned (see Anderson
1980 for review). Data from anatomy, cy-
tology, and chemistry suggest it is a poly-
phyletic assemblage. Nevertheless, a suitable
taxonomic reorganization of the group has
not been achieved. Therefore, I choose to de-
scribe new taxa under the name Haplopappus
even though the species will very probably
be placed in some other genus at a later date.

Some years ago I found plants in southern
Utah that appeared to be H. bloomeri Gray
ssp. compactus Hall; a chromosome count for
the collection, Anderson 3358, was published
under that name (Anderson et al. 1974).
Other collectors had identified similar plants
as H. suffruticosus (Nutt.) Gray. The com-
bined collections actually represent two new
species of section Macronema, and, addition-
ally, H. bloomeri ssp. compactus should be
elevated to species level.

Methods and Materials

Fresh and dried materials were processed
for anatomical study as in Anderson (1970a).
Five heads from personal collections, along
with one to five heads from other collections,
were measured (as in Anderson 1964) for in-
volucral and floral data. Cytological methods
are those of Anderson (1966).

Voucher specimens for anatomical (a) and
morphological (m) studies are: H. bloomeri:
Anderson 1620, m (FSU), Anderson 2018, a
and m (FSU), Anderson 2943, a (FSU), Ander-
son 4539, m (FSU), Reveal 1070, m (FSU); H.

compactus: Ackerman 30797, m (FSU), An-
derson 6186, a and m (FSU), Clokey 8570, m
(UTC); H. crispus: Anderson 5504, a and m
(FSU), Cottam 1526, m (BRY), Maguire
13386, m (UTC), Stanton in 1927, m (BRY);
H. suffruticosus: Anderson 1023, m (FSU),
Anderson 2920, a (FSU), Anderson 2970, a
and m (FSU), Goodrich 10133, m (BRY),
Shultz 3738, m (UTC), Yoder-Williams 1311,
a (FSU), Wiggins 9298, m (UTC); and H. zi-
onis: Anderson 3358, m (FSU), Anderson et
al. 5094, a and m (FSU), Arnow 107, m (UT).

Taxonomy

Haplopappus compactus (Hall) L. C. Ander-
son, comb. nov.

Basionym: Haplopappus bloomeri Gray ssp.
compactus Hall. Carnegie Inst. Publ.
389:199, fig. 68, 1928. Type. - Nevada:
Clark Co., Charleston [Spring] Mountains, E.
C. Jaeger on 12 Sep 1925 (holotype: POM!;
isotype: UC!).

Synonymy: Haplopappus bloomeri Gray
var. compactus (Hall) Blake in Clokey. Univ.
Calif. Publ. Bot. 24:231. 1951.

The only description of this taxon is that
given by Hall (1928). It is amplified and
emended with the following: woody shrubs to
5 dm tall; leaves oblanceolate-spatulate,
2-3.5(4) cm long, (2)2.5-3(5) mm wide; heads
(involucres) 12-14.8 mm long, 4.5-5.5 mm
wide, phyllaries 18-24; disk flowers 11-16,
golden yellow (fading lighter), corollas
9.1-11.4 mm long, tubes glandular, strongly
dilated at point of staminal departure (at 50
percent of total corolla length), lobes 0.8-1.6
mm long, styles 12.5-14.5 mm long.

'Department of Biological Science, Florida State University, Tallahassee, Florida 32306.

358

April 1983

Anderson: New Haplopappus

359

In addition to specimens cited by Hall
(1928) and Clokey (1951), the following rep-
resent this species: NEVADA: Clark Co.,
above McWilliams Campground at 8500 ft,
Lee Canyon, Spring Mountains, L. C. Ander-
son 6186 (BRY, FSU, UNLV, UTC), A. D.
Blauer & £. D. McArthur N-144, N-145 (FSU,
SSLP), W. E. Niles 3160 (FSU, UNLV); top
of ridge, Charleston Mountain, 11,500 ft, T.
Dawson 7 (UC); top of Sheep Mountains,
9300 ft, T. L. Ackemian 30797 (FSU, NY).

The last collection is a range extension for
H. compactus; the taxon was previously
thought to be endemic to the Spring (or
Charleston) Mountains.

Haplopappus crispus L. C. Anderson, sp.
nov.

Frutices lignosi 3-4(5) dm alti; caules fo-
liosi, graciles, erecti, tenuiter glandulosi; folia
spatulata vel oblanceolata et acuminata, mar-
ginibus undulatis et crispis, (1.5)2-2.5(30) cm
longa, (3)5-6(8) mm lata, in ramis altioribus
solum parvo minora; inflorescentia cyma so-
lute paniculata vel aliquanto congesta, tem-
poribus paucis solum uno capitulo vel duobus
per ramum; capitula campanulata,
(12.5)13-14(15) mm longa, (5)6.5-7(9) mm
lata, foliis superioribus saepissime separata;
phyllaria (24)26-30(35); nuUi radii flosculi;
disci flosculi (14)15-20(24), luridi, coroUis
(9.5)10-10.8 mm longis, tubulo aliquanto di-
lato, lobis 1-1.5 mm longis, stylis 14-18 mm
longis, lineis stigmaticis multo brevioribus
quam appendicibus; achenia 6.5-8.5 mm
longa et raro pubescentia.

Type.— Utah: V^ashington Co., weathered
andesite with manzanita in mountain mahog-
any-fir woods with few pine and aspen along
Whipple Valley Trail at 8100 ft, above Pine
Valley, 19 air mi NE of St. George in Pine
Valley Mountains, T39S, R14W, El/2 Sec
29, 18 Sep 1981, L. C. Anderson 5504 (holo-
type: BRY; isotypes: FSU, MO, NY, RSA,
UC, UTC).

Woody shrubs, much branched at base,
3-4(5) dm tall (or taller?); leafy stems slender,
erect, covered with short-stalked glands;
leaves entire, alternate, green, spatulate to
oblong-oblanceolate, acuminate, margins
wavy-crispate, glutinous with low glands (not
prominently stalked as in H. suffruticosus),
(1.5)2-2.5(3) cm long, (3)5-6(8) mm wide.

Fig. 1. Haplopappus crispus (Anderson 5504): a.,
flowering branch with crispate leaves and rather naked
peduncles; b., involucral bracts, outermost with large
green tips, inner ones with narrowed tips, innermost
without green tips; c, disk flower with gradually flaring
corolla and sparsely strigose achenes. A., x 1; b. and c, x
4.

only slightly reduced toward the in-
florescence, but generally not crowding the
heads as in H. zionis; inflorescence a loosely
paniculate to somewhat congested cyme, oc-
casionally reduced to one or two heads per
branch (Fig. la); heads campanulate,
(12.5)13-14(15) mm long, (5)6.5-7(9) mm
wide; phyllaries (24)26-30(35), 1-nerved,
finely glandular, outermost bracts mostly
green, leaflike, inner bracts stramineous,
abruptly or gradually narrowed into a slender
green tip, innermost bracts stramineous,
erose-ciliate apically, but not as pronounced
as in H. zionis; ray flowers absent; disk flow-
ers (14)15-20(24), pale yellow, corollas
(9.5)10-10.8 mm long, tubes glandular pu-
bescent, not abruptly dilated as in H. com-
pactus and H. zionis, lobes 1-1.5 mm long,

360

Great Basin Naturalist

Vol. 43, No. 2

styles 14-18 mm long, branches slender, stig-
matic lines much shorter than appendages
(22-36 percent of total branch length);
achenes cylindric, 6.5-8.5 mm long, sparsely
but evenly pubescent, pappus 8-9.5 mm
long; n = 9. Infrequent, generally in moder-
ately open settings in spruce-pine associ-
ations, (3000?) 8100-10,000 ft, apparently en-
demic to mountains of southwestern Utah.
August- September .

Additional specimens examined.— Utah:
Millard Co., Pine Valley, W. D. Stanton in
1927 (BRY); Washington Co., south end Pine
Valley Mountains, 9200 ft, B. Albee 2911a
(UT), Anderson Valley area, north end Pine
Valley Mountains, 8400 ft, B. Albee 2911b
(UT), Pine Valley Mountains, T39S, R14W, /.
L. Gentry & E. Jensen, Jr., 2245 (BRY, NY,
UTC), R. K. Gierisch 542 (UC, UTC), Pine
Valley Mountains, 8500 ft, W. P. Cottam
5699 (UT), 10,000 ft, B. Maguire & B. L.
Richards, Jr., 13386 (UC, UTC), Santa Clara,
3000 ft, W. P. Cottam 1526 (BRY, NY, UT).
The last collection is so far out of range alti-
tudinally that it may represent a chance in-
troduction; the population has not been
found again to determine its persistence. The
collection is of further interest for the nota-
tion "along ditch banks, 8-10 ft tall"; also, it
has heads with greater numbers of phyllaries
and flowers than the other collections.
Haplopappus zionis L. C. Anderson, sp. nov.

Frutices humiles et diffundentes 1-3 dm
alti; corpulenti caules decumbentes vel as-
cendentes, foliosi in inflorescentiam et
glandulosi-hispiduli; folia spatulata et acumi-
nata, (2.5)3-3.5(4) cm longa, 2.4-4(7) mm
lata; inflorescentia cyma paniculata et con-
gesta et foliosa; capitula turbinata vel ang-
uste campanulata, (14)16-19(22) mm longa,
6-8 mm lata, bracteis exterioribus in forma
folii apicibus longis et acuminatis, interior-
ibus apice villosis, phyllariis (17)20-22(23);
nulli radii flosculi; disci flosculi
(10)12-18(21), flavi, corollis (9.5)9.8-
10.8(11.5) mm longis, tubulis subito dilatis,
lobis 1.2-2 mm longis, stylis 18-19 mm long-
is, lineis stigmaticis multo brevioribus quam
appendicibus; achenia 7-8 mm longa et
ferme glabra.

Type.— Utah: Iron Co., moderately bare,
weathered pink limestone member of
Wasatch Formation in aspen, spruce, limber

pine, bristlecone pine association near top
of Cedar Canyon, 9800 ft, 13.5 air mi SE of
Cedar City, 30 Aug 1980, L. C. Anderson, S.
L. Welsh, and M. Chatterley 5094 (holotype:
BRY; isotypes: ASU, DS, FSU, MO, NY, RSA,
UC, UTC).

Low, branching, spreading shrubs, 1-3 dm
tall; stems stout, decumbent or ascending,
glandular-hispidulous, leafy into the in-
florescence (Fig. 2a); leaves entire, alternate,
green, spatulate, acuminate, margins not un-
dulate or crispate, glandular but less glu-
tinous than H. crispus, (2.5)3-3.5(4) cm long,
2.5-4(7) mm wide; inflorescence a congested,
leafy paniculate cyme; heads turbinate to
narrowly campanulate, (14)16-19(22) mm
long, 6-8 mm wide, lateral heads fewer flow-
ered than central ones; phyllaries
(17)20-22(23), 3-nerved, finely glandular,
outermost bracts leaflike and much longer
than other bracts, inner bracts stramineous
with long green tips, innermost bracts stra-
mineous with praemorse, prominently villous
tips (Fig. 2b); ray flowers absent; disk flowers
(10)12-18(21), golden yellow, corollas
(9.5)9.8-10.8(12) mm long, tubes abruptly di-
lated at point of staminal departure (at 40
percent of total corolla length), sparsely
glandular, lobes 1.2-2 mm long, styles 18-19
mm long, branches slender, stigmatic lines
much shorter than appendages (23-31 per-
cent of total branch length); achenes cylin-
dric, 7-8 mm long, essentially glabrous with
few hairs apically (Fig. 2c), pappus 9-9.5 mm
long; n = 9. Infrequent, usually on gravelly
sandy clay from limestones, often with man-
zanita on rather barren slopes in spruce, fir,
pine associations, 8,000-10,000 ft, apparently
endemic to mountains of southern Utah. July-
September.

Additional specimens examined: Utah,
Garfield County, vie. summit between Esca-
lante and Widtsoe, Escalante Mountains,
9000 ft, W. P. Cottam 6562 (UT), 9100 ft, L.
C. Anderson 3358 (BRY, FSU, KSC, NY, UC,
UTC), 10,000 ft, E. Neese & S. White 3964
(BRY), 2 miles south of Pine Lake, Escalante
Moimtains, E. Neese & S. White 3844; Iron
County, Cedar Canyon, southeast of Cedar
City, 8000 ft, L. Arnow 107 (UT), R. Foster
5297 (BRY), vie. Midway Summit, west of
Cedar Breaks, 10,000 ft, B. Maguire & B. L.

April 1983

Anderson: New Haplopappus

361

Fig. 2. Haplopappus zionis (Anderson et al. 5094): a.,
flowering branch with foHage overtopping flowering
heads; b., involucral bracts, outermost leaflike and 3-
nerved, inner ones with elongate green tips, innermost
with prominently villous tip; c, disk flower with abrupt-
ly dilated corolla tube, long style, and nearly glabrous
achene with very few hairs just below pappus, a., x 1; b.
and c, X 4.

Richards, Jr. 13389 (UTC); Co. unknown,
Dixie National Forest, R. K. Gierisch 217

(UTC).

Cytology and Anatomy

Chromosome numbers of section Mac-
ronema of Haplopappus are monotonously
uniform at n = 9 with an apparent absence
of polyploidy and aneuploidy (Anderson et
al. 1974). The new species are also n = 9.
Original counts for H. compactus and H.
crispus were made from root tip squashes
from Niles 3160 and Anderson 5504, respec-
tively. An earlier count for H. zionis (Ander-
son 3358) was listed as H. bloomeri ssp. com-
pactus in Anderson et al. (1974).

Meiotic behavior has not been observed for
any of the new species, but all three have
pollen fertilities above 98 percent (as deter-
mined by staining in cotton blue in lactophe-
nol). Embryo sac development appears nor-
mal (Polygonum type) in H. compactus, H.
crispus, and H. suffruticosus; it was not stud-
ied in H. bloomeri and H. zionis. Mature em-
bryo sacs are long and narrow (280-380 jam
long) and 8-nucleate without multiplication
of antipodals as found in many Chryso-
thamnus (Anderson 1970b).

Aspects of floral anatomy for the new spe-
cies and possibly related taxa are summarized
in Table 1 (following format in Anderson
1970a); frequency categories are: -I- + , abun-
dant; -I- , frequent; -, rare; and 0, absent.
General information on the other taxa of
Haplopappus is in Nelson (1982). Achenes of
H. zionis have the greatest amount of vas-
culature, whereas those of H. crispus have
the least. Secretory canals vary from abun-
dant to absent at selected levels in the ach-
enes and corollas. They are always absent in
the styles of these species but present in H.
macronema (Anderson 1970a).

Achenes of H. bloomeri, H. compactus, and
H. zionis are essentially glabrous but with a
few twin hairs just below the pappus attach-
ment (the last being the "most glabrous"). A
few short glandular trichomes (60-66 jum
long) are hidden among the abundant twin
hairs on H. suffruticosus achenes, and larger
glandular hairs (190 jum) occur rarely on
achenes of H. bloomeri. Glandular trichomes
on achenes were so rare that they were
omitted from Table 1, and they do not occur
in the new species. Pappus bristles are posi-
tioned in three closely spaced rows on ach-
enes of H. zionis and in two in the other four

362

Great Basin

Naturalist

Vol. 43, No. 2

Table 1. Comparisons for range and aspects of morphology of selected taxa oi

F Haplopappus

, section Macronema.

H. bloomeri

H. compactui

( H. crispus

H. suffruticosus

H. zionis

Range

Calif., W Nev.,

SNev.

SW Utah

Calif., Nev., ]

[daho.

SUtah

Oregon, S Wash.

W Wyo., SW Mont

Elevation, ft

3500-9600

7900-11,500

(3000)8100-10,000

7500-12,300

8000-10,000

Habit of growth

tall, 4-5 dm

medium, 3-4 dm

low, 1-3 dm

Leaf size, cm x mm

2-6 X 0.5-3

2-4 X 2-5

1.5-3 X 3-8

1-3 X 1.5-5

2.5-4 X 2.5-7

Leaf shape

filiform-narrowly
oblanceolate

oblanceolate-
spatulate

■ spatulate-
oblanceolate

oblong-spati

ilate

spatulate

Leaf margins

smooth

crispate

smooth

Head length, mm

7.8-12.2(16)

12-15

12.5-15

8-16

14-22

Head width /length ratio .29- .35

.36-.40

.45-.48

.45-.60

.38-.41

Phyllary number

17-36

18-24

24-35

17-31

17-23

Ray flower number

(0)1-3

(0)1-8

Disk flower number

4-8(12)

11-16

14-24

15-40

10-21

Disk flower length, mm

7.3-9.4(10.9)

9.1-11.4

9.5-10.8

8.5-11

9.5-12

Corolla tube

straight

dilated

± dilated

straight

dilated

Corolla lobe length, mm 0.9-1.9

0.8-1.6

1.0-1.5

1.0-1.7

1.2-2.0

Style branch, mm

2.5-4.3

3.8-4.4

3.8-4.8

2.6-4.6

4.2-5.2

Achene length, mm

6-7

7.5-8.5

6.5-8.5

7.5-8

7-8

Achene pubescence

glabrous

sparsely strigose

villous-sericeus

glabrous

species (contrary to the meaning of the ge-
neric name).

The five species have glandular corolla
tubes with the trichomes, "g" in Table 1,
being similar to those in Chrysothamnus (fig.
20-22, Anderson 1970a); however, those in
H. zionis are less abimdant, are longer, and
have narrower glandular heads as in Figure
19. Additionally, corollas of H. crispus have a
few very long, nonglandular villi like Fig. 17
of Anderson (1970a).

Nectaries at the base of the style are most
prominent in H. zionis and least so in H.
crispus. The new species have somewhat
thicker corollas than do H. bloomeri and H.
suffruticosus as determined by cell number
radially through the tubes between adjacent
vascular bundles.

Nodal anatomy is trilacunar, three trace
for the species. In leaf anatomy, they are sim-
ilar in that leaves are isolateral with three
rows of palisade cells facing each epidermis.
Bundle sheath extensions are present only on

the midvein— a feature characteristic of all
taxa of Haplopappus with a chromosome
base of X = 9, whereas those with x = 4, 5,
or 6 have leaves with bimdle sheath exten-
sions on lateral veins as well. A possible ex-
ception would be H. parryi, (x = 9 group),
which has many bundle sheath extensions on
bifacial leaves, but it is better placed in Sol-
idago (Anderson and Creech 1975).

Some variation exists in leaf thickness. Of
leaves studied, those of H. compactus and H.
zionis have blades averaging 336 jum thick
with midveins 480 and 350 jum, respectively.
Leaves of H. crispus are thicker, 355 /xm in
the blade and 400 at the midvein. Much thin-
ner leaves occur in H. suffruticosus with
blades 202 and 250 {Anderson 2970 and 2920,
respectively), and both with midveins about
260 jLim thick.

Glandular trichomes on leaves of H. com-
pactus and H. crispus average 77 jum long
with nearly spherical heads 50 jum and 38 /xm
wide, respectively. Glands on H. zionis

Table 2. Features of floral anatomy for selected taxa of Haplopappus, section Macronema.

Ovarian bundle
number

Secretory

canal distribution

Achene

Corolla

Taxon

Average

Range

III

H. bloomeri
H. compactus
H. crispm

H. suffruticosus
H. zionis

5.5
5.5
5.0

5.2
7.0

5-6

5-6
5-9

+
-1- +
+ +

+
+

+ -1-
+ +

+ +

-1-
+ +

+
+ +

+ +

April 1983

Anderson: New Haplopappus

363

leaves, thoughsomewhat less abundant, have
larger buttressed bases and average 100 /xm
long with heads 38 jum wide. For H. suffruti-
cosus, glands on Anderson 2970 leaves aver-
age 80 jum long with spherical heads, whereas
those on Anderson 2920 have long slender
stalks and heads, averaging 220 jum long with
the elongate heads 48 jum long and 29 jum
wide. Glands on H. hhomeri are nearly ses-
sile, only 38-42 ju,ni long with spherical heads
29 jLim in diameter.

Relationships

Table 2 gives ranges of the species and
summarizes some aspects of their morphol-
ogy based on my observations; some mea-
surements differ from those of Hall (1928).
Reveal 1070 is an unusual form of H. bhom-
eri, with exceptionally large heads with many
long disk flowers (features listed paren-
thetically in Table 2).

The new species do not occur close geo-
graphically to H. hhomeri or H. suffruti-
cosus, species with which they have been
confused. Further, the new species are dis-
tinct from the other two in their constant ab-
sence of ray flowers. Ray flowers are almost

always present in heads of H. bloomeri and H.
suffruticosus, and, although occasionally in-
dividual plants may be eradiate, some plants
of the population will always have rays.

Haplopappus compactus, once considered
a subspecies of H. bloomeri, differs from that
species in involucral length and width, disk
flower number, corolla shape, and achene
size as well as the absence of ray flowers and
some aspects of leaf shape and size. Hap-
lopappus compactus and H. zionis appear to
be closely related. Both have dilated corolla
tubes. They can be distinguished in habit,
head size, and style length, and generally so
in leaf shape and size. In addition, H. zionis
usually has leafier involucres. The two also
differ anatomically.

Haplopappus crispus is intermediate geo-
graphically between H. compactus and H. zi-
onis, but it is closer to H. suffruticosus in
relationship even though it is eradiate. In ad-
dition to absence of rays, H. crispus can be
further distinguished from H. suffruticosus
by its habit, leaf size and thickness, degree of
glandularity (and odor), narrower heads, few-
er flowers, and sparsely strigose achenes.

A key to the new species and related taxa
is presented here.

1. Ray flowers present (at least in the population); widespread but not in S

Nevada or S Utah 2

— Ray flowers always absent in each head; plants of S Nevada or S Utah 3

2(1). Achenes densely pubescent (sericeus); heads campanulate with more than 15

flowers H. suffruticosus

— Achenes essentially glabrous; heads cylindric-turbinate with less than 15
flowers H. bloomeri

3(1). Achenes sparsely strigose with hairs distributed evenly; leaves crispate .. H. crispus

— Achenes essentially glabrous but with few hairs near the pappus; leaves not
crispate 4

Table 2 continued.

Trichomes

Corolla
thickness

Nectary
length, jum

Achene

Corolla tube

Index of

Freq.

Length, mm

Type

Freq.

Length,

jLtm

specialization

144

0.5

+ +

150

6.5

168

0.43

-1-

168

5.6

115

0.53

g
n

-1- -1-

165

432

6.8

140

+ +

0.8

+ +

160

6.9

240

0.52

270

5.0

364

4(3).

Great Basin Naturalist

Vol. 43, No. 2

Heads mostly less than 14 mm long; styles less than 15 mm long; woody

shrubs 3-5 dm tall H. compactus

Heads mostly more than 14 mm long; styles over 17 mm long; low shrubs

1-3 dm tall H. zionis

In the Asteraceae, amounts of secretory tis-
sue and floral venation bear evolutionary sig-
nificance with a reduction in both usually in-
dicating advancement. In Chrysothamnus,
these features (correlated with other charac-
teristics such as karyotypes) were used to de-
velop a phylogenetic index of specialization
on a scale of 0-10 for the taxa (Anderson and
Fisher 1970).

Section Macronema is close to Chryso-
thamnus with natural hybridization known to
occur between H. macronema and C. nau-
seosus (Anderson and Reveal 1966). There-
fore, the index of specialization developed
for Chrysothamnus should be applicable for
showing relationships among the new species.
Index values are listed in Table 2. Reduction
in involucral bract vasculature is apparently
correlated with that of the ovary wall; phyl-
laries of H. zionis have three vascular bundles
each; those of H. crispus have one.

Of the taxa studied here and in Anderson
(1970a), H. macronema (the most widespread
species in the section) has the lowest index
number of 4.5, and H. ophitidus (clearly a de-
rived species that is specialized for serpen-
tine) has the highest at 8.5. Hall (1928) con-
sidered H. suffruticosus the most primitive
member of section Macronema, but H. mac-
ronema is probably a better candidate. Hap-
hpappus compactus, and, especially, H. zion-
is, though much more restricted in range
than H. bloomeri and H. suffruticosus, seem-
ingly represent more primitive stock
phy logenetically .

Acknowledgments

Appreciation is expressed to Stanley L.
Welsh for field trip support and to Wesley

Niles for seed of H. compactus. Melanie
Darst prepared the line drawings; Walter
Forehand is thanked for assistance with the
Latin diagnoses. This study was supported by
National Science Foundation Grant DEB
76-10768.

Literature Cited

Anderson, L. C. 1964. Taxonomic notes on the Chryso-
thamnus viscidiflorus complex (Astereae, Com-
positae). Madrono 17:222-227.

1966. Cytotaxonomic studies in Chrysothamnus

(Astereae, Compositae). Amer. J. Bot. 53:
204-212.

1970a. Floral anatomy of Chrysothamnus (As-
tereae, Compositae). Sida 3:466-503.

1970b. Embryology of Chrysothamnus (Astereae,

Compositae). Madroiio 20:337-342.

1980. Haplopappus alpinus (Asteraceae): a new

endemic species from Nevada. Great Basin Nat.
40:73-77.

Anderson, L. C, and J. B. Creech. 1975. Comparative
leaf anatomy of Solidago and related Asteraceae.
Amer. J. Bot. 62:486-493.

Anderson, L. C, and P. S. Fisher. 1970. Phylogenetic
indicators from floral anatomy in Chrysothamnus
(Astereae, Compositae). Phytomorph. 20:
112-118.

Anderson, L. C, and J. L. Reveal. 1966. Chryso-
thamnus bolanderi, an intergeneric hybrid. Ma-
drono 18:225-233.

Anderson, L. C, D. W. Kynos, T. Mosquin, A. M.
Powell, and P. H. Raven. 1974. Chromosome
numbers in Compositae. IX. Haplopappus and
other Astereae. Amer. J. Bot. 61:665-671.

Clokey, I. W. 1951. Flora of the Charleston Mountains,
Clark County, Nevada. Univ. California Publ.
Bot. 24:1-274.

Hall, H. M. 1928. The genus Haplopappus, a phyloge-
netic study in the Compositae. Carnegie Inst.
Publ. 389:1-391.

Nelson, J. B. 1982. Floral anatomy of North American
Haplopappus (Asteraceae): systematic consid-
erations. Unpublished dissertation. Florida State
Univ. Library, Tallahassee.

A BOUQUET OF DAISIES (ERIGERON, COMPOSITAE)

Stanley L. Welsh'

Abstract. — Described as new are six critical species and one variety of Erigeron from Utah. They are E. awa-
pensis Welsh, £. canaani Welsh, E. carringtonae Welsh, £. goodrichii Welsh, E. maguirei Cronq. var. harrisonii
Welsh, E. tintemuinnii Welsh and Goodrich, and E. wahwahensis Welsh.

Erigeron is perhaps the most complex of all
genera within the Compositae of Utah. The
genus is allied to both Aster and Conyza, but
most of the problems lie within the genus it-
self. To make matters more difficult Erigeron
is one of the largest genera in Utah, with
more than 60 taxa. Morphological features
used as diagnostic characters often involve
pubescence and its position on the plants.
Other vegetative features have been relied
upon also, because of the uniformity of flow-
ers and bracts. Shape of leaves, whether
mainly basal, mainly cauline, or distributed
along the stem constitute important diagnos-
tic characters. Pubescence type on the in-
volucral bracts is considered as diagnostic in
many cases. The technical nature of the dis-
tinguishing characters have led to wide mis-
understanding of the genus, and most bota-
nists have avoided the members of the group,
relying, where possible, on specialists for crit-
ical determinations.

Preparation of a manuscript of the Com-
positae for the Utah flora project has led me
to study Erigerons of Utah and the surround-
ing states in some detail. That study has in-
dicated the presence of several taxa whose
descriptions appear to be beyond those pre-
viously known to occur within Utah. The
specimens have been compared to all other
materials at BRY, and duplicates of many
have been examined by Arthur Cronquist at
NY. I wish to acknowledge his suggestions
and consideration, but wish not to share
blame for problems created by this author
alone in the following interpretations.

Erigeron awapensis Welsh, sp. nov. A E.
abajoensis Cronq. in caulis erectis foliorum

basalis non vaginatis et floribus plus numer-
osis differt.

Perennial herbs from a branching caudex,
the caudex branches clothed with ragged
brown marcescent leaf bases; stems erect or
nearly so, 10-24 cm long, strigose, the hairs
ascending; basal leaves 1.5-7 cm long, 2-8
mm wide, not especially sheathing; cauline
leaves well developed, oblong to linear, most-
ly 1-4 cm long, 2-4 mm wide; heads 2-4,
rarely solitary; involucres 3-9 mm wide,
3.7-4.5 mm high, the bracts more or less im-
bricate, thickened near the base dorsally,
greenish, strigulose, the hairs multicellular;
rays 35-45, pink purple to pink (or white?),
5-6 mm long, 0.9-1.8 mm wide; pappus ap-
parently simple, of 15-20 slender bristles,
and with a few inconspicuous shorter setae in
some; achenes 2-nerved, hairy. Pinyon-juni-
per and sagebrush communities at 2135 to
2260 m in Garfield and Wayne counties;
endemic.

Type.- USA. Utah. Garfield County,
T32S, R2W, S23, igneous bouldery slope in
canyon, in Artemisia community, at milepost
26.5 south of Antimony, 4 September 1969,
S. L. & S. L. Welsh 9388 (Holotype BRY; Iso-
types, 4 distributed previously as E. divergens
T. & G.). Additional specimen: Utah. Wayne
County, T29 S, R4E, S20, 1 mi SE of Teas-
dale, sandy slope in pinyon-juniper commu-
nity, 7 August 1980, M. E. Lewis 6657 (BRY).

The Awapa daisy is most similar to E. aba-
joensis, but stands apart from that taxon,
which has become a catchall for all speci-
mens that run to the end of the key. It seems
apparent that the assemblage will be clarified
only upon examination of much additional
material not now available in collections.

'Life Science Museum and Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602.

365

366

Great Basin Naturalist

Vol. 43, No. 2

Erigeron canaani Welsh, var. nov. Similis
E. eatonii Gray in aspectum et capitulum
amplitudem sed in foliis caulino et radi-
calibus ambo linearibus et involutis folio
basalibus ampliato et ciliato et radiis paucior-
ibus differt.

Perennial herbs from a simple (or
branched?) caudex, this clothed with brown
marcescent leaf bases, the taproot prominent;
stems 7-20 cm tall, decumbent to ascending,
sometimes pm-plish at the base, sparingly
hairy with ascending hairs; leaves pubescent
Uke the stem, the basal ones tufted, 1 -nerved,
1.4-9 cm long, 0.7-1 mm wide, linear, in-
volute, sharply acute, conspicuously ex-
panded and long-ciliate basally; cauline
leaves numerous, reduced upwards; heads
1-3; involucres 9-13 mm wide, 5.3-6.5 mm
high; bracts imbricate, conspicuously glandu-
lar and sparingly to moderately villous-pilose
with multicellular hairs, green or variously
suffused with purple; rays 15-22, white or
pinkish, 3.5-5 mm long, 1.8-2.1 mm wide;
pappus single, of ca 20 slender bristles;
achenes 2-nerved, hairy. Ponderosa pine
community at 1585 to 2075 m in Washington
County; endemic.

Type.— USA. Utah. Washington County,
with Petrophytum and Panicum, in crevices
in Navajo Sandstone, Canaan Mountain,
T43S, RlOW, S4, 11 June 1980, John Ander-
son sn (Holotype BRY; Isotype NY). Addi-
tional specimen: Utah. Washington County,
Zion National Park, Clear Creek Canyon,
T41S, RlOW, S24, ponderosa pine commu-
nity, on Navajo Sandstone, 25 Sept. 1982, S.
L. Welsh 21398 (BRY).

The Canaan daisy is similar in general as-
pect to E. eatonii. The involute linear
glaucous leaves and few ray flowers appear
to be diagnostic.

Erigeron carringtonae Welsh, sp. nov. Af-
finis E. untermannii in foliis crassis sed in
pilis involucrorum longioribus et tenuioribus
et radiis longioribus differt.

Pulvinate perennial herbs with a plurici-
pital caudex, the branches clothed with con-
spicuous brown to straw colored or ashy mar-
cescent leaf bases; leaves mainly basal,
thickish, 0.6-3.5 cm long, 1-5 mm wide,
spatulate to oblanceolate, strigose to pilosu-
lose, obtuse to rounded apically; scapes 2.5-8
cm tall; heads solitary; involucres 9.8-15 mm

wide, 5.8-7 mm high, the bracts imbricate,
suffused with purple or green, the inner
greenish with scarious margins, spreading-vil-
lous with long multicellular hairs; rays 18-30,
pink to pink purple, 6.8-8.2 mm long,
1.4-2.3 mm wide; pappus double, the inner
of 25-35 barbellate bristles, the outer of short
setae; achenes 2-nerved, pilose. Meadows and
escarpment margins, commonly on Flagstaff
Limestone at 3050 to 3355 m in Emery and
Sanpete counties; endemic.

Type.— USA. Utah. Sanpete County, T18S,
R4W, S33, Manti-La Sal NF, 10 mi 115 de-
grees from Manti, 3220 m, 3 July 1981, S.
Goodrich 15780 (Holotype BRY; Isotypes 14,
distributed as E. simplex). Additional speci-
mens: Utah. Sanpete County. T20S, R4E,
S33, ca 17 mi W of Ferron, 3355 m. Flagstaff
Limestone barrens, 13 July 1977, S. L. Welsh
and S. Clark 15393 (BRY); do, T18S, R4E,
S22, 7 mi due NNW of Ferron Reservoir, 19
July 1977, E. Neese and S. White 3708
(BRY); do. Heliotrope Mountain, 13 July
1977, M. E. Lewis 4915 (BRY). Emery Coun-
ty, Big East Mountain, head of Rilda Canyon,
13 July 1979, M. E. Lewis 6009 (BRY); do,
Big East Mountain, above Upper Joes Valley,
8 August 1977, M. E. Lewis 5129 (BRY).

The Carrington daisy has been identified
with E. simplex by previous workers, but ap-
pears to be more clearly allied to E. unter-
mannii, at least superficially. It has long,
spreading multicellular hairs similar to those
of E. simplex, but the pulvinate caespitose
habit and thick obtuse to rounded leaves of
E. carringtonae appear to be diagnostic.

The species is named in honor of the mem-
ory of Jane Carrington, evidently the first
woman to collect Utah plant materials desig-
nated as types (Welsh 1982).

Erigeron goodrichii Welsh, sp. nov. Similis
et affinis E. clokeyi Cronquist sed in caudi-
cibus tenuioribus et capitulis majoribus
differt.

Perennial herbs from a stout taproot and
caudex, the caudex branches with dark brown
marcescent leaf bases; stems 3-10 cm tall, de-
cumbent-ascending to erect, spreading hairy;
basal leaves 0.4-6 cm long, 1.2-5 mm wide,
narrowly oblanceolate, the veins not appar-
ent, pilosulose, obtuse apically; cauline leaves
more or less developed, but much reduced
upwards; heads solitary; involucres 10.5-18

April 1983

Welsh: Erigeron

367

mm wide, 6.4-7.8 mm high; bracts imbricate,
spreading villous-pilose with multicellular
hairs, thickened basally, green or the apices
suffused purplish, the inner with scarious
margins, the attenuate apices more or less
glandular and sometimes spreading; rays
40-65, pink purple to pink or white, 6.8-10.4
mm long, 1.5-2 mm wide; pappus apparently
single, of 20-30 minutely barbellate bristles;
achenes 2-nerved, pilose. Engelmann spruce
krummholz and meadow communities, often
on rock outcrops or talus at 3050 to 3400 m
in Duchesne, Summit, and Utah counties;
endemic.

Type.— USA. Utah. Duchesne County,
T2N, R8W, S22, Ashley NF, Uinta Moun-
tains, S rim of South Fork of Rock Creek, 14
mi n of Tabiona, 3264 m, with clumps of
Engelmann spruce krummholz, on gravelly
ground, 29 July 1981, S. Goodrich & R. Jep-
son 15907 (Holotype BRY; Isotypes NY, CAS,
POM, MO, US, UT, UTC, RM). Additional
specimens: Utah. Duchesne County, divide
between Log Hollow and Rock Creek, 12 mi
355 degrees from Tabiona, 31 July 1979, S.
Goodrich 13550 (BRY); do, head of Log Hol-
low, T2N, R7W, S30, 12 mi N of Tabiona, 3
July 1978, S. Goodrich and L. Hart 11696
(BRY); do, above Wedge Hollow, 11.5 mi and
354 degrees from Tabiona, 1 August 1979, S.
Goodrich 13579 (BRY). Summit County,
T2N, R12 E, S24, 11.5 mi NW of Kings Peak,
East Fork of Blacks Fork, 31 August 1981, S.
Goodrich 16203 (BRY). Utah County, Mt.
Timpanogos, Emerald Lake, 7 August 1941,
E. Castle 101c (BRY).

This dwarf alpine species has been recog-
nized as having affinities with E. asperu-
gineus (D.C. Eaton) Gray and E. clokeyi
Cronquist, each of them dwarf alpine species.
The Goodrich daisy differs from the former
by its more slender leaves and from the latter
by the caudex which lacks the distinctive
thatch of gray -brown marcescent leaf bases.

The specific epithet honors the enthusias-
tic collector of the type and other materials,
Sherel Goodrich, 'student of western botany,
whose energetic pursuit of Utah and Nevada
plants is unsurpassed.

Erigeron maguirei Cronq. var. harrisonii
Welsh, var. nov. Similis var. maguirei sed in
capitulis plus numerosis, radiis angustioribus,
et disci brevioribus differt.

Type.- USA. Utah. Wayne County, T29S,
R6E, S14, ca 1 mi ENE of Fruita, ca 5700 ft
elev., Navajo Sandstone, juniper community,
2 June 1982, S.L. & E.R. Welsh 21178 (Holo-
type BRY; Isotypes NY; CAS). Additional
specimens: Wayne County, canyon near Nat-
ural Bridge, 6 April 1934, B.F. Harrison 7385
(BRY); do, Fruita (Hickman Bridge trail), 10
June 1938, D E. Beck s.n. (BRY). The harri-
sonii phase of the E. maguirei differs in minor
technical ways from materials of the type va-
riety. Both phases are plants of sandy canyon
bottoms, and perhaps they represent nothing
more than ecological variants of a common
theme.

The variety is named in honor of its dis-
coverer, Bertrand F. Harrison, collector,
teacher, and student of Utah botany.

Erigeron untermannii Welsh & Goodrich,
sp. nov. Similis E. compactus sed in foliis la-
tioribus pilis ascendentibus vel patentibus et
radius brevioribus differt.

Perennial pulvinate herbs with an in-
tricately branched caudex, the caudex
branches mainly basal, 0.8-3.3 cm long, 1-4
mm wide, narrowly oblanceolate to spatu-
late, pilosulose with ascending, often curved,
hairs; scapes 2-6 cm tall; heads solitary; in-
volucres 7-11 mm wide, 5-5.7 mm high, the
bracts more or less imbricate, green, or the
inner somewhat chartaceous, the margins
hyaline, the tips suffused with purple (some-
times throughout), densely hispidulous with
short spreading hairs; rays 14-26, white,
4-6.5 mm long, 1.5-2.1 mm wide; pappus
apparently single, of ca 20 slender fragile-
bristles; achenes 2-nerved, pilose. Pinyon-
juniper community on calcareous shales and
sandstones of the Uinta and Green River for-
mations at 2135 to 2380 m in Duchesne and
Uintah counties; endemic.

Type.— USA. Utah. Duchesne Co., T5S,
R6W, S22, Indian Canyon, 17 mi S Du-
chesne, pinyon-juniper community, on Green
River Shale, 4 June 1980, N. D. Atwood 7554
(Holotype BRY; Isotype NY). Additional
specimens: Utah. Duchesne County, Tava-
puts Plateau, Uinta Formation, junction of
Right Fork and Left Fork of Indian Canyon,
ca 10 mi SW of Duchesne, steep slopes and
narrow ridge tops, shale and marly limestone,
26 May 1976, S. Goodrich 5317 (BRY); do, 13
mi 220 degrees from Duchesne, T5S, R6W,

368

Great Basin Naturalist

Vol. 43, No. 2

S21, ridge between Right Fork and Left Fork
of Indian Canyon, 31 May 1979, S. Goodrich
12402 (BRY). Uintah County, Vernal District,
Ashley National Forest, dry ridge top, S rim
of Red Pine Canyon, 5 June 1976, S. Good-
rich 5652 (BRY).

The Untermann daisy is compared to E.
compactus Blake in the diagnosis, and it is
probably allied to that taxon. However, the
similarity to E. nematophyllus Rydb. cannot
be ignored. It differs from that taxon in the
broader leaves, generally harsher and more
spreading hairs of the leaves, involucre, and
achenes. The plant is named to honor the
memory of the late G. E. and B. R. Unter-
mann, an amazing husband and wife team
who worked throughout their lives to under-
stand the geology, natural history, and an-
thropology of the Uinta Basin. They in-
fluenced the lives of all persons whom they
contacted.

Erigeron wahwahensis Welsh, sp. no v.
Herbis similis E. eatonii in aspectus sed ro-
bustioribus caudicibus crassioribus pilis pat-
entibus pro parte et bracteis crassioribus ad
basim et dense vel sparse pilis patentibus
differt.

Perennial herbs, from a branching caudex,
the caudex branches with conspicuous fibrous
brown to ash-colored marcescent leaf bases;
stems 15-40 cm long, decumbent to ascend-
ing; basal leaves 3-18 cm long, 4-13 mm
wide, linear-oblanceolate to oblanceolate or
elliptic, 3-nerved, petiolate, appressed to
spreading hairy with curved hairs; cauline
leaves reduced, sessile, and bracteate above;
heads solitary or 2 or 3; involucres 13-17 mm
wide, 6-7 mm high, spreading-villous with

multicellular hairs, glandular apically; bracts
imbricate, green, the tips reddish, thickened
basally; rays 30-40, pink or white, 5.5-7 mm
long, 1.7-2.2 mm wide; disk corollas 3.5-4.2
mm long, the tube ca 2 mm long, the lobes
0.4 mm long; pappus of 15-20 bristles, with
inconspicuous outer setae; achenes 2-nerved,
short-hairy. Sagebrush, oak-maple, and pin-
yon-juniper communities at 1670 to 2440 m
in Beaver and Washington counties; endemic.

Type.- USA. Utah. Beaver County, T28S,
R15W, S31, Wah Wah Mountains, ca 12 mi
SSW of Wah Wah Spring, Pine Grove Pass,
2450 m elev, sagebrush and pinyon-juniper
community, 12 June 1982, S.L. Welsh 21229
(Holotype BRY; Isotypes NY, CAS, POM,
RM, UT, UTC, US, GH, MO, and others to
be distributed). Additional specimens: Utah.
Beaver County, T29S, R16W, S12. Willow
Creek, Wah Wah Mountains, 8 June 1978, K.
Ostler & D. Anderson 1274 (BRY); do, T28S,
R15W, S31, Wah Wah Mountains, divide be-
tween Quartz Creek and Pine Grove Canyon,
22 May 1981, S.L. Welsh 20520 (BRY).

The Wah Wah daisy is more or less inter-
mediate between phases of E. eatonii and E.
jonesii, but has features not shared by either.
The pubescence is similar to E. jonesii, but
the general aspect is more like E. eatonii.
The plants tend to be larger than either, and
the thick caudex appears to be diagnostic.

Literature Cited

Cronquist, a. 1947. Revision of the North American
Species of Erigeron north of Mexico. Brittonia 6:
121-302.

Welsh, S. L. 1982. Utah Plant Types— historical per-
spective 1840 to 1981— annotated list, and bibli-
ography. Great Basin Nat. 42:129-195.

NEW TAXA IN THELESPERMA AND TOWNSENDIA (COMPOSITAE) FROM UTAH

Stanley L. Welsh'

Abstract. — Three new taxa are named and described: Thelesperma subnudum Gray var. alpinum Welsh, Towns-
endia jonesii (Beaman) Reveal var. lutea Welsh, and Townsendia montana var. caelilinensis Welsh.

Preparation of a manuscript of the sun-
flower family for the Utah flora project dem-
onstrated the existence of diversity within
Thelesperma and Townsendia that seems best
treated at taxonomic rank within existing
species in those genera. In both genera the
plants named are ecological specialists, with
plants of each taxon occurring on calciferous
or gypsiferous substrates of the Carmel, Flag-
staff, Arapien, or other peculiar formations.
Each of them belongs to the mound-forming
group of substrate specialists of barren or
semibarren habitats where water relations are
controlled by the parent material. Generally
the habitats are arid, despite the large
amount of rainfall in the higher elevation
sites. In all habitat examples cited there are
other similarly restricted specialists, either
species or varieties.

Thelesperma subnudum Gray var. alpi-
num Welsh, var. nov. Similis Thelesperma
subnudo Gray var. subnudo sed in habito hu-
milioribus et capitulis parvioribus differt.

Perennial herbs from a taproot and less
commonly with a caudex and creeping root-
stock; stems 3-7 cm tall, subscapose; leaves
mainly basal on the stem, 1.5-4 cm long, pin-
nately lobed, or the upper entire; petioles
ciliate and the blades puberulent; involucres
6.3-9 mm high, 9-14 mm wide; outer bracts
oblong to lanceolate, with narrow scarious
margins, to half as long as the inner; inner
bracts united to below the middle; rays lack-
ing; disk flowers yellow; pappus a toothed
crown; achenes glabrous or hairy apically.

Type.- USA. Utah. Wayne County, T28S,
R4E, S13 (NEi/4;l. 3 mi due N of Bicknell,
bristlecone pine forest on multicolored clay
hills, 2745 m, 20 July 1980, D. Atwood and

B. Thompson 7646a (Holotype BRY). Addi-
tional specimen: Utah. Wayne County, S of
Teasdale, 25 July 1978, D. Atwood 6924
(BRY).

This dwarf phase of Thelesperma sub-
nudum occurs about 500 m above the up-
permost elevations known for the typical va-
riety. Discoid plants are not uncommon for
the widespread phase of the species, includ-
ing at least a portion of the type (taken in
Red Canyon, near Paragonah; isotype BRY),
but the typical phase is ordinarily radiate.
The dwarf alpine plants have involucres that
are both shorter and narrower on the aver-
age. The plants are apparently restricted to
the Carmel Limestone, on the peculiar vari-
colored phase of that formation as it occurs
at the east margin of Rabbit Valley.

Townsendia jonesii (Beaman) Reveal var.
lutea Welsh, var. nov. Affinis T. jonesii var.
jonesii sed in coroUis luteis ventralis differt et
in substratis gypsiferis confinis.

Subacaulescent to acaulescent caespitose
herbs, the caudex commonly branched; stems
not conspicuously white-strigose, mainly 2-3
cm tall, forming clumps to 5 cm wide; leaves
10-30 mm long, 1-4 mm wide, oblanceolate'
to spatulate or almost linear, strigose; heads
mostly solitary; involucres 9-12 mm high,
8-13 mm wide; bracts in 4 or 5 series, lan-
ceolate, green or suffused purple, sparsely
strigose; rays 13-21, yellow ventrally, often
suffused reddish dorsally, glandular, 4-7 mm
long; disk corollas yellow, ca 3.5 mm long;
achenes 3-5 mm long, pubescent with glochi-
diate hairs; ray pappus 2-4.5 mm long; disk
pappus 5-8 mm long.

Type.— USA. Utah. Sevier County, Ara-
pien shale outcrop with scattered juniper, ca

'Life Science Museum and Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602.

369

370

Great Basin Naturalist

Vol. 43, No. 2

4.2 km E of Sigurd, T23S, RIW, S8, at mile-
post 14, Utah Hwy 24, 8 May 1975, S.L.
Welsh 12700 (Holotype BRY; 2 isotypes dis-
tributed previously as T. aprica Welsh &
Reveal).

Additional specimens: Utah. Sevier Coun-
ty, Rainbow Hills area, 2 mi NE of Glen-
wood, 7 May 1981, L. Greenwood 5013; do,
4 mi S of Salina, 24 Mar. 1981, L. Green-
wood 5003; do, 2 mi E of Sigurd, 7 May
1981, L. Greenwood 5012. Piute County, NE
side of Piute Reservoir, 26 Apr. 1979, E.
Neese et al. 7149; do 30 Apr. 1981, D. At-
wood and E. Neese 7794 (all BRY).

The presence of yellow flowers in T. aprica
was considered to be noteworthy for the
genus, although corollas that dried yellowish
were admitted for T. pnesii (Reveal 1970).
The existence of populations with yellow co-
rollas when fresh in Piute and western Sevier
counties at first indicated placement of those
plants with T. aprica, but the long pappus of
the ray flowers indicates relationships with T.
pnesii, in which the corollas sometimes fade
yellowish.

Most of the localities for this variety are
situated on the Arapien Shale, which is noted
for its deposits of commercial gypsum. The
exception involves those plants from near the
Piute Reservoir in Piute County, where the
plants grow on gypsiferous or calciferous sub-
strates amidst igneous gravels.

Toumsendia montana Nutt. var. caelili-
nensis Welsh, var. nov. A var. montana
differt in foliis spatulatis late capitulis major-
ibus et bracteis latioribus.

Type:- USA. Utah. Sanpete Co., T20S,
R4E, S33, ca 24 km W of Perron, 3050 m.
Flagstaff Limestone, barrens, 13 July 1977, S.
Welsh & S. Clark 15385 (Holotype BRY; 3

isotypes distributed previously as T. montana
Nutt.). Additional specimens: Utah. Duchesne
County, Argyle Canyon, 27 June 1978, E.
Neese & L. England 5864; do, T6S, R6W,
SW of Duchesne, 2 June 1978, E. Neese
5305. Sanpete County, 8 mi E of Spring City,
12 July 1977, E. Neese & S. White 3552; He-
liotrope Mountain, 26 June 1977, M.E. Lewis
4254; do, 17 July 1981, D. Atwood 7998; do,
head of Bacon Rind Canyon, J. W. Hum-
phrey 52958, 22 July 1927; do, 11 mi 134 de-
grees from Manti, 3 July 1981, 22 July 1927;
South Tent Mountain, 27 July 1976, M. E.
Lewis 4365; Skyline at head of Duck Fork,
22 July 1976, M. E. Lewis 4246; do, E of
Manti, 28 July 1977, S. Clark 2877. Wasatch
County, 27 mi 98 degrees east of Spanish
Fork, 19 July 1982, S. Goodrich 17303 (all
BRY).

This is the large-headed phase of the spe-
cies, with broadly rounded spatulate leaves,
which lies intermediate with the typical
montane materials and the dwarf plants of
var. minima. The broad leaves and large
heads are diagnostic in segregation of this en-
tity from both the typical montane phase and
the more southern var. minima.

The substrate occupied by var. caelili-
nensis on the Wasatch Plateau is composed
of weathered Flagstaff Limestone. The Du-
chesne County populations occur on calcif-
erous members of the Green River Formation.

Literature Cited

Beaman, J. H. 1957. The systematics and evolution of

Townsendia Compositae. Contr. Gray Herb. 183:

1-151.
Reveal, J. L. 1970. A revision of the Utah species of

Townsendia (Compositae). Great Basin Nat.

30:23-52.

NEW HAPLOPAPPUS VARIETY IN UTAH (COMPOSITAE)

Stanley L. Welsh' and Frank J. Smith-

Abstract. — Reported as a new variety from the Green River Formation of Uintah and Duchesne counties, Utah,
is Haplopappus armerioides (Nutt.) Gray var. gramineus Welsh & F. J. Smith.

The arid Green River Formation, with its
pecuhar soil relations, has been the spawning
ground for numerous narrow endemics. The
taxon described herein is another belonging
to that unique group of mound-forming, low-
growing taxa that grow on the peculiar shale
substrates. Relatively little variation occurs
within the typical variety of H. armerioides.
Thus, the variation represented by the taxon
proposed below falls well outside that of the
typical variety and substantiates its recogni-
tion. The main features that distinguish this
variety from the main body of the species in-
clude the short stature and slender leaves, but
the heads are also smaller, on the average,
and the involucral bracts tend to be narrow-
er. This latter feature seems to indicate an af-
finity with the related H. acaulis (Nutt.)
Gray, but the bulk of the characteristics seem
to represent an extension of the basic mor-
phology of H. armerioides; hence, the place-
ment with that taxon. The numerous speci-
mens cited below have resulted from the
laborious efforts of students of the Utah flora,
whose contributions are gratefully ac-
knowledged.

Haplopappus armerioides (Nutt.) Gray
var. gramineus Welsh & F. J. Smith, var.
nov. Similis var. armerioides sed in staturis
humilioribus et folius gracilioribus differt.

Perennial caespitose herbs from a thick lig-
neus pluricipital caudex and stout taproot,
the caudex branches clothed with brown to
ashy marcescent leaf bases and leaves; herb-
age resinous-glandular, otherwise glabrous or
with scabrous leaf margins; stems mainly 3-8
cm tall; basal leaves 1.5-4 cm long, 1-3 mm
wide, rigid, linear, sharply mucronate, 1- or

obscurely 3-nerved; cauline leaves few, re-
duced upwards; heads solitary; involucres
campanulate, 8-11 mm high, 10-14 mm
wide; bracts in 3 or 4 series, imbricate, ob-
long to oval or obovate, obtuse, sometimes
lobed below the apex, greenish near the apex,
glabrous; rays 8-12, 10-12 mm long, yellow,
3-4 mm wide; pappus white; achenes silky-
villous.

Type.- USA. Utah. Uintah County, T13S,
R25E, S8, Atchee Ridge road, 1.4 km N of
Boulevard Ridge, 2727 m elev, Green River
Formation, pinyon-juniper, serviceberry,
mountain mahogany, and sagebrush commu-
nity, 29 May 1982, K. Thome & B. Neely
1836 (Holotype BRY; Isotypes NY; CAS).

Additional specimens: Utah. Duchesne
County, Dry Canyon, 40 km due NE of
Price, 25 May 1978, E. Neese 4916. Uintah
County, along Watson-Ouray rd, 8 km W of
Bitter Creek, 26 May 1935, E. H. Graham
8988 (UTC); do. Big Pack Mtn., 16 May
1978, E. Neese & J. S. Peterson 4628; do, ca
45 km S of Ouray, 18 May 1978, E. Neese &
J. S. Peterson 4729; do, Johnson Draw, be-
tween Hill and Willow creeks, 27 Apr. 1978, '
E. Neese & J.L. England 4288; do. Long
Draw, 2 km S of Ouray-Rainbow road, 18
May 1979, J. L. England 1777; do, Long
Draw, ca 4.5 km SW of Rainbow, 16 May
1982, E. Neese & F. J. Smith 11388, 11389;
do. Long Draw, ca 4 km W of Rainbow road,
26 May 1982, K. Thome and B. Neely 1793;
do. Bitter Creek and West Fork of Asphalt
Wash jet., 26 May 1982, K. Thome and B.
Neely 1798; do, T13S, R25E, Sll, 30 May
1982, R. Kass & J. Trent 862; do, Atchee
Ridge road, near East Seep Canyon, 7 June

'Life Science Museum and Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602.
'26 North 100 East, Smithfield, Utah 84335.

371

372

Great Basin Naturalist

Vol. 43, No. 2

1982, F. J. Smith 1639, 1640; do, 4.5 km S of
Rainbow, 15 May 1982, F. J. Smith 1596b,
1597; do. Asphalt Wash, 28 May 1982, R.
Kass & J. Trent 842 (all BRY, except as
noted).

There is a tendency for some plants to
have acutish bracts, and the broad subapical
spot is only more or less developed. Despite

the presence of these features, which are
somewhat intermediate with phases of H.
acaulis, the taxon stands with few inter-
mediates to H. armerioides in a strict sense.

Literature Cited

Hall, H. M. 1928. The genus Haplopappiis. Carnegie
Institution of Washington. 391 pp.

NEW SPECIES OF HYMENOXYS AND PERITYLE (COMPOSITAE) FROM UTAH

Stanley L. Welsh' and Elizabeth Neese^

Abstract. — Named as new species are Hymenoxys lapidicola Welsh & Neese, from Uintah County, Utah, and
Perityle specuicola Welsh & Neese, from Grand County, Utah.

Revision of the sunflower family, as part of
the Utah Flora project, has led to the recog-
nition of several taxa that were previously
known but had been placed within other en-
tities. Additionally, some of the taxa had ap-
parently not been collected previously, but
were similar to previously described taxa.
Careful observation of each of the numerous
specimens in the herbarium at Brigham
Yoimg University has led to recognition of
tiie great diversity in our flora. The two taxa
described below each occur on sandstone,
where they grow in crevices. They are part
of a cadre of species with crevice habitats,
and both are narrowly restricted endemics.
Much more work remains to be done in the
areas of the state occupied by massive sand-
stones, limestones, and other geological strata
that produce cliffs and escarpments. The fol-
lowing species are named to indicate the pe-
culiarities of their places of growth.

Hymenoxys lapidicola Welsh & Neese, sp.
nov. Ab H. torreyana (Nutt.) Parker in
bracteis recuvatis et capitulis parvioribus
differt et a H. depressa (T. & G.) Welsh &
Reveal in bracteis recurvatis et marginem
scariosis differt.

Pulvinate caespitose herbs from a multi-
cipital caudex, this densely clothed with
brown marcescent leaf bases, acaulescent;
leaves all basal, 0.3-1.2 cm long, 0.8-2 mm
wide, narrowly oblanceolate, the inner con-
spicuously glandular-punctate, the blades
glabrous, the axils long-villous; heads solitary,
immersed in the leaves; disks 5.5-9 mm wide;
involucres 5-8 mm high; bracts distinct, in 2
or 3 subequal series, sparingly villous and suf-
fused reddish, the margins scarious, the tips
more or less squarrose-spreading and some-

what thickened; rays 5 or 6, yellow, 5-6 mm
long; pappus scales lance-acuminate, 2.3-3
mm long; achenes 2-2.5 mm long, pilose.

Type.- USA. Uintah County, T5S, R25E,
S20, Point of Pines camp area, S edge of Blue
Mt. Plateau, at 2485 m, in ponderosa pine-
manzanita community, sandy loam and
standstone outcrop, 12 June 1982, E. Neese
and C. Fullmer 11734 (Holotype BRY; Iso-
type NY).

Additional specimens: Utah. Uintah Coun-
ty, Blue Mountain, cliff face, 11 June 1982,
F. Smith & J. Trent 1653 (BRY); do. Point of
Pines campground, Weber Sandstone, 11
June 1982, R. Cass & E. Neese 919 (BRY); do,
10 mi E of Jensen in draw at base of Blue
Mountain, 14 May 1982, E. Neese et al,
11370 (BRY).

The habit of H. lapidicola simulates that of
H. depressa, but the features of the involucral
bracts and the proportionally broader non-
cuspidate leaves indicate a closer relationship
with H. torreyana. From that entity it differs
in the recurved outer involucral bracts with
thickened reddish tips and much smaller
heads.

Perityle specuicola Welsh & Neese, sp.'
nov. Similis Perityle congesta (Jones) Shinners
sed in aspectis et pappo setarum 3 vel 4 (nee
1 vel 2), et in pappo palearum sigmoidis et
expansis differt.

Perennial suffruticose herbs, mainly 50-75
cm tall; stems sprawling or pendulous, much
branched; herbage glandular-hispidulous;
leaves mostly alternate, short-petiolate, the
blades 3-6 mm long, 1.5-3 mm wide, ovate-
elliptic, entire, hispidulous; heads few to
many in a branching corymbose in-
florescence; involucres 3.5-5 mm high, 5-6

'Life Science Museum and Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602.
'Life Science Museum, Brigham Young University, Provo, Utah 84602.

373

374

Great Basin Naturalist

Vol. 43, No. 2

mm wide; bracts 11-16, oblong to elliptic,
keeled; ray flowers lacking; disk flowers nu-
merous, ca 2.5 mm long, whitish (?); pappus
of 3 imequal scabrous bristles and often with
1 apically flattened and sigmoid scale;
achenes 3-3.8 mm long, the faces flattened,
glabrous, the margin thickened and with
short ascending hairs.

Type.- USA. Utah. Grand County, T25S,
R21E, S24, 2 mi due N of Moab, 1220 m,
hanging garden community, 29 Sept. 1977,
S.L. Welsh 16283 (Holotype BRY; 6 isotypes
distributed previously as Laphamia congesta
Jones).

Additional specimens: T25S, R22E, S29,
Castle Valley, 2 mi due NNE of Moab, Negro

Bill Canyon, 1220 m, Navajo sst, hanging gar-
den community, 6 Oct. 1977, S.L. Welsh
16365 (BRY).

The technical features of this taxon in-
dicate an affinity with P. congesta, but the
differences in the pappus features, inter alia,
suggest the need for recognition of this dis-
tinctive plant. It seems to be more than an
etiolated shade form of its near ally, from
which it is remote geographically by more
than 300 km.

Literature Cited

Powell, A. M. 1973. Taxonomy of Perityle section Lap-
hamia (Compositae-Helenieae-Peritylinae). Sida
5:61-128.

NEW VARIETY OF STEPHANOMERIA TENUIFOLIA (COMPOSITAE) FROM UTAH

Sherel Goodrich' and Stanley L. Welsh'
Abstract. — Named as new is Stephanomeria tenuifolia (Torr.) Hall var. uintaensis Goodrich & Welsh.

Identification of plants taken in prepara-
tion of a flora of the Uinta Basin demon-
strated the presence of a Stephanomeria
whose description is beyond that of other
taxa in the genus. Specimens were compared
with all other taxa in the genus, and with
published keys and descriptions of all con-
temporary floras. The plants are situated at
the margin of the range of S. tenuifolia, and
are similar in general aspect to other phases
of that species that are ecologically stressed.
Plants taken on the West Tavaputs Plateau at
2745 m elevation (Welsh & Clark 15923
BRY) are similar to the aberrant materials
from the Uinta Basin in general aspect and in
leaf features, but the involucres are within
the normal size range for typical S.
tenuifolia.

The material described below grows in one
small isolated stand in a ponderosa pine com-
munity and shows variation in size of heads
and shape of the involucral bracts. The cau-
dex is not well developed, producing solitary
or few stems. Recognition at varietal level
seems justified. More work in the genus is
indicated.

Stephanomeria tenuifolia (Torr.) Hall var.
uintaensis Goodrich & Welsh, var. nov. A
var. tenuifolia imprimis in involucris long-
ioribus bracteis attenuatis radiis longioribus
et foliis basalibus bipinnatifidis differt.

Perennial herbs from a woody caudex; cau-

dex sparingly branched or the branches lack-
ing, with few marcescent leaf bases, not
hairy; stems solitary or 2, 25-40 cm tall;
herbage puberulent; leaves 1-11 cm long,
1-8 mm wide, the lower and sometimes the
middle cauline ones runcinate-pinnatifid, the
upper (at least) entire, linear, finally brac-
teate; heads solitary at the ends of stems or
on branches; involucres 10-16 mm high, 3-5
mm wide; main bracts lance-attenuate, green
or suffused with purple, puberulent; outer
bracts very short; rays 5, pink, 7-10 mm
long; pappus white, plumose to the base;
achenes 5-6 mm long, longitudinally ribbed,
the angles smooth.

Type.- USA. Utah. Uintah County, T2S,
R19E, S14, NWi/4, 18 mi NW 319 degrees of
Vernal, Ashley N.F., Uinta Mountains,
Brownie Canyon, 2486 m; ponderosa pine,
Douglas fir, Juniperus scopulorum, and sage-
brush, south exposure, 20 Aug. 1982, Good-
rich 17708 (Holotype BRY; isotypes NY, US,
CAS, POM, MO, UT, UTC, RM, GH).

Additional specimens: Utah. Uintah Coun-
ty, Brownie Canyon, 13 Sept. 1982, S. Good-
rich & D. Atwood 17968 (BRY).

This variety approaches S. parryi in bract'
size, but has the appearance of phases of S.
tenuifolia at the edge of their ecological tol-
erance. The tall involucres appear to be diag-
nostic when taken with the other features
mentioned in the diagnosis.

'USDA Forest Service, Intermountam Forest and Range Experiment Station, Ogden, Utah 84401, stationed in Provo, Utah, at the Shrub Sciences
Laboratory.
'Life Science Museum and Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602.

375

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TABLE OF CONTENTS

Utah flora: Compositae (Asteraceae). Stanley L. Welsh 179

Haplopappus crispus and H. zionis (Asteraceae): new species from Utah. Loran C.

Anderson 358

A bouquet of daisies (Erigeron, Compositae). Stanley L. Welsh 365

New taxa in Thelespetma and Townsendia (Compositae) from Utah. Stanley L.

Welsh 369

New Haplopappus variety in Utah (Compositae). Stanley L. Welsh and Frank J.

Smith 371

New species of Hymenoxys and Perityle (Compositae) from Utah. Stanley L. Welsh

and Elizabeth Neese 373

New variety of Stephanomeria tenuifolia (Compositae) from Utah. Sherel Goodrich

and Stanley L. Welsh 375

HE GREAT BASIN NATURALIST

}lume 43 No. 3

July 31,1983

Brigham Young University

LIPOAQY

UNiVERSITV

I'M;

LIBRARY

.„\V

/.u/..^ . HARVARD

GREAT BASIN NATURALIST

Editor. Stephen L. Wood, Department of Zoology, 290 Life Science Museum, Brigham Young

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9-83 650 66769

ISSN 017-3614

The Great Basin Naturalist

Published at Provo, Utah, by
Brigham Young University

ISSN 0017-3614

Volume 43

July 31, 1983

No. 3

A RE-EVALUATION OF THE POSTGLACIAL VEGETATION
OF THE LARAMIE BASIN, WYOMING-COLORADO

Deborah L. Elliott-Fisk', Betty S. Adkins-, and Jeanine L. Spaulding'

Abstract.— Previous work by Wells in the Laramie Basin suggested that a coniferous forest /woodland covered
the basin floor in the recent past (until the latest Holocene). We have found no evidence for this woodland and sug-
gest instead that these scattered woodlands along sandstone outcrops and their immediate margins are outliers of the
montane forest of the Medicine Bow Mountains, existing in their apparently anomalous locations due to favorable
microenvironments set up by the topography and substrate.

During the past two years, we worked at
various sites in and surrounding the Laramie
Basin in an attempt to collect sufficient infor-
mation with which to reconstruct postglacial
vegetation (and environment) change for the
region. The basin's unique geography as one
of the few high-altitude basins surrounded by
the Rocky Mountains, and the knowledge
that a periglacial climate existed here in pre-
Holocene times (Mears 1981), encouraged us
to seek postglacial paleoclimatic data from
not only upper elevations in the mountains
(as done traditionally in the Rocky Moun-
tains), but for the basin itself. Faunal, soil,
and geomorphic studies here (Hager 1972,
Reider et al. 1974, Grasso 1979, Reider and
Burgess, in prep.) point to late Pleistocene
and Holocene environmental change. In ad-
dition, a 116-year-old record of mete-
orological data is available for the basin,
which will aid any climatic reconstructions
attempted.

Upon visiting the Sand Creek region of the
Laramie Basin (Fig. 1), we were delighted to

find an abundance of fossiliferous material,
including in situ dead trees (macrofossils) and
Neotoma (woodrat) middens, as well as living
trees that appeared to be at least several cen-
turies old. Both living and dead trees were
restricted to the sandstone outcrops and their
immediate margins, these outcrops principal-
ly lithified sand dunes of the Pennsylvanian
Casper Formation (Fig. 2).

It later surprised us when we read in the
papers of Wells (1970a, 1970b) on the post-
glacial vegetation of the Laramie Basin tRat
he believed these trees had in recent times
(the latter half of the Altithermal and per-
haps part of the Neoglacial) covered the en-
tire floor of the Laramie Basin. Knowing
both the ecological requirements of these
conifer species {Juniperus scoptilorum Sarg.—
Rocky Mountain juniper; Pinus flexilis
James— limber pine; and P. ponderosa Laws.—
ponderosa pine) and the relatively minor de-
gree of climatic change in the Holocene
compared to the Pleistocene, we considered
Wells's hypothesis not plausible. Instead,

'Department of Geography, University of California, Davis, California 95616.
'Department of Geography, University of Alberta, Edmonton, Alberta, Canada T6G 2H4.
'Department of Geography, University of Wyoming, Laramie, Wyoming 82071.

377

378

Great Basin Naturalist

Vol. 43, No. 3

Fig. 1. Map of Sand Creek study area in the Laramie Basin of Wyoming-Colorado.

topography and microenvironmental condi-
tions could have favored a more vigorous
woodland (as evidenced by population and
individual sizes) on the outcrops sometime in
the past.

We therefore decided to critically reeval-
uate the paleoenvironmental data for the
Sand Creek area. This involved examining a
larger area than apparently done by Wells
(pers. comm. with local ranchers) and also an
attempt to tie the botanical data in with
other paleoenvironmental studies conducted
in the region. The completion of this project
awaits radiocarbon dates and further den-
droclimatic reconstructions. It is our hypoth-
esis that these scattered woodland stands are
outliers of the montane forest of the Medi-
cine Bow Mountains, existing in their appar-
ently anomalous locations due to favorable
microenvironments set up by the topography

and substrate, and not relicts of a former for-
est over the basin floor as hypothesized by
Wells.

Methods

The methods we have used to solicit infor-
mation as to the nature and degree of change
of past environments at Sand Creek involve
gathering various paleoecological materials.
As we have yet neither found fossil soils in
this immediate area, nor analyzed animal re-
mains found in the Neotoma middens, we
confine our discussion to the botany and pa-
leobotany of the area.

Extensive hiking and collecting was done
in the area outlined in Figure 1 to determine
the spatial and temporal extent of conifers in
this region. Dead and living individuals were
mapped, along with Neotoma middens (both

July 1983

Elliott-Fisk et al.: Postglacial Vegetation

379

Fig. 2. Study site 1 at Sand Creek (Wells' primary site). The landscape here is characterized by lithified sand
dunes of the Pennsylvanian Casper Formation. The tree in the middle of the photo is a Rocky Mountain juniper
(Jiiniperiis scopuhrtim) approximately 1000 years of age.

fossil and contemporary). Although collecting
has been completed at many of these sites,
materials are still being obtained. In addition,
the vegetation, regenerative capacity, micro-
climatic setting, geology, and soils of the sites
were recorded.

Initial tree-ring and midden analyses, fol-
lowing the traditional methods of Stokes and
Smiley (1968), Fritts (1976), and Wells
(1976), have been conducted. Studies on the
regenerative capacity of the conifer popu-
lations follow the methods outlined by Elliott
(1979). We present our initial findings here.
We believe these findings are important for
those attempting to reconstruct Holocene cli-
matic change for the Western USA, because
they offer a radically different conclusion
than diat presented by Wells (1970a, 1970b).
Also, for anyone undertaking recent pa-
leoenvironmental studies, it becomes increas-
ingly important to have a thorough under-
standing of the life strategy and ecological
requirements of the species investigated, and
also of the local geography of a site, if one is
to accurately reconstruct a climatic record.

Results

The present distribution of trees and tree
species is shown in Figure 3. Multiple mono-
specific and mixed species stands occur along
this semicontinuous sandstone ridge extend-
ing out from the southwestern Medicine Bow
Mountains. The lower forest limit here is
2,500 m, with woodland patches reaching
elevations of 2,300 m.

It is important to note that the lowest trees
surviving here are not on the floor of the ba-
sin proper, but are restricted to the sandstone
outcrops (Fig. 4) or their immediate margins.
The dead trees (macrofossils) reported by
Wells and found by us occupy the same topo-
graphic position as the living conifers, where
winter snowdrift and summer runoff can sup-
ply them with the necessary moisture for sur-
vival in this marginal environment (Fig. 5).

It is interesting to note that two of the fall-
en fossil junipers plus two standing dead
stumps in the area are much larger than any
living junipers in the area today (Fig. 6). As
this species is apparently at its lower mois-

380

Great Basin Naturalist

Vol. 43, No. 3

Juniperus scopulorum
PInus tiexilis
Pinus ponderosa

Sites
Outcrops

CONIFER POPULATIONS:
SAND CREEK

Fig. 3. Distribution of living and fossil (in situ dead)
trees in Sand Creek study area. Findings in woodrat
middens are not included here. Sandstone outcrops ex-
tend from the Medicine Bow Mountains to their south-
west, decreasing in elevation toward the northeast.

tiire, but not temperature, limit today (Po-
wells 1965), it seems reasonable to speculate
that more moisture (either due to increased
precipitation or decreased temperatures/
evapotranspiration) was available during at
least part of the past life spans of these trees.

This hypothesis is also supported by the
fact that no mixed stands of ponderosa pine
and juniper (as evidenced in some of the fos-
sil middens) exist here today. Stands of limber
pine and juniper do exist, along with mixed
stands of all three species and pure stands of
juniper or ponderosa pine. Fossil midden
equivalents of all stand types mentioned here
except the monospecific stands are found.

All three species are successfully sexually
regenerating today in each stand in which
they occur. This is witnessed by the presence
of both viable seed (cones) and juveniles.

It proves difficult to make any type of rig-
orous statement in reference to the age struc-
tures of the tree populations here. This is due
not only to a lack of radiocarbon dates for

each dead tree (macrofossil), but primarily
because both dead and apparently also living
trees have been removed from this area for
use as fence posts, firewood, etc. Many of the
cut junipers we have found were harvested
for "cedar table tops" in the 1920s (pers.
comm., F. Lilly to B. Mears 1980), a fact that
is distressing to paleoecologists trying to re-
construct the history of a site. Cross-dating of
the tree rings in mature junipers is also ex-
tremely difficult; however, limber and pon-
derosa pine are easily cross-datable.

Juveniles of all three species, ranging from
a few to 50 years in age, are abundant. All
age classes (standard 10-year groups) of pon-
derosa pine are found for the last few hun-
dred years, with no individuals over 350
years old (as yet found) surviving today. Pon-
derosa pine macrofossils (both in situ dead
trees and smaller macrofossils in middens) are
infrequent, though we have found cones and
seeds at site 1. Limber pine is found up to
several hundred years old, with some individ-
uals approaching 1,000 years in age. In situ
limber pine macrofossils are common at sites
2 and 3; some of these dead trees were also
much larger than those living today. Sam-
pling of these macrofossils is not yet
complete.

Juniper appears to have had considerable
difficulty through at least the last few thou-
sand years (as documented by Wells's radio-
carbon dates) with the establishment of an
equilibrium population in this area. The liv-
ing individuals today are primarily either
several hundred to 1,000 years old or under
100 years in age. Juveniles tend to be clus-
tered around singular dead or living mature
individuals, which most likely served as the
seed (mother) trees. Layering of juveniles
from each other (but not from the larger
trees) is also found. Occasionally, juveniles
are found a great distance (over 1 km) from
any possible seed trees. Few young adults are
found, an age group in which the two pine
species are abundant. Further dendroeco-
logical work with response functions for a
species may help explain this difference.

One of the most intriguing facets of the
Sand Creek sites is the tremendous variation
in average tree-ring widths as one looks at
living versus dead individuals. The ponderosa
pines seem to have found a couple of sites at

July 1983

Elliott-Fisk et al.: Postglacial Vegetation

381

Fig. 4. Trees at site 2. The majority ot trees in this stand are situated on top of the sandstone outcrop, with indi-
viduals occasionally found along the margins. Junipcrus scopulontrn, Pinits flexilis, and Pinus ponderosa are all pres-
ent here as both living individuals and fossils.

which they can exist with only occasional
stress (as evidenced by very narrow rings
every 30 to 40 years which are cross-datable);
these individuals show wide, symmetrical
crowning with consistently good growth and
low mean sensitivities. In contrast, limber
pine and juniper appear to have been under
considerable environmental (climatic) stress,
though this has not precluded the estab-
lishment of juveniles in recent times. Narrow
rings and high mean sensitivities are common
for both species here.

For comparative purposes, we present av-
erage ring-width data for several individuals
of Juniperus scopulorum that we believe are
representative of our samples. Whereas two
measured, undated fossil trees have average
ring widths of 1.09 and 2.38 mm, respective-
ly, the largest living tree at Wells's site (our
site 1) has an average ring width of 0.19 mm.
One section of a large dead juniper cited by
Wells with a radiocarbon date of 940 ± 105
BP (Gx-140F) on the outer wood has an aver-
age ring width of 0.17 mm; however, this val-
ue is derived from measurements on a branch

cross-section and may therefore be expected
to be smaller than those from the primary
trunk. (All of the other measurements given
are from main trunk cores and cross-sections.)
The above data show almost a 25-fold dif-
ference in ring-widths. As the living and fossil
trees are found in the same habitats, this dif-
ference in growth must be attributed to a
change in climate, with other environmental
factors (topography, etc.) remaining constant.
It is not attributable to intrapopulation or
age differences.

Conclusion

It appears that Holocene climatic change
in the Laramie Basin has been sufficient to
trigger the death of conifers in the most mar-
ginal low-elevation sites. This is suggested by:
(1) change in species composition of some of
the lowest woodland stands, with only the
most xeric tree species surviving this deterio-
ration, (2) the death of individuals at the most
marginal microclimatic/topographic loca-
tions, and (3) the decrease in ring width

382

Great Basin Naturalist

Vol. 43, No. 3

Fig. 5a. Remains of fossil (dead) Juniperus scopulorum at the margin of a sandstone outcrop. This individual, like
many in the area, has been partly harvested for lumber. Site 1.

Fig. 5b. Immature (not yet bearing seeds) Juniperus scopulorum along the margin of a sandstone outcrop between
sites 1 and 2. This is a typical habitat for juniper regeneration. This individual is approximately 80 years old.

July 1983

Elliott-Fisk et al.: Postglacial Vegetation

383

Fig. 6. One of two large, dead, fallen junipers at site 1. A cross-section through the midpart of this tree has been
cut out, most likely for a table top. Presence of small branches on this individual attest to the fact that it has prob-
ably not been dead for more than a few hundred years.

(average growth rate) of the mature conifer
populations.

We have found no evidence that a con-
iferous woodland or forest ever existed over
the floor of the Laramie Basin in the Holo-
cene (or even Pleistocene, though this latter
time period is out of the scope of this study).
It does appear, however, that more and occa-
sionally larger individuals of the extant con-
ifer species did exist on the sandstone out-
crops and at their immediate margins in the
early Neoglacial and perhaps late
Altithermal.

Acknowledgments

This research was initiated while the au-
thors were faculty (Elliott-Fisk) and students
(Adkins and Spaulding) in the Department of
Geography at the University of Wyoming;
we appreciate the use of laboratory facilities
there. We are also indebted to several local
ranchers in the Laramie Basin for permission
to work on their property and for various
types of information and to Richard G. Rei-
der, Brainerd Mears, Jr., and Thomas P. Har-

lan for helpful suggestions. Tree-ring mea-
surements were done by Elliott-Fisk at the
Laboratory of Tree-Ring Research, Univer-
sity of Arizona; the use of this facility is very
much appreciated. We also thank the many
students and faculty who have accompanied
us in our field work at Sand Creek.

Literature Cited

Elliott, D. L. 1979. The stability of the northern Cana-
dian tree limit: current regenerative capacity.
Unpublished dissertation. Univ. of Colorado,
Boulder. 192 pp.

FowELLS, H. A. 1965. Silvics of forest trees of the United
States. USDA Agr. Handbk. 271. Washington,
D.C. 762 pp.

Fritts, H. C. 1976. Tree rings and climate. Academic
Press, New York. 567 pp.

Grasso, D. N. 1979. Paleoclimatic significance of fossil
ice-wedge polygons in the Laramie Basin,
Wyoming. Unpublished thesis. Univ. of Wyo-
ming, Laramie, 105 pp.

Hager, M. W. 1972. A late Wisconsin-Recent vertebrate
fauna from the Chimney Rock animal trap, Lari-
mer County, Colorado. Contrib. to Geol.
2:6.3-71. (Univ. of Wyoming, Laramie).

Mears, B., Jr. 1981. Periglacial wedges and Late Pleisto-
cene environment of Wyoming's intermontane
basins. Quat. Res. 15:171-198.

384

Great Basin Naturalist

Vol. 43, No. 3

Reider, R. G., N. J. KuNiANSKY, D. M. Stiller, and P. J.
Uhl. 1974. Preliminary investigation of com-
parative soil development on Pleistocene and
Holocene geomorphic surfaces of the Laramie
Basin, Wyoming. Pages 27-.33 in M. Wilson, ed..
Applied geology and archaeology: the Holocene
history of Wyoming. Geol. Surv. of Wyoming,
Rept. of Investigations 10.

Stokes, M. A., and T. L. Smiley. 1968. An introduction
to tree-ring dating. Univ. of Chicago Press, Chi-
cago. 73 pp.

Wells, P. V. 1970a. Postglacial vegetational history of
the Great Plains. Science 167:1574-1582.

1970b. Vegetational history of the Great Plains: a

post-glacial record of coniferous woodland in
southeastern Wyoming. Pages 185-202 in W.
Dort and K. J. Jones, eds.. Pleistocene and Recent
environments of the Central Great Plains. Univ.
of Kansas Press, Lawrence.

1976. Macrofossil analysis of woodrat (Neotoma)

middens as a key to the Quaternary vegetational
history of arid America. Quat. Res. 6:223-248.

COMPARATIVE LIFE HISTORY AND FLORAL CHARACTERISTICS
OF DESERT AND MOUNTAIN FLORAS IN UTAH

Patrick D. Collins', Kimball T. Harper', and Burton K. Pendleton-

Abstract.— Life forms and floral characteristics of plants at Arches National Park (desert communities), the Mt.
Nebo complex, and a subalpine meadow in the Uinta Mountains (montane and subalpine communities) were com-
pared. Characteristics observed were (1) life form, (2) longevity, (3) pollination system, (4) flower structure, (5) flower
symmetry, and (6) flower color. Common families in each flora were also compared. Results showed that there is a
significant overrepresentation of shnib species at Arches, and an underrepresentation of perennial forbs. Relative
number of perennial forb species was significantly higher at Mt. Nebo and the subalpine meadow than at Arches Na-
tional Park. Native annuals and wind-pollinated species were significantly overrepresented at Arches. Flowers with
open stnicture that permit free access of most insects to nectar and pollen were overrepresented at Mt. Nebo and in
the subalpine meadow. The distribution of flower colors also differs significantly among these ecologically con-
trasting floras, with yellow being best represented in the desert and white in the mountains.

Because of accelerating development of
energy and other natural resources, plant
communities of the world are constantly
being altered. In the United States, law re-
quires that such disturbed areas be restored
to their natural condition (Public Law 95-87,
1977), but little is known of the relative pro-
portion of life histories and floral character-
istics that enhance coexistence and self-
perpetuation of a variety of wild plant spe-
cies on common sites. What controls the rela-
tive success of pollen transfer by wind or ani-
mal in various natural communities? Do the
contrasting climatic conditions of certain en-
vironments affect the success of species of
various life forms, longevity, and/or floral
characteristics?

In this paper, we compare characteristics
of three Utah floras: the floras are from
Arches National Park (desert), the Mt. Nebo
complex (midelevation montane vegetation),
and a subalpine meadow at high elevation in
the Uinta Mountains. Arches National Park is
a semiarid, cold desert region in southeastern
Utah with an average elevation of about
1,220 m above sea level. Its topography con-
sists of rolling hills and sandstone outcrops.
The mean annual precipitation at Moab, near
Arches, was 21.7 cm with a standard devia-
tion of 4.3 cm (Nat. Oceanic and Atmosph.

Admin. 1971-1979). The average annual tem-
perature was 13.5 C. Mt. Nebo and adjacent
mountains form a montane habitat with ele-
vations between 1,829 and 3,621 m, but aver-
age elevation is in the neighborhood of 2,500
m. Average annual temperatures at Tim-
panogos Cave (1,720) 73.3 km north of Mt.
Nebo was 9.4 C. The mean annual precipi-
tation at Timpanogos Cave was 55.3 cm for
the 1971-1979 period and that at the Payson
Guard Station (2,454 m) was 73.3 cm with a
standard deviation of 11.9 cm (Whaley and
Lytton 1978). The Payson Guard Station is
17.4 km north of the crest of Mt. Nebo. The
series of subalpine meadows selected for this
study are located just below Bald Mountain
Pass at the 28-mile marker (45.1 km) on the
Mirror Lake Highway (Utah Highway 150) in
Summit County, Utah. Average elevation at
these meadows is 3,216 m, whereas mean an-
nual precipitation at Trial Lake (near the
meadows) was 102.8 cm with a standard de-
viation of 18.3 cm (Whaley and Lytton
1978). The mean annual temperature of this
site is approximately -2.4 C (Callison and
Harper, in review).

Jaccard's community coefficient (1912)
shows the Arches and Nebo floras to be 90
percent dissimilar, Arches and Bald Mountain
98 percent dissimilar, and the Nebo and Bald

'Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602.
'Department of Biological Sciences, Wayne State University, Detroit, Michigan 48202.

385

386

Great Basin Naturalist

Vol. 43, No. 3

Mountain floras 90 percent dissimilar. Be-
cause these floras are so dissimilar and oc-
cupy such physically different ecological situ-
ations, it was hypothesized that there would
be statistically significant differences in re-
productive strategies of the plant species of
the three floras. This paper compares the dis-
tribution of life form, longevity, and floral
characteristics in the three floras. The com-
parisons evaluate the relative success of vari-
ous reproductive strategies of plant species
native to these three contrasting envi-
ronments.

Methods

Floristic checklists furnish the data on
which this study is based. The list for Mt.
Nebo and adjacent mountains, Utah County,
was compiled from Collins (1979) and Allred
(1975). Checklists for Arches National Park
were prepared by Harrison et al. (1964) and
Allan (1977). The Bald Mountain meadow
checklist was taken from Pendleton (1981).

Life history and floral characteristics were
determined for all species from preserved
specimens in the Brigham Young University
Herbarium. There were 734 species (647 na-
tive; 12.0 percent introduced) on the Mt.
Nebo complex checkhst, 356 (322 native; 9.6
percent introduced) on the desert list (Arches
National Park), and 134 (all native) on the
high elevation meadow list (Bald Mountain).
The following information was obtained for
each of the native species: (1) longevity, (2)
life form, (3) likely pollinating agent, (4)
flower symmetry, (5) flower structure, and (6)
flower color. All analyses reported in this pa-
per are based on native species only.

Longevity was simply recorded as annual
or perennial. Species described in keys as
biennials were treated here as annuals, except
those species listed as "biennials to short-
lived perennials" were considered perennials.
Plant life form was noted as tree, shrub, forb,
or grass. In respect to pollination system,
plant species were classified as anemophilous
or zoophilous. It is realized that some of the
species may be self-pollinated, but this could
not be determined without independent re-
search on each species. Thus, no attempt was
made to identify self-pollinated taxa.

Flower symmetry was regarded as either
zygomorphic or actinomorphic. Species were
also classified according to flower structure.
Structure of zoophilous flowers was de-
scribed as restricted when access to nectar or
pollen was difficult for unspecialized pollina-
tors. Restricted access flowers had long co-
rolla or calyx tubes or had nectaries at the
base of long spurs, thus limiting access to pol-
len or nectar. Flowers classified as open-
structured were saucer or bowl shaped and
appeared incapable of mechanically exclud-
ing any pollinator. Moreover, some plants
were considered to be only partially restric-
tive, having short calyx tubes or deeply lobed
and/or widely flaring sympetalous corollas:
such flowers were listed as semi-restricted.
Flower colors were listed as red, violet, blue,
yellow, pink, white, or greenish.

Important families for each location were
summarized by the number of species found
in each flora. Jaccard's similarity index was
employed to test compositional similarity be-
tween floras. The Chi-square statistic was
used to identify departures from random ex-
pectations. In the Chi-square analyses, ran-
dom expectations are based on the propor-
tion of the species in the pooled floras that
share a particular trait (e.g., the proportion
of the species in the combined floras from the
desert and montane environments that have
red flowers). If the trait is randomly dis-
tributed between the two floras, the propor-
tion of species having the trait in each flora
should not differ significantly from the pro-
portion having that trait in the pooled flora.
The Chi-square statistic was used to test
whether the observed and expected numbers
of species (tests use absolute numbers, not the
proportions) sharing a trait in the individual
floras differed significantly. If there was a
significant departure, the trait was consid-
ered to be under- or overrepresented in a giv-
en flora. Introduced species were omitted
from all analyses on the assumption that they
may not have achieved stable reproductive
characteristics in their new home.

Results and Discussion

A total of 70 families, 307 genera, and 734
species occurred in the Mt. Nebo flora. There
are 60 families, 203 genera, and 356 species

July 1983

Collins et al.: Desert and Mountain Floras

387

reported for the Arches flora. Thirty-six fami-
hes, 86 genera, and 134 species appear on the
Bald Mountain meadows checklist. Jaccard's
index (Jaccard 1912) shows the Arches and
Nebo floras to be 10.0 percent similar on the
basis of species, 32.5 percent similar on the
basis of genera, and 75.5 percent similar on
the basis of families. The Arches and Bald
Mountain floras are 1.5 percent similar on
the basis of species, 12.0 percent similar on
the basis of genera, and 39.1 percent similar
on the basis of families. The Mt. Nebo and
Bald Mountain floras show 10.2 percent sim-
ilarity by species, 23.2 percent by genera,
and 45.2 percent by families. Because the
families contributing species to these flora
are so similar, most of the observed differ-
ences in plant adaptations in the three areas
can be attributed to ecological selection
rather than to differences in basic phylogeny.
Families contributing most of the species
in the three floras are reported in Table 1.
The families Asteraceae and Poaceae domi-
nate the three floras: Fabaceae holds third
place in the Arches flora, Cyperaceae claims
that position at Bald Mountain, and Brassi-
caceae takes that slot on Mt. Nebo. The fam-
ily Chenopodiaceae contributes over three
times as many species in relative terms in the
desert as in the mountains. In contrast, spe-
cies of Rosaceae, Caryophyllaceae, Polemo-
niaceae, Saxifragaceae, and Scrophulariaceae
are twice or more as common in our moun-
tains as in the desert flora considered.

Life Form.— In all three floras, the pre-
dominant life form (as represented by num-
ber of species) is the broad-leaved herb (forb).
At Arches National Park, 64.3 percent of the
total flora is contributed by forbs; on Mt.
Nebo, 73.0 percent of the species are forbs;
and at Bald Mountain, 64.9 percent of the
species are forbs (Table 2). Analysis shows
that forbs are significantly overrepresented in
the midmontane flora, whereas they are un-
derrepresented in the high-elevation mead-
ows and in deserts (Table 3). The shrub life
form contributes proportionally over twice as
many species in the desert (14.3 percent) as
in the mountain floras at Mt. Nebo (7.0 per-
cent) and at Bald Mountain (6.8 percent).
That difference is statistically highly signifi-
cant (2X2 = 30.46, P < 0.005, Table 3A).

The results support the hypothesis that un-
der dry conditions, shrubs are more successful
than forbs. Deserts are notorious for unpre-
dictable climatic patterns, and many forbs do
not tolerate moisture deficits for long periods
(Hironaka 1963, Mueggler 1972, Harner and
Harper 1973). Shrubs can tolerate such con-
ditions. They exhibit a variety of adaptations
to dry environments, such as deep root sys-
tems and reduced reliance on turgor pressure
to keep leaves expanded to collect light and
carbon dioxide (Sharif and West 1968).
Shrubs also have leathery or firm leaves that
reduce breakage from heavy winds and are
resistant to herbivory; and, finally, shrubs
have well-developed secondary meristems

Table 1. A comparison of important families showing the number of species and percent (in parenthesis) of thg
total floras of Arches National Park, the Mt. Nebo complex, and Bald Mountain meadows, Utah.

Family

Arches

No. (%)

Mt. Nebo

No. (%)

Bald Mtn.

No. (%)

Asteraceae

Poaceae

Fabaceae

Chenopodiaceae

Brassicaceae

Scrophulariaceae

Boraginaceae

Cyperaceae

Rosaceae

Liliaceae

Caryophyllaceae

Ranunculaceae

Polemoniaceae

Saxifragaceae

Salicaceae

Other

Totals

(22.5)

(14.3)

(7.0)

(6.2)

(4.8)

(2.8)

(2.5)

(2.0)

(0.6)

(1.1)

(1.4)

(0.3)

(1.7)

100

(28.0)

115

(15.7)

105

(14.3)

(4.1)

(1.9)

(6.8)

(4.9)

(2.5)

(2.2)

(4.6)

(1.9)

(2.5)

(3.8)

(2.6)

(2.5)

(2.0)

203

(27.7)

356(100.0)

734 (100.0)

(14.9)

(13.4)

(1.5)

(0.0)

(3.0)

(6.0)

(0.7)

(9.0)

(3.0)

(4.5)

(3.7)

(0.7)

(2.2)

(0.7)

(33.6)

134 (

100.0)

388

Great Basin Naturalist

Vol. 43, No. 3

that probably permit individual roots to be
longer lived than is possible for species that
lack secondary meristems (as in grasses and
sedges). During dry periods, shrubs persist
and maintain root systems in both lateral and
vertical space; when better moisture condi-
tions do return, herbs attempting to colonize
barren spaces between shrubs experience ex-
treme competition from the already estab-
lished root systems of shrubs. Even in moist
years, however, the barren interspaces be-
tween shrubs are only sparsely clothed with
annual plants, but nearby areas that have
been deprived of their shrub cover by abu-
sive grazing or mechanical disturbance sup-

port a nearly complete cover of annual plants
(Hutchings and Stewart 1953).

Graminoides tend to be better represented
in the Arches flora (grasses) and at Bald
Mountain (sedges) than at Mt. Nebo. Trees
are best represented in the midelevation
mountain flora (Table 2).

Longevity.— Longevity of species in the
desert and montane floras also show signifi-
cant differences. The deserts have more an-
nual species than one would expect by
chance (2X2 = £1.27, P < 0.005, Table 3B).
Native annuals contribute 18.9 percent of the
322 species at Arches National Park, 10.5
percent of the 647 species on the Mt. Nebo

Table 2. Characteristics of the native floras of Arches National Park, the Mt. Nebo complex, and the Bald Moun-
tain meadows. The table shows the number of species and the percent of the native flora (in parentheses).

Arches

Mt. Nebo

Bald Mtn.

No. Sp. (% flora)

Life form

Trees

13 (4.0)

39 (6.0)

3 (2.2)

Shrubs

46 (14.3)

45 (7.0)

9 (6.7)

Forbs

207 (64.3)

472 (73.0)

87 (64.9)

Graminoides

56 (17.4)

91 (14.0)

35 (26.1)

Total

322 (100.0)

647 (100.0)

134 (100.0)

Longevity

Annual

61 (18.9)

68 (10.5)

7 (5.2)

Perennial

261 (81.1)

579 (89.5)

127 (94.8)

Total

.322 (100.0)

647 (100.0)

134 (100.0)

Pollination system (annuals

excluded)

Anemophilous

101 (38.7)

162 (28.0)

39 (30.7)

Zoophilous

160 (61.3)

417 (72.0)

88 (69.3)

Total

261 (100.0)

579 (100.0)

127(100.0)

Flower symmetry

(zoophilous species)

Radial

188 (88.7)

395 (84.4)

77 (83.7)

Bilateral

24 (11. .3)

73 (15.6)

15 (16.3)

Total

212 (100.0)

468 (100.0)

92 (100.0)

Flower structure (zoophilous species)

Open

55 (25.9)

179 (38.2)

42 (45.7)

Restricted

59 (27.8)

128 (27.4)

14 (15.2)

Semirestricted

98 (46.2)

161 (34.4)

36 (39.1)

Total

212 (100.0)

468 (100.0)

92 (100.0)

Flower color (zoophilous species)

White

50 (23.6)

165 (35.3)

30 (32.6)

Yellow

88 (41.5)

148 (31.6)

23 (25.0)

Blue

20 (9.4)

57 (12.2)

8 (8.7)

Violet

27 (12.7)

36 (7.7)

12 (13.0)

Pink

8 (3.8)

34 (7.2)

12 (13.0)

Red

14 (6.6)

15 (3.2)

1 (1.1)

Green

4 (1.9)

8 (1.7)

5 (5.4)

Other

1 (0.5)
212 (100.0)

5 (1.1)

1 (1.1)

Total

468 (100.0)

92 (100.0)

July 1983

Collins et al.: Desert and Mountain Floras

389

complex, and only 5.2 percent of the 134 spe-
cies from the Bald Mountain flora.

Climatic unpredictability enhances the
success of annuals in deserts (Schaffer and
Gadgil 1975), where precipitation is sporadic
and scarce. The annual strategy seems well
suited for such conditions, whereas perennial
forbs consistently contain high levels of tissue
moisture (Sharif and West 1968). Our results
show that perennial forbs are under-
represented in the desert (Table 3C).

Pollination Systems.— Because shrubs have
been shown to be overrepresented at Arches,
and because anemophily is heavily favored
among woody species (Ostler and Harper
1978, Freeman et al. 1980), we anticipated
that anemophily would be most prevalent at

Arches. At Arches National Park, 38.7 per-
cent of the perennial flora is anemophilous;
on Mt. Nebo only 28.0 percent and at Bald
Mountain only 30.7 percent of the species
are wind pollinated (annuals have been
omitted from this analysis to minimize the
possible confusing effect of self-pollinated
species, which are believed to be especially
common among annuals [Solbrig 1977]). The
differences in modes of pollination in the
three floras are statistically significant (SX^
= 9.64, P < 0.005, Table 3D).

The reason that there are more anemo-
philous species in the desert is not that there
is more wind movement there. The Arches
area receives only half as much wind (1,590.5
km at Moab) as the Mt. Nebo area (2,984.4

Table 3. Chi-square analyses comparing life histories and floral characteristics of the native plant species of
Arches National Park, the Mt. Nebo complex, and the Bald Mountain complex of Utah. Observed and expected
numbers of species (in parentheses) are shown. Asterisks indicate significance level: single < .05, double < .010,
triple < .005 probability.

Summation

Chi-square

Arches

Mt. Nebo

Bald Mtn.

values

Life form

Trees

13 (16.1)

39 (32.3)

I (6.7)

Shrubs

46 (29.2)

45 (58.7)

(12.1)

Forbs

207 (223.6)

472 (449.2)

(93.1)

Grasses

56 (53.1)

91 (106.8)

(22.1)

30.46°°°

Growth cycle

Annuals

61 (39.7)

68 (79.8)

(16.5)

Perrenials

261 (282.3)

579 (567.2)

127

(117.5)

21.27°°°

Growth cycle/life form

Perennial forbs

150 (184.5)

406 (370.7)

(76.8)

Other native spec

•ies

172 (137.5)

241 (276.3)

(57.2)

22.99°°° •

Pollination system (annuals (

excluded)

Anemophilous

101 (81.5)

162 (180.8)

(39.7)

Zoophilous

160 (179.5)

417 (398.2)

(87.3)

9.64°°°

Flower structure

(for zoophilous species)

All species considered (

annuals included)

Open

55 (75.8)

179 (167.3)

(32.9)

Restricted

59 (55.2)

128 (121.8)

(24.0)

Semirestricted

98 (81.0)

161 (178.8)

(35.2)

19.11°°°

Only perennial species

considered (semirestricted taxa

ignored)

Open

46 (59.2)

160 (156.6)

(28.2)

Restricted

61 (47.8)

123 (126.4)

(22.8)

14..35°°°

Flower color (for

zooph

ilous

species)

Red

14 (8.2)

15 (18.2)

(3.6)

Other

198 (203.8)

453 (449.8)

(88.4)

6.70°

White

50 (67.3)

165 (148.5)

(29.2)

Other

162 (144.7)

303 (319.5)

(62.8)

9.21°°

Yellow

88 (71.1)

148 (157.0)

(30.9)

Other

124 (140.9)

320(311.0)

(61.1)

9.82°°

390

Great Basin Naturalist

Vol. 43, No. 3

km at Lehi) in the April-September period
(Whaley and Lytton 1979). Conditions that
may favor wind-pollinated species at Arches
include dominance of most perennial covers
by a few woody species that have large pop-
ulations, low-growing, open vegetation, and
severe, unpredictable periods of drought. Ac-
curate wind movement readings were not
available for the Bald Mountain area.

When only perennial species are consid-
ered, woody species are much better repre-
sented in the Arches flora (22.6 percent) than
at Mt. Nebo (14.5 percent) or Bald Mountain
(9.4 percent). Furthermore, shrubs dominate
all major communities at Arches (Allan
1977). Diversity of perennial species as mea-
sured by number of species per 1.0 m^ (a var-
iable known to reduce the success of wind-
pollinated taxa [Ostler et al. 1982]) is shown
by Allan (1977) to be 2.6 at Arches (10 com-
munities considered) as compared with 4.3 in
the Wasatch Moimtains (of which Mt. Nebo
is a part [Ostler and Harper 1978, 25 com-
munities reported]) and 6.9 at Bald Mountain
(Pendleton 1981, 4 communities reported).
Wind pollination is further facilitated at
Arches by a plant cover that is more open
than that at Nebo. Allan (1977) reported an
average of 41.3 percent living cover at
Arches, but, considering the fact that 70 per-
cent of the Mt. Nebo study area is dominated
by oak woodland or forests of aspen and/ or
conifer, plant cover there undoubtedly aver-
ages well over 65 percent (see Allan 1962,
Crowther and Harper 1965, Kleiner 1966,
and Harper 1981 for cover estimates for sim-
ilar vegetations in northern Utah). Vegetative
cover in subalpine meadow in the Uinta
Moimtains averages about 76 percent (Ostler
et al. 1982). Finally, annual precipitation is
more likely to fall below a level sufficient to
support flowering of many species at Arches
than in the mountain study areas. Frequent
years of sparse or no flowering distributed at
random through time should reduce the like-
lihood that insect pollinators can maintain
large enough populations to pollinate all the
flowers produced in years of adequate soil
moisture. Wind-pollinated species should be
favored in such situations provided individual
plants are large enough to intercept a re-
liable flow of air, foliage cover is not so dense

that it seriously interferes with pollen move-
ment in the wind, and conspecific individuals
are close enough together to insure that most
stigmas will receive pollen. On all counts, the
Arches area is better suited for wind pollina-
tion than the two mountain locations.

Flower Structure.— If reproduction of
animal-pollinated species is to be successful,
floral structure should encourage the likeli-
hood of sequential visits by specific pollina-
tors. When flowers of coexisting species com-
pete for pollinators, species having flowers
that mechanically exclude many kinds of pol-
linators should be able to conserve more nec-
tar or pollen for adapted visitors than species
whose flowers can be worked by any visitor.
Thus visits by such adapted pollinators
should be reinforced by more dependable
nectar or pollen rewards, thus encouraging
the pollinator to seek out other flowers of the
same type. As a result, flowers with restricted
access should be at a reproductive advantage
in diverse assemblages of plants that flower
simultaneously.

Ostler and Harper (1978) show that flowers
that have restricted access to the nectar
and/ or pollen supply are positively corre-
lated with the diversity of animal-pollinated
species per unit area in the Wasatch Moun-
tains of Utah and Idaho. Thus, one might be
tempted to hypothesize that, because diver-
sity is lower at Arches than in Utah moun-
tains as noted above, one could expect an
overrepresentation of open-structured flow-
ers in the desert. The data show, however,
that when all species are considered, the
mountain floras have relatively more species
with open flowers than the desert (ZX^ =
19.11, P < 0.005, Table 3E). Even among
perennial species only, that pattern continues
to hold (2X2 = 14.35, P < 0.005, Table 3E).

The difference in flower structure among
the floras of Arches, Bald Mountain, and Mt.
Nebo may be attributable to differences in
flowering phenology. Species must flower
when moisture conditions are favorable. In
the desert, moisture conditions for most spe-
cies are apparently optimal in the spring, be-
cause it is at that time that most desert spe-
cies flower. Accordingly, although there are
fewer species per imit area in the desert,
more species may actually flower simultane-
ously than in the mountain zone. Thus, at

July 1983

Collins et al.: Desert and Mountain Floras

391

certain times, there may be greater com-
petition among plant species for pollinating
animals in the desert than in at least mid-
elevation moimtains.

In contrast to deserts, midelevation moun-
tain commimities have favorable moisture
conditions throughout much of the growing
season. It is therefore possible for coexisting
species to partition the available time by
flowering out of synchrony. Such out-of-
phase flowering should decrease competition
for pollinators and allow for more open flow-
ers (Mosquin 1971). This argument probably
does not hold at high elevations where grow-
ing seasons are short and flowering of all spe-
cies is confined to that brief season. In such
environments, simultaneous flowering of
many species is undoubtedly commonplace.
The profusion of open-flowered taxa in the
subalpine meadows (Table 2) thus cannot be
attributed to low flowering-plant diversity.

It has become clear in recent years that
pollinating insect faunas are larger and more
diverse in warmer and lower elevation envi-
ronments than in cold and/ or high elevation
habitats (Arroyo et al. 1982, Warren et al.
1982). Hymenopteran insects especially ap-
pear to decline in colder and higher elevation
communities, and Dipteran and Lepidopte-
ran pollinators become relatively more im-
portant. It has long been recognized that
Hymenopteran pollinators are the most ef-
ficient of all insects at manipulation of com-
plex flowers (Faegri and Pijl 1971). It is thus
possible that the observed predominance of
open flowers in our mountain floras is related
to changes in composition of the pollinator
fauna along the altitudinal gradient. Open
flowers may be favored when there is com-
petition among flowers for an impoverished
and less efficient guild of pollinating insects.

Flower Symmetry.— The native flora of
Arches National Park consists of 88.7 percent
radially symmetrical and 11.3 percent zygo-
morphic-flowered species. The Mt. Nebo
flora includes 84.4 percent radial and 15.6
percent zygomorphic flowers. Finally, the
Bald Mountain Flora consists of 83.7 percent
radial and 16.3 percent zygomorphic flower
(Table 2). Although differences between the
three floras are not statistically significant
(2X2 = 2.42), there is a trend toward a great-
er nmnber of zygomorphic flowers as eleva-
tion and diversity increases. Zygomorphic

structure is believed to force insects to ap-
proach flowers in more stereotyped ways.
Thus, in zygomorphic flowered taxa, muta-
tions that result in loss of stamens and stigmas
off the regular access route of the insect to
the floral reward (nectar and/or pollen)
could be accommodated without loss in fe-
cundity. In harsh environments where car-
bohydrate gains are marginal, zygomorphy
and the energy economy associated with re-
duced numbers of reproductive parts and
lower pollen production could be expected to
have selective advantages.

Flower Color.— Is there any difference in
the distribution of colors among these floras?
Ostler and Harper (1978) showed that floral
color diversity was positively correlated with
species diversity of the communities studied.
Our data demonstrate that red and yellow
flowers are overrepresented at Arches Na-
tional Park (Table 3F), with the differences
being statistically significant for both (P <
0.05 or better). White flowers are significant-
ly overrepresented at Mt. Nebo and Bald
Mountain. White seems to be favored in
moist and wooded environments (Ostler and
Harper 1978, Del Moral and Standley 1979),
whereas yellow flowers are consistently bet-
ter represented in dry environments (Kevan
1972, Tikhomirov 1966, and Weevers 1951).
The results for white flowers were expected
in the Nebo flora (but not at Bald Mountain
meadow), because white color had been
shown to be more abundant in forest under-
stories by both Ostler and Harper (1978) and
Del Moral and Stanley (1979). The Mt. Nebo
complex is largely dominated by forest or
woodland commimities, but there is only a
minor amount of woodland at Arches Nation-
al Park and Bald Mountain. Ostler and Har-
per (1978) speculated that white flowers re-
flected more light and were more easily
found by pollinators in forest understories.
Baker and Hurd (1968) noted that moths tend
to replace bees as major pollinators in shaded
habitats, and moths show a preference for
white flowers. Finally, it is possible that the
insect eye, like that of the human, is unable
to perceive color at low light intensities
(Proctor and Proctor 1978). Thus, the insect
may react to flower color in shaded environ-
ments in terms of contrast alone rather than
in terms of different hues per se.

392

Great Basin Naturalist

Vol. 43, No. 3

The abundance of white flowers in the sub-
alpine meadows cannot be explained by the
foregoing argument. At this point, we can
only hypothesize that the importance of
white flowers and the diminished abundance
of yellow-flowered species in high elevation
meadows is somehow related to the cooler,
more moist environment or to altered com-
position of the pollinating insect community
(or both) at high elevations.

The proportionally greater number of red
and, to a lesser extent, yellow flowers in the
Arches flora seems attributable to the near-
ness of the park to areas of high diversity of
hummingbird species in Arizona (up to nine
species, see Crosswhite and Crosswhite 1981).
Hummingbirds are believed to have been the
selective force responsible for the evolution
of many orange- and red-flowered species in
the American Southwest (Crosswhite and
Crosswhite 1981). At the mountain sites, such
"hot"-colored species are relatively less com-
mon (Table 2). Only four species of hum-
mingbirds occur in the Mt. Nebo area, and
three hummingbird species occur in-
frequently at the high subalpine meadows
considered here.

Conclusion

The results demonstrate that there are def-
inite differences in the distribution of plant
longevity, life form, mode of pollination, flor-
al structure, and flower colors in the desert
and moujitain floras compared in this study.
Annuals and shrubs are overrepresented in
the Arches National Park flora, and zoophily
is significantly more abundant in the moun-
tain floras. Open-structured flowers are sig-
nificantly overrepresented on the mountain
floras. White flowers are most common in
the mountain floras, whereas red and yellow
flowers are best represented in the desert.

Acknowledgments

Sincere appreciation is extended to C. Da-
vidson, Arthur Holmgren, and others who re-
viewed this manuscript and made valuable
suggestions. This paper was funded in part by
the U.S. Forest Service, Uinta National For-
est, Provo, Utah.

Literature Cited

Allan, J. S. 1977. The plant communities of Arches Na-
tional Park. Unpublished dissertation. Brigham
Young Univ., Provo, Utah.

1962. The plant communities of the Big Cotton-
wood Canyon drainage. Unpublished thesis.
Univ. of Utah. Salt Lake City.

Allred, K. W. 1975. Timpanogos flora. Unpublished
thesis. Brigham Young Univ., Provo, Utah.

Arroyo, M. T. K., R. Primack, and J. Armesto. 1981.
Community studies in pollination ecology in the
high temperate Andes of central Chile, I. Pollina-
tion mechanisms and altitudinal variation. Amer.
J. Bot. 69:82-97.

Baker, H. C, and P. H. Hurd, Jr. 1968. Intrafloral ecol-
ogy. Ann. Rev. Entomol. 13:511-523.

Callison, J., and K. T. Harper. In review. Temper-
ature in relation to elevation in the Uinta Moun-
tains, Utah. Brigham Young Univ., Provo, Utah.

Collins, P. D. 1979. Checklist of the plants on Mt.
Nebo and adjacent mountains. Unpublished re-
port to the Uinta National Forest, Provo, Utah.

Collins, P. D., and K. T. Harper. 1980. Vegetation
map of the Mt. Nebo complex and San Pitch
Mountains, Utah. BYU Printing Service, Provo,
Utah.

Crosswhite, F. S., and C. D. Crosswhite. 1982. Hum-
mingbirds as pollinators of flowers in the red-
yellow segment of the color spectrum, with spe-
cial reference to Pensteinon and the "open habi-
tat." Desert Plants 3:156-170.

Crowther, E. C, and K. T. Harper. 1965. Vegetational
and edaphic characteristics associated with aspen
"strips" in Big Cottonwood Canyon. Proc. Utah
Acad. Sci., Arts, Lett. 42(II):222-230.

Del Moral, R., and L. A. Standley. 1979. Pollination
of angiosperms in contrasting coniferous forests.
Amer. J. Bot. 66:26-35.

Faegri, K., and L. van der Pijl. 1971. The principles of
pollination ecology, 2d ed. Pergamon Press,
Oxford.

Freeman, D., K. T. Harper, and W. K. Ostler. 1980.
Ecology of plant dioecy in the intermountain re-
gion of western North America and California.
Oecologia 44:410-417.

Harner, R. F., and K. T. Harper. 1973. Mineral compo-
sition of grassland species in the eastern Great
Basin in relation to stand productivity. Canadian
J. Bot. 51:2037-2046.

Harper, K. T. (ed.) 1981. Potential ecological impacts of
snowpack augmentation in the Uinta Mountains,
Utah. Final report to the Bureau of Reclamation,
Engineering and Research Center, Federal Cen-
ter, Denver, Colorado.

Harrison, B. F., S. L. Welsh, and G. Moore. 1964.
Plants of Arches National Monument. BYU Sci.
Bull., Biol. Ser. 5(1): 1-23.

HiRONAKA, M. 1963. Plant environment relations of ma-
jor species in sagebrush-grass vegetation of south-
ern Idaho. Unpublished dissertation. Univ. of
Wisconsin, Madison.

Hutchings, S. S., and G. Stewart. 1953. Increasing for-
age yields and sheep production on inter-
mountain winter ranges. USDA Circ. No. 925.

July 1983

Collins et al.: Desert and Mountain Floras

393

Jaccard, p. 1912. The distribution of the flora of the al-
pine zone. New Phytol. 11:37-50.

Kevan, p. G. 1972. Floral colors in the high arctic with
reference to insect-flower relations and pollina-
tion. Canadian J. Bot. 50:2289-2316.

Kleiner, E. F. 1966. A study of the vegetational com-
munities of Red Butte Canyon, Salt Lake County,
Utah. Unpublished thesis. Univ. of Utah, Salt
Lake City.

MosQUiN, T. 1971. Competition for pollinators as a stim-
ulus for the evolution of flowering time. Oikos
22:398-402.

MuEGGLER, W. F. 1972. Plant development and yield on
mountain grasslands in southwestern Montana.
U.S. Forest Serv. Paper INT-124.

National Oceanic and Atmospheric Administration.
1971-79. Annual .summaries for Utah. Climatolo-
gical Data 73-81(13):pp. variable.

Ostler, W. K., and K. T. Harper. 1978. Floral ecology
in relation to plant species diversity in the
Wasatch Mountains of Utah and Idaho. Ecology
59:848-861.

Ostler, W. K., K. T. Harper, K. B. McKnight, and D.
C. Anderson. 1982. The effects of increasing
snowpack on a subalpine meadow in the Uinta
Mountains, Utah, USA. .\rctic and Alpine Res.
14:20.3-214.

Pendleton, B. K. 1981. Aspects of the floral biology of a
subalpine meadow of the Uinta Mountains, Utah.
Unpublished thesis. Brigham Young Univ., Provo,
Utah.

Proctor, J., and S. Proctor. 1978. Color in plants and
flowers. Everest House Publishers, New York.

ScHAFFER, VV. M., and M. D. Gadgil. 1975. Selection
for optimal life histories in plants. Pages 142-157
in M. L. Cody and J. M. Diamond, eds.. Ecology
and evolution of communities. Belknap Press,
Cambridge, Massachusetts.

Sharif, C. M., and N. E. West. 1968. Forage moisture
variations on mountain summer range. J. Range
Mgt. 21:228-2.35.

SoLBRiG, O. T. 1977. On the relative advantages of cross-
and self-fertilization. Ann. Missouri Bot. Card.
63:262-276.

TiKHOMiRov, B. A. 1966. The interrelationships of the
animal life and vegetational cover of the tundra.
Academy of Sciences of the USSR, Botanical In-
stitute (Translated from Russian by the Israel
Program for Scientific Translation, Jerusalem,
1966).

U.S. Congress. Public Law 95-87. 91 Stat. 445. 95th
Cong., 3 Aug. 1977.

Warren, S. D., G. M. Booth, and K. T. Harper. 1982.
Elevational distribution of insect pollinators. Un-
published manuscript.

Whaley, B. L., and L. Lytton. 1979. Summary of pre-
cipitation measurements for Utah, 1930-1978.
Report distributed by the Snow Survey Section,
Soil Conservation Service, USDA, Salt Lake City,
Utah. 168 pp.

Weevers, T. H. 1952. Flower colours and their frequen-
cy. Acta Bot. Neerl. 1:81-92.

FLORA OF THE LOWER CRETACEOUS CEDAR MOUNTAIN FORMATION

OF UTAH AND COLORADO, PART I.

PARAPHYLLANTHOXYLON UTAHENSE

G. F. Thayne', W. D. Tidwell", and W. L. Stokes'

Abstract.— ParapItyUantJwxijIon utahense, sp. nov., is described from the Cedar Mountain Formation and com-
pared with similar fossil and modern woods. Fossil angiosperm woods from the Early Cretaceous are of great interest
because very few have been reported from strata of this age. This species demonstrates that the angiosperms had de-
veloped many of their modern characteristics by Early Cretaceous time.

The Lower Cretaceous Cedar Mountain
Fonnation is fossiliferous at several localities.
Fossils reported from this formation include
the wood of conifers, Tempskya, and cyca-
deoids, charophytes, pelecypods, gastropods,
ostracods, and fish scales (Stokes 1952, Young
1960, Thayn et al. 1973, Tidwell et al. 1976),
as well as dinosaur bones (Bodily 1969).

A species of dicotyledonous wood assigned
to the genus Paraphyllanthoxylon Bailey
1924, is described in this report from the Ce-
dar Moimtain Formation. This is the first re-
port of petrified dicotyledonous wood from
the diverse flora in this formation. These
angiosperm woods are of great interest in
that very few Early Cretaceous angiosperm
woods have been previously reported. Since
the Cretaceous Period is the assumed time
for the origin of the angiosperms, a tax-
onomic study of Early Cretaceous angio-
sperm wood is significant in that it expands
oiu- knowledge of the early members of this
division. The petrified wood described in this
study was collected from two localities. Lo-
cality 1 is 6 road miles (3.7 km) east of Castle
Dale, Utah, and Locality 2 is 9 road miles
(5.6 km) east of Ferron, Utah (Figs. 1, 5, 6, 7).

The Cedar Mountain Formation at Local-
ity 1 is composed of brown to grey shales. It
contains at least one horizon of nearly coali-
fied material from which Tempskya has been
collected in growth position (Tidwell and
Hebbert 1976). The dicotyledonous woods
studied here were collected from a horizon

between 10 (3.1 m) and 30 (9.2 m) feet below
the overlying Dakota Sandstone, which is
represented by 10 (3.1 m) to 20 (6.2 m) feet
of coarse brown sandstone that forms a cap
rock in the area.

Fig. 1. Index map of collection sites.

'Bureau of Land Management, Salt Lake City, Utah 84112.

'Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602.

'Department of Geology, University of Utah, Salt Lake City, Utah 84112.

394

July 1983

Thayne et al.: Cretaceous Flora

395

Ulah
0 Prove

Arizono

Fig. 2. Geographic extent of the Cedar Mountain
Formation (after Young 1960).

Specimens were collected from seven dif-
ferent logs at Locality 2, where the Cedar
Mountain Formation consists of a bed of
coarse white sandstone underlaid by channel
fills of yellow conglomeritic sandstones alter-
nating with grey-green shales. These are, in
turn, underlaid by a dark green nodular
weathering shale (Figs. 5-7). The Dakota
Sandstone is missing at this particular site,
but reappears in the section about 3 miles
(1.9 km) to the southeast. Dicotyledonous
woods were found associated with Tempskya
and fossil conifer wood at this locality.

Stratigraphy of
the Cedar Mountain Formation

Stokes (1944) defined the Cedar Mountain
Formation as those sediments lying between
the Brushy Basin Member of the Upper Juras-
sic Morrison Formation and the Lower Cre-
taceous Dakota Formation. These strata were
formerly considered part of the Morrison
Formation. Cedar Mountain deposits are
present over much of eastern Utah, western
Colorado, and northwestern New Mexico

Fig. 3. ParaphijUanthoxijlon i/ta/iense— Ilhistration of
the transverse section showing the relative abundance of
ray tissue (dark areas) and the size, shape, and arrange-
ment of the vessel elements (open circles). Note the radi-
al pore multiples.

(Fig. 2). Stokes (1952) defined two members
of the formation, the Buckhorn Con-
glomerate at the base and the Cedar Moun-
tain Shale at the top. At the type locality
near Castle Dale, Utah, the Buckhorn Con-
glomerate is thick and massive, but it gener-
ally thins to the east and is absent on the
eastern side of the San Rafael Swell. East of
the Colorado River a mudstone and con-
glomeritic sandstone unit occupies the same

Fig. 4. ParaphijUanthoxijlon lita/iense— Illustration of
the tangential section showing the size, shape, and ar-
rangement of the rays (dark lines) and a vessel element
(center).

396

Great Basin Naturalist

Vol. 43, No. 3

W" -*

Fig. 5. Ferron collecting site, showing the lithology.
A is white sandstone cap, B is the surface of the yellow
conglomeritic channel fill from which the specimens
were recovered, and C is the underlying grey-green
shale.

relative position as the Cedar Mountain For-
mation on the west side of the river. These
rocks were termed the "Post-McElmo" beds
(Coffin 1921), but were later renamed the
Burro Canyon Formation (Stokes and Phoe-
nix 1948). Young (1960) proposed that the
Burro Canyon Formation and the Cedar
Mountain Formation are a physically contin-
uous unit and should both be referred to as
Cedar Mountain Formation.

Based on the presence of the pelecypods
Protoelliptio douglassi, Unio farri, and the
conifer Frenelopsis varians, as well as the
stratigraphic position of the Cedar Mountain
Formation, Young (1960) as well as Stokes
(1952), suggested that it is Lower Cretaceous
in age. Another indication of its age is the
presence of Tempskya, which Read and Ash
(1961) considered to be an index fossil to the
Lower Cretaceous (Albian). Fisher et al.

Fig. 6. Overview of the Ferron collecting site. Snow-
capped mountains in the background are in the Wasatch
Plateau. Dicotyledonous logs along with Tempskya were
collected from tlie uppermost layer of sediment shown
in the foreground.

Fig. 7. Petrified dicotyledonous wood shown as it is
found weathered upon the surface of the Ferron site.

(1960) listed the formation as Aptian, but it
may be only Albian or, most probably, may
include rocks of both ages.

Paraphyllanthoxylon utahense sp. nov.
Figs. 3-4, 8-18

Description.— This species is described
from several pieces of black petrified second-
ary wood. The preservation is excellent, and
fine stRictural detail can be observed.

Growth rings: Lacking.

Vessels: Diffuse porous, approximately
12 /mm 2, solitary or more commonly in radial
rows (pore multiples) of 2-3 up to 5 cells
long; individual vessels range from 204 jum
radial by 165 jum tangential diameter to 58
jum radial by 48 jum tangential, average 105
jum radial by 93 jum tangential diameter; per-
forations exclusively simple, located on
oblique end walls; thin-walled tyloses abun-
dant, obscuring the vessel length; vessel walls
3 ium-5 jum thick; tangential pitting with nu-
merous, often appressed, 6 jum-10 /xm diame-
ter; alternate bordered pits with slitlike aper-
tures and occasionally up to 12 |u,m long,
slightly bordered pits with large elliptic aper-
tures probably representing the vessel to pa-
renchyma pitting; radial intervascular pitting
similar to tangential; vessel to ray inter-
vascular pitting similar to tangential; vessel
to ray pitting consisting of small circular or
large, up to 24 jum, scalariform, elliptic to an-
gular slightly bordered pits; 3-6, occasionally
more, pits per crossover field.

Axial Parenchyma: Rare, apotracheal dif-
fuse or scanty paratracheal.

Rays: 12/mm2, heterogeneous with both
uniseriate and multiseriate present; uniseriate

July 1983

Thayne et al.: Cretaceous Flora

397

Fig. 8. Transverse section illustrating solitary vessels
and vessel chains with tyloses. Note that the axial paren-
chyma is scarce (65X).

rays rare, many partially biseriate, with both
procmnbent and upright cells, uniseriate rays
range from 2 cells (80 jum) to 6 cells (300 jum)
high (average 5 cells, 200 jam-220 jum); mul-
tiseriate rays range from 9 cells (380 jum) to
33 cells (1360 [xm) high and 2 cells (30 jum) to
5 cells (100 jum), wide with 106 rows (com-
monly 2) of upright border cells; procumbent
cells range from 25 jam-40 jum vertical, 50
jum-80 jLim radial, and 25 jum-45 jUm tan-
gential diameter; some cubodial cells present,
approximately 40 jum in diameter; upright
cells approximately same size as procumbent
but radial and vertical dimensions reversed;
ray cells' walls 2.5 jum thick, pitted and ap-
pearing beaded in radial section.

Fibretracheids: Septate, libriform, round to
square in cross section, approximately 36 /xm
in diameter, with approximately 2.4 jum thick
walls.

Repository: Brigham Young University,
2190 (Holotype)

Horizon: Cedar Moimtain Formation

Age: Early Cretaceous

Discussion

Twelve species of ParaphyUanthoxylon
have been described in the past. The features
generally constant in all of these reported
species are as follows:

Fig. 9. Transverse section illustrating distribution of
vessels and multiseriate rays (30X).

Diffuse porous wood; vessels in radial rows (pore multi-
ples); exclusively simple perforations; alternate inter-
vascular pitting; elongate vessel to ray pitting; rays of
two sizes, 1-7 seriate, heterocellular with 107 rows of
upright border cells, rays commonly over I mm high, ax-
ial parenchyma lacking or scanty apotracheal diffuse,
scanty paratracheal, or combination of both; septate
fibretracheids; vessels commonly with tyloses.

Paraphijllanthoxijlon utahense fits well with-
in the boundaries of this genus.

Comparison with
Described North American Species

Three species of ParaphyUanthoxylon have
been described from Cretaceous strata in
North America.

ParaphyUanthoxylon arizonense Bailey
1924.— ParaphyUanthoxylon utahense differs
from the upper Cretaceous P. arizonense in
several ways. The most obvious variations are
the size of the vessels and rays. Although
Bailey (1924) gave no measurements and
merely stated that the vessels of P. arizonense
are large, it can be seen that the vessels
shown in his figures at 35X are almost as
large as those of P. utahense at 65X. Also, the
rays shown at 35X in his figures are approx
imately twice as high and wide and more
parallel in outline than those of P. utahense
at a comparable magnification. Another dif-
ference is that P. arizonense has slitlike pits
on the fiber walls that P. utahense lacks. On
the basis of these differences, the Utah speci-
mens have been determined to be distinct
from P. arizonense.

ParaphyUanthoxylon idahoense Spackman
1948.— ParaphyUanthoxylon utahense is
closer to P. idahoense, which was reported

398

Great Basin Naturalist

Vol. 43, No. 3

Fig. 10. Transverse section illustrating a closeup view
of fibretracheids and a heterocellular multiseriate ray

(495X).

from the Lower Cretaceous Wayan Forma-
tion of Idaho, than to other Para-
phyllanthoxylon species. The diameter of the
vessels in P. idahoense is 60 jum-160 jum.

Fig. 11. Radial section with irregularly shaped vessel
to ray cell crossover pits inked in. The beaded nature of
the ray cell walls can also be seen (lOOX).

Fig. 12. Radial section illustrating the relative size
and distribution of the vessels and cells of the hetero-
cellular rays (33X).

However, in comparing the holotype slides of
this species with those of P. utahense, it can
be seen that the vessels of the former are dis-
tinctly larger than those of the latter, which
are 48 ]u,m-165 /xm in tangential diameter.
The pitting is similar in both species, and the
intervascular pits are also alternate, circular
elliptical, and sometimes compacted and an-
gular. The bordered pits of P. idahoense are
10 ju,m-12 jum in diameter, whereas, those in
P. utahense vary from 6 /xm to 10 /xm in di-
ameter. The major differences between these
two species are the compaction of the vessels
and the size of the rays. The number of ves-
sels per square millimeter was not given for
P. idahoense, but its vessels are more tightly
compacted than those in P. utahense. The
rays in P. idahoense are made up of smaller
cells and are narrower than those of P. uta-
hense, although both have multiseriate rays
from two to five cells wide. Since P. utahense
has smaller vessels that are fewer in number
per square millimeter, and larger rays than P.
idahoense, these two species are considered
distinct from one another.

Paraphyllanthoxylon alabamense Gaboon
1972.- As described by Gaboon (1972) from
the Upper Gretaceous Tuscaloosa Formation,
this species has a wide range of variation,
which can be seen by comparing Figures 5,
11, and 14 of her paper. These photos are all

July 1983

Thayne et al.: Cretaceous Flora

399

Fig. 13. Radial section illustrating the irregularly
shaped, narrowly bordered vessel to parenchyma pits

(495X).

listed as being magnified 55X, but the rays in
Figure 5 are approximately five times wider
than those in Figure 14. She stated that this
species was described from 11 different type
specimens that are all similar but show some
variation. The most obvious variation is in
the size of the rays. Barghoorn (1941) has
shown that such variation could conceivably
be found within a species or even within the
trunk of an individual tree, but, since pa-
leobotanists are often restricted to working
with fragments, they have traditionally de-
scribed such fragments as form genera and
hence form species. Spackman (1948) distin-
guished P. idaJioense from P. arizonense be-
cause P. idahoense has smaller vessels, less
abundant pitting, and smaller rays and ray
cells. He stated that:

The magnitude and nature of these variations are well
within the range of variability found in individuals of
many living species, and thus the differences in the two
fossils might be accounted for on the basis of the part of
the tree from which the specimen was derived, differ-
ences in growth rate, etc. In spite of this, however, it
seems appropriate, because of these differences to de-
scribe this new wood as a new species with the hope
that the tnie relationship of these two fossils will be
demonstrated in the hiture. (Spackman, 1948, p. 108).

We agree with Spackman's reasoning, and
therefore believe that P. alabamense as it
now stands includes at least two or three

Fig. 14. Radial section illustrating oblique simple per-
foration plates and oppositely arranged bordered inter-
vascular pits with slitlike apertures on the radial vessel
wall (495X).

form species. Therefore, P. utahense cannot
be accurately compared to it at this time.
The holotype specimen shown by Gaboon
(1972) appears to differ from P. utahense by
having larger rays. The other specimens re-
ported by her appear similar to P. utahense,
although one has larger rays and the other
has smaller. Before any conclusions can be
drawn as to the species boundaries and rela-
tionship between P. utahense and the Ala-
bama specimens, more detailed measure-
ments and comparisons need to be made.

Paraphyllanthoxylon pfefferi Platen
1908.— This species was collected from the
Tertiary of California. It was originally de-
scribed as Carpinoxylon pfefferi (Platen
1908), but was combined with Para-
phyllanthoxylon by Madel (1962). Para-
phyllanthoxylon utahense has larger vessels
(up to 204 jum radial diameter as opposed to
137 jLim radial diameter), which are fewer per
square millimeter (12 compared to 44), and
broader rays (100 jum compared to 50 jum)
than P. pfefferi.

Comparison With Other Paraphyllantho-
xylon Species.— Paraphyllanthoxylon uta-
hense differs from species of Para-
phyllanthoxylon described from geographical
areas beyond the boundaries of North Ameri-
ca (Table 1) in such features as the size and
density of the vessels and dimensions of the

400

Great Basin Naturalist

Vol. 43, No. 3

Fig. 15. Tangential section illustrating the relative
size and distribution of multiseriate rays, fibretracheids,
and vessels with tyloses (30X).

rays. Paraphyllanthoxylon utahense is most
similar to P. capense but differs in having
fewer vessels per square millimeter, and ves-
sels that are larger and fewer per pore
multiple.

Affinities of Paraphyllanthoxylon.— The
original species, Paraphyllanthoxylon arizo-
nense, was described by Bailey (1924) from
silicified wood fragments of the Colorado
Group in Arizona. He proposed the name to
indicate a relationship to Bridelia and Phyl-
lanthus in the section Phyllanthoidea of the
Euphorbiaceae. Madel (1962) combined
woods which had been described as Phyllan-
thinium and Glochidioxylon into the genus
Paraphyllanthoxylon and reserved the genus
for woods with general structure of the Glo-
chidion wood group of the Euphorbiaceae.
Other authors have compared their species to
a number of genera in several other families.

Although Paraphyllanthoxylon alahamense
may be an aggregation of species, further in-
formation concerning the affinities of the
genus may be inferred by the leaf com-
pressions that occur along with it in the Tus-
caloosa sediments. Gaboon (1972) reported

Fig. 16. Tangential section ilkistrating the hetero-
cellular rays and septa in the fibretracheids (lOOX).

that of the families with wood similar to Par-
aphyllanthoxylon only the Sapindaceae, Eu-
phorbiaceae, and Lauraceae are represented
by fossil leaves from the Tuscaloosa.

Phylogenetic Considerations.— The pro-
cesses of convergent and divergent evolution
have obscured the genealogy of even modern
genera and species. Pax and Hoffman (1931)
considered the Euphorbiaceae to be poly-
phyletic in origin, making it imlikely that the
Lower Gretaceous Paraphyllanthoxylon spe-
cies are ancestral to the various groups with-
in the family. The possibility does exist that
they are ancestral to at least some members
of the Glochidion group. Considering the
large number of genera that are similar to the
genus, Paraphyllanthoxylon could be related
to the taxon from which several genera in
many families originated.

By comparing the features of Para-
phyllanthoxylon with Tippo's (1946) list of
primitive and advanced wood characteristics
(Table 2), it can be seen that the anatomy of
the Lower Gretaceous members of the genus
supports evidence from fossil leaf com-

Table 1. Paraphyllanthoxylon species from outside of North America.

Species

Author

Occurrence

pseudohohash iraish i

sahnii

tertiunim

bangalamodense ■

keriense

capense

yvardi

teldense

Ogura, 1932
Prakash, 1958
Ramanujam, 1956
Navale, 1960
Dayal, 1968
Madel, 1962
Koeniguer, 1967
Prive, 1975

Tertiary of Japan

Tertiary of India

Upper Cretaceous of S. Africa

Neogene of France

Oligocene of France

July 1983

Thayne et al.: Cretaceous Flora

401

Fig. 17. Tangential section illustrating various sizes
and shapes of rays and dark cell contents in many of the
ray cells (SOX).

pressions that the angiosperms had developed
many of their modern characteristics by
Early Cretaceous times.

Acknowledgments

The authors express appreciation to S. R.
Rushforth, J. D. Brotherson, and J. Keith Rig-
by of Brigham Young University and S. R.
Ash of the Department of Geology, Weber
State College, Ogden, Utah, who graciously
reviewed the manuscript. We also express
appreciation to Naomi Hebbert who helped
in preparation of the illustrations.

Table 2. Comparison with primitive and advanced
features.

Primitive

Advanced

4 ^

ci- 3 J- ^ «

_ - T3 e

Us ^

c 2 o So.^ o i;

c/2 H &< < <! -J i'i

Paraphyllanthoxylon X* X X X X X X*'

*(X) indicates that species of Paraphyllanthoxylon possesses that feature.
"(X) has low heterogeneous rays.

a^ > iZ 0; ._

15 "3 ;s Sfi "5 5P §
c/5 O c/2 £-4 c/i 3- Z

Fig. 18. Tangential section showing appressed, op-
positely arranged, bordered pits with slitlike apertures
on the tangential vessel wall (495X).

Literature Cited

Bailey, I. W. 1924. The problem of identifying the
wood of Cretaceous and later dicotyledons: Para-
phifUanthoxylon arizonense. Ann. Bot. .38:
4.39-451.

Barghoorn, E. S. 1941. The ontogenetic development
and phylogenetic specialization of rays in the xy-
lem of dicotyledons. II. Modification of the mul-
tiseriate and uniseriate rays. Amer. J. Bot.
28:273-282.

Bodily, N. M. 1969. An armored dinosaur from the
Lower Cretaceous of Utah. Brigham Young Univ.
Geol. Studies 16(.3):35-60.

Cahoon, E. J. 1972. Paraphyllanthoxylon alabamense: a
new species of fossil dicotyledonous wood. Amer.
J. Bot. 59(1):5-11.

Coffin, R. C. 1921. Radium, uranium and vanadium de-
posits of southwestern Colorado. Colorado Geoft
Surv. Bull. 16.

Dayal, R. 1968. A new fossil wood of the Eu-
phorbiaceae from the Deccan Intertrappean beds
of Madhva Pradesh. Palaeobotanist 16(2):
148-150.

Fisher, D. J., D. E. Erdmann, and J. B. Reeside. 1960.
Cretaceous and Tertiary formations of the Book
Cliffs, Carbon, Emery, and Grand counties, Utah,
and Garfield and Mesa counties, Colorado. U.S.
Geol. Surv. Prof. Paper .3.32. 1-80.

KoENiGUER, J. C. 1967. Etude paleoxylologique de la
Fouraine: Paraphyllanthoxylon yvardi, n. sp. C.
R. 92c Congress Nat. Soc. Sav., Strasbourg,
3:71-75.

Madel, E. 1962. Die fossilen Euphorbiaceen-Holzer niit
besonderer Benicksichtigung neuer Funde aus
der Oberkreide Sud-Afrikas. Senkenberg. Leth
43(4):283-321.

Navale, G. K. B. 1960. Phyllanthinium bangalamo-
dense, a new species of fossil Euphorbiaceous
wood from the Cuddalore Series of India. Pa-
laeobotanist 9:11-16.

402

Great Basin Naturalist

Vol. 43, No. 3

Ogura, Y. 1932. On the stnicture of a silicified wood
found near hobashiraishi at Najima near Fukuoka
City. Japanese J. Bot. 6(2): 18.3-190.

Pax, F., and K. Hoffman. 1931. Euphorbiaceae. In A.
Engler and K. Prantl: die naturlichen pflanzenfa-
milien, 2Aufl. 19(c): 11-233.

Platen, P. 1908. Untersuchungen fossiler Holzer aus
deni Westen der Vereinigten Staaten von Nor-
dainerika. Diss: 1-55.

Prakash, U. 1958. Studies in the Deccan Intertrappean
Flora, 4: two silicified woods from Madhya Pra-
desh. Palaeobotanist 7(1): 12-20.

Prive, C. 1975. Sur la presence dun bois fossile D Eu-
phorbiaceae dans loligocense du Cantal. Act.
95th Natl. Cong. Soc. Savantes 1970(,3):111-125.

Ramanujam, C. K. G. 1956. Fossil woods from the Eu-
phorbiaceae from the Tertiary rocks of south Ar-
cot District, Madras. J. Indian Bot. Soc.
35(3):284-307.

Read, C. B., and S. R. Ash. 1961. Stratigraphic signifi-
cance of the Cretaceous fern Tempskya in the
western conterminous United States. U.S. Geol.
Surv. Res. 8.32:D-250-D-254.

Spackman, W., Jr. 1948. A dicotyledonous wood found
associated with the Idaho Tempskyas. Ann Mo.
Bot. Card. .35:107-116.

Stokes, W. L. 1944. Morrison and related deposits in
and adjacent to the Colorado Plateau. Ceol. Soc.
Amer. Bull. 55:951-992.

1952. Lower Cretaceous in Colorado Plateau.

Amer. Assoc. Pet. Geol. Bull. .36(9): 1774- 1796.

Stokes, W. L., and D. A. Phoenix. 1948. Geology of the
Engas-Gypsum Valley Area, San Miguel and
Montrose counties, Colorado. U.S. Geol. Surv.
Prelim. Map 93, Oil and Gas Inv. Ser.

Thayn, G. F., W. D. Tidwell, and W. L. Stokes. 1973.
Petrified angiosperm wood from the Lower Cre-
taceous Cedar Formation of Utah. Geol. Soc.
Amer. Abstr. with programs 5(6):519.

Tidwell, W. D., and N. E. Herbert. 1976. Tempskija
from the Lower Cretaceous Cedar Mountain For-
mation, Utah. Amer. J. Bot. Abstr. 63:33.

Tidwell, W. D., G. F. Thayn, and J. L. Roth. 1976.
Cretaceous and Early Tertiary floras of the Inter-
mountain Area. Brigham Young Univ. Geol. Stud-
ies 22:77-98.

Tippo, O. 1946. The role of wood anatomy in phylogeny.
Amer. Midi. Nat. 36:362-372.

Young, R. G. 1960. Dakota Group of Colorado Plateau.
Amer. Assoc. Pet. Geol. Bull. 44(1): 1,56-194.

ACULEATA HYMENOPTERA OF SAND MOUNTAIN
AND BLOW SAND MOUNTAINS, NEVADA

R. W. Rust', L. M. Hanks,i2, and R. C. BechteP

Abstract.— There were 198 species of aculeata Hymenoptera in 15 families collected from Sand Mountain and
Blow Sand Mountains, Nevada. Four species are considered new to science and none are considered endemic to ei-
ther dune area.

Sand Mountain and Blow Sand Mountains
were visited 19 times in a 13-month period
for the purpose of surveying selected groups
of arthropods. Here we report on the acu-
leate Hymenoptera collected during the
study. Over 2,000 specimens were obtained,
representing 198 species in 15 families. Four
species are considered new to science and 21
species were identified as "species" in un-
studied genera. Most of the unknown species
were bees (Apoidea) in the genera Perdita,
Dialictus, Sphecodes, and Hesperapis. None
of the species is considered sand obligate or
endemic to either dune.

Previous studies on Hymenoptera in Ne-
vada include the faunistic inventory of the
Nevada Test Site conducted by Brigham
Young University from 1958 to 1966 that
produced almost 8,000 adults and 1,100 im-
matures (Beck and Allred 1968). Of these, the
ants (Cole 1966), mutillid wasps (Ferguson
1967, Allred 1973), tiphiid wasps (Wasbauer
1973), and bees (G. E. Bohart, pers. comm.)
have been identified. Wheeler and Wheeler
(1978) studied the mountain ants of Nevada
and they have produced a manuscript on the
ants of Nevada (to be published by Los An-
geles Coimty Museum). Pretruszka (1980) ob-
tained almost 2,000 specimens, identified to
family, from Fairview Valley, Nevada, from
pitfall and malaise trapping.

Study Sites

Sand Mountain (SM) dunes and Blow Sand
Mountains (BSM) dunes were sampled from

June 1979 through July 1980. Sand Mountain
is approximately 46 km ESE of Fallon,
Churchill County, Nevada (39°20'N-
118°20'W) and is about 1,250 m in elevation.
Blow Sand Mountains are approximately 52
km SE of Fallon, Nevada (39°10'N-
118°35'W) and are about 1,400 m in eleva-
tion. The dunes are separated by 25 km air
distance. Sand Mountain is a star dune of ap-
proximately 3.2 km^ and Blow Sand Moun-
tains are complex star and linear dunes of ap-
proximately 9.2 km^. However, both dunes
result from the same eolian sand deposited
during the Turupah and Fallon formations of
about 4,000 years B.P. (Morrison and Frye
1965).

The floras of the two dunes were similar.
The dominant vegetation was Atriplex con-
fertifolia (Torr. & Frem.), Tetradymia tetra-
meres (Blake), Chrysothamnus viscidiflorus
(Hookl), Astragalus lentiginousus Dougl., and
Psoralea lanceolata (Pursh.), and at Sand
Mountain only Eriogonum kearneyi Tidestr.
and Psorotharnnus polyadeniiis (Torr.). The*
common grass was Oryzopsis hyrnenoides (R.
&S.).

Methods

Several collecting techniques were used to
obtain specimens (see Bechtel et al. 1981).
Permanent pitfall traps were 0.95 L plastic
cartons. They were operative for 30-day peri-
ods. Temporary pitfall traps were 15 cm di-
ameter cereal bowls placed level with the
sand surface. Temporary pitfall traps were

'Biology Department, University of Nevada, Reno, Nevada 89557.

^Present address; Department of Entomology, University of Maryland, College Park, Maryland 20742.

'Nevada State Department of Agriculture, .350 Capitol Hill Avenue, Reno, Nevada 89502.

403

404

Great Basin Naturalist

Vol. 43, No. 3

used for 12-18 hr during a survey period.
Two UV light traps were operated from dusk
to dawn. Hand-held UV lamps were used in
searching the dimes for fluorescing arthro-
pods. Sand was sifted through two screens
12 X 12 cm and 1.5 X 1.5 cm mesh to recov-
er subsurface arthropods. General collecting
involved the use of aerial nets, plant in-
spection, and walking the dunes at night with
lamps to obtain specimens. During a survey
period, four or five different sites on the dune
were visited and the sites were varied each
survey period.

Data are presented in the following man-
ner: For each species its known geographic
distribution, present location, numbers ob-
tained, dates of occurrence, and, for the bees,
their flower visitation as determined from
pollen-load analysis (Hanks and Rust 1983).
Once a genus and species of plant is given as
a pollen source, subsequent references to it is
by use of the first two letters of the genus
and species, e.g. Astraglus lentiginosus is
Asle. Geographic distributions were obtained
from the Hymenoptera Catalog (Krombein
and Hurd 1979) and are presented as North
America (NA), western United States (WUS),
southwestern United States (SWUS), Great
Basin (GB), or by individual state. For the
ants, Formicidae, the number given repre-
sents collections and not individual ants.

The following taxonomists identified the
material being presented: R. C. Bechtel (Mu-
tillidae), G. E. Bohart (Apoidea), R. M. Bo-
hart (Chrysididae, Vespidae, Sphecidae), A.
A. Grigarick (Megachilidae), L. Kimsey
(Chrysididae), W. E. LaBerge (Apiodea), A.
S. Menke (Sphecidae), F. D. Parker (Sphe-
cidae), J. G. Rosen (Andrenidae), R. W. Rust
(Apoidea), R. R. Snelling (Apiodea, Formi-
cidae), R. W. Thorp (Andrenidae), M. S.
Wasbauer (Tiphidae, Scoliidae, Pompilidae),
and G. C. and J. Wheeler (Formicidae). All
specimens, except new specimens, are in the
collection of the University of Nevada, Reno.

Results

The most numerous species obtained was
the California harvester ant, Pogonomyrmex
californicus (Buckley), with 26 collections
containing hundreds of specimens. One per-
manent pitfall trap contained over 500 indi-
viduals. It was present at both dunes from

May through October. Two other ants were
common at SM Vero7nessor lariversi M. R.
Smith and Conomyrma insana (Buckley).
Glyptacros new species and Xeroglypta egr-
egia Mickel and Krombein were perhaps the
most interesting wasps collected. They both
have wingless females and were collected by
sifting sand. Other wingless females obtained,
mutillids, were found active on the surface or
attracted to UV light traps. Next to the ants,
the most numerous species were Parnopes
ftilvicornis Cameron (Chrysididae), Para-
nistrocerus toltecus (Saussure) (Vespidae),
Acanthetropis aequalis (Fox) (Tiphiidae),
Sphaerophthalma sp. (Mutillidae), Micro-
benibex argyropletira Bohart and Eucerceris
nevadensis (Dalla Torre) (Sphecidae), and
Agapostemon melliventris Cresson and Dia-
lictus sp. (Halictidae).

Chrysididae

Elampus nitidus Aaron (WUS) SM 3 Aug.

Oct.
Holopyga hora Aaron (WUS) SM 1 May.
Hedychridium amahile Gockerell (WUS) SM

5 Aug. Sept.
Hedychridium arietinum Bohart (CA) SM 1

Sept., BSM 1 Sept.
Hedychridium mancopae Bohart (CA) SM 2

April May.
Hedychridium mirum Bohart (CA) SM 14

June July, BSM 1 June.
Hedychridium species (?) BSM 1 June.
Chrysis inflata Aaron (WUS) SM 1 Sept.
Spintharosoma trochilus (duBuysson) (WUS)

SM 1 April.
Parnopes fulvicornis ftilvicornis Cameron

(WUS) SM 132 June July Aug. Sept.

Tiphiidae

Brachysistis lacustris lacustris Malloch (Mo-

jave desert) SM 3 July Aug. Sept., BSM

9 July Aug.
Brachycistis triangularis Fox (SWUS) SM 4

June, BSM 10 June July.
Brachycistis agama (Dalla Torre) (NA) SM 12

July Aug. Sept., BSM 57 July Aug.
Brachycistis species (?) BSM 9 June.
Colocistis crassa (Bradley) (SWUS) BSM 3

June.

July 1983

Rust et al.: Nevada Hymenoptera

405

Acanthetropis aequalis (Fox) (WUS) SM 30

July Aug. Sept., BSM 31 July Aug. Sept.
Acanthetropis noctivaga (Bradley) (SWUS)

SM 16 July Aug., BSM 6 July Aug.
Glyptacros new species (?) SM 5 Sept. Oct.,

BSM 16 Jime Aug. Sept.
Xeroglypta egregia Mickel and Krombein

(SoCA) SM 1 Aug.

Mutillidae

Sphaeroptluilma species (?) SM 57 June July

Aug., BSM 26 Jime July Aug.
Dasymutilla gloriosa (Saussure) (SWUS) BSM

I Aug.

Dasymutilla satanas Mickel (SWUS) SM 2
June July.

Scoliidae

Crioscolia alcione (Banks) (SWUS) SM 3 July
Aug.

Formicidae

Pogonomyrmex californicus (Buckley)

(SWUS) SM 26 May to Oct. BSM 14.
Veromessor lariversi M. R. Smith (WUS) SM

II June July Aug. Sept., BSM 2 June
Aug.

Creniatogaster species (?) SM 1 Sept., BSM 1

Aug.
Conornyrrna hicolor (W. M. Wheeler) (WUS)

SM 2 July Aug.
Conornyrrna insana (Buckley) (WUS) SM 15

June July Aug. Sept., BSM 2 Aug.
Camponotus vicinus Mayr (US) SM 5 Aug.

Sept.
Myromecocystus kennedyi Cole (WUS) SM 4

Feb. Aug., BSM 2 Aug.
Myromecocystus pyramicus M. R. Smith

(WUS) SM 3 Aug. Sept., BSM 2 Aug.
Myromecocystus new species (?) SM 4 April

May.

Vespidae

Pterocheilus crispocornis Bohart (SWUS) SM

1 June, BSM 1 Aug.
Pterocheilus diversicolor Rohwer (SWUS)

BSM 1 Aug.
Pterocheilus hurdi Bohart (CA) BSM 1 June.
Pterocheilus hirsutipennis Bohart (SWUS) SM

1 May.

Pterocheilus laticeps Cresson (WUS) BSM

1 Jime.
Pterocheilus tricoloratus Bohart (SWUS) BSM

1 June.
Leptochilus species (?) BSM 1 Aug.
Maricopodynerus maricoporum (Viereck)

(SWUS) SM 4 April July.
Stenodynerus percampanulatus (Viereck)

(WUS) SM 6 June July Aug. Sept., BSM

8 Aug. Sept.
Parancistrocerus toltecus (Saussure) (WUS)

BSM 95 June July Aug. Sept.
Euodynerus annulatus sulphureus (Saussure)

(WUS) SM 1 June.
Ancistrocerus acatskill halophila Viereck

(WUS) SM 2 Oct.

Pompilidae

Pepsis pallidolimbata pallidolimbata Lucas

(WUS) SM 2 Sept., BSM 1 Aug.
Hemipepsis ustulata ochroptera Stal. (SWUS)

SM 1 June.
Aporus hirsutus (Banks) (WUS) BSM 2 June

Aug.
Evagetes padrinus padrinus (Viereck) (WNA)

BSM 9 June Aug. Sept.
Agenioideus biedermani (Banks) (SWUS) SM

1 Aug.
Episyron oregon Evans (WNA) SM 1 May,

BSM 1 Aug.
Anoplius relativus (Fox) (NA) SM 5 June,

BSM 10 June July Aug. Sept. Oct.
Anoplius tenebrosus (Cresson) (NA) SM 1

Sept.
Pompilus orophilus Evans (NA) SM 3 May,

BSM 3 Aug.
Pompilus phoenix Evans (WUS) BSM 3 June.
Aporinellus borregoensis Evans (SWUS) SM 1

June.
Aporinellus fasciatus (Smith) (NA) SM 1

June, BSM 1 June.
Aporinellus medianus Banks (NA) SM 12

June July Aug. Sept.
Aporinellus taeniatus taeniatus (Kohl) (NA)

Sm 1 Sept.
Aporinellus yucatanensis (Cameron) (NA) SM

1 Sept., BSM 1 June.

Sphecidae

Prionyx canadensis (Provancher) (NA) SM 1
June, BSM 6 June July Aug.

406

Great Basin Naturalist

Vol. 43, No. 3

Prionyx subatratus Bohart (WUS) SM 1 Aug.
Podahnia communis (Cresson) (WUS) SM 12

June July, BSM 6 July.
Ammophila aberti Haldeman (WUS) SM 2

May June.
Ammophila polita Cresson (WUS) BSM 4

June.
Ammophila pruinosa Cresson (WUS) SM 2

May Sept.
Ammophila wrightii (Cresson) (WUS) BSM 1

June.
Mimesa coquilletti (Rohwer) (CA NV) SM 13

May Sept., BSM 2 June.
Astata bechteli Parker (SWUS) SM 3 June

July Aug.
Astata occidentalis Cresson (WUS) BSM 9

June July Aug.
Larropsis waslioensis Bohart and Bohart (NV)

SM 3 May June, BSM 11 June July Aug.
Ancistromma granulosa (Bohart and Bohart)

(WUS) SM 1 Aug., BSM 1 Aug.
Tachytes ermineus Banks (SWUS) SM 2 June.
Tachytes nevadensis Bohart (WUS) SM 3

Aug. Sept.
Tachytes new species SM 7 June.
Tachysphex apicalis fusiis Fox (NA) BSM 3

June.
Tachysphex ashmeadii Fox (WUS) SM 7 Aug.
Tachysphex spinosus Fox (WUS) SM 1 June.
Tachysphex species (?) BSM 14 June.
Plenoculus boregensis Williams (SoCA) SM 1

June.
Pisonopsis species (?) SM 3 May June.
Miscophus species (?) SM 2 May June.
Oxybelus abdominalis Baker (WUS) SM 10

Aug. Sept., BSM 6 June.
Oxybelus pitanta Pate (SWUS) SM 12 Aug.

Sept., BSM 2 June.
Crabro opalenscens Bohart (WUS) SM 2 May.
Bicyrtes ventralis (Say) (NA) SM 3 Aug.
Bicyrtes capnopteris (Handlirsch) (NA) SM 3

Aug.
Microbembex argyropleura Bohart (SWUS)

SM 39 June July Aug. Sept., BSM 15

July Aug.
Microbembex californica Bohart (SWUS) SM

1 June, BSM 1 July.
Bembix rugosa Parker (AZ) SM 18 Aug.
Bembix stenobdoma Parker (AZ) SM 1 July.
Bembix occidentalis Fox (SWUS) SM 2 June.
Bembix americana comata Parker (SWUS)

SM 2 Aug. Sept.

Stictiella corniculata Mickel (WUS) SM 3

Sept.
Stictiella nubilosa Gillaspy (SoCA) SM 2 June

July.
Stictiella speciosa (Cresson) (WNA) SM 1

Sept.
Glenostictia argentata (Fox) (SoCA) SM 12

Aug.
Glenostictia megacera (Parker) (WUS) SM 1

Aug.
Glenostictia tenuicornis (Fox) (SWUS) SM 2

Aug.
Philanthus crotoniphilus Viereck and Cock-

erell (WUS) SM 19 Aug. Sept.
Philanthus multimaculatus Cameron (WNA)

SM 13 Aug. Sept.
Philanthus pacificus pacificus Cresson (NA)

SM 11 July Aug. Sept., BSM 1 June.
Philanthus pulcher Dall Torre (SWUS) SM 1

May.
Philanthus ventralis (Mickel) (PCS) SM 17

Sept.
Philanthus zebratus Cresson (WUS) SM 1

Sept.
Clypeadon evansi Bohart (SWUS) SM 4 July

Aug.
Clypeadon laticinctus (Cresson) (WUS) SM 8

Aug.
Clypeadon utahensis (Baker) (SWUS) SM 7

Aug. Sept.
Clypeadon species (?) SM 1 July.
Cerceris bridwelli ScuUen (SoCA AZ) SM 8

Aug.
Cerceris californica Cresson (WUS) SM 5 July

Aug.
Cerceris conifrons Mickel (WNA) SM 4 Aug.

Sept.
Cerceris crotonella Viereck and Cockerell

(WUS) SM 5 June July Aug., BSM 1

June.
Cerceris echo echo Mickel (WUS) SM 3 July

Aug. Sept.
Cerceris species (?) SM 1 Aug.
Eucerceris arenaria Scullen (SWUS) SM 9

Aug. Sept.
Eucerceris nevadensis (Dalla Torre) (WUS)

SM 74 July Aug. Sept.

Colletidae

Colletes rnandibularis Smith (EUS) SM 3
Sept., Chrysothamnus viscidiflorus.

July 1983

Rust et al.: Nevada Hymenoptera

407

CoUetes slevini Cockerell (WUS) SM 4 June
July Sept., Chvi, Psorothamnus
polijdeniiis.

CoUetes stepJiani Timberlake (SoCA) SM 1
June.

CoUetes species (?) SM 30 June Aug. Sept.,
BSM 4 June Sept., Chvi, Eriogonum
keameyi, Pspo, Tetradymnia coniosa.

Andrenidae

Andrena (Diandrena) makwothricidis Thorp

(SoCA) BSM 2 May, Malacothrix

sonclioides.
Andrena (Onagrandrena) chyUsmiae Linsley

and MacSwain (ECA) SM 15 April,

BSM 2 May.
Andrena (Onagrandrena) Unsleyi Timberlake

(SWUS) SM 5 April.
Andrena (Thysandrena) vierecki Cockerell

(GB) BSM 2 June, Mentzelia albicauUs.
Nornadopsis (Nomadopsis) pueUae (Cockerell)

(WUS) SM 7 May June, SM 32 May

June, Maso.
Nornadopsis (Micronomadopsis) phaceliae

Timberlake (ECA) SM 2 May, BSM 1

June, PhaceUa sp.
Nornadopsis new species (?) BSM 35 June

July, Psorotliamnus kingii.
Perdita (Cockerellia) utahensis Cockerell

(SWUS) SM 16 July Aug., HeUanthus

deserticola.
Perdita (Perdita) lepidosparti Timberlake (GB)

SM 11 July Aug. Sept., Cleome

sparsifolia.
Perdita (Perdita) hirticeps Timberlake (SWUS)

SM 16 June July Aug., Chvi, BSM 5

July.
Perdita (Perdita) phymatae Cockerell (SWUS)

SM 2 Sept., Chvi.
Perdita (Procockerellia) albonotata Tim-
berlake (SoCA) SM 1 July.
Perdita species 1 (?) SM 84 Aug. Sept., Chvi.
Perdita species 2 (?) SM 1 Aug., Pspo.
Perdita species 3 (?) SM 51 July, Tiguilia

nuttaUii.
Perdita species 4 (?) SM 15 July Aug., Pspo.

Halictidae

Nomia (Acunomia) howardi
(SWUS) SM 2 Aug., Erke.

Crawford

Agapostemon femoratiis Crawford (WNA)
BSM 1 June, Clsp, Sphaeralcea
atnbigua.

Agapostemon meUiventris Cresson (WUS) SM
73 June July Aug., BSM 2 Aug. Sept.,
Atriplex sp, Amaranthus sp, Camissonia
clavaefonnis, Chvi, Clsp, Hede, Erke,
Psoralea lanceolata Pspo.

Lasioglosswn sisymbrii (Cockerell) (WUS)
SM 16 April June July Aug. Sept., As-
tragalus lentiginosus, Chvi.

Evylaeus aberrans (Crawford) (WUS) SM 9
May June July, BSM 2 May June.

DiaUctus species 1 (?) BSM 5 April.

DiaUctus species 2 (?) SM 78 April May June
July Aug. Sept., BSM 37 July Aug.
Sept., Chvi, Erke, Hede, Psla, Step-
hanomeria exigiia, Tetradymia
tetrameres.

DiaUctus species 3 (?) SM 9 Aug. Sept., Abr-
ionia turbinata, Asle, Chvi, Erke, Pen-
stemon acuniinatus, PhaceUa sp.

Sphecodes species (?) SM 1 Sept.

Melittidae

Hesperapis species (?) SM 22 May, PhaceUa
sp, Oenothera deUoides.

Megachilidae

Anthidium rodecki Schwartz (CO NV) SM 15
May June July Aug., BSM 44 July Aug.,
Psla, Pski, Pspo.

Dianthidium subparviim Swenk (WNA) SM
4, Chaenactis xantiana, Chvi.

AnthidieUiim notatum robertsoni (Cockerell)
(WNA) SM 2 Aug., Psla.

SteUs species (?) BSM 1 June.

Proteriades (HolpUtina) bidUfacies (Michener)
(ECA) SM 3 April May, BSM 5 May
June, (Asle) PhaceUa.

Anthocopa (Eremosmia) robustula (Cockerell)
(SWUS) SM 4 April May, BSM 5 June,
PhaceUa sp.

Anthocopa (Eremosmia) timberlakei (Cock-
erell) (SoCA) SM 28 May June, BSM 15
May June, Asle, Gael, Meal, Peac, Pha-
ceUa sp, Pski, Pspo.

AshmeadieUa (Ashmeadiella) aridula Cock-
erell (WUS) SM 5 Aug., BSM 3 Aug,
Clsp, PhaceUa sp, Psla, Pski, Pspo.

408

Great Basin Naturalist

Vol. 43, No. 3

Ashmeadiella (Ashmeadiella) bticconis den-

ticulata (Cresson) (WUS) SM 1 June,

Pspo.
Osmia (Nothosmia) titusi Cockerell (SoCA)

SM 2 April June, Asle, Phacelia sp,

Pspo.
Megachile (Derotropis) xerophila Cockerell

(SoCA AZ) SM 6 May, BSM 3 June,

Chxa, Maso.
Coelioxys (Coelioxys) mitchelli Baker (SUS)

SM 2 May.

Anthophoridae

Diadasia australis (Cresson) (WUS) BSM 12

June, Opiintia pidchella, Mentzelia al-

bicaidis, Hede.
Diadasia vallicola Timberlake (AZ CA) SM

16 May.
Synhalonia albescens Timberlake (SWUS)

SM 3 May.
Synhalonia primaveris Timberlake (SWUS)

SM 30 April May, BSM 4 May June,

Asle, Meal, Peac, Phacelia.
Synhalonia speciosa (Cresson) (WUS) SM 1

May.
Melissodes (Eumelissodes) biniatris LaBerge

(WNA) BSM 2 Sept.
Melissodes (Eumelissodes) lutulenta LaBerge

(WNA) SM 14 June, BSM 2 June July,

Chaetadelpha wheeleri, Hede, Pspo,

Spam, Teco.
Melissodes {Eumelissodes) montana Cresson

(WUS) SM 39 Sept, BSM 1 Sept, Chvi,

Erke.
Anthophora (Anthophora) affabilis Cresson

(WUS) SM 31 April May, BSM 3 May,

Asle, Cacl, Peac, Phacelia.
Anthopfiora (Anthophora) porterae Cockerell

(WUS) SM 39 April May, BSB 9 May

June, Asle, Peac.
Anthcrphora (Anthoplwra) urbana Cresson

(WUS) SM 35 Aug. Sept., BSM 7 Aug.

Chvi, Clsp, Erke, Stex.
Anthophora (Micranthophora) hololeuca

Cockerell (SWUS) SM 18 June July

Aug, BSM 38 June July Aug, Pski, Pspo.
Anthophora (Micranthophora) petrophila

Cockerell (SWUS) SM 11 Aug. Sept.,

BSM 3 Aug, Chvi.
Centris (Xerocentris) californica Timberlake

(CA AZ) SM 34 Aug. Sept., Clsp.

Ceratina (Zadontomerus) neomexicana Cock-
erell (SWUS) SM 10 May June July
Aug., BSM 3 June July, Erke, Hede,
Maso, Pspo, Stex, Teco.

Apidae

Apis mellifera Linnaeus (world wide) SM 7
April, Asel.

Acknowledgments

We thank G. E. Bohart, R. M. Bohart, A.
A. Grigarick, L. S. Kimsey, W. E. LaBerge,
A. S. Menke, F. D. Parker, J. G. Rozen, R. R.
Snelling, R. W. Thorp, M. S. Wasbauer, and
G. C. and J. Wheeler for the identification of
specimens used in this study. Dave Goi-
coechea, BLM State of Nevada, made the
study possible.

Literature Cited

Allred, D. M. 1973. Additional records of mutillid
wasps from the Nevada Test Site. Great Basin
Nat. 33:156-162.

Bechtel, R. C, L. M. Hanks, and R. W. Rust. 1981.
Orthopteroids of Sand Mountain and Blow Sand
Mountains, Nevada. Entoniol. News 92:125-129.

Beck, D. E., and D. M. Allred. 1968. Faunistic inven-
tory—BYU ecological studies at the Nevada Test
Site. Great Basin Nat. 28:1.32-141.

Cole, A. C. 1966. Ants of the Nevada Test Site. BYU
Sci. Bull., Biol. Ser. 7(3): 1-26.

Ferguson, W. E. 1967. Male Sphaeropthalmine wasps
of the Nevada Test Site. BYU Sci. Bull., Biol. Ser.
8(4): 1-26.

Hanks, L. M., and R. W. Rust. 1983. Bee pollinators in
a sand dune community. Submitted to Ecology.

Krombein, K v., and p. D. Hurd. 1979. Catalog of
Hymenoptera in America north of Mexico.
Smithsonian Institution Press, Washington, D.C.

Morrison, R. B., and J. C. Frye. 1965. Correlation of
the middle and late quaternary successions of the
Lake Lahontan, Lake Bonneville, Rocky Moun-
tains (Wasatch Range), southern Great Plains,
and eastern midwest areas. Nevada Bureau Mines
9:1-45.

Pietruszka, R. D. 1980. Observations on seasonal varia-
tion in desert arthropods in central Nevada.
Great Basin Nat. 40:292-297.

Wasbauer, M. S. 1973. The male Brachycistidine wasps
of the Nevada Test Site (Hymenoptera: Tiph-
iidae). Great Basin Nat. 33:109-112.

Wheeler, G. C, and J. Wheeler. 1978. Mountain ants
of Nevada. Great Basin Nat. .38:379-396.

STATUS AND LIFE HISTORY NOTES ON THE NATIVE FISHES
OF THE ALVORD BASIN, OREGON AND NEVADA

Jack E. Williams' and Carl E. Bond-

.\bstract.— Three fishes, two species of Gila, and an undescribed subspecies of cutthroat trout, are endemic to
the Alvord Basin. Historically, the Alvord cutthroat trout, Salmo clarki ssp., inhabited the larger creeks of the basin
but has been extirpated in pure form because of introgression with introduced rainbow trout, Salmo gairdneri. Gila
boraxobitis is restricted to the thermal waters of Borax Lake and its outflows in the northern part of the basin. This
species is endangered because of alteration of its fragile habitat. The Alvord chub, G. alvordensis, is recorded from
16 localities throughout the basin, including springs, creeks, and reservoirs. Although G. alvordensis as a species is
not in jeopardy, many populations are small and could be easily eliminated by habitat destruction or by the in-
troduction of exotic fishes. Competition with exotic guppies, Poecilia reticulata, has extirpated the Thousand Creek
Spring population of Alvord chubs.

Both species of Gila are opportunistic omnivores, consuming primarily chironomids, microcrustaceans, and dia-
toms. Tlie Borax Lake chub also consumed large numbers of terrestrial insects, but specialized feeding on molluscs
was noted in the West Spring population of Alvord chubs. Borax Lake chubs spawn throughout the year; however,
most spawning occurs in early spring. Borax Lake chubs mature at a small size, occasionally less than 30 mm stan-
dard length, and seldom live more than one year. Alvord chubs are typically much larger than the Borax Lake spe-
cies and live at least into their fifth year.

The Alvord Basin of southeastern Oregon
and northwestern Nevada is an endorheic
part of the Great Basin province. Aquatic
habitats are sparse and consist primarily of
Trout Creek in Oregon, the Virgin-Thousand
Creek system in Nevada, as well as several
small streams and springs (Fig. 1). During the
late Pleistocene, a lake of over 1,200 km^
covered much of the valley floor (Snyder et
al. 1964). As pluvial waters dried, fishes were
restricted to remaining permanent springs
and creeks. Three native fishes are endemic
to the Alvord Basin. Chubs, genus Gila, oc-
cupy many of the isolated waters in the Al-
vord Basin and have diverged into two spe-
cies. The Alvord chub, Gila alvordensis, is
the most common fish in the basin and occurs
in a variety of springs and creeks. The Borax
Lake chub, G. boraxobitis, is restricted to Bo-
rax Lake and its outflows in Oregon. The Al-
vord cutthroat trout, Salmo clarki ssp., is the
third fish native to the basin. Historic habitat
for the Alvord cutthroat trout consisted of
the larger streams in the basin, such as Trout
and Virgin creeks. Another undescribed sub-
species of cutthroat trout occurs in Willow

and Whitehorse creeks just east of the Trout
Creek Mountains in a separate basin. Al-
though exhibiting affinities for the Alvord
cutthroat trout, the subspecies foimd in Wil-
low and Whitehorse creeks will not be
treated further in this report.

Our knowledge of the native fishes of the
Alvord Basin is limited. The monograph of
Great Basin fishes by Hubbs and Miller
(1948) provided the first detailed account of
the Great Basin ichthyofauna and included a
brief discussion of the native Alvord Basin
fishes and their isolation. However, all the
Alvord Basin fishes remained undescribed un-
til 1972, when Hubbs and Miller (1972) diag-
nosed the Trout Creek population of Gila as
G. alvordensis. Our studies have resulted in
the description of Gila boraxobitis (Williams
and Bond 1980) and a further description of
G. alvordensis with a taxonomic analysis of
seven disjunct populations of the species
(Williams 1980, Williams and Bond 1980).
Characters of the Alvord cutthroat trout have
been provided by Behnke (1979), but the sub-
species remains undescribed. The only pub-
lished life history information on Alvord

'Endangered Species Office, U.S. Fish and Wildlife Service, 1230 "N" Street, 14th Floor, Sacramento, California 95814.
'Department of Fisheries and Wildlife, Oregon State University, Corvallis, Oregon 97331.

409

410

119

mJk

Great Basin Naturalist
o

Vol. 43, No. 3

^Dufurrena

Area ^og Hot

Thou Cr . I T.- . Res

/^ Borax Lake. ,^

:^J

50
I 1 I 1

Fig. 1. Map of major aquatic habitats in the Alvord Basin, Oregon and Nevada.

Basin Gila consists of a study of feeding ecol- of the native fishes of the Alvord Basin and
ogy by Williams and Williams (1980). This presents additional information on their life
paper documents the distribution and status history.

July 1983

Williams, Bond: Native Fishes

411

Materials and Methods

The distribution and status of fishes was
determined by field surveys, museum records,
and testimony of local residents. Many of the
habitats in the Oregon part of the basin were
known prior to this work. On the other hand,
the Nevada part of the basin had received
little attention by ichthyologists, and there-
fore most of our survey efforts were focused
in the southern one-half of the basin. Surveys
were conducted from 12 June 1978 to 26 Au-
gust 1979 and from 13 to 15 April 1982.
Habitats were sampled with 3 m seines (9.5
mm mesh), dip nets, fish traps, backpack
electroshocker, and 15 m gill nets (51 mm
mesh). Fishes utilized in this study are depos-
ited at Oregon State University (OS), The
University of Michigan Museum of Zoology
(UMMZ), Tulane University (TU), and the
University of Nevada, Las Vegas (UNLV).

Information concerning reproduction, lon-
gevity, and adult sex ratio was determined
for Borax Lake chubs collected monthly from
March 1978 to January 1979. Fish were col-
lected from the southwest one-quarter of Bo-
rax Lake. Specimens were preserved in 10
percent formalin and transferred to 45 per-
cent isopropanol after one week. Standard
length (SL) of specimens was measured to the
nearest 0.1 mm with dial calipers. After
blotting fish dry on paper towels, wet weight
was measured to the nearest 0.01 g. A gona-
dosomatic index was calculated by weighing
the left ovary or testis to the nearest 0.001 g,
multiplying by two, thereby accounting for
the right gonad, and dividing by fish weight.
Three classes of ova were identified: class 1—
mature ova, yellow color, 0.7 to 1.2 mm di-
ameter; class II— immature ova, opaque
white color, 0.4 to 0.6 mm diameter; and
class III— immature ova, transparent, 0.1 to
0.3 mm diameter. The number of ova was
enumerated in females that possessed only
class I and/ or class II ova. By this method,
accurate counts could be obtained and these
numbers are probably more indicative of the
actual number of eggs deposited during
spawning. In females shorter than 35 mm SL,
all ova were counted in both ovaries. In fe-
males 35 to 50 mm SL, all ova were counted
in the left ovary, then multiplied by two to
derive the total number of ova. Ova were

enumerated in larger females by multiple
subsamples of the ovary. Age was determined
by examining annuli of scales taken from the
left side of the body above the lateral line.
Monthly collections were grouped into sea-
sons as follows: spring (March-N4ay), summer
(June-August), fall (September-November),
and winter (December-February).

Population Accounts

Alvord Cutthroat Trout, Salmo clarki ssp.

The Alvord cutthroat trout, Salmo clarki
ssp., is now extinct in pure form. This native
trout was known from Virgin Creek in Ne-
vada and Trout Creek in Oregon, but prob-
ably existed in several of the larger Alvord
Basin creeks during recent times (Hubbs and
Miller 1948).

Trout Creek (Harney County, Oregon).—
The Alvord cutthroat trout occurred in the
headwater canyon area of Trout Creek,
where it flows through the Trout Creek
Mountains. Introgression of introduced rain-
bow trout, Salmo gairdneri, with the native
cutthroat was already noticeable in 1934 col-
lections of fish made in Trout Creek by Carl
L. Hubbs, although trout from more head-
water localities were quite similar in appear-
ance to pure Alvord cutthroat trout (Behnke
1979). Cutthroat trout pigmentation was evi-
dent in many specimens collected by one of
us (CEB) in 1953 and 1957, but none had
basibranchial teeth. Trout collected from*
Trout Creek in 1972 (Behnke 1979) and 1978
to 1980 (our collections) exhibited only rain-
bow trout characteristics and we conclude
that the native trout has been extirpated
from this creek. Collections made from Cot-
tonwood Creek and other streams draining
the Trout Creek Mountains yielded only rain-
bow trout.

Virgin Creek (Humboldt County, Ne-
vada).—Thirty small (<15 cm) Alvord cut-
throat trout were collected from Virgin
Creek in or near Virgin Creek Gorge by Carl
L. Hubbs in 1934 (UMMZ 130532). Behnke
(1979) considers these specimens to be pure
native cutthroat trout and provides their de-
scription as follows: body with fewer than 50
relatively large round spots, spots concen-
trated posteriorly and above lateral line; few

412

Great Basin Naturalist

Vol. 43, No. 3

spots on caudal fin. Gill rakers 20 to 26. Lat-
eral series scales 122 to 152. Scales above lat-
eral line 33 to 37. Pelvic fin rays 8 or 9.
Branchiostegal rays 8 or 9. Vertebrae 59 to
63. Trout collected in 1971 (OS 3832, OS
3834) from approximately the same region of
Virgin Creek exhibited typical rainbow trout
features (Behnke 1979). During 1978 surveys,
we found only rainbow trout or introgressed
trout in Virgin Creek at and upstream of the
north end of Virgin Creek Gorge. No fish
were found in Virgin Creek Gorge at the
nexus of Hell and Virgin creeks. The up-
stream sections of Virgin Creek, in Virgin
Creek Gorge near Alkali Ranch, also are fish-
less. Although we received reports of large
trout from beaver ponds in Virgin Creek
Gorge downstream of Wilson Ranch, none
could be secured for examination. The large
number of introduced rainbow trout in Vir-
gin Creek Gorge would seem to preclude the
survival of pure Alvord cutthroat trout in
Virgin Creek. An extensive survey of Hell
Creek, the only permanently flowing tribu-
tary of upper Virgin Creek, revealed only a
single fish, which was typically rainbow in
character. This individual was apparently
able to ascend the falls separating Hell and
Virgin creeks during a flood. The negative
survey of Hell Creek in 1978 and 1979 causes
us to consider the Alvord cutthroat trout to
be extinct.

Borax Lake Chub, Gila boraxobius

The Borax Lake chub, Gila boraxobius
Williams and Bond, is restricted to the ther-
mal waters of Borax Lake and its outflows.
Gila boraxobius was described in 1980 and is
considered a dwarf relative of G. alvordensis
(Williams and Bond 1980).

Borax Lake (T37S, R33E, Sec 14; Harney
County, Oregon).— Borax Lake is a relatively
shallow 4.1 ha natural lake that receives wa-
ter from several thermal springs. These
springs issue into the bottom of the southwest
portion of the lake at approximately 35 to 40
C. Lake temperature is typically 29 to 32 C
but can vary from 17 to 35 C depending on
season, weather, and distance from the spring
sources. The water is clear. Substrates range
from rocky outcroppings in the southeast
portion of the lake to gravels in the north

and soft, easily roiled silt in the remainder of
the lake. The lake shoreline consists of salt
crusts, which have been deposited by the lake
waters. These salt deposits have built up over
hundreds or, more probably, thousands of
years until the lake is now 10 m higher in
elevation than the surrounding land. Histori-
cally, the lake waters overflowed along the
south and southwest shoreline, creating a
marsh. These outflows also provided water
for Lower Borax Lake, a reservoir southwest
of Borax Lake. Adjacent to Borax Lake are
two small pools, one about 25 m southwest of
Borax Lake and an artificial pool about 75 m
west of the lake.

Borax Lake chubs occur throughout Borax
Lake except in the hot spring inflows. Obser-
vations at Borax Lake indicated that Borax
Lake chubs avoided water with a temper-
ature above 34 C. These observations are
supported by unsuccessful attempts to chase
the chubs into hot spring inflow areas. In
aquaria. Borax Lake chubs lost equilibrium
when water temperature was raised to 34.5
C, indicating a critical thermal maximum
near this temperature. Borax Lake chubs also
occurred in small numbers in the two small
pools near Borax Lake. The pools are appar-
ently formed from Borax Lake overflow
waters.

The Borax Lake chub is a dwarf species
that typically reaches maturity at 30 to 35
mm SL. Males as small as 28.6 mm SL are
highly tuberculate and females as small as
31.8 mm SL have been found with mature
eggs. Typical adult size is 33 to 45 mm SL.
The largest male collected from Borax Lake
was 50.6 mm SL, whereas two exceptionally
large females, 90.4 and 93.0 mm SL, have
been collected from Borax Lake.

Most spawning probably occurs in early
spring, although some spawning can occur
year around. The gonadosomatic index was
highest in females during March and April,
with mature, class I ova present during
March, April, and January (Table 1). Ovaries
were usually poorly developed during May
through August. In males, the gonadosomatic
index was highest in April and September,
when testes averaged 0.97 and 1.11 percent
body weight, respectively. A search of mu-
seum specimens disclosed large females with
mature ova collected on 17 June (OS 4137)

July 1983

Williams, Bond: Native Fishes

413

and 11 September (OS 4106). Thus, spawning
may occur at any time of the year. However,
a major spawning in early spring is supported
by observation of numerous larval chubs dur-
ing April, May, and early June. Young fish,
eight to 15 mm SL, typically inhabit the shal-
low cove areas along the west and south mar-
gins of Borax Lake. Water is cool, only a few
cm deep, and vegetation is common in the
coves.

Ova number increases dramatically with
fish length. The number of ova was deter-
mined in eight females 32.7 to 93.0 mm SL,
that contained only class I and/ or class II
ova. The smallest females examined, 32.7 and
34.5 mm SL, contained 75 and 82 ova, re-
spectively. Larger females, 39.0, 39.3, 44.6,
and 49.4 mm SL contained 252, 246, 380,
and 362 ova, respectively. The exceptionally
large females, 90.4 and 93.0 mm SL con-
tained 2,143 and 6,924 ova, respectively. Al-
though females larger than 60 mm SL are
very rare in Borax Lake, their contribution to
recruitment may be substantial.

Most Borax Lake chubs live one year, with
few age I and II fish present. Annuli devel-
opment, although difficult to discern, in-
dicated that the 90.4 and 93.0 mm SL fe-
males were probably age III. This appears to
be the maximum age achieved by chubs in
Borax Lake. A length frequency analysis of
113 individuals collected 5 August 1977 ap-
pears in Figure 2. Because most spawning oc-
curs in spring, with young of approximately
10 mm SL prominent in May and June, most
fish in the August collection are probably
young-of-the-year. Some age I fish, 33 mm to
51 mm SL are present, whereas the two
larger individuals are probably age II (Fig. 2).

Most older fish are females. Adults (>33 mm
SL) typically comprised less than 25 percent
of specimens collected during March, April,
and May. The percentage of adults in month-
ly collections then increased until reaching a
peak of 82 percent during November.

Based on monthly collections made
throughout the year, the sex ratio of 190 indi-
viduals greater than or equal to 30 mm SL
was 1.0(5 :1.3 ? . Seasonally, the sex ratio was
as follows: spring (n = 23) 1.0(5 :1.3 ? , sum-
mer (n = 67) 1.0(5:1.9$, fall (n = 50)
1.0 5:11?, and winter (n = 50) 1 <5 :1 ? .
The larger number of females may be in-
dicative of higher survivorship following
spawning.

The feeding ecology of Borax Lake chubs
has been reported by Williams and Williams
(1980). The relative importance of foods var-
ied seasonally, but diatoms, microcrustaceans,
and chironomid larvae were often the pri-
mary foods consumed. Terrestrial insects
were important foods during summer and fall
(Williams and Williams 1980).

Lower Borax Lake (T37S, R33E, Sec 15;
Harney County, Oregon).— Lower Borax
Lake is a reservoir that receives water from
the southwest outflow creek of Borax Lake.
Water levels of Lower Borax Lake fluctuated
seasonally, often holding little water during
summer. Unfortunately, alteration to Borax
Lake during 1979 diverted water away from
the reservoir. Prior to the diversions, the res-
ervoir occasionally harbored Borax Lake
chubs that entered from the southwest out-
flow creek. It is doubtful that Borax Lake
chubs ever spawned in Lower Borax Lake*
and the population was probably dependent
on an influx of fish from Borax Lake via the
outflow creek.

Table 1. Monthly reproductive characteristics of female Borax Lake chubs longer than 30 mm SL.

Classes

Gonadosomatic index

of ova

Month

xSL

Range

present

March

38.0

0.91-5.01

2.89

2.05

LII,III

April

33.1

0.78-10.56

3.45

4.13

i,n,ni

May

34.8

0.45-0.55

0.51

0.05

III only

June

35.9

0.53-2.38

1.01

0.66

II,III

July

44.0

0.50-1.54

1.06

0.37

III only

August

37.4

0.16-1.60

0.95

0.41

IIJII

September

38.3

0.70-4.56

2.48

1.61

II,III

November

39.3

1.28-2.13

1.61

0.34

IIJII

December

42.5

1.20-4.45

2.02

1.03

II,III

January

41.4

0.99-2.27

1.59

0.48

I,II,III

414

Great Basin Naturalist

Vol. 43, No. 3

>«
u

3
O"
0>

n I n

57 63 6<

Standard Length in mm

Fig. 2. Length frequency of 113 Borax Lake chubs, Gila boraxobius, collected 5 August 1977 from Borax Lake,
Oregon.

Alvord Chub, Gila alvordensis

The Alvord chub, Gila alvordensis Hubbs
and Miller, was found in 16 habitats located
throughout much of the basin in Oregon and
Nevada. The species was collected from a va-
riety of habitats, including springs, creeks,
and reservoirs. The species has been collected
from Jimiper Lake, Oregon (Bond 1974), al-
though its presence appears attributable to
an introduction because the lake dries during
drought years.

Serrano Pond (T36S, R33E, Sec 1; Har-
ney County, Oregon).— Serrano Pond is a 0.1
ha reservoir that receives water from a cool-
water spring approximately 60 m distant.
Water flows from the spring at approx-
imately 17 C and water temperature in the
pond is typically 16 to 21 C during the sum-
mer. The substrate of the relatively shallow
pond is primarily silt. The water is somewhat
turbid and aquatic vegetation is abundant.
Recent alteration to this area has resulted in
a diversion canal draining part of the flow
away from the pond. Alvord chubs are absent
from the spring, but are abundant in remain-
ing waters of the pond and in the diversion
creek. More than 100 fish can be easily col-
lected from the pond in a single seine haul

during the summer. Adult males are typically
about 50 mm SL and adult females average
approximately 65 mm SL, but females great-
er than 80 mm SL are occasionally collected.

Alvord chubs from Serrano Pond are high-
ly opportunistic feeders. Bottom in-
vertebrates are grazed extensively, as are
midwater crustaceans and diatoms. Very few
foods are consumed from the water's surface.
Chironomid larvae, diatoms and cladocerans
were the principal foods during summer, and
ostracods, harpacticoid copepods, and chi-
ronomid pupae were of secondary impor-
tance (Williams and Williams 1980). Eighty-
nine percent of the intestines examined by
Williams and Williams (1980) contained one
food that accounted for more than 50 per-
cent of intestinal volume. Thirty-nine per-
cent of the intestines contained one food, chi-
ronomid larvae, diatoms, or cladocerans,
almost exclusively.

Trout Creek and Alvord Lake (Harney
County, Oregon).— Trout Creek is the largest
stream in the Alvord Basin and discharges an
average of 15 cfs, as measured in the canyon
area 8 km east of Trout Creek Ranch (Libbey
1960). The creek heads in Trout Creek
Mountains just north of the Nevada border.
In the headwaters. Trout Creek flows

July 1983

Williams, Bond: Native Fishes

415

through canyon areas where its waters are
clear and fast-flowing. Water temperature in
the canyon is cool during summer, near 15 C,
and colder during winter months. As Trout
Creek leaves the canyon and enters the val-
ley floor, flows decrease and water temper-
atures increase. Naturally lower summer
flows and irrigation diversions often reduce
the lower portions of Trout Creek to an in-
termittent stream during late summer. Turbi-
dity is often high (visibility 1 cm) in lower
reaches during summer. Substrate type
changes from mostly gravel in upstream areas
to silt in downstream sections. Trout Creek
eventually empties into Alvord Lake, a rem-
nant of the large lake that covered the valley
during pluvial times. Alvord Lake varies
greatly in size and occasionally dries com-
pletely during drought years.

Alvord chubs are common, although not
abundant, in upstream canyon areas, and
abundant in downstream sections. Introduced
rainbow trout, Salrno gairdneri, also occur in
upstream regions of Trout Creek. This is the
only habitat where Gila occur sympatrically
with another fish in the Alvord Basin. Alvord
chubs in the canyon area are large, the long-
est measuring 122 mm SL. Three specimens
113 to 122 mm SL are all age class IV.
Downstream areas also produce large Alvord
chubs, although maximum length appears
somewhat less. Color differences between Al-
vord chubs in the canyon and downstream
areas are striking. Alvord chubs from the can-
yon are very dark, nearly black, dorsally,
with golden sides possessing some black
speckles, and a silver belly. In sections of
Trout Creek in the valley floor, Alvord chubs
are lighter in color, exhibiting a light green
color on the dorsal part of the head and
body, silver sides without speckles, and a
white belly. All fins of Gila from Trout
Creek are translucent red or orange in color
except at the tips, which are white.

Pueblo Slough (T40S, R35E and T41S,
R35E; Harney County, Oregon).— Pueblo
( = Denio) Slough is a wetland area approx-
imately 13 km long, extending from Tum
Turn Lake in the north to just north of the
Nevada border in the south. Various marsh,
spring, and creek areas south of Tum Tum
Lake provide most of the habitat in the
slough. Water in Pueblo Slough is provided

by Van Horn and Colony creeks, which drain
the Pueblo Mountains, as well as at least 25
springs in the slough itself. These springs are
mostly cool and shallow. Red Point School
(T40S, R35E, Sec 14) is located in approx-
imately the center of the slough. In August,
Alvord chubs were collected from a shallow,
clear pool at Red Point School where water
and air temperatures were 15 and 17 C, re-
spectively. Alvord chubs were abundant in
the pool. Despite searches for chubs in
streams draining the Pueblo Mountains, none
could be found— although small rainbow
trout were collected in lower Van Horn
Creek.

Bog Hot Reservoir (T46N, R28E, Sec 17;
Humboldt County, Nevada).— Bog Hot Res-
ervoir is a relatively small impoundment fed
by thermal waters flowing from Bog Hot
Springs. Water issues from Bog Hot Springs
at approximately 44 C and flows for 1.2 km
before entering Bog Hot Reservoir. Water
from Bog Hot Springs enters the reservoir at
about 30 C. Typical water temperatures in
the reservoir are 20 to 21 C during early
summer. The waters of Bog Hot Reservoir
are slightly turbid (visibility 31 cm), and the
substrate is mostly silt with some gravel. Al-
vord chubs are abundant in Bog Hot Reser-
voir but are absent in Bog Hot Springs and in
practically all the inflow creek between the
springs and the reservoir. No Alvord chubs
were found upstream of the 31.1 C boundary,
where water from the inflow creek enters
Bog Hot Reservoir.

Alvord chubs collected from Bog Hot Res-
ervoir on 13 June 1978 were in spawning
condition. On that date, water and air tem-
peratures were 20.4 and 20.7 C, respectively.
The fish were collected from open, slightly
turbid water, 30 to 40 cm deep. Young-of-
the-year fish 10 to 15 mm SL were abundant
in the reservoir during June 1978, indicating
a spawning season from at least April imtil
July. Most young were observed in the shal-
low northwest end of the reservoir near the
warm inflow creek.

Bog Hot Creek (T46N, R28E; Humboldt
County, Nevada).— Bog Hot Creek flows for
approximately 5.2 km below Bog Hot Reser-
voir before entering Thousand Creek. Parts
of Bog Hot Creek below the reservoir have

416

Great Basin Naturalist

Vol. 43, No. 3

been diverted or otherwise modified by agri-
cultural practices. Alvord chubs were not
collected in Bog Hot Creek except in the
lower reaches near Thousand Creek. One
poeciliid, probably a guppy, was observed in
the highly modified section of the creek
about halfway between the reservoir and
Thousand Creek. In the downstream part of
Bog Hot Creek, Alvord chubs were rare to
common during an April 1982 svirvey and oc-
curred primarily in pools 20 to 46 cm in
depth. Water and air temperatures on 15
April 1982 were 2.3 C and 1.5 C, respective-
ly. The water was clear, but appeared brown
and quite acidic.

Unnamed spring (T46N, R26E, Sec 31;
Humboldt Covmty, Nevada).— This spring,
measuring 2.4 m wide and 5 m long at its
maximiun extent, is by far the smallest habi-
tat supporting fish in the Alvord Basin. Max-
imum depth is 77 cm. The water is clear and
the substrate is an easily roiled silt. Water
temperature was 11.4 C (air 7.5 C) during
April and 18.2 C (air 20.4 C) during June.
The unnamed spring is well isolated from
nearby Thousand Creek by approximately
100 m of greasewood flat. The closest waters
of Thousand Creek do not support Alvord
chubs. The nearest population occurs approx-
imately 1 km away in Dufurrena Pond 19.

A very small population of Alvord chubs,
estimated at slightly less than 100 individuals,
inhabits the spring. Many seine hauls, each
encompassing the entire spring, yielded 52
fish during an April 1982 survey. The Alvord
chubs ranged in size from 32 mm SL to, con-
sidering the small size of the habitat, an ama-
zingly large 87 mm SL individual. No juve-
niles were seen during April, but young-of-
the-year were observed on 14 June, when wa-
ter temperature had risen to 18.2 C. Juveniles
occurred among rushes, Juncus sp., in water
only a few cm deep. Adults were occasionally
observed darting across the open center of
the spring, but spent most of the daylight
hours under a narrow band of floating algal
mats along the periphery of the spring.

Thousand Creek Spring (T46N, R26E,
Sec 31; Humboldt County, Nevada).— Thou-
sand Creek Spring forms the headwaters of
Thousand Creek. The clear spring waters
achieve a maximum depth of approximately
31 cm. Substrates are mostly fine gravels

with some silt. Water and air temperatures
during June were 27.1 and 18.2 C, respec-
tively. Currently, the spring is inhabited by
swarms of exotic guppies, Poecilia reticulata.
Guppies have become established here and in
the spring pool at nearby Dufurrena Camp-
ground. Competition from introduced gup-
pies probably extirpated Alvord chubs from
Thousand Creek Spring. Alvord chubs have
not been recorded from Thousand Creek
Spring, but their historic presence is in-
dicated by the occurrence of Alvord chubs in
downstream areas of Thousand Creek and in
a nearby spring. Except for the presence of
guppies. Thousand Creek Spring appears to
provide a suitable habitat for Alvord chubs.

Thousand Creek and Continental Lake
(Humboldt County, Nevada).— Thousand
Creek heads at Thousand Creek Spring, flows
through the Dufurrena area, where it re-
ceives Virgin Creek, and then enters Thou-
sand Creek Gorge. Below the gorge. Thou-
sand Creek becomes braided and receives
water from Bog Hot Creek before eventually
emptying into Continental Lake. In the Du-
furrena area, Thousand Creek is dammed at
several locations to create reservoirs. Thou-
sand Creek is usually turbid (visibility 7 cm),
shallow, and about 1 to 2 m wide. The sub-
strate is mostly silt. In Thousand Creek
Gorge, the creek is surprisingly deep (>300
cm) and cool, near 15 C during late summer.
Below the gorge. Thousand Creek is often in-
termittent during summer, when water tem-
peratures can reach 27 C. Typical summer
and fall water temperature is 16 to 18 C. Al-
vord chubs are abundant in Thousand Creek
except in some upstream areas where guppies
have been introduced or habitat has been al-
tered by reservoir construction. Guppies are
abundant in Thousand Creek Spring and oc-
cur sporadically in Thousand Creek between
Thousand Creek Spring and Dufurrena Pond
19. None were found downstream of Dufur-
rena Pond 19. Thousand Creek contains
many large Alvord chubs; the largest mea-
sured 104.9 mm SL. During summer, Alvord
chubs concentrate in deep pools in down-
stream areas of Thousand Creek. Continental
Lake usually dries completely during sum-
mer, but harbors Alvord chubs during winter
months.

July 1983

Williams, Bond: Native Fishes

417

The sex ratio of 23 adults greater than 35
mm SL was 1.1 5 :1.0 ? . Food habits of Al-
vord chubs collected during Jime from Thou-
sand Creek were reported by Williams and
Williams (1980). They found 10 foods in the
intestines, of which chironomid larvae, cla-
docerans, copepods, and ostracods were of
greatest importance. Chironomid larvae oc-
curred in all intestines examined and ac-
counted for approximately 26 percent mean
volume (Williams and Williams 1980). Mi-
crocrustaceans comprised almost 45 percent
mean volimie of intestines, whereas diatoms
accoimted for only 5 percent mean volume.
No terrestrial insects were observed in the in-
testines. Alvord chubs in Thousand Creek ap-
pear to feed primarily on bottom in-
vertebrates and midwater crustaceans,
avoiding surface foods.

DuFURRENA PoND 19 (T46N, R26E, Sec
32; Humboldt County, Nevada).— Dufurrena
Pond 19, approximately 1 km downstream of
Thousand Creek Spring, is the first reservoir
on Thousand Creek. The reservoir is shallow
(typically < 50 cm) and moderately turbid
(visibility 14 cm). Alvord chubs occur in the
reservoir but are not abundant. Young-of-the-
year (<20 mm SL) were abundant during
June in the inflow diversion creek feeding the
reservoir. At this time, the inflow creek was
shallow (< 8 cm) and clear, with water and
air temperatures of 14.5 and 13.0 C,
respectively.

Dufurrena Pond 22 (T45N, R26E, Sec 2;
Humboldt County, Nevada).— Dufurrena
Pond 22 is a reservoir fed by waters of Thou-
sand and Virgin creeks. The water is very
turbid (visibility 2 cm) and shallow (typically
15 to 20 cm deep). Water level fluctuates
greatly with season. During late summer, the
reservoir is reduced to a small pool. Water
and air temperatures during June were 17.6
and 13.2 C, respectively. Alvord chubs are
abundant and achieve a large size in the
reservoir.

Virgin Creek (Humboldt County, Ne-
vada).— Virgin Creek heads near the southern
extent of the Alvord Basin, flows north
through the more than 300-m-deep Virgin
Creek Gorge, and then flows east until reach-
ing Thousand Creek. Springs scattered along
most of the length of Virgin Creek provide
its flow. Alvord chubs are abundant in Virgin

Creek from the north end of Virgin Creek
Gorge to the nexus of Virgin and Thousand
creeks. Alvord chubs are absent in Virgin
Creek Gorge, where introduced rainbow
trout are common. Virgin Creek below the
gorge is cool, relatively shallow, and moder-
ately turbid (visibility 10 cm). Below the
gorge, water temperature is typically 15 to
18 C during summer months and maximum
depth is usually less than 75 cm. The sub-
strate is mostly silt.

Warm Spring (T45N, R25E; Humboldt
County, Nevada).— Warm Spring and its out-
flow are tributary to Virgin Creek approx-
imately 2.5 km downstream from Virgin
Creek Gorge. The outflow creek is a small,
clear-water stream with an easily roiled silt
bottom. Summer water and air temperatures
of the creek just below the spring were 26.0
and 26.4 C, respectively. Alvord chubs are
abundant in the outflow creek. The spring it-
self was not sampled, but Carl Hubbs col-
lected Alvord chubs from the spring in 1934
("Italian Camp Spring," UMMZ 130533).
The steep gradient and low flow of the
Warm Spring crenon as it approaches Virgin
Creek inhibits mixing between the Virgin
Creek and Warm Spring populations of Al-
vord chubs. Although Warm Spring flows
through Virgin Valley Ranch, the spring and
outflow creek have been only slightly altered
and no exotic fish were present during a 1978
survey of the spring system.

Dufurrena Pond 13 (T45N, R26E, Sec
17; Humboldt County, Nevada).— Dufurrena
Pond 13 is the only reservoir on Virgin Creek
between Virgin Creek Gorge and Thousand
Creek. Water characteristics are typically
those of Virgin Creek except that maximum
depth is greater (almost 2 m) and aquatic
vegetation is abundant. Alvord chubs are
common to abundant just upstream and
downstream of the reservoir but are rare in
the pond itself.

Gridley Springs (T44N, R27E, Sec 22;
Humboldt County, Nevada).— Gridley
Springs is a series of approximately 17 cool-
water springs located on an alkali flat just
south of the Gridley Lake playa. Many of the
springs are little more than seeps, but a few
have spring pools nearly 2 m deep with out-
flow creeks 30 or 40 m in length. During
April 1982, most of the spring waters were

418

Great Basin Naturalist

Vol. 43, No. 3

10-

■R

27 33 39 45 51 57 63 69

Standard Length in mm

Fig. 3. Length frequency of 44 Alvord chubs, Gila alvordensis, collected 14 April 1982 from Gridley Springs,
Nevada.

clear, with a temperature of 12 C (air 10 C).
Rushes, Juncus sp., were the dominant plants
around the springs, with some larger springs
also harboring pondweed, Potamogeton sp.,
and cattails, Tijpha sp. Only one of the 17
springs examined contained Alvord chubs.
This spring is located near the northwestern
margin of the Gridley Springs series. Alvord
chubs were found in the outflow creek,
which extended approximately 40 m and was
3 m wide at its greatest extent. Only a trace
of current could be detected in the creek.
The water was imusually turbid, visibility 4
cm, with a maximimi depth of 30 cm. Water
temperature was 11.5 C (air 6.8 C) on 14
April 1982. The bottom was silt. Unlike most
of the larger springs in the area, this spring
contained only rushes along its margin. The
Gridley Springs area is overgrazed by cattle
and horses, but it is not known to what extent
this is detrimental to the Alvord chub
population.

Alvord chubs were not abundant in the
outflow creek, but were common enough to
collect 50 fish in three short seine hauls.
Forty-four individuals collected in April 1982
ranged in size from 27 to 91 mm SL, but
were mostly 30 to 38 mm SL (Fig. 3). The sex
ratio of 32 individuals greater than 35 mm
SL was 1 5 :3 ? .

West Spring (T44N, R27E, Sec 20; Hum-
boldt County, Nevada).— West Spring issues
from the base of Big Mountain and forms
West Creek, which flows into the alkali flat
south of Gridley Springs. The waters of West
Spring are clear and shallow. Maximum
depth is 12 cm over a gravel and sand sub-
strate. Water and air temperatures recorded
during summer were 21.8 and 20.4 C, respec-
tively. Alvord chubs are common in the

spring and its outflow creek. The largest of
33 individuals collected on 18 August 1978
was 62.8 mm SL. The sex ratio of 30 adults
greater than 40 mm SL was 1^:1$. Alvord
chubs collected during August from West
Spring fed exclusively or almost exclusively
on small hydrobiid snails. This snail, which
occurs in great abundance, apparently repre-
sents an undescribed species endemic to West
Spring (Jerry Landye, pers. comm.).

West Creek (T44N, R27E, Sec 20, 28, 29;
Humboldt County, Nevada).— West Creek
flows for nearly 3 km before emptying into
the alkali flat approximately 2 km south of
Gridley Springs. Because West Creek is
formed by West Spring, water characteristics
are similar for both. The creek is quite small,
often 1 to 2 m in width and less than 15 cm
deep. Current is moderate in the upper
reaches but slows considerably upon reaching
the flat. Alvord chubs occur throughout the
creek, but are somewhat smaller than those
in West Spring.

Discussion

The Alvord cutthroat trout, Alvord chub,
and Borax Lake chub are all restricted in dis-
tribution to waters of the Alvord Basin. The
Alvord cutthroat trout is now extinct but for-
merly occurred in larger creeks of the basin.
Hybridization with introduced trout caused
the demise of the native form. The Borax
Lake chub has the most restricted natural dis-
tribution of the three fishes, occurring only in
Borax Lake and adjacent lake outflows. The
Alvord chub is relatively widespread in the
basin and was recorded from 16 localities, in-
cluding Bog Hot Reservoir, Bot Hot Creek,
Thousand Creek Spring, an unnamed spring.

July 1983

Williams, Bond: Native Fishes

419

Dufurrena Pond 19, Dufiirrena Pond 22, Du-
furrena Pond 13, West Spring, and West
Creek as new locality records.

Because of the fragility of the small aquat-
ic habitats and the overall paucity of water in
the basin, the two extant native fishes are
easily threatened by the activities of man.
The naturally restricted range of the Borax
Lake chub and threats from geothermal
energy development prompted the American
Fisheries Society to list the species as threat-
ened in 1979 (Deacon et al. 1979). Also dur-
ing 1979, several portions of the north and
east shoreline of Borax Lake were altered so
that overflow waters exited the lake to the
north and east rather than to the south and
west, as was the historical condition. This al-
teration caused Lower Borax Lake as well as
the marsh and pools to the south and west to
dry, thus eliminating Borax Lake chubs from
these waters. Leasing of surrounding lands
for geothermal exploration and alteration of
the shoreline caused the U.S. Fish and Wild-
life Service temporarily to list the Borax
Lake chub as an endangered species on 28
May 1980. As a result of the listing, geother-
mal exploration was prohibited from a one-
mile buffer zone around Borax Lake. The
emergency listing has since been supple-
mented by a final rulemaking that designated
the species as endangered pursuant to the
Endangered Species Act. The Alvord chub
has fared better than the Borax Lake species
because of its wider distribution. Never-
theless, competition with exotic guppies has
extirpated the Thousand Creek Spring popu-
lation of Alvord chubs, and other populations
are threatened by habitat alteration. The Al-
vord chub appears easily eliminated by the
presence of exotic fishes. Thousand Creek
reservoirs stocked with game fish, such as
Dufurrena Ponds 20 and 21, lack Alvord
chubs. White crappie, Pomoxis annularis,
pumpkinseed, Lepomis gibbosiis, and large-
mouth bass, Micropterus sahnoides, were col-
lected from Dufurrena Ponds 20 and 21.

Borax Lake chubs are dwarf and typically
mature at 30 mm SL. Adults are usually 33
mm to 45 mm SL and typically live for one
year. A few Borax Lake chubs, mostly fe-
males, live more than one year. Adult Alvord
chubs are larger, achieving more than 100
mm SL in Trout and Virgin creeks. Even in

very small springs, such as the unnamed
spring and Gridley Springs, Alvord chubs
achieve 90 mm SL. The presence of large
chubs in the cool springs and creeks indicates
a longer life span for the Alvord chub than
typically occurs for the Borax Lake species.
Borax Lake chubs spawn year around in their
thermal lake habitat, but a spring spawning
peak is indicated. Alvord chubs appear to
spawn only once a year in their thermally
fluctuating habitats. Both species of Gila are
opportunistic omnivores, consuming primari-
ly chironomids, microcrustaceans, and dia-
toms. The Borax Lake species also consumed
large quantities of terrestrial insects during
summer and fall. The Alvord chubs in West
Spring are unusual in that they are greatly
dependent on the endemic hydrobiid snail for
food.

Acknowledgments

Funds to study the Alvord Basin fishes
were provided by U.S. Fish and Wildlife Ser-
vice contract 14-16-0001-78025 to inventory
the fishes of the Sheldon National Wildlife
Refuge and by the Department of Fisheries
and Wildlife at Oregon State University. The
study greatly benefited from the field and
editorial assistance of Cynthia D. Williams.
Ray S. Taylor, J. J. Long, K. M. Howe, G.
DeMott, B. Boccard, A. Tiehm, J. E. Deacon,
M. S. Deacon, D. E. Deacon, and E. M. Lo-
rentzen assisted with field collections. Robert
R. Miller lent museum specimens and collec-
tion notes of our late mentor, Carl Leavitt
Hubbs. Reviews of this paper were provided
by C. D. Williams, J. E. Deacon, E. P. Pister,
and S. V. Gregory. This contribution is Tech-
nical Paper 6510 of the Oregon Agricultural
Experiment Station.

Literature Cited

Behnke, R. J. 1979. Monograph of the native troiits of
the genus Salmo of western North America. Re-
port to U.S. Fish and Wildlife Service. 215 pp.

Bond, C. E. 1974. Endangered plants and animals of
Oregon I. Fishes. Agricultural Exp. Station, Ore-
gon State Univ. Spec. Rept. 205. 9 pp.

Deacon, J. E., G. Kobetich, J. D. Williams, S.
Co.NTRERAS, et al. 1979. Fishes of North America
endangered, threatened, or of special concern:
1979. Fisheries 4:29-44.

Hubbs, C. L., and R. R. Miller. 1948. The zoological
evidence: correlation between fish distribution

420

Great Basin Naturalist

Vol. 43, No. 3

and hydrographic history in the desert basins of
western United States. Pages 17-166 in The
Great Basin, with emphasis on glacial and post-
glacial times. Bull. Univ. Utah, Vol. .38.

HuBBS, C. L., AND R. R. Miller. 1972. Diagnoses of new
cyprinid fishes of isolated waters in the Great Ba-
sin of western North America. Trans. San Diego
See. Nat. Hist. 17:101-106.

LiBBEY, F. W. 1960. Boron in Alvord Valley, Harney
County, Oregon. Ore-Bin. 22:97-105.

Snyder, C. T., G. Hardman, and F. F. Zdenek. 1964.
Pleistocene lakes in the Great Basin. U.S. Geol.
Surv., Misc. Geol. Investigations, Map 1-416.

Williams, J. E. 1980. Systematics and ecology of chubs
{Gila: Cyprinidae) of the .\lvord Basin, Oregon
and Nevada. Unpublished dissertation. Oregon
State Univ. 175 pp.

Williams, J. E., and C. E. Bond. 1980. Gila boraxohius,
a new species of cyprinid fish from southeastern
Oregon with a comparison to G. alvordensis
Hubbs and Miller. Proc. Biol. Soc. Washington
93:293-298.

Williams, J. E., and C. D. Williams. 1980. Feeding
ecology of Gila boraxohius (Osteichthyes: Cypr-
inidae) endemic to a thermal lake in southeastern
Oregon. Great Basin Nat. 40:101-114.

KRAMER PALOUSE NATURAL AREA

Del W. Despain^'^ and Grant A. Harris^

Abstract.— The 27-acre Kramer Palouse Natural Area located in southeastern Washington State represents the
best remaining example of what was once the most productive portion of the original Palouse Prairie. This area is
being maintained in its pristine condition as a key to the past and as a memory to a unique and once extensive
prairie land by the Department of Forestry and Range Management at Washington State University.

Many of the most productive rangelands of
the past are now agricultural lands. The Pa-
louse Prairie of the inland Pacific Northwest
is no exception. Once part of a vast prairie-
land that extended throughout eastern Wash-
ington and Oregon, as well as adjacent Idaho,
most of this region has since been turned un-
der by the plow, to become some of the most
productive unirrigated farmland in the
world.

The Palouse Prairie occupies a region of
relatively gentle topography, with the princi-
pal relief being low hills having the general
appearance of dunes. These wind-deposited
loesal materials originated in the arid lands
and volcanoes to the west, and were depos-
ited on a basalt rock foundation. Fertile xe-
rolls of silty and clay loam texture have de-
veloped under the influence of a semiarid
climate. Average armual precipitation is from
about 400 to 600 mm (16-24 inches), coming
mostly as rain or snow during fall, winter,
and spring. Prefarming era vegetation was
characteristic of a true grassland region, and
was composed of dense stands of caespitose
perennial grass species (Daubenmire 1970).

The climax vegetation, though palatable
and nutritious, apparently developed without
significant grazing use. Dominant species are
easily injured by close cropping, and under
poor grazing management are replaced by
introduced annual grasses (chiefly Bromiis
tectorum). Archeologists estimate that the
small bison poulation of the region became
extinct about 2,000 years ago, and large her-
bivore grazing was practically nil from that
time until horses from early Spanish missions

of the southwest were introduced in about
1730 (Osborne 1953). The region is so re-
markably adapted to intensive wheat and pea
cropping that today livestock grazing never
has become an important land use, except in
waste places.

With the development of a side-hill com-
bine and other technologically advanced
farm equipment, only the very steepest "eye-
brows" and slopes of the loesal deposits of
the Palouse have not been tilled. Many of
these small islands of native prairie have
been used for other purposes and have
changed dramatically over the past. Most re-
maining uncultivated segments are so small
as to have been changed to dense stands of
brush or weeds through activities on adjacent
land including tillage, herbicide applications,
and fertilization.

Probably the best remaining example of
the more productive portions of this region is
the Kramer Palouse Natural Area. Ownership
has been obtained by Washington State Uni-
versity in an effort to retain this unique spec-
imen in its natural condition as a reference
point to the past for demonstration and
research.

Historical Background

The 27-acre Kramer Palouse Natural Area
was once part of a producing wheat farm,
but it was held out of production by virtue of
steep topography and shape of the ownership
boundaries of the farm. It reaches over the
crest of a high ridge, too steep for even the
intrepid Palouse area farmers to cultivate.

'Department of Forestry and Range Management, Washington State University, Pullman, Washington 99163.
'Present address: School of Renewable Natural Resources, University of Arizona, Tucson, Arizona 85721.

421

422

Great Basin Naturalist

Vol. 43, No. 3

The ownership boundaries included the
steepest part of the ridge top, with neighbors
on three sides, and no access to the back side
without crossing their land. Consequently,
the area was not plowed until 1961 when the
owner decided to cultivate the lower north-
em extent of the parcel. However, after hav-
ing made one pass with the plow, the farmer
says he didn't have the heart to continue and
left the tract untouched by further dis-
tubance. The path of that one pass, half-
circle in shape along the base of the ridge,
can still be faintly seen today, but the vegeta-
tion is now similar to adjacent sites.

The area was occasionally used as a pas-
ture for farm milk cows, but only a small part
was noticeably changed where the cattle con-
gregated for resting. A road that once tra-
versed the area for movement of farm equip-
ment and animals is the most evident scar of
disturbance from the past, but even this has
healed over with native vegetation similar to
the undisturbed portions.

Dr. Rexford F. Daubenmire is reported to
have found the area in about 1955 during his
search for bench mark natural areas in sup-
port of his ecological studies. He established
permanent study plots there at that time and
continued to make observations as needed.
He later showed the area to Dr. Grant A.
Harris, who pursued the possibility of pur-
chasing the tract through John P. Nagle, then
chairman of the Department of Forestry and
Range Management at Washington State
University. The parcel, previously owned by
Caroline Kramer, was not immediately avail-
able for purchase. However, following pro-
bate settlements for the Kramer estate, the
University was able to purchase the 27-acre
tract. The deed was filed in the imiversity's
name on 28 March 1962. The Department of
Forestry and Range Management, with spe-
cific assignment to Dr. Harris, was given the
responsibility for administration and mainte-
nance of the Natural Area, and this assign-
ment continues at present.

Description

The Kramer Palouse Natural Area, 27
acres in size, is located about 5 miles west of
the farming community of Colton, Whitman
County, in the southeastern portion of the

state of Washington (N %, SE V4, NE V4 of
Section 25, Township 13 North, Range 44
East, WPM). Aspects are generally steep on
the south and north, with elevations of 805 to
869 m. Precipitation at the site averages ap-
proximately 550 mm (22 inches) annually.

Zonal vegetation in this environment is ex-
pressed in the Festuca idahoensis/
Symphoricarpos alhus association (Dauben-
mire 1970). A major part of the north slopes
of the Natural Area supports this habitat
type. Major species include the caespitose
grasses, Idaho fescue {Festuca idahoensis),
bluebunch wheatgrass {Agropijron spicatum),
and June grass (Koelaria cristata), in associ-
ation with shrubs dominated by snowberry
{Symphoricarpos alhus) and wild rose {Rosa
nutkana and Rosa woodsii). The type has a
rich diversity of perennial forbs, creating a
virtual sea of flowers across the Natural Area
during much of the growing season. The
snow-catching boundary fences along the
north slope have sufficiently altered the mi-
croclimate in places to increase the normal
coverage of the tall shrub phase {Sym-
phoricarpus), creating a border of dense
shrubs including chokecherry {Frunus virgi-
nana) and occasionally bittercherry {Frunus
emarginata). There are also small patches of
the black hawthorn/cow parsnip {Crataegus
douglassii/ Heracleum kinatum) habitat type
at the base of the steep north slopes, typical
of bottom lands that are more moist than lo-
cal zonal soils. The south face of the Natural
Area supports a topographic climax cover of
the bluebunch wheatgrass/ Sandberg's blue-
grass {Agropyron spicatum /Foa sandhergii)
habitat type typical of the more droughty
vegetation zone to the west.

Detailed soil and vegetation studies by Al-
ler et al. (1981) on the site have revealed a
"perched water table" situation in the solid
steep north exposures. On this site, as well as
two other similar locations, they found an un-
described topoedaphic climax plant as-
sociation dominated by Carex geyeri, as well
as a Festuca phase and Symphorocarpos phase
of Daubenmire's Festuca idahoensis/
Symphorocarpos alhus association that had
not previously been reported.

Soils on the site have been classified into
the Palouse and Calouse series, which are rel-
atively deep, well-drained soils that formed

July 1983

Despain, Harris: Kramer Palouse Area

423

in loess deposits containing varying amounts
of volcanic ash (Donaldson 1980). Approx-
imately 80 percent of the surface is classified
as Palouse-Thatuna silt loam, 10 percent Pa-
louse silt loam (SE comer), and 10 percent
Calouse silt loam (SW corner). The Palouse-
Thatima series is a complex of approximately
50 percent Palouse and 50 percent Thatuna
silt loam, mapped together because of their
intermingled occurrence in the field. The Pa-
louse silt loams are generally found on con-
vex slopes and Thatuna on concave slopes,
examples of which are found on south and
north exposures of the natural area, respec-
tively. All the soils found there are deep,
well-drained, and fertile, and, in cultivation,
produce excellent crops of wheat, lentils,
peas, barley, and alfalfa.

An abimdance of wildlife inhabits the area,
including coyotes, badgers, and occasionally
whitetail deer.

Management philosophy encourages non-
destructive scientific study of entire ecosys-
tems found in the Natural Area. Scientists
from several imiversity departments, includ-
ing the Universities of Idaho and Washing-
ton, as well as from state and federal
agencies, have established short-term biologi-
cal studies there. In addition to vegetation,
studies have investigated natural status of
soils, rodents, insects, birds, and atmosphere,
all in considerable detail. Interest in the Pa-
louse Natural Area as a bridge to the past
continues to increase as research interest in
biological subjects increases, and the area
will become ever more valuable as time
passes.

Access to the Natural Area is somewhat
hampered because it is isolated from the lo-
cal road system and is completely surrounded
by cultivated land. The lack of easy access,
however, simplifies maintenance of the un-
disturbed characteristics of the area. Ar-
rangements have been made with the neigh-
bor on the west to walk one-half mile along
his fence line southward from Rim Road near
its junction with the Colton— Wawawai
Road. This provides the principle access for
observation and study.

Management Problems

Maintaining a natural area in its natural
state, especially one as small as the Kramer

Area, is not easy. Most of the problems stem
from the "unnatural" interface of the area
with adjacent cultivated farmland.

There is some indication that the dense
cover of tall shrubs that has developed along
some of the fence lines is migrating inward
and increasing its extent over the area. The
potential loss of the grass-forb- and low-
shrub-dominated communities may become a
concern in the future.

Because of good cover and protection pro-
vided by the shrubs along the fence line, and
because of availability of adjacent cultivated
crops as forage, the population of Columbian
ground squirrels (Citellus columhianus) has
increased dramatically around the perimeter
of the track. In the past, these rodents have
stripped adjacent field crops as far as 100
feet out from the boundary. Following com-
plaints from neighboring farmers, attempts
were made to control excessive populations
of the squirrels, but most methods considered
or tried proved to be either largely unsuc-
cessful or involved equipment that was diffi-
cult to get to the inaccessible area or that
would cause undue disturbance. Recently,
however, a very successful control effort has
been implemented using a product with the
trade name "Ramik Green," made by Velsi-
col Chemical Corporation. This rodenticide
was placed in bait stations located around the
perimeter of the area during early spring.
The immediate question that has been raised
is whether or not this control of rodents
changes the natural balance of the ecosystem.
It is our opinion that rodent populations pri-
or to control were artificially high due to the
adjacent crops as a forage source. No attempt
has been made to completely destroy rodent
populations, with control efforts directed
only at critical locations around the perime-
ter of the Natural Area. The alternative, legal
action taken by neighboring farmers due to
obvious crop damage, would clearly endan-
ger the status of the Natural Area.

Another problem of the intensive agricul-
ture interface is herbicide drift from adjacent
fields, primarily 2-4-D. The most obvious im-
pact is damage to shrubs along the bound-
aries. What changes in species composition
herbicide drift causes is unknown at this
time, but it does not appear to be a serious
problem as far as maintaining natural condi-
tions over most of the area.

424

Great Basin Naturalist

Vol. 43, No. 3

Except for cheatgrass [Bromus tectorum)
along the ridge top, noxious, introduced
weeds now common to the region have
largely been excluded from the unit by the
well-established stands of native vegetation.
Canada thistle {Cirsium arvense) has ap-
peared in a couple of places and could possi-
bly become a problem to deal with in the
future.

Research use is encouraged, but care is
taken to limit the kinds of uses to those that
can be made without significant disturbance
to its natural character. A system of approval
has developed in which the applicant submits
a copy of a detailed proposal for review by
the administrator. If destructive procedures
are found, negotiations usually modify these
so that the work can be completed. Visits by
undergraduate classes are not encouraged.

Due to limited access to the Natural Area,
disturbance by the general public has not
been a problem. Public use has been limited
primarily to upland game-bird hunters.

Literature Cited

Aller, a. R., M. a. Fosberg, M. C. LaZelle, and A. L.
Falen. 1981. Plant communities and soils of
north slopes in the Palouse region of eastern
Washington and northern Idaho. Northwest Sci.
55(4):248-262.

Daubenmire, R. F. 1970. Steppe vegetation of Washing-
ton. Washington Agr. Res. Center Tech. Bull. 62,
Pullman, Washington 99164.

Do.NALDSON, N. C. 1980. Soil survey of Whitman Coun-
ty, Washington. Soil Conser. Serv. U.S. Dept.
Agr. in cooperation with Washington State Univ.

Osborne, D. 1953. Archaeological occurrence of prong-
horn antelope, bison, and horses in the Columbia
Plateau. Sci. Monogr. 77:260-269.

WINTER FOOD HABITS OF COUGARS FROM NORTHEASTERN OREGON

Chris Maser' and Ronald S. Rohweder-

Abstract.— Sixty-four cougar {Felis concolor) stomachs and 41 intestinal tracts were examined for food items in
northeastern Oregon from 1976 through 1979. Food items, in order of decreasing frequency, were mule deer {Odo-
coileus hemionus). North American elk (Cervus elaphus), porcupine (Erethizon dorsattim), snowshoe hare [Lepus
americanus), and deer mouse {Peromijsciis maniciilatus).

Bounties existed on cougars in Oregon
from 1843, when the Oregon Territorial Gov-
ernment offered them on most "predators,"
until the boimty system was repealed by the
1961 Oregon Legislature (Ebert 1971, Kebbe
1961). Once found throughout most of Ore-
gon, the decline of the cougar during the
1950s and early 1960s aroused concern for its
continued existence within the state. The
cougar was classified as a game animal in
September 1967 (Oregon State Game Com-
mission 1967). The hunting season was imme-
diately closed and remained closed until De-
cember 1971, when the first controlled
hunting season was opened. Twenty-two cou-
gars were killed by hunters during the De-
cember 1971 and December 1972 himting
seasons.

Little is generally known about food habits
of the cougar (Young and Goldman 1964),
particularly in Oregon, where indiscriminate
boimty hunting kept cougar populations too
low for such studies. Maser et al. (1981) and
Toweill and Meslow (1977) discussed cougar
food habits in general; Toweill and Meslow
(1977) also discussed the food habits of those
cougars killed during the 1971 and 1972
hunting seasons. The purpose of this paper is
to present data on the winter food habits of
cougars from northeastern Oregon and to of-
fer some tentative interpretations of these
data.

Methods

Sixty cougars were obtained from hunters
during four one-month hunting seasons (De-

cember 1976, 1977, 1978, 1979) in north-
eastern Oregon counties: Baker (5), Union
(15), and Wallowa (40). An additional four
cats were obtained from Umatilla (1) and
Wallowa (3) counties; one was killed illegally
and three were killed because of their prox-
imity to livestock.

Cats, killed by himters, were brought into
an Oregon Department of Fish and Wildlife
office within 48 hours of being killed. Most
cats were received intact, but a few had been
field dressed.

Each individual was sexed, weighed, mea-
sured, and, if intact, eviscerated. Each cat's
heart, lungs, liver, stomach and intestinal
tract, and reproductive organs were placed in
separate plastic bags, labeled, and quick-
frozen for later analysis. The present food
habit study was done in conjunction with a
study of endoparasites, which necessitated
separately examining the stomach, small in-
testine, and large intestine. Each cougar thus
had three separate analyses for food items.
This procedure worked well because we
could determine what appeared to be tlie
contents of two meals for each cat that con-
tained food in its alimentary canal— one meal
in the stomach and a different meal in the co-
lon. The two meals "mixed" in the small in-
testine. Thus, by identifying the stomach con-
tents first, the colon contents second, and the
small intestine contents third, we had a cross-
check on the content determinations. Mate-
rials from the small intestine usually con-
tained elements of both stomach and colon
contents and have not been included in the
discussion. Further, by identifying the plant

'U.S. Department of the Interior, Bureau of Land Management, Forestry Sciences Laboratory, 3200 Jefferson Way, Corvallis, Oregon 973.3L
■Oregon Department of Fish and WildUfe, Route 2, Box 2283, La Grande, Oregon 97850.

425

426

Great Basin Naturalist

Vol. 43, No. 3

material in the alimentary canal, we could
determine, by tree associations and by infer-
ence, the general habitat within which the
cougar had taken its meal.

Results and Discussion

Results of the food habit study are given in
Tables 1 and 2. The sexes of cougars analyzed
during this study probably approximates a
random sample (Table 3). Although cougars
are, in some sense, opportunistic predators
(Robinette et al. 1959, Sitton 1977) their
main diet was mule deer. North American
elk, and porcupine (Table 1), which concurs
with studies reviewed in Toweill and Meslow
(1977). Stomachs ranged from being empty to
containing 3.5 kilograms of mule deer.

Mule deer was the most frequently con-
sumed prey; North American elk was second
(Table 1). In eight cases, the consumed elk
could be classified as adult or calf. From the
limited sample, the five male cougars that
had eaten, and presumably killed, adult elk
were in the upper size-limit of the overall
sample: 68 kg (150 lbs)— the largest cougar—
66 kg (146 lbs), 64 kg (140 lbs), 64 kg (140
lbs), and 62 kg (137 lbs). The average weight
of the five male cats was 64.8 kg (142.7 lbs).
On the other hand, the three cats that had ea-
ten known calf elk were a female (32 kg— 70
lbs), a male (41 kg— 91 lbs), and another fe-
male (50 kg— 110 lbs). The average weight of
these three cats was 41 kg (91 lbs). The 50-kg
female that killed a calf elk was 12.3 kg (27
lbs) lighter than the smallest of the males that
killed an adult elk. Thus it seems that the
larger a cougar, the larger a prey animal it
can kill, and the more energy efficient such a
kill will be.

Porcupine would seem to be an energy-
efficient meal as soon as a young cougar is
old enough to kill because these large rodents
are slow, easily caught, and seem to be read-
ily dispatched by cougars. Although porcu-
pines occurred in the diet with only 10.6 per-
cent frequency in stomachs and 5.2 percent
frequency in colons in our study, Robinette et
al. (1959) found them to account for 19 per-
cent of the cougar's diet, based on scat analy-
sis, in Utah and Nevada. Evidence— in the
form of quills embedded in and around the
gum lines, the skinned shoulders and feet, and
embedded in stomach walls— indicated that
most cougars encounter porcupines at some
time during their life. Such quills, repre-
sented by their embedded tips, appear as
dark streaks. Apparently, a cougar's body
readily absorbs the softer, light shaft of a
quill but not the harder, dark tip.

Cougars seem to be variously adept at eat-
ing porcupines. For example, some cougars
appear to avoid the quills as much as possible
and have only a few hairs mixed with the
porcupine flesh in their digestive tract,
whereas others eat almost everything. In ad-
dition, a cougar killed in 1973 had eaten a
porcupine about an hour prior to being shot.
It had consumed the entire porcupine, except
the head and digestive tract. The quills had
already begun to soften in the cat's stomach.

The proportion of a cougar's diet that is
composed of porcupine is probably related to
the availability of the prey. Connolly (1949
cited in Robinette et al. 1959), for example.

Table 2. Miscellaneous associated items consumed by
60 cougars killed in December.

Content

Stomach
% frequency

Colon
% frequency

Table 1. Prey species

consumed by b

0 cougars killed

Grass

20.0

23.8

in December.

Douglas-fir needles

17.1

4.8

Grand fir needles

14.3

19.0

Stomach

Colon

Ponderosa pine needles
Engelmann spruce
needles

14.3

Content

% frequency

11.4

Mule deer

55.3

42.1

9.5

Elk

21.3

15.8

Twigs

5.7

4.8

Porcupine

10.6

5.2

Soil

5.7

23.8

Unidentified hair

6.4

10.5

Alder leaf

2.9

—

Snowshoe hare

4.3

Lichen (Alectoria

Bird

2.1

fremonti)

2.9

—

Lagomorph

—

5.2

Larch needles

2.9

Cougar hair

21.1
99.9

Pebbles
Total

2.9

—

Total

100.0

100.1

100.0

July 1983

Maser, Rohweder: Winter Food of Cougars

427

indicated that, in his Utah study area, cougars
killed one porcupine per week in winter. Of
the 64 cougars examined in this study, plus
97 cougars examined prior to this study, none
showed ill effects from encounters with por-
cupines, even when quills remained em-
bedded in a cat's tissues. Such lack of serious
damage or infection from porcupine quills
has also been noted in the fisher {Maries pen-
nati) and spotted skunk {Spilogale piitorius)
(Maser et al. 1981).

The snowshoe hare occurred fourth (of the
identifiable items) in the cougars' stomach
contents (Table 1). Because these hares were
relatively abundant in the coniferous forests
of northeastern Oregon during our study,
their low frequency (4.3 percent) indicates
that they were taken incidentally by the
cougars.

The lagomorph remains in the colon
(Table 1) were either snowshoe hare or
mountain cottontail (Sylvilagus nuttalli), but
they could not be identified to species once
they reached the colon. The bird (Table 1)
was probably a grouse.

Miscellaneous items associated with food
are given in Table 2. Other than grasses,
some of which were intentionally eaten, iden-
tifiable vegetation gave clues to the habitats
in which the cats presumably had been hunt-
ing and had consumed their prey. Of the five
stomachs that contained elk and vegetation,
Engelmarm spruce {Picea engelmannii) oc-
curred in 20 percent, western larch (Larix oc-
cidentalis) in 20 percent, grand fir {Abies
grandis) in 40 percent, and ponderosa pine
{Pinus ponderosa) in 20 percent. Of the 13
stomachs that contained deer and vegetation,
Engelmann spruce occurred in 15 percent,
grand fir in 8 percent, ponderosa pine in 38
percent, and Douglas-fir {Pseudotsuga men-
ziesii) in 38 percent. From the conifer nee-
dles in the stomachs, it seems that elk were
killed primarily in denser, moister forests be-
cause the Engelmann spruce, western larch,
and grand fir accoimted for 80 percent of the
needles, whereas ponderosa pine, character-
istic of more open habitat, accounted for
only 20 percent. With respect to mule deer,
on the other hand, ponderosa pine-Douglas-
fir, which occiu-s as a drier, more open forest,
accounted for 76 percent of the conifer nee-
dles, as opposed to the moister, denser forests

of grand fir and Engelmann spruce, which
represented 23 percent of the needles. Thus,
it seems that the elk were usually killed in
dense forest where the advantage would lie
with the stalking cougar and the smaller,
more easily subdued mule deer was most of-
ten hunted in more open habitats.

In addition to vegetation, several cats had
eaten much soil— evidence of having cleaned
up a kill.

Conclusions

Of the 60 cougars killed during the De-
cember hunting season, 31.6 percent had vir-
tually or totally empty stomachs, and 30 per-
cent had empty colons. The 4 cougars killed
because of their proximity to livestock also
had virtually empty stomachs. Thus, if the
stomachs or the colons are used independent-
ly as the sole source of food habit data, a
large sample is needed. If, on the other hand,
both the stomach and colon contents are used
independently but together as dietary sam-
ples, the chances of getting adequate food
habit data are good because it is unlikely that
both stomach and colon are simultaneously
empty.

Acknowledgments

Murray L. Johnson (Puget Sound Museum
of Natural History, University of Puget
Sound, Tacoma, Washington), Donald K.
Grayson (Department of Anthropology, Uni-
versity of Washington, Seattle, Washington),
Maurice Hornocker (USDI, Fish and Wildlife
Service Cooperative Wildlife Research Unit,
University of Idaho, Moscow, Idaho), and
Paul E. Ebert (Oregon Department of Fish
and Wildlife, Portland, Oregon) read and im-
proved the paper. Phyllis Taylor-Hill (USDI

Table 3. Year of capture, number, and sex of cougars
studied in northeastern Oregon.

No. of

Year

cougars

males

females

1976

1977

1978

1979

Total

428

Great Basin Naturalist

Vol. 43, No. 3

Bureau of Land Management, Forestry Sci-
ences Laboratory, Corvallis, Oregon) typed
the various drafts of the manuscript.

The following Oregon Department of Fish
and Wildlife personnel helped collect the
cougar viscera: Ronald Bartels, Vic Coggins,
Paul Ebert, Mark Henjum, Dick Humphreys,
Mike Kemp, and Walt Van Dyke. Laboratory
space and partial financing for this study
were provided by the USDA Forest Service,
Pacific Northwest Forest and Range Experi-
ment Station, Project 1701, Range and Wild-
life Habitat Laboratory, La Grande, Oregon.
We sincerely appreciate their help.

Literature Cited

Connolly, E. J., Jr. 1949. Food habits and life history of
the mountain Hon, Felis concolor hippolestes. Un-
pubUshed thesis, Univ. of Utah, Salt Lake City.
176 pp. (Cited in Robinette et al, 1959).

Ebert, P. W. 1971. The status and management of the
felids of Oregon. Pages 69-71 in S. E. Jorgensen

and L. D. Mech, eds., Proceedings of a sym-
posium on the native cats of North America. U.S.
Department of the Interior, Fish and Wildl.
Serv., Bur. Sport Fish and Wildl, Region 3.

Kebbe, C. E. 1961. Bounties. Oregon State Game
Comm. Bull. 16:3, 6-7.

Maser, C, B. R. Mate, J. F. Franklin, and C. T.
Dyrness. 1981. Natural history of Oregon Coast
mammals. U.S. Department of Agricutlure, For-
est Service Gen. Tech. Rept. PNW-1.33. Pacific
Northwest Forest and Range Expt. Sta., Portland,
Oregon. 496 pp.

Oregon State Game Commission. 1967. Cougar. Ore-
gon State Game Comm. Bull. 22.

Robinette, W. L., J. S. Gashwiler, and O. W. Morris.
1959. Food habits of the cougar in Utah and Ne-
vada. J. Wildl. Mgt. 23:261-272.

SiTTON, L. W. 1977. California mountain lion in-
vestigations with recommendations for manage-
ment. State of California Resour. Agency, Dept.
Fish and Game Wildl. Restoration Proj. W-51-R,
Big Game Invegations. 35 pp.

Toweill, D. E., and E. C. Meslow. 1977. Food habits
of cougars in Oregon. J. Wildl. Mgt. 41:576-578.

Young, S. P., and E. A. Goldman. 1964. The puma,
mysterious American cat. Dover Publ., Inc., New
York. 358 pp.

A NEW SPECIES OF PENSTEMON (SCROPHULARIACEAE)
FROM THE UINTA BASIN, UTAH

Elizabeth Neese' and Stanley L. Welsh'

Abstract.— Described as a new species is Penstemon flowersii Neese & Welsh from Utah. An illustration is pro-
vided. Tlie species is named in honor of the late Seville Flowers, professor of botany at the University of Utah.

The genus Penstemon is large and complex
within the state of Utah. The flowers of
plants of this genus are among the most beau-
tiful within the state. Several taxa have been
described as new from the intermountain re-
gion during recent years. It is not surprising
that another such novelty should occur in the
region. The species was discovered during in-
vestigations of the rare plants in the Uinta
Basin, a region noted for its narrowly dis-
tributed endemics.

Penstemon flowersii Neese & Welsh, sp. nov.

Species haec ab P. immanifesto N. Holm-
gren in staminodiorum barbis multo breviore,
et ab P. carnoso Pennell in floribus roseo non
violaceo-caeruleis, ab uterque foliis basalibus
nullis differt.

Perennial glabrous glaucous herbs, with
simple ascending stems arising from a
branching woody caudex, 8-25 (32) cm tall,
the basal rosette lacking; cauline leaves all
entire, fleshy-thickened, (1.5) 2-5.5 cm long,
(4) 10-25 mm broad, the lower shortly petio-
late, spatulate, the middle ones larger, sessile,
lanceolate or elliptic, obtuse, the upper re-
duced, broadly ovate, acute; thyrsus cylindric
(not secund), with 4-9 verticils, the cymes
many flowered; calyx glabrous, 5-6.5 mm
long, the lobes broadly lanceolate, acumi-
nate, the margin scarious, suffused with rose;
corolla 15-18 mm long, rose within, the
striae dark rose-pink, the limb ampliate,
10-12 mm in diameter; staminode equaling
the tube, not exserted, the apex shortly bar-
bellate (to 0.1 mm long); stamens included,
the anthers glabrous, dehiscent throughout
and in the connective, not explanate, the sacs

opposite, 1-1.2 mm long; capsules 7-10 mm
long.

Type: USA. Utah: Uintah Co., T3S RIE
S9-10, 5.6 km W of Randlett, 12 May 1980,
Neese & White 8609 (Hoiotype: BRY; Iso-
types: NY, US, RM, CAS, UTC, MINN).

Paratypes: Utah. Uintah Co., T3S RIE
SIO, 4.8 km W of Randlett, 12 May 1898,
Neese & White 8600 (BRY, UT, NY, MO); do
T3S RIW S3, 5.8 km S of U.S. Hwy 40, 14
km W of Randlett, 12 May 1980, Neese &
White 8606 (BRY, NY, CAS); do T3S RIE
SIO, 4.8 km W of Randlett, 16 May 1979, E.
Neese & B. Welsh 7212 (BRY, NY, GH, MO).
Duchesne Co., T3S R2W S21, 3.2 km WNW
of Myton, 16 May 1979, Neese & B. Welsh
7218 (BRY, NY); do T3S R2W S12, 4 km due
N of Myton, 15 May 1980, Neese & White
8662 (BRY, NY, UC).

The Flowers beardtongue grows in shad
scale communities on pale-colored clay
slopes and benches between 1,500 and 1,600
m, where old terraces of the Uinta Formation
in the Duchesne River drainage are mantled
with Pliocene or Pleistocene pedimen,tal
gravels. It is common on such habitat in an
area of about 8 X 25 km between Randlett
and Myton. The species is remarkable in its
uniformity in regard to both morphology and
habitat. The plants, with their dusty pink
flowers and pale gray-green foliage are in-
conspicuous against the gray clay on which
they grow. The near congener, P. immani-
festus, of central eastern Nevada and western
Utah possesses a more prominently bearded
staminode. Penstemon carnosus Pennell is
similar in diagnostic characteristics, but the
Flowers beardtongue is quite distinctive in its
smaller stature, tufted, usually numerous

'Life Science Museum and Department of Botany and Range Science, Brigham Young University, Prove, Utah 84602.

429

430

Great Basin Naturalist

Vol. 43, No. 3

Fig. 1. Penstemon floiversii Neese & Welsh: A, Habit. B, Flower. C, Anther-sacs. D, Staminode.

July 1983

Neese, Welsh: A New Penstemon

431

stems, absence of a basal rosette, and pink
(not lavender-blue) flowers. Penstemon car-
nosus is a species of the western Colorado
Plateau, from the San Rafael Swell and the
Henry Mountains westward to Aquarius
Plateau (Holmgren 1978).

The plant is named to honor the memory
of Dr. Seville Flowers, late professor of bot-
any at the University of Utah. Dr. Flowers
was a student of lichens, mosses, and higher
plants, and his imtimely passing has left a
void in the understanding of the plants of
Utah and the West.

Acknowledgments

We express thanks to Noel H. Holmgren
for his examination of the type materials, and
for his comments regarding relationships of
this taxon. Kaye Thome provided the illustra-
tions, and for this we are grateful.

Literature Cited

Holmgren, N. H. 1978. An overlooked new species of
Penstemon (Scrophulariaceae) from the Great Ba-
sin. Brittonia 30:334-339.

A COMPARATIVE STUDY OF COYOTE FOOD HABITS ON TWO UTAH DEER HERDS

Jordan C. Pederson' and R. Gary Tiickfield'

Abstract.— Coyote {C-anis latrans) scats from two southern Utah deer herd units were collected and analyzed to
establish diet selection. The category showing the most consistent frequency of occurrence was mule deer Odo-
coileus hemionus; lagomorphs were next. Formal statistical analysis revealed that the only significant difference in
coyote food habits between herd units was in the frequency of rabbits eaten. These data suggest that coyotes in this
region of southern Utah show a comparatively higher preference for mule deer but, at the same time, do not eat deer
in proportion to the frequency of their occurrence.

Documented reductions in deer popu-
lations in most southern Utah mule deer
(Odocoileus hemionus) herds have led to
speculation concerning the cause or causes
for these declines (Workman and Low 1976).
This paper investigates the hypothesis that
coyote {Canis latrans) predation may reflect
differential selection for deer. This was done
by assessing coyote food habits in two adja-
cent deer herd units in southern Utah's San
Juan Coimty. Areas studied included the Blue
Moimtain (31 A) and Elk Ridge (3 IB) herd
units. Since the deer population is known to

be larger within the Blue Mountain unit
(Jense 1981), an examination of coyote scats
from both areas could indicate whether deer
occur in coyote diets in relationship to herd
size. If this relationship was positive at a high
level of significance, it would lend some cre-
dence to the coyote predation hypothesis.

Study Area

The San Juan-Blue Mountain deer herd
unit (31 A) is, for the most part, that portion
of San Juan County east of the North and

Table 1. Relative frequency of occurrence of food items in coyote diets as determined from 460 scats collected
from September 1977 to December 1979.

(n)

Blue

Mountain

Perioc?

Vegetation

Rodent

Deer

Lagoniorph

Cattle

Bird

Carrion

(18)

11.1

44.4

48.9

0.0

5.6

0.0

(15)

26.7

20.0

46.7

33.3

0.0

(105)

23.8

3.8

61.0

15.2

1.0

6.7

(41)

4.9

0.0

14.6

87.8

0.0

2.4

0.0

(12)

16.7

91.7

0.0

(62)

21.0

16.1

58.1

33.9

1.6

.30.6

(37)

59.5

40.5

37.8

13.5

10.8

21.6

(12)

14.3

50.0

46.4

25.0

17.0

10.7

0.0

Totalis

(318)

mean'^

26.8

19.8

40.7

42.4

3.9

4.0

7.4

Standard

deviation*^

16.60

17.27

17.17

30.56

4.81

4.52

12.04

^Period

Dates

Sep-Dec 1977

Jaii-Jiin

1978

Jul- Sep

1978

Oct-Dec 1978

Jan-Mar 1979

Apr-Jun

1979

Jul-Sep

1979

Oct-Dec 1979

'Utah Wildlife Resources, 115 North Main Street, Springville, Utah 84663.

^Department of Statistics, Brigham Young University, Provo, Utah 84602. Present address: Department of Biology Indiana University, Bloomington, In-
diana 47405.

432

July 1983

Pederson, Tuckfield: Coyote Food Habits

433

South Cottonwood drainages. Its highest
point is Abajo Peak at 11,360 ft (3,463 m),
and it ranges to a low elevation at Bluff City
of 4,473 ft (1,363 m). The summer range area
of this unit is 153 mi^ (396 km^), and the area
of the winter range is 1,394 mi 2 (3,610 km 2).
Major vegetational types within this unit are
conifer, aspen, mountain brush, sagebrush,
pinyon-jimiper, and blackbrush (Coles and
Pederson 1968, 1969).

The San Juan-Elk Ridge deer herd unit
(3 IB) is that area of San Juan County west of
the North and South Cottonwood Wash
drainages. Horse Mountain, at 9,320 ft (2,840
m) elevation, is the highest point; and the
lowest is also at Bluff City, which divides
these two herd units. The area of the summer
range is 195 mi^ (505 km^), and that of the
winter range is 1,132 mi^ (2,932 km2). Major
vegetational complexes include conifer, as-
pen, mountain brush, sagebrush, pinyon-
juniper, and salt desert shrub (Coles and Pe-
dersen 1968, 1969).

During the period from 1976 to 1979, the
number of deer harvested per 1000 ha of
summer range was 2.9 for the Blue Mountain

unit and 1.10 for the Elk Ridge unit. The
number of deer harvested per hunter day (ef-
fort) for the same time period was 0.061 and
0.049, respectively.

Materials and Methods

Data on dietary selection were obtained
from analyses of coyote scats collected along
established roads. Scat analysis was chosen
over stomach content analysis because a
larger sample size could be collected during
specific time periods and at specified local-
ities without diminishing the predator popu-
lation (Knowlton 1964, Meinzer et al. 1975).
Scats were collected every three months dur-
ing a 27-month period from 1 September
1977 to 31 December 1979, with the excep-
tion of a 6-month lapse during period 2. Scats
were air dried for a minimum of 30 days and
then analyzed after thoroughly crumbling.
All remains were identified with the aid of a
binocular dissecting microscope, hair (Moore
et al. 1974), and feather keys, as well as a ref-
erence collection of skeletons and vegetation.

Table 1 continued.

Elk Ridge

(n)

Vegetation

Rodent

Deer

Lagomorph

Cattle

Bird

Carrion

(0)

—

(4)

0.0

50.0

25.0

50.0

0.0

(26)

19.2

15.4

57.7

19.2

3.8

0.0

3.8

(5)

0.0

80.0

20.0

0.0

(22)

9.1

0.0

18.2

81.8

4.5

0.0

(18)

16.7

72.2

33.3

0.0

5.6

(31)

77.4

58.1

25.8

9.7

6.5

0.0

3.2

(36)

8.3

9.3

27.8

86.1

5.6

2.8

0.0

(142)

18.7

14.1

47.4

39.3

10.1

1.2

26.92

20.71

24.13

31.33

17.80

2.20

2.36

''63.2 and 74.6 percent of all scats contained unidentifiable material from the Blue Mountain and Elk Ridge herd units, respectively.
'Computed as the average overtime periods

434

Great Basin Naturalist

Vol. 43, No. 3

Food habits are reported as relative frequen-
cy of occurrence.

Comparisons between the two coyote pop-
ulations were made using three statistical
procedures, viz., normal approximation to
two sample binomial data (Snedecor and
Cochran 1967), stepwise logistic regression
(Fienberg 1980), and stepwise discriminant
analysis (Morrison 1976). The statistical com-
puting programs PIF, PLR, and P7M, re-
spectively, were employed from the BMDP
series (Brown 1977).

In the first statistical procedure, each scat
was considered to represent a bernoulli trial
for each category of remains identified.
Hence the total number of scats from each
herd unit was treated as a binomial random
sample, of which a certain proportion con-
tained remains but the complement did not.

In the second procedure, we treated the lo-
cation (herd unit) category as a "response"
variable and all other dichotomous categories

Table 2. Summary of coyote dietary studies

of identified remains as "design" or explana-
tory variables. The logic of the response vari-
able was then regressed on the explanatory
variables.

In the final procedure, each scat was con-
sidered to be a multivariate observation, i.e.,
a vector of remains categories. Discriminant
analysis was then used to determine which
variables (categories) best discriminated be-
tween the two groups (herd units).

Results and Discussion

We collected and analyzed 460 coyote
scats: 318 from the Blue Mountain imit and
142 from the Elk Ridge unit. Equal search ef-
fort was not expended on both areas, and scat
numbers are not indicative of coyote num-
bers. The major food items found in the scats
from both areas were mule deer, birds, car-
rion, lagomorphs (black-tailed jackrabbit
[Lepus calif ornicus], mountain cottontail

Authority

Study area

Sample size

Source

Bond

1939

California

282

S and S

Murie

1940

Yellowstone National Park

5,086

Scats

Sperry

1941

8,339

Stomachs

Murie

1945

British Columbia

311

Scats

1945

Montana

286

Scats

1945

Montana

Scats

Fitch

1948

California

1,173

Scats

Ferrel at al.

1953

California

2,222

Scats

Fichter et al.

1955

Nebraska

747

Stomachs

1955

Nebraska

2,500

Scats

Korschgen

1957

Missouri

770

Stomachs

Korschgen

1957

Missouri

326

Scats

Ozoga** et al.

1966

Michigan

Scats

Gier

1968

Kansas

1,451

Stomachs

Clark

1972

Utah and Idaho

186

Stomachs

Hawthorne

1972

California

384

Scat

Mathwig

1973

Iowa

151

Stomachs

Richens et al.

1974

Maine

Stomachs

Gipson

1974

Arkansas

168

Stomachs

Meinzer et al.

1975

Texas

514

Scats

1975

Texas

Stomachs

Niebauer et al.

1975

Wisconsin

3,353

S and S

Nellis et al.

1976

Alberta, Canada

344

Stomachs

Johnson et al.

1977

Arizona

224

Scats

Ribic^

1978

Colorado

Scats

Neffetal.

1979

Arizona

65
102

Scats
Scats

Litvaitis et al.

1980

Oklahoma

361

Scats

Springer and Smith'^

1981

Wyoming

404

Scats

^Percent could not be determined from data presented.

"Winter study only.

'^Summer study only.

"Largely carrion; irmards. heads, and feet.

July 1983

Pederson, Tuckfield: Coyote Food Habits

435

[Sylvilagus nuttallii]), rodents (rock squirrel
[Spertnophilus variegatus], least chipmunk
[Eutomius 7Jiinimiis], Apache pocket mouse
[Perognathtis opoche], and deer mouse [Per-
omyscus riianiculatus]), and vegetation
(Table 1).

When results of our study are compared to
data collected in 23 previous studies of coy-
ote diets (Table 2) dating from 1939 through
1981, only two show deer occurring in the
diets with greater relative frequency (Ozoga
and Harger 1966, Hawthorne 1972). Coyote
diets from both our study areas also showed a
higher relative frequency of carrion than
most other studies reported (Table 2). How-
ever, since it was difficult to postively identi-
fy carrion during the winter months, this cat-
egory was not included in the statistical
analyses reported hereafter. The greatest
amoimt of fluctuation from one time period
to another occurred in the category of la-
gomorph remains. Mule deer were the diet-
Table 2 continued.

ary item showing the most consistent use
(highest relative frequency) across collection
periods occurring in four out of eight and
four out of seven collection periods for the
Blue Mountain and Elk Ridge herd units, re-
spectively. Lagomorphs were the second
most consistently used food item identified in
scats, occurring in two of eight and two of
seven collection periods, respectively. Analy-
sis suggests coyotes could be a factor in the
fluctuations of deer populations in these
southeastern Utah herd imits. These results
do not constitute evidence for a cause and ef-
fect relationship. Mule deer may be killed
and eaten by coyotes or they may be eaten as
carrion. Deer carrion could occur as a result
of winter stress, other predators, disease, par-
asites, or other factors, but the reason for
these mortality factors warrants further
investigation.

Table 3 contains the single category com-
parisons of binomial proportions between

Percentage of specimens in which item occurred

Lagomorphs

Rodents

Carrion

Livestock

Birds

Deer

Vegetation

38.8

62.5

4.2

8.8

2.5

26.1

16.9

4.0

59.9

0.0^

0.1

3.1

1.0

2.0

4.3.0

32.0

25.0

20.0

13.0

6.0

4.0

69.4

6.1

9.2

6.6

7.4

4.6

1.8

3L8

40.1

12.6

0.0

3.5

1.1

2.4

52.7

1.7

8.9

6.4

12.9

1.1

3.2

45.4

43.7

1.0

2.0

0.0

3.0

29.3

49.1

0.0'

23.2

18.1

18.5

0.0*

58.2

0.0»

0.0^

26.1

44.1

0.4

3.6

23.0

0.0»

30.5

33.7

7.6

16.0

55.3

36.3

8.6

13.8

22.0

2.9

7.9*

80.4

33.3

o.oa

2.8

14.7

0.0

23.5

17.0

69.8

83.0

2.1

.5.1

91.4

19.8

54.3

41.5

37.7

0.0*

24.8

0.0*

3.1

84.0

15.0

0.0»

10.3

2.1

2.0

1.6

5.7

74.2

0.0*

1.5

3.7

35.2

45.3

6L0

37.7

0.0*

31.1

21.2

0.0

64.2

19.6

0.0*

11.8

19.6

15.9

78.4

7.0

9.0

30.0

13.0

44.0

5.0

36.0

10.5

24.5

6.0

0.0

1.1

0.0

48.5

10.8

20.2

21.1

0.0

4.5

0.3

20.5

28.0

21.0

0.0*

0.0

12.1

26.9

36.2

3.0

22.0

0.0*

44.0^

11.0

0.0*

7.0

27.0

19.4

0.0*

0.0

18.6

0.0

4.5

24.0

45.0

0.0

30.0

13.0

78.0

26.2

43.0

0.0

15.4

1.5

12.3

32.8

0.0

38.2

0.0

6.9

34.3

2.0

39.5

11.0

53.0

0.0

6.0

19.0

20.0

32.0

63.0

45.0

0.0*

5.0

42.0

436

Great Basin Naturalist

Vol. 43, No. 3

herd units. Normally we would conclude a
significantly higher (p = 0.27) proportion of
coyote scats from the Elk Ridge herd unit
contained lagomorph remains than did that
of the Blue Mountain unit, but such a con-
clusion would be somewhat misleading. All
tests reported in Table 3 are not independent
of one another since the information in each
came from the same sample of coyote scats.
One generally acknowledged and conserva-
tive interpretation of these kinds of results
uses a Bonferroni procedure (Neter and Was-
serman 1974) in which the level of accept-
able Type I error is divided by the number of
simultaneous tests (six in this investigation).
Hence, the "appropriate" significance level
for the results in Table 3 (assuming P = .05)
is .008, in which case none of the test results
are significant. It is interesting to note that
the only other category approaching the
point of demonstrating even weak evidence
in favor of a difference in coyote diets be-
tween herd units was deer. The results of the

stepwise discriminant analysis indicated the
most important variable (category) to signifi-
cantly discriminate between groups was lago-
morphs (approx. F-statistic at 1st step =
4.941, p = .027). Similarly, the results of the
stepwise logistic regression analysis indicated
lagomorph remains were the only variable to
account for a significant (x^ = 4.859 at 1st
step, p = .028) amount of variability in the
logit (response) variable.

These results suggest coyotes do not in-
clude deer in their diets based on the poten-
tial frequency of occurrence of this food
item. However, we did not conduct any si-
multaneous census of deer numbers in either
of the areas where scats were collected. Fur-
ther investigation is warranted.

Acknowledgments

We thank Marlene and Robert Hasenya-
ger, A. Ray Johnson, John C. Kimball, and
Aurelia, Mary Ann, and Linda Pederson for

Table 3. Cell frequencies and .statistical test results when each category of coyote scat material is considered to
be a normal approximation to a two-sample binomial problem.

Lagomorph;

Deer

Present

Absent

Total

Present

Absent

Total

Deer herd

111

207

318

151

167

318

142

Total

176

284

460

207

253

460

Pi" = .349

P.'' =

.458

P,"

= .475

P,b =

.394

z = 2.216

Prob.

= .027

z =

1.603

Prob.

= .109

Birds

Cattle

Present

Absent

Total

Present

Absent

Total

Deer herd

308

318

306

318

140

142

134

142

Total

448

460

440

460

P," = .031

P,b =

.014

Pi"

= .038

P,b =

.056

z = 1.079

Prob.

= .280

z =

.904

Prob.

= .366

Rodents

Vegetation

Present

Absent

Total

Present

Absent

Total

Deer herd

266

318

241

318

114

142

106

142

Total

380

460

113

347

460

Pi" = .164

P,b =

.197

p,"

= .242

P,b =

.254

z = .880

Prob.

= .379

z =

.263

Prob.

= .793

1 = Blue Mountain

2 = Elk Ridge

"Proportion of scats collected from the Blue Mountain Range containing the indicated remains
"Proportion of scats collected from the Elk Ridge Mountain Range containing the indicated remains

July 1983

Pederson, Tuckfield: Coyote Food Habits

437

their help with this study. We also acknowl-
edge the help of Leonard Newlin, Albert W.
Heggen and Norman V. Hancock of the Utah
Division of Wildlife Resources. This study
was supported by the Utah Division of Wild-
life Resources and the Brigham Young Uni-
versity Department of Statistics. Critical re-
view of this manuscript was provided by
Dwight Bimnell, Jerran T. Flinders, K. T.
Harper, Clyde L. Pritchett, Alvin C. Ren-
cher, and Bnice L. Welsh.

Literature Cited

Bond, R. M. 1939. Coyote food habits on the Lava Beds
Monument. J. Wildl. Mgt. .3:180-198.

Brown, M. B. 1977. BMDP-77 Biomedical Computer
Programs, P-Series. Univ. of California Press,
Berkeley. 880 pp.

Clark, F. W. 1972. Influence of jackrabbit density on
coyote population change. J. Wildl. Mgt.
36:343-356.

Coles, F. H., and J. C. Pederson. 1968. Utah big game
range inventory. Utah Div. Fish Game Pubi. 68-
2. 120 pp.

1969. Utah big game range inventory. Utali Div.

Fish Game Publ., 69-2. 164 pp.

Ferrel, C. M., H. R. Leach, and D. Tillotson. 1953.
Food habits of the coyote in California. Califor-
nia Fish and Game .39:301-341.

Fighter, E., G. Schildman, and J. H. Sather. 1955.
Some feeding patterns of coyotes in Nebraska.
Ecol. Monogr. 25(l):l-.37.

Fienberg, S. E. 1980. The analysis of cross-classified
categorical data. MIT Press, Cambridge, Eng-
land. 198 pp.

Fitch, H. S. 1948. A study of coyote relationships on
cattle range. J. Wildl. Mgt. 12:7.3-78.

Gier, H. T. 1968. Coyotes in Kansas. Kansas State Univ.
Agr. Exp. Stn.Bull. 393. 118 pp.

GiPsoN, p. S. 1974. Food habits of coyotes in Arkansas. J.
Wildl. Mgt. 38:848-853.

Hawthorne, V. M. 1972. Coyote food habits in Sagehen
Creek Basin, northeastern California. California
Fish and Game 58:4-12.

Jense, G. K. 1981. Utah big game investigations and
management recommendations 1980-81. Utah
State Div. of Wildl. Publ. No. 81-4.

Johnson, M. K., and R. M. Hansen. 1977. Foods of coy-
otes in the lower Grand Canyon, Arizona. J. Ari-
zona Acad. Sci. 12:81-83.

Knowlton, F. F. 1964. Aspects of coyote predation in
south Texas with special reference to white-

tailed deer. Unpublished dissertation. Purdue
Univ. 189 pp.

Korsghgen, L. J. 1957. Food habits of the coyote in Mis-
souri. J. Wildl. Mgt. 21:424-435.

Litvaitis, J. A., and J. H. Shaw. 1980. Coyote move-
ments, habitat use and food habits in south-
western Oklahoma. J. Wildl. Mgt. 44:62-68.

Mathwig, H. J. 1973. Food and population character-
istics of Iowa coyotes. Iowa State J. Research
47:167-189.

Meinzer, W. p., O. N. Clegkert, and J. T. Flinders.
1975. Food niche of coyotes in the rolling plains
of Texas. J. Range Mgt. 28:22-27.

Moore, T. D., L. E. Spenge, and C. E. Dugnolle. 1974.
Identification of the dorsal guard hairs of some
mammals of Wyoming. Wyoming: Game and
Fish Dept. Bull. 14. 177 pp.

Morrison, D. F. 1976. Multivariate statistical methods.
2d ed. McGraw-Hill, New York.

Murie, a. 1940. Ecology of the coyote in Yellowstone,
Fauna Series No.' 4, 1940. U.S. Govt. Print. Off.,
Washington, D.C. 206 pp.

Murie, O. J. 1945. Notes on coyote food habits in Mon-
tana and British Columbia. J. Mammal. 26:33-40.

Neff, D. J., AND N. WooLSEY. 1979. Effects of predation
by coyotes on antelope fawn survival on Ander-
son Mesa. Spec. Rept. No. 8. Fed. Aid in Wildl.
Restoration Proj. W078-R. Phoenix, Arizona. Ari-
zona Game and Fish Dept.

Nellis, C. H., and L. B. Keith. 1976. Population dy-
namics of coyotes in central Alberta, 1964-68. J.
Wildl. Mgt. 40:389-399.

Neter, J., and W. M. Wasserman. 1974. Applied linear
statistical models. Richard D. Irwin, Inc., Home-
wood, Illinois.

Niebauer, T. J., and O. J. Rongstad. 1975. Coyote food
habits in northwestern Wisconsin. Proc. 1975
Predator Symp. Montana Forest and Conser.
Exp. Stn., University of Montana, Missoula.

OzoGA, J. J., AND E. M. Harger. 1966. Winter activities
and feeding habits of northern Michigan coyotes.
J. Wildl. Mgt. 30:809-818.

Ribic, C. a. 1978. Summer foods of coyotes at Rocky
Flats, Colorado. Southwestern Nat. 23:152-153.

RiCHENS, V. B., AND R. D. HuGiE. 1974. Distribution,
taxonomic status, and characteristics of coyotes
in Maine. J. Wildl. Mgt. 38:447-454.

Snedecor, G. W., and W. G. Cochran. 1967. Statistical
methods. Iowa State Univ. Press, Ames. 593 pp.

Sperry, C. C. 1941. Food habits of the coyote. U.S. Fish
Wildl. Ser. Res. Bull. 4. 70 pp.

Springer, J. T., and J. S. Smith. 1981. Summer food
habits of coyotes in central Wyoming. Great Ba-
sin Nat. 41:449-456.

Workman, G. W., and J. B. Low, eds. 1976. Mule deer
decline in the West: a symposium. Utah State
Univ., Logan. 134 pp.

A SMALL CARNIVORE SURVEY TECHNIQUE

Tim W. Clark' and Thomas M. Campbell III-

Abstract.— a track station survey method for small, nocturnal mammalian carnivores is described. The method
has been field tested under a wide varietv of conditions. Stations were visited by rodents, mustelids, and canids, as
well as insects and birds.

Small carnivores, because they often range
relatively widely, occur at low densities, and
tend to be solitary and nocturnal, are one of
the more difficult mammal groups to study.
Five basic categories of carnivore survey and
census methods were described by Linhart
and Knowlton (1975): (1) direct counts or
capture-recapture, (2) counts of sign (dens,
tracks, or droppings), (3) questionnaires and
boimty payments, (4) catch per imit of effort
(trap-nights), and (5) elicited responses to
man-made stimuli (frequency of visitations to
scent stations, howl responses to sirens).
These methods vary in their effectiveness by
species and habitat type.

We used a track recording method em-
ploying two varieties of lures (olfactory and
acoustic) to elicit small carnivore responses
between 1978 and 1982. Field-use deter-
mined: (1) efficacy, (2) ease and thrift of con-
struction and setup, and (3) durability and
amount of maintenance required. Elicited re-
sponses sought included: (1) deposition of
tracks and scats at stations and (2) drawing
animals to these stations to make them more
visible during spotlight surveys. Our track-
recording technique was a combination and
modification of methods used by Mayer
(1957) and Justice (1961) to determine small
mammal presence and by Linhart and
Knowlton (1975) to index coyote populations.
We also compared our track stations accord-
ing to the above objectives with Linhart and
Knowlton's (1975) station design. We were
particularly interested in small carnivores on
prairie dog colonies, and therefore placed
track stations of both types on white-tailed

{Cynomys leucurus), Gunnison's (C. gun-
nisoni), and black-tailed prairie dog (C. hi-
dovicianus) colonies, although they could be
placed anywhere.

Track Stations

Our track station consisted of track-
sensitive smoked kymograph paper covering
a base of 4 mm thick plywood (0.6 m^) (Fig.
1). The kymograph paper was smoked in the
field by burning kerosene-soaked cotton in-
side an aluminum can which had a long, 1 cm
wide slit cut in one side. Track impressions
were preserved by spraying with quick
drying shellac.

Scent and bait attractants were placed on a
stake in the center of each track station (Fig.
1). Scents consisted of a variety of com-
mercial mustelid and other lures (i.e., weasels
Mustela frenata, mink M. visoji, marten Mar-
ies americana, fisher M. pennanti, badger
Taxidea taxus, wolverine Gulo giilo, and
black bear Ursus americanus. Baits consisted
of an aged liver and sardine mixture.

The acoustic attractant, a modified elec-
tronic bird call producing a "chirping"
sovmd, was used in association with track sta-
tions. This call was developed by the U.S.
Fish and Wildlife Service (electronic sche-
matic available from the authors). The call
was capable of fimctioning for a couple of
weeks on a single 6-volt battery. The call was
placed inside a camouflaged paint can (one-
pint, 0.5 liters) with holes in the lid to emit
the sounds.

Location of track stations was determined
in the field to minimize wind damage and to

'Department of Biology, Idaho State University, Pocatello, Idaho 8.3201 and Biota Research and Con.sulting, Inc., Box 2705, Jackson, Wyoming 83001.
•Biota Research and Consulting, Inc., Box 2705, Jackson, Wyoming 83001.

438

July 1983

■<'*!fjc^''

Clark, Campbell: Survey Techni9ue

439

^Ca-'T*!*'--" ^"'Mii^f^^^^^

Fig. 1. Photograph of kymograph-smoked track station used by a prairie dog. Scent stake is coated with
hver/sardines.

maximize dispersal of odors and sounds. Each
station was checked early each morning for
nocturnal tracks and again in evenings for
diurnal tracks for several consecutive days.
Baits, scents, and kymograph paper were re-
placed and soil resifted as needed.

Linhart and Knowlton's (1975) track sta-
tions consisted of a 1-m diameter circle of
freshly sifted fine dirt 1 cm deep and scented
and baited with specially prepared tabs or
our lures.

Results

We accumulated 264 kymograph and 112
sifted dirt track stations days (1 track station
day was 1 track station in operation for 24
hrs) on 24 different prairie dog colonies. A
variety of animals left track impressions on
both kinds of stations; they were: long-tailed
weasels, skunks {Mephitis mephitis), kit foxes
{Viilpes velox), coyotes {Canis latrans), badg-
ers, mice {Peromyscus sp., Dipodomys sp.,
OnycJiomys sp.), ground squirrels (Sperrno-
phihis sp.), prairie dogs, unidentified passe-
rine birds, and numerous insects.

As a substitute for the rarest carnivore on
prairie dog towns, the black-footed ferret
{Mustela nigripes), we presented kymograph
track stations to Steppe ferrets (M. evers-
manni) in laboratory conditions and they
locomoted on them.

Discussion

The utility of the kymograph and sifted
dirt track stations depended on the target
species sampled, the quality and permanence
of visitation record sought, and field condi-
tions encountered. Kymograph stations re-
quired about 4 minutes each to prepare, in a
sheltered location, and about 5 minutes to set
out. This type of station was most effective in
low humidity and low to moderate winds—
where stations might be effective 3-5 days
before needing new smoked paper. Concerns
that the kerosene odor and unfamiliar sub-
strate texture might deter all wild species
was not substantiated.

The sifted dirt stations required less time
and equipment to establish and were easier

440

Great Basin Naturalist

Vol. 43, No. 3

to replace, but tracks were sometimes in-
definite and harder to positively identify than
with kymograph tracks. Photography was the
only method to permanently record these
tracks.

No scats or urinations were found within
the track station areas. We never observed a
carnivore at a track station during nearly
continuous nighttime surveys, even though
tracks indicated their visitations.

We think our track station method has
utility in surveying site-specific areas, espe-
cially if camera monitoring of animal visits as
described by Dodge and Synder (1960) and
Loveless et al. (1963) were added to our sys-
tem. Our technique, as suggested by Clark
(1977, 1978) and Clark and Campbell (1980),
may offer a valuable addition to traditional
survey methods for some rare species like the
black-footed ferret. Since its efficacy has
been demonstrated, needed now are follow-
up quantitative experimental studies on the
value of the kymograph track station.

Acknowledgments

We thank those organizations who sup-
ported our field studies during which our
track station method was tried— National
Audubon Society, National Geographic So-
ciety, Defenders of Wildlife, U.S. Bureau of
Reclamation, Cleveland Cliffs Iron Co., and
Mid-American Pipeline Co. Denise Casey

and John Weaver provided critical advice on
the manuscript. Wildlife Preservation Trust
International and the New York Zoological
Society Animal Research and Conservation
Center provided support for manuscript
preparation.

Literature Cited

Clark, T. W. 1977. A research proposal to develop sur-
vey techniques and search for the black-footed
ferret in Wyoming. Submitted to U.S. Fish and
Wildlife Service— Office of Endangered Species,
Washington, D.C., and Denver Regional Office.
23 pp.

1978. Current status of the black-footed ferret in

Wyoming. J. Wildl. Mgt. 42:128-1.34.

Clark, T. W., and T. M. Campbell. 1980. Suggested
guidelines for black-footed ferret surveys. Avail-
able from: Box 2705, Jackson, Wyoming 8.3001. 7
pp.

Dodge, W. E., and P. Snyder. 1960. An automatic cam-
era device for recording wildlife activity. J.
Wildl. Mgt. 24:341-344.

Justice, K. E. 1961. A new method for measiuing home
ranges of small mammals. J. Mammal.
42:462-470.

Llnhart, S. B., and F. F. Knowlton. 1975. Determin-
ing relative abundance of coyotes by scent station
lines. Wildl. Soc. Bull. 3:119-124.

Loveless, C. M., J. D. Coffelt, D. E. Medin, and L. E.
Yeager. 1963. A photoelectric-cell device for use
in wildlife research. .'Vmer. Instit. Biol. Sci. Bull.
13:55-57.

Mayer, W. V. 1957. A method for determining the ac-
tivity of burrowing mammals. J. Mammal.
.38.5.31.

EVALUATION OF DRABA OLIGOSPERMA, D. PECTINIPILA,
AND D. JUNIPERINA COMPLEX (CRUCIFERAE)

Robert W. Lichvar'

Abstract.— Since Dniha pectinipihi Rollins was described in 1953, it has been assigned to several different tax-
onomic categories. It has be n recognized at the species and variety level and has also been placed in synonymy un-
der D. oUgospemui Hook. Then Dorn (1978) described D. junipenna and contrasted it to D. pectinipila and D. oligos-
perma. To clarify the status of these three taxa, the evaluation included field and herbarium observations and
scanning electron microscope studies.

A proposal to give Draba pectinipila Rol-
lins protection under the Endangered Species
Act prompted extensive field studies on this
and two closely related taxa. Rollins (1953)
described D. pectinipila from the alpine habi-
tat of Clay Butte, Park County, Wyoming. It
is now known from two other alpine loca-
tions in British Columbia and Colorado.
Since this taxon was described, it has been as-
signed to several different taxonomic cate-
gories. Before further action could be taken
to protect the plants, the taxonomic status
had to be reevaluated.

Hitchcock (1964) gave Draba pectinipila
varietal status imder D. oligospenna Hook.
His only comment was that this variety was
the only fairly distinct variant for the species.
Mulligan (1972) placed it in synonomy under
D. oligospenna without making field observa-

tions. Then Dorn (1978) described plants
from the low elevation areas near the Utah-
Wyoming border, mentioned by Rollins
(1953) under D. pectinipila, as D. juniperina.
To clarify the taxonomy of this species com-
plex, D. pectinipila and D. juniperina are
compared to one another and to D.
oligospenna.

Methods

This analysis of Draba oligosperma, D. pec-
tinipila, and D. juniperina included field and
herbarium observations and scanning elec-
tron microscope (SEM) studies. Field obser-
vations included habitat, plant aspects, and
flowering dates. Herbarium specimens were
studied for shape, and scape characteristics.
The herbarium analysis was done at the

Table 1. Character differences between the three Draba species.

Character

D. oligosperma

D. pectinipila

D. juniperina

Habitat

Scape pubescence
Scape height
Mature fruit pedicel

length
Petal color
Silique tip
Silique base
Style length
Flowering dates
Basal leaf trichomes
Valve trichomes
Distribution

exposed rocky slopes and

ridges
glabrous
1-4 (9) cm

0.1-0.5 (1.0) cm

yellow

mostly rounded

rounded

0.1-1 mm

May-July

fine

simple (90% + )

Western U.S. and Canada

alpine slopes

pubescent

(4) 5-9 (11) cm

0.5-1.2 (1.4) cm
yellow
tapered
rounded
0.3-0.8 mm
July-August
medium

doubly pectinate (90% + '
British Columbia,
Wyoming, Colorado

pinyon-juniper woodlands

pubescent

(5) 7-15 cm

0.5-1.0 cm

yellow

tapered

0.6-1.5 mm

April-May

coarse

doubly pectinate (90% + )

Wyoming, Colorado, Utah

'Wyoming Natural Heritage Program, Nature Conservancy. 1603 Capitol .Avenue. Room 325, Cheyenne, Wyoming 82001.

441

442

Great Basin Naturalist

Vol. 43, No. 3

Fig. 1. Leaf trichome of Draha pectinipila X200 (Lichvar 2066).

Rocky Mountain Herbarium, Laramie,
Wyoming; the Gray Herbarium, Cambridge,
Massachusetts; and the University of Colo-
rado Museum, Boulder, Colorado. The fol-
lowing specimens were most pertinent to this
study: At RM: Dawson, s.n.; McCoun, s.n.;
Dorn, 897; Nelson, 1223; Lichvar, 2066;
Johnston, 1424, 1442A, 1434; at GH: Rollins
and Porter 51269; Williams, 476; at COLO:
Beaman and Erbisch, 1208, 1276; Johnston,
1273, 2314; Calder, Savile, and Ferguson,
13780.

Results and Discussion

Comparison of field observations, herba-
rium, and SEM analysis showed considerable
differences between taxa in this complex.
These taxa have overlapping ranges with
areas of sympatry. Of the three taxa, Draha
oligospemia has the widest range and is the
most variable in habitat specificity. Draha
oligospemia occurs from lower basin areas to

high alpine ridges on either sandstone or
limestone formations. Draha pectinipila is al-
ways found on limestone in an alpine habitat,
and D. jiiniperina is found on sandstone for-
mations in association with a pinyon-juniper
woodland at lower elevations. When D. oli-
gospemia and D. pectinipila are syinpatric,
D. pectinipila is readily identifiable in the
field by its taller, more erect stature and the
high percentage of pectinate hairs on the
fruit valves. Near the Flaming Gorge area of
Wyoming, Utah, and Colorado, D. juniperina
is not found in the same habitat as D. oligo-
spemia. Draha oligospemia occurs on sand-
stone or gravelly outwash ridges and out-
crops, but D. juniperina is almost always
found in association with pinyon-juniper
woodlands or adjacent sagebrush that overlies
a sandstone formation.

Certain characteristics separate Draha pec-
tinipila and D. juniperina from D. oligos-
pemia (Table 1). The correlation of charac-
ters that are similar between D. pectinipila

July 1983

Lichvar: Draba

443

Fig. 2. Leaf trichonie of Draba oligosperma X200 {Lichvar 1981).

and D. jiiniperina are the tapered fruit tips
with doubly pectinate hairs on the valves of
the fruits and the taller scapes that are pu-
bescent. Draba oligosperma has rounded fruit
tips with mostly simple hairs and shorter
scapes that are glabrous. To distinguish be-
tween the two taxa with a more limited dis-
tribution, D. pectinipila and D. jiiniperina,
habitat, style length, and leaf trichomes may
be used. Draba pectinipila is an alpine taxon
found on limestone and has tapered fruits
above witli styles 0.3-0.8 mm long, but D. jii-
niperina is found at lower elevations in the
basins on sandstone in a pinyon-juniper
woodland and has tapered fruits above and
below, and styles 0.6-1.5 mm long.

Rollins (1953) noted that the lower eleva-
tion taxon, Draba juniperina, had not only ta-

pered fruits above and below but also coarse
hairs on the basal leaves. Scanning electron
micrographs (Figs. 1, 2, 3) show that D. ju-
niperina (Fig. 3) has doubly pectinate leaf
hairs twice the diameter of D. pectinipila
(Fig. 1). Draba oligosperrna (Fig. 2) is inter-
mediate in leaf trichome diameter.

Recognition of each of these species can be
justified at the species level for three reasons.
First, specimens or plants of each of these
taxa can consistently be separated in either
the field or herbarium. Second, when any
combination of these species occurs sympa-
trically in the field they are easily and con-
sistently identifiable. Finally, based upon tri-
chome characters, those used to distinguish
these species are consistent with those used
throughout the genus of Draba.

Key to the Species

1. Scapes glabrous; 1-4 (9) cm long; fruits with 90 percent or more simple hairs ...

D. oligosperma

444

Great Basin Naturalist

Vol. 43, No. 3

Fig. 3. Leaf trichome of Draba jtiniperina X200 {Lichvar 2821).

— Scapes pubescent, 4-12 cm long; fruits with 90 percent or more doubly
pectinate hairs 2

2(1). Plants alpine with fine hairs on basal leaves; siliques tapered above, rounded

below; styles 0.3-0.8 mm long D. pectinipila

— Plants of pinyon-juniper woodlands with coarse hairs on basal leaves; siliques
tapered above and below; styles 0.6-1.5 mm long D. juniperina

Acknowledgments

Thanks are extended to Robert Dorn for
reviewing the manuscript, to Robert Bow-
men for doing the SEM work, and to the
curators of the collections mentioned above
for allowing me to study specimens in their
herbaria.

Literature Cited

Dorn, R. D. 1978. A new species of Draba (Cruciferae)
from Wyoming and Utali. Madrono 25:101-103.

Hitchcock, C. L., A. Cronquist, M. Ownby, and J. W.
Thompson. 1964. Vascnlar plants of the Pacific
Northwest. Part 2. Univ. of Washington Press,
Seattle. 496 pp.

Mulligan, G. A. 1972. Cytotaxonomic studies of Draba
species in Canada and Alaska: D. oligosperma
and D. incerta. Canadian J. Bot. 50(8): 1763-1766.

Rollins. R. C. 1953. Draba on Clay Butte, Wyoming.
Rhodora 55:229-235.

PRESENCE OF MAXILLARY CANINE TEETH IN MULE DEER IN UTAH

Jordan C. Pederson'

Abstract.— A hunter-harvested adult female mule deer {Ock>coiletis hemionus) was found to have canine teeth
present along both rows of teeth of the maxillae.

The reported occurrence of erupted canine
teeth in mule deer {Odocoileus hemionus) has
been observed in CaHfornia (Nordquist 1941),
Utah (Robinette 1958), and Colorado (Robin-
ette et al. 1977). The Utah incident occurred
in a three-year-old male having a single
erupted canine in the right maxilla (Robin-
ette 1958).

On 27 November 1982, a hunter-harvested
4-year-old female mule deer was checked and
collected by me. She was killed near In-
dianola in the Little Clear Creek drainage in
Utah County. The presence of upper max-

illary canine teeth on both sides was observed
when the jaw was cut for field aging (Fig. 1).
The head was removed from the carcass and,
after cleaning and photographing, donated to
the Brigham Young University Life Science
Museum, Provo, Utah, Catalog no. 7225.

The following skull measurements were
taken: total length 291 mm, condylobasal
length 272 mm, zygomatic breadth 128 mm.
Both maxillary canines are found posterior to
the nasomaxillary suture identical to the posi-
tion of canines found in elk {Cervus cana-
densis). Further measurements showed each

Fig. 1. Palatal view of a mule deer skull showing canine teeth on maxillae.

'Utah Division of Wildlife Resources, 1115 North Main, Springville, Utah 84663.

445

446

Great Basin Naturalist

Vol. 43, No. 3

canine is positioned 46 mm posterior of the
anterior point of the premaxilla and 43 mm
anterior of the premolar. Measurements of
the left and right canine, respectively, are:
length of outside (labial), exposed part, 9.2,
9.9 mm; anteroposterior diameter of crown,
4.9, 5.1 mm; labial lingual diameter of
crown, 3.8, 4.0 mm.

Acknowledgments

Thanks are extended to Dr. Jerran T. Flin-
ders, Dr. Kent M. Van De Graaff, Dennis L.

Shirley, and Laura John for their help in the
preparation of this manuscript.

Literature Cited

NoRDQuiST, G. 1941. Elk teeth in California deer. Cali-
fornia Fish and Game 27:39-40.

RoBiNETTE, W. L. 1958. Unusual dentition in mule deer.
J. Mammal. .39:1,56-157.

RoBiNETTE, W. L., N. V. Hancock, and D. A. Jones.
1977. The Oak Creek mule deer herd in Utah.
Publ. No. 77-15. Utah Wildlife Resources, Salt
Lake City. 148 pp.

COMPARATIVE SUCCESSIONAL ROLES OF TREMBLING ASPEN AND
LODGEPOLE PINE IN THE SOUTHERN ROCKY MOUNTAINS

Albert J. Parker' and Kathleen C. Parker'

Abstract.— A review of the ecological distribution and siiccessional roles of lodgepole pine and trembling aspen
in the Southern Rocky Mountains suggests that the two species have different strategies for occupying disturbed
sites. Lodgepole pine's easily dispersed seeds and faster growth from seed in unsuppressed conditions allows it to col-
onize severe bums, even from remote seed sources. Aspen appears to compensate for ineffective development from
seed by vegetative reproduction from durable root stocks, which promotes geographic persi.stence. Such persistence
is achieved by the maintenance of a forest structure conducive to light surface fires, which stimulate suckering and
retard conifer invasion, and by the accumulation of soil organic matter, which improves site nutrient retention and
water availabilitv.

Empirical studies of the dynamics of
trembling aspen {Popiihis tremiiloides) and
lodgepole pine (Pinus contorta ssp. latifolia)
forests show that both tree species commonly
colonize open sites following disturbance
(Clements 1910, Ives 1941, Stahelin 1943,
Marr 1961, Langenheim 1962). In portions of
the upper montane and subalpine zone
(2,400-3,000 m) of the Southern Rocky
Mountain region, the geographic and habitat
ranges of these two important colonizers
overlap, so that either species (or both) might
be encountered on a disturbed site. Within
this zone of cooccurrence, neither the site
preferences of nor the successional relation-
ship between these two species is satisfac-
torily detailed (Marr 1961, Peet 1981). Re-
garding habitat range, early workers thought
that aspen more frequently occurred on mes-
ic sites, and lodgepole more commonly occu-
pied drier settings (Bates 1924, Daubenmire
1943). More recently, Marr (1961) and Peet
(1978) have questioned the simplicity of this
arrangement. Peet (1978) asserts that both
species possess a comparable ecological opti-
mum on mesic sites in the lower/ middle sub-
alpine zone, as evidenced by the distribution
of aspen in mountainous regions where
lodgepole is absent. He observed that, in re-
gions where both species occur, lodgepole is
a better competitor than aspen on prime sites
and therefore tends to preempt aspen from
optimal settings. Aspen maintains popu-

lations in this region of cooccurrence by pos-
sessing a broader environmental tolerance
range, often being restricted to a variety of
both wetter and drier sites at higher and
lower elevations than lodgepole.

The successional relationship of the two
species in this region of cooccurrence is com-
plex (Moir 1969, Reed 1971, 1976, Whipple
and Dix 1979, Peet 1981). Differences in
their respective patterns of colonization are
likely related to a number of factors, chief
among which is the fundamental dissimilarity
in their reproductive strategies. Lodgepole is
a prolific seeder, depending on widespread
wind dispersal of its light seeds to facilitate
invasion of disturbed sites. Aspen, although it
is capable of reproduction by seed, more of-
ten reproduces by vegetative suckering. Marr
(1961) observed that aspen roots often sur-
vive fire, thus providing a stock for vegeta-
tive propagation on burned sites. Further-
more, both Marr (1961) and Peet (1981)
noted that aspen is often found in the under-
story of a variety of different forest covers,
including dense, mature conifer forests. Thus,
aspen is able to maintain a suppressed but vi-
able population on a site through long peri-
ods of time, and is capable of colonizing
burned sites by the release of the persi.stent
rock stock. Horton and Hopkins (1965), in an
examination of fire ecology in aspen groves,
found that light burns (i.e., low temperatures)
stimulate aspen suckering (probably through

'Department of Geography, University of Georgia, Athens, Georgia 30602.

447

448

Great Basin Naturalist

Vol. 43, No. 3

both the reduction of competition with the
thick ground layer and mobilization of the
nutrient supply tied up in the ground layer
vegetation), but heavy burns (i.e., high tem-
peratures) inhibit aspen suckering (presum-
ably through damage to perennating organs
in the root stock). Heavy burns are likely to
enhance the establishment of lodgepole pine
on disturbed sites, because they create a min-
eral seedbed and eliminate much of the
ground layer vegetation that might normally
inhibit development from seed of lodgepole
pine. Hence, postburn colonization of sites by
either aspen or lodgepole in their zone of
cooccurrence is influenced by their respec-
tive reproductive modes and ecological toler-
ances of environmental factors, by burn in-
tensity, and by a chance element (Marr 1961)
associated with the probability/proximity of
a lodgepole seed source or an aspen
rootstock.

The present study presents a review and
interpretation of both the habitat ranges and
successional relationship of lodgepole pine
and trembling aspen in the Colorado Front
Range. This discussion is accompanied by a
data set examining tlie age/size structure and
community characteristics of an abrupt
aspen /lodgepole ecotone on the south flank
of Bierstadt Moraine in Rocky Moimtain Na-
tional Park, Colorado.

Bierstadt Moraine is a lateral moraine of
approximately 200 m relief, deposited by al-
pine glaciers during the late Wisconsin gla-
cial maximum (Pinedale stage, Richmond
1960). Bierstadt Moraine trends slightly north
of east, extending for approximately 6 km
along the northern margin of the Glacier
Creek valley from Bear Lake to near Glacier
Basin Campgroimd. The dominant particle
size in the till is sand (exceeding 70 percent
of the total fine earth fraction); soils devel-
oped under both aspen and lodgepole forests
are immature (typic Cryorthents), although
there are distinct differences in the A-horizon
under each cover type. A transect down the
south-facing slope of Bierstadt Moraine from
top to bottom reveals the following sequence
of plant communities: lodgepole pine forest
on the gently rolling upland, sagebrush {Arte-
misia ssp.) scrub on the steep upper slope, as-
pen forest on the middle slope, and lodgepole

pine forest on the lower slope and through-
out the adjacent valley bottom. The ecotone
studied is between the aspen forest and the
lower lodgepole forest, at an elevation of
2,700 m.

The study area is located within a much
larger region (perhaps 10 km^) which was
burned by the Bear Lake fire of 1900 (Peet
1981). None of the trees cored on the study
site is older than this burn, so that the mod-
ern forest is representative of postburn recov-
ery dating three-quarters of a century from
this extensive fire.

Methods

Seven 4 X 60 m belt transects were placed
with their long-axis oriented normal to eleva-
tion contours and the aspen /lodgepole eco-
tone. Each transect was subdivided into six 4
X 10 m quadrats and placed so that three of
these quadrats were under aspen cover and
three were under lodgepole cover. Although
precise location of the "boundary" between
types is subjective, in this case abrupt differ
ences in both litter type and ground cover
were used to determine the midpoint of the
belt transect. Belt transects were spaced
along the flank of the moraine at intervals of
60 m. In each quadrat, all living trees (stem
DBH > 6.25 cm) were identified to species
and their diameter recorded. All saplings (0
< stem DBH < 6.25 cm) and seedlings
(stems less than breast height) in each quad-
rat were counted by species. All standing
dead stems in each quadrat were counted by
species. Along the central long axis of each
belt transect, the coverage of all herbs and
shrubs was determined in 10-m intervals by
the line-intercept method (Canfield 1941). In
two of the seven belt transects a radial core
was extracted from each tree (at 0.3 m
height) with an increment borer, and the
tree's age determined. Measurements of slope
aspect, steepness, and configuration were
taken for each 4 X 10 m quadrat. In two of
the belt transects, the type and depth of litter
was measured, using a point-frame, at the
center of each 4 X 10 m quadrat. The point-
frame was 1 m wide, with a 5 cm recording
interval; hence, there are 21 litter measure-
ments per quadrat. In addition, two soil pits
were dug, one under each cover type, and
the soil profiles were described.

July 1983

Parker, Parker: Succession

449

Table 1. Vegetation data by forest type.

Lodgepole cover
.odgepole Aspen

Aspen coyer

Characteristic L

Lodgepole

Aspen

Tree density
(stems-ha"l)

2420

430

100

2800

Stand basal area
dm2-ha-l)

2280

170

140

1910

Mean diameter
(cm'stem"^)

15.5

10.2

19.5

13.2

Sapling density

(sfha-i)

290

190

540

Seedling density
(sfha-l)

210

3870

Standing dead density
(st-ha-i) 250

920

2510

Results

The aspen forest on the south flank of
Bierstadt Moraine is on a slightly steeper
slope than the lodgepole forest (15° vs. 12°),
fingering to lower elevations along local con-
vexities of the slope face.

The principal difference in the soil profiles
under each cover type is the presence of a
dark brown (10 YR 3/2) surface mineral hori-
zon, approximately 10 cm thick, under the
aspen forest. This darker topsoil, which is ab-
sent under lodgepole, is indicative of higher
organic matter content associated with humi-
fication of the deciduous aspen leaves and lit-
terfall from the thick ground layer vegetation
(cf Hoff 1957, Tew 1968). Hydrometer analy-
sis (Bouyoucos 1962) of soil material col-
lected from varying depths in both profiles
revealed no meaningful differences in texture

with depth or between profiles, all samples
being coarse sandy loams. This texture, which
normally would permit only low water reten-
tion and rapid leaching of cations, imparts in-
creased significance to the organic buildup in
the A-horizon under aspen (Morgan 1969), in
that organic colloids increase the water re-
tention capacity and the cation (or nutrient)
exchange capacity of the aspen soil relative
to the lodgepole soil.

The ground layer of the lodgepole forest
was poorly developed, with 3 percent ground
coverage divided evenly between herbs and
woody shrubs. The aspen forest ground layer
was well developed (58 percent ground cov-
er) and almost exclusively herbaceous. Such
differences between aspen and conifer
ground layer development have been re-
ported elsewhere (Hoff 1957, Marr 1961,
Peet 1981) and apparently result from a com-
bination of the high acidity of conifer litter
(Daubenmire 1953) and increased light pene-
tration to the forest floor under aspen (par-
ticularly in the spring prior to aspen leaf
flush).

Under the lodgepole cover, tree density
was 2,850 stems-ha-^; lodgepole pine ac-
counted for 84.9 percent of this total (Table
1). Under the aspen cover, tree density was
2,900 stems'ha'; aspen accounted for 96.6
percent of this total. Similarly, under lodge-
pole cover, stand basal area .was 2,450
dm^-ha' (93.1 percent lodgepole); under as-
pen cover, stand basal area was 2,050dm^ha-i
(93.2 percent aspen).

Little evidence of invasion by more toler-
ant tree species over the last 75 years exists.
The only other tree species encountered were

Table 2. Cross transect patterns of mean tree basal area and understory representation by species.

Cover/Quadrat/Species

Mean tree

Basal area

(dm^'stem"^)

Number of saplings
and seedlings

Number of
standing dead

. Lodgepole
Aspen
Lodgepole _^ Lodgepole
cover Aspen

^Lodgepole

— Ecotone-

Aspen

■ Lodgepole
Aspen
Aspen -Lodgepole

cover Aspen

^Lodgepole
Aspen

1.74
0.47
1.68
0.37
2.34
0.89

10
9
5
10
10
15

9
12

2.52
1.21
1.47
1.45
4.91
1.47

3
121

2
125

4
124

0
92

0
60

1
59

450

Great Basin Naturalist

Vol. 43, No. 3

Lodgepole Forest
Cover

Lodgepole Pine

UJ
CD

Aspen

IV d cvj id f^' O CO lo K d ro

C£)CD<MCX)rO00';l-(T)ipOiO

i^dcviidf^dcJidcxid
•— --^ -— cJcvjcvJcvjro

9
>

Aspen Forest
Cover

Lodgepole Pine

72—1—^2 I r / J / /P^

1^" d c\J in h-' d oJ in h-^

' ^.— ■— ■— CJOJC\JCM

ro I I I I I I I I
i^'dcJidi^doJid

'"•—'"'" CJ CVJ CVJ

DIAMETER -CLASS (cm)

Fig. 1. Diameter-class distribution by forest cover. Histograms of stem number by diameter class for both lodge-
pole pine and aspen under each forest cover are depicted (diameter class interval = 2.54 cm). The number of stems
in the smallest diameter class (6.3-7.5 cm) has been doubled to adjust for its half-interval width.

willow {Salix spp.), with two saplings present
in a quadrat adjacent to the valley bottom,
and Douglas-fir {Psendotsiiga menziesii), with
a single seedling found under aspen.

Cross transect trends in the number and
size of aspen and lodgepole (Table 2) demon-
strate that lodgepole pine is infrequently en-
countered under aspen cover, but, where

July 1983

Parker, Parker: Succession

451

Lodgepole Pine

Aspen

< 15

^ 5

tiii

9
>

UJ
CO

w I I ' I I I I I I

^ OinOLOQi^omo

w I I I I I I I I I

^ oioOinoioOinO

AGE-CLASS (yr.)

Fig. 2. Age-class distribution of lodgepole pine and aspen in cored transects. Histograms of the stem number by
five-year age class intervals are shown for each species.

present, trees are generally large. Conversely,
aspen is commonly encountered beneath
lodgepole, although these trees are generally
small. This pattern is more emphatically ex-
pressed in the understory; aspen saplings,
seedlings, and standing dead stems (chiefly
aborted suckers) are much more common un-
der lodgepole cover than lodgepole is under
aspen cover. The high number of aspen seed-
lings and standing dead stems under lodge-
pole cover is indicative of a successional
strategy that relies on maintenance and grad-
ual spread of the aspen root stock into the
understory of adjacent conifer forests. The
trend in mean tree diameter and in the num-
ber of both understory and dead stems across
the transect clearly demonstrates the progres-
sive spread of aspen across the ecotone
(Table 2). The mean tree diameter of aspen
and the total number of understory and dead

aspen stems steadily decline away from the
aspen grove.

The collection of 72 lodgepole pine and 51
aspen tree cores was used to correlate age
with stem diameter. Both species exhibited a
comparable age-diameter relationship, the
coefficient of variation of age (lodgepole =
0.22, aspen = 0.25) being less than the
coefficient of variation of diameter (lodge-
pole = 0.38, aspen = 0.41) in each case.
Furthermore, age was significantly correlated
with diameter for each species (for aspen r =
0.654, p < 0.001; for lodgepole r = 0.665, p
< 0.001). Lodgepole pine displayed a some-
what stronger tendency toward even-
agedness than aspen, although both species
exhibited unbroken representation in age
ranges between 30 and 75 years. Fifty per-
cent of the lodgepole stems were in the 60-to
75-year age range, suggesting colonization

452

Great Basin Naturalist

Vol. 43, No. 3

(1)

Aspen Cover

Trees

Soil/Ground layer

Annual nutrient-
rich litter fall

Open canopy
branctiing pattern

Nonflammable
plant parts

Reproduction
by suckering

Pronounced biocycling
(good nutrient reserve)

Thick

herbaceous

layer

Low-temperature
surface fires

Fig. 3. Diagrammatic representation of the interactions among the dominant tree species, soil, ground layer vege-
tation, and fire characteristics under aspen and lodgepole pine forest cover.

immediately following the Bear Lake burn,
and the maximum concentration of aspen
stems (42 percent) occurred in the 45- to 60-
year range (Fig. 1). The unsuppressed diame-
ter growth rate of each species was deter-
mined using the 10 largest lodgepole and as-
pen trees cored. Our data reveal that
following successful establishment unsup-
pressed lodgepole pine grows more rapidly
than aspen (0.37 cm yr' vs. 0.32 cm yr') on
the study site. It should be cautioned that the
period of establishment is generally several
years longer for lodgepole pine produced
from seed, which may require from 3 to 20
years to reach 0.2 m height (Romme and
Knight 1981), than for aspen suckers, which
may reach 3 m or greater height in 6 to 8
years (Jones and Trujillo 1975).

The composite diameter class diagram
(Fig. 2), which is based on all seven belt
transects and presented by cover type, does
not show a tendency for concentration of
lodgepole pine in larger size-classes even
though many lodgepole trees are relatively

old, suggesting that older lodgepole stems
may persist as suppressed individuals follow-
ing postburn colonization for a lengthy peri-
od. Examination of these diameter-class dia-
grams reiterates that aspen is a fairly
common understory tree beneath lodgepole
forests, but only a few generally large-sized
lodgepole individuals are scattered through-
out the aspen canopy.

Discussion

The soil profile and age structure differ-
ences between aspen and lodgepole pine
stands suggest that each species, where domi-
nant, reinforces a distinct group of vegeta-
tion-soil-fire interactions (Fig. 3). Further-
more, the persistence of these cover types
appears to be more closely tied to stand his-
tory than to direct environmental gradients.

Under aspen cover, the deciduous, nutri-
ent-rich foliage of aspen (Daubenmire 1953)
and the dense herbaceous understory com-
bine to enhance nutrient cycling and humifi-

July 1983

Parker, Parker: Succession

453

(2)

Lodgepole Cover

Trees

Soil/Ground layer

Nutrient-poor

litter fall

Dense canopy
branching pattern

Flammable
plant parts

Reproduction by seeds

a) requires high
light regime and
mineral seedbed.

b) serotinous cones.

c) widely dispersed seeds

Slow biocycling
(low nutrient reserve)

sparse

ground cover,

shrub

dominated

High-temperature
crown fires

Fig. 3 continued.

cation, resulting in an increased cation ex-
change capacity and nutrient concentration
in surface mineral horizons (Hoff 1957). The
increased soil water retention conferred by
the humus accumulation acts in concert with
the ground cover (which buffers soil surface
temperature and decreases windspeed, thus
reducing evaporation) to increase the avail-
ability and effective use of soil moisture.
Hence, aspen maintains a broad habitat range
by direct enhancement of soil nutrient and
moisture status (Lutz and Chandler 1946). On
the contrary, the acidic lodgepole pine nee-
dles promote leaching, and the paucity of
ground cover under lodgepole pine limits
biocycling of nutrients; consequently, soils
under lodgepole pine are often impoverished
relative to adjacent aspen substrates (Hoff
1957). In addition, the mechanical resistance
of pine needles to decomposition results in a
decrease in himiification and the buildup of
pine needle litter.

Fire plays a prominent role in the perpetu-
ation of discrete populations of both aspen
and lodgepole pine (Fig. 3). Aspen domin-
ance is maintained on a site through stimu-

lation of vegetative propagation following
low-temperature surface fires, presumably
through reduction of apical dominance (Dan-
iel 1980). The buildup of surface fuels by the
thick herbaceous layer, the mesicness of the
ground layer, and the relative openness of the
aspen canopy favor light-burning surface
over crown fires (Horton and Hopkins 1965).
Because of its suckering habit, aspen can sus
tain and is capable of slowly expanding local
populations vegetatively into adjacent favor-
able sites. The ability of aspen to sucker in
relatively dense shade facilitates this spread.
Continued aspen dominance on a site re-
quires the perpetuation of a surface fire re-
gime that releases advance regeneration
(Marks 1974, Oliver 1981) and stimulates vig-
orous reestablishment of aspen suckers. In the
absence of fire, eventual ascendance of more
tolerant conifer species often does not pre-
clude the persistence of the aspen root stock
in a suppressed condition (Marr 1961). De-
pendence on the maintenance of a "parental"
aspen root stock is necessary to offset the
competitive superiority of lodgepole pine
(and other conifers) when both species are es-

454

Great Basin Naturalist

Vol. 43, No. 3

tablished from seed, and favors persistent re-
colonization by aspen following light surface
fires.

In lodgepole forests a denser canopy
branching network, greater leaf area index,
and a high resin production combine with a
paucity of imdergrowth to favor hot crown
fires. By creating high surface light levels and
mineralizing the litter layer, crown fires cre-
ate a favorable seedbed for lodgepole pine
establishment and trigger a wave of lodge-
pole pine regeneration that manifests itself in
a tendency toward even-agedness in Rocky
Mountain lodgepole forests. This contagious
postbum colonization pattern is facilitated
by lodgepole pine's lightweight, easily wind-
dispersed seeds and rapid growth rate follow-
ing seedling establishment on disturbed sites
(allowing colonization from a remote seed
source). Furthermore, colonization of severe-
ly burned sites by lodgepole pine is rein-
forced locally by partial core serotiny (Po-
wells 1965). In addition to favoring lodgepole
establishment, severe burns inhibit aspen
suckering, because most suckers develop
from roots which are within 5 cm of the soil
surface, and hence are easily killed in a hot
fire (Daniel 1980).

The persistence of both aspen and lodge-
pole pine populations on sites with little evi-
dence of successional alteration suggests that
both species can be expected to maintain do-
minance for extended periods, in accordance
with Egler's (1954) view of vegetation devel-
opment. Only with prolonged fire exclusion
are stands likely to be invaded and replaced
by more tolerant conifers. Moreover, changes
in dominance on a site appear to be related
to the character of initiating disturbances
(Henry and Swan 1974, Anderson and Holte
1981) and the differential reproductive habits
of each species. A low-temperature surface
fire regime favors the maintenance and grad-
ual spread of aspen dominance by aggressive
suckering. Stand-destroying crown fires open
sites to rapid colonization by lodgepole pine,
and repeated crown fires reinforce lodgepole
pine dominance.

In summary, both trembling aspen and
lodgepole pine are successful colonizer spe-
cies in the southern Rocky Mountain region,
although they accomplish colonization and

persistence in different ways. Aspen perpetu-
ates itself through time on suboptimal sites
by its suckering habit that rapidly recolonizes
lightly burned sites, by promotion of a favor-
able surface fire regime, and by enhancement
of site quality through improved soil water
and nutrient retention capacity. Lodgepole
pine, which develops more rapidly from seed
than aspen and has easily dispersed seeds, is
able to preempt aspen on optimal sites, and
exhibits a relatively even-aged, contagious
colonization pattern following stand-
destroying crown fires.

Literature Cited

Anderson, J. E., and K. E. Holte. 1981. Vegetation de-
velopment over 25 years vvithont grazing on sage-
brush-dominated rangeland in southeastern
Idaho. J. Range Mgt. 34:25-29.

B.\TEs, C. G. 1924. Forest types in the central Rocky
Mountains as affected bv climate and soil. USDA
Bull. 1233. 152 pp.

BouYoucos, G. J. 1962. Hydrometer method improved
for making particle size analysis of soils. Agron. J.
54:464-465.

Canfield, R. H. 1941. Application of the line intercept
method in sampling range vegetation. J. For.
39:388-.394.

Clements, F. E. 1910. The life history of lodgepole burn
forests. USDA Forest Service Bull. 79:7-56.

Daniel, T. W. 1980. The middle and southern Rocky
Mountain region. Pages 277-.340 in J. W. Barrett,
ed.. Regional silviculture of the United States.
Wiley-Interscience, New York.

Daubenmire, R. F. 1943. Vegetation zonation in the
Rocky Mountains. Bot. Rev. 9:325-393.

1953. Nutrient content of leaf litter of trees in

the northern Rocky Mountains. Ecology
34:786-793.

Egler, F. E. 1954. Vegetation science concepts I. Initial
floristic composition, a factor in old-field vegeta-
tion development. Vegetatio 14:412-417.

Fowells, H. a. 1965. Silvics of forest trees in the United
States. USDA Agr. Handbook 271. 762 pp.

Henry, J. D., and J. M. A. Swan. 1974. Reconstnict-
uring forest history from live and dead plant ma-
terial—an approach to the study of forest succes-
sion in southwest New Hampshire. Ecology
55:772-783.

HoFF, C. C. 1957. A comparison of soil, climate, and
biota of conifer and aspen communities in the
central Rocky Mountains. Amer. Midi. Nat.
58:115-140.

HoRTON, K. W., AND E. J. HoPKiNs. 1965. Influence of
fire on aspen suckering. Canadian Dept. Forestry
Publ. 1095:1-19.

Ives, R. L. 1941. Forest replacement rates in the Colo-
rado headwaters area. Bull. Torrev Bot. Club.
68:407-408.

Jones, J. R., and D. P. Trujillo. 1975. Development of
some young aspen stands in Arizona. USDA For-
est Service Research Paper RM-151. 11 pp.

July 1983

Parker, Parker: Succession

455

Langenheim, J. H. 1962. Vegetation and environmental
patterns in the Crested Butte area, Gunnison
County, Colorado. Ecol. Monogr. 32:249-285.

LuTZ, H. J., AND R. F. Chandler. 1946. Forest soils.
Wiley & Sons, New York.

Marks, P. L. 1974. The role of pin cherry (Pruniis pen-
stjlvanica L.) in the maintenance of stability in
northern hardwood ecosystems. Ecol. Monogr.
44:7.3-88.

Marr, J. W. 1961. Ecosystems of the east slope of the
Front Range in Colorado. Univ. Colo. Stud., Biol.
8. 134 pp.

MoiR, W. H. 1969. The lodgepole pine zone in Colo-
rado. Amer. Midi. Nat. 81:87-98.

Morgan, M. D. 1969. Ecologv of aspen in Gunnison
County, Colorado. Amer. Midi. Nat. 82:204-228.

Oliver, C. D. 1981. Forest development in North Amer-
ica following major disturbances. Forest Ecol.
Mgt. 3:153-168.

Peet, R. K. 1978. Latitudinal variation in southern
Rocky Mountain forests. J. Biogeogr. 5:275-289.

1981. Forest vegetation of the Colorado Front

Range: composition and dvnaniics. Vegetatio
45:3-75.

Reed, R. M. 1971. Aspen forests of the Wind River
Mountains, Wyoming. Amer. Midi. Nat.
86:327-343.

1976. Coniferous forest habitat types of the Wind

River Mountains, Wyoming. Amer. Midi. Nat.
95:159-173.

Richmond, G. M. 1960. Glaciation of the east slope of
Rocky Mountain National Park, Colorado. Geol.
Soc. Amer. Bull. 71:1.371-1382.

Rom.me, W. H., and D. H. Knight. 1981. Fire frequency
and subalpine forest succession along a topo-
graphic gradient in Wyoming. Ecology
62:319-326.

Stahelin, R. 1943. Factors influencing the natural re-
stocking of high altitude burns by coniferous
trees in the central Rocky Mountains. Ecology
24:19-30.

Tew, R. K. 1968. Properties of soil under aspen and
herb-shrub cover. USDA Forest Service Res. Note
INT-78. 4 pp.

Whipple, S. A., and R. L. Dix. 1979. Age structure and
successional dynamics of a Colorado subalpine
forest. Amer. Midi. Nat. 101:142-158.

DIFFERENTIAL UTILIZATION OF BAT BOXES BY HOUSE WRENS
{TROGLODYTES AEDON)

Hal L. Black'

Abstract.— Pine boxes nailed to trees in four habitat types in southeastern Utah as roosting sites for bats proved
ineffective for bats but were utilized bv house wrens in all but one habitat. Boxes were most often utilized in the as-
pen habitat. Hypotheses to account for differential utilization are presented.

In late Fall of 1977, 35 rough-sawn, pine-
wood, day-roosting boxes for bats, con-
structed after the design of Stebbings (Quar-
terly J. Deven Trust for Nature 6:114-119,
1974) were placed in each of four habitat
types on the Abajo Mountains of southeastern
Utah in the hope of attracting several mon-
tane species of vespertilionid bats. The aspen
forest habitat was within a watershed rela-
tively ungrazed by livestock and had a rich
understory of low-growing shrubs, forbs, and
grasses. The oak-pine habitat, which is
grazed annually by cattle, had, relative to the
aspen, a poor understory and more hetero-
geneous structure. The cottonwood habitat
was within a rather broad stream bed and
was not grazed by livestock. The spruce-fir
forest was also ungrazed and consisted of
large mature trees, which formed a fairly
closed canopy with essentially no understory
vegetation. Permanent water was available
within each habitat. Boxes were attached to
tree tnmks with nails at heights of 3-4 m and
arranged in a zig-zag fashion within each
habitat at 12-15 m intervals. Unlike tradi-
tional bird boxes, these had a slit entrance on

the ventral side that measured about 25 X
175 mm.

As roosting sites for insectivorous bats the
boxes were disappointing, but as nesting sites
for house wrens {Troglodytes aedon) the boxes
were immediately successful. Table 1 in-
dicates by habitat and year the number of
boxes from which young wrens were fledged.
Hypotheses to explain the differential utiliza-
tion of these boxes include: (1) differences in
productivity of the understory vegetation
and, therefore, insect biomass on which
wrens feed and (2) differences in the avail-
ability of natural cavities for nesting. The as-
pen habitat appears to have the most produc-
tive understory and the most homogeneous
forest structure. The importance of nest
boxes in the aspen habitat suggests that re-
moval of understory vegetation by grazing or
through competitive processes may have a
rather severe effect on the abundance and
distribution of foliage gleaning species like
the house wren. These data suggest that the
addition of nest boxes to habitats may selecti-
vely enhance population densities.

Table 1. Summary of patterns of utilization of bat-roosting boxes by house wrens. The open number under each
habitat indicates the number of boxes out of .35 that were used. The number in parenthesis represents percent of
total.

Habitat

Type

Year

Aspen
(El. 2700m)

Oak-Pine

(El. 2400m)

Cottonwood
(El. 2400m)

Spruce-Fir
(El. 3200m)

1977
1978

10 (29)
12 (34)

2(6)
2(6)

1(3)

0
0

'Department of Multicultural Education and Department of Zoology, Brigham Young University, Provo, L'tah 84602.

456

PLANT AND SOIL RELATIONSHIPS IN
TWO HYDROTHERMALLY ALTERED AREAS OF THE GREAT BASIN

N. M. Milton' and T. L. Purely'

Abstract.— In two areas of hydrothermally altered rocks in the Great Basin, the native vegetation differs in com-
position and areal cover from imaltered to altered sites on the same geologic formations. Analysis suggests that phys-
ical rather than chemical factors may be the cause of the vegetation differences, especially permeability of bedrock,
depth and texture of soils, and, possibly, amounts and types of clay minerals present. These characteristics influence
the ability of soils to absorb and retain water.

In the East Tintic Moimtains, Utali, the soils from argillized or mixed argillized and silicified parent materials
have more characteristics associated with dryness and support sparser vegetation and more species especially
adapted to dry conditions than do soils from unaltered or silicified parent materials.

In Battle Mountain, Nevada, unaltered areas have greater vegetation cover and have soil depth and texture that
are more favorable for plants than do altered areas. Soil pH is higher in altered areas than in unaltered areas.

Where geology is obscured by vegetation,
as in the humid regions of the world, vegeta-
tion can be used as a clue to the underlying
rocks and minerals. Changes in the vegeta-
tion along zones of mineralization have been
recognized since ancient times and have been
well docimiented in several recent reviews
(Malyuga 1964, Rommel 1968, Nesvetailova
1970, Cannon 1971, Brooks 1972). The rea-
sons for the vegetation differences should be
found in the physical and chemical proper-
ties of the soils that develop on hydro-
thermally altered, locally mineralized rocks
which differ from those of soils that develop
on unaltered rocks.

Two previous workers have studied the
vegetation in hydrothermally altered areas in
the Great Basin of the western United States.
Billings (1950) concluded from greenhouse
experiments that acid conditions and defi-
ciencies in nitrogen and phosphorus were re-
sponsible for the lack of sagebrush on altered
sites in the Virginia Range near Virginia
City, Nevada. Salisbury (1954, 1964) per-
formed similar experiments at Big Rock Can-
dy Mountain, Utah, and concluded that nutri-
ents in acid soils (pH 3.3) were chemically
bound by iron and aluminum and therefore
were unavailable to plants. These studies de-
scribe extreme states, in which the soil pH is
very low and the vegetation is drastically dif-
ferent from that on nearby unaltered rocks.

In many other hydrothermally altered areas,
altered soils have pH ranges comparable with
those of unaltered soils, and vegetation differ-
ences are more subtle.

The most striking* vegetation differences in
the study areas described in this report are on
sites of intermediate alteration intensity,
whereas the unaltered and the most intensely
altered sites have more similar vegetation.
Since intensely altered sites have been sub-
jected to more leaching than intermediate
sites, nutrient deficiencies and toxicities
would seem to be ruled out as likely causes of
the vegetation differences. The most common
limiting factor of plant growth in arid and
semiarid areas is water. Infiltration and water
retention are closely related to soil depth, soil
texture, i.e., the distribution of different-sized
particles in the soil, and the type of clay min-
erals in the soil (Black 1968, Foth and Turk
1972). For the present study, the hypothesis
of drier soil conditions in areas of inter-
mediate alteration intensity was tested by ex-
amining soil characteristics on altered and
imaltered sites and by comparing the vegeta-
tion distribution with that found by other
workers in similar terrain.

Study Areas

Two study areas in Utah and Nevada were
chosen to coincide with areas of ongoing re-

'Mailstop927, U.S. Geological Survey, Reston, Virginia 22092.

457

458

Great Basin Naturalist

Vol. 43, No. 3

search in the use of remote sensing tech-
niques to identify hydrothermally altered
rocks. The Utah study area has been used as a
test site for mapping hydrothermally altered
rocks from high altitude and satellite imagery
(Rowan and Abrams 1978a, b, Rowan and
Kahle 1982). Krohn et al. (1978) used Landsat
imagery to detect hydrothermally altered
rocks at the Nevada study area. Both areas
were found to be at the limit beyond which
limonitic hydrothermally altered rocks could
not be detected through the plant cover by
the remote sensing techniques used. For this
reason they were considered ideal sites for
devising a method to detect altered rocks by
using differences in vegetation.

Emplacement of intrusive bodies during
the Tertiary resulted in alteration of host
rocks to form several types of altered rocks.
Argillic and silicic alteration are the most
common types in the study areas, and are the
only types included in this study. In general,
the acidic hydrothermal fluids followed faults
and fractures in the rocks. The mineralizing
fluids changed in composition away from the
source, which resulted in a gradual decrease
in alteration along the path of the fluid. An-
other change that took place outward from
the path resulted in a gradation of alteration
away from the conduit. These changes pro-
duced a zonation of altered rocks. Silicified
rocks are closest to the conduit and the
source and are surrounded by argillized
rocks, beyond which is an area of gradual
transition to unaltered rock. Widths of zones
vary and can range from a few meters to sev-
eral hundred meters (Lovering 1949, 1960,
Lovering and Shepard 1959).

Degrees of alteration are reflected in the
different clay minerals produced. Moderate
alteration and supergene weathering under
mild conditions favored the formation of
montmorillonite. More intense alteration and
a more acidic weathering environment re-
sulted in the formation of kaolinite. In addi-
tion, supergene weathering resulted in the
conversion of ferrous iron to limonite, caus-
ing the weathering environment to be more
acidic in altered areas than in unaltered areas
(Lovering 1949).

The study includes unaltered, argillized,
and silicified sites. The argillized rocks are
bleached, limonite stained, and friable. They

are formed by cation leaching, addition of
water, and formation of clays. Kaolinite,
mixed-layer clays, and montmorillonite are
present. Kaolinite decreases away from the
conduit. Initial porosity is greater in the al-
tered rock than in the host rock. In the Utah
study area, however, gravity compacts the
argillized rocks, causing the pores to close,
and a nearly impermeable rock results (H. T.
Morris, oral comm. 1979). Silicified rocks are
limonite stained and very hard, composed
largely of silica, and containing some kaoli-
nite and mixed-layer clays. Porosity remains
higher than in the fresh and argillized rock
because of the rigidity of the matrix sur-
rounding the pores (H. T. Morris, oral
comm., 1979). The SiOa content of silicified
rocks can be as high as 90-95 percent, mak-
ing the original texture of the rock difficult
or impossible to determine.

East Tintic Mountains, Utah

The East Tintic Mountains are located in
Juab and Utah counties, west central Utah,
near the eastern edge of the Great Basin (Fig.
1). The area is classified as semiarid desert,
having an average annual precipitation of
about 30 cm. The range is made up of Paleo-
zoic sedimentary rocks partly overlain by
Tertiary volcanic rocks. The Tertiary rocks
include quartz latite and latite tuffs and
flows. Intrusive bodies associated with the
tuffs and flows are numerous, though not
large, and range from monzonite to quartz
monzonite. The Paleozoic rocks were exten-
sively folded and faulted prior to the Tertiary
volcanism (Morris 1957, 1964a, Morris and
Lovering 1961, 1979, Lovering 1960).

The study was confined to two Tertiary
units, the Packard Quartz Latite and the
overlying Laguna Springs Volcanic Group.
The Packard Quartz Latite consists of quartz
latite tuffs and flows that contain phenocrysts
of andesine, sanidine, quartz, and biotite in a
fine-grained to glassy groundmass (Morris
and Lovering 1961).

The Laguna Springs Volcanic Group con-
sists of latite tuffs and flows. The tuffaceous
member ranges from fine to coarse grained
and in some areas is agglomeratic. The flow
is a medium- to coarse-grained latite that
contains phenocrysts of orthoclase, plagio-

July 1983

Milton, Purdy: Plant-Soil Relationships

459

120^

115^

WYOMING

0 100 200 Kilometers
L I I ■ J '

Fig. 1. Location of study areas in the Great Basin.

clase, hornblende, biotite, augite, magnetite,
and quartz (Morris and Lovering 1961).

Battle Mountain, Nevada

Battle Mountain is located in Humboldt
and Lander counties, north central Nevada

(Fig. 1). The climate is arid, having about 15
cm average annual precipitation. The area is
made up of Cambrian to Tertiary sedimen-
tary and volcanic rocks. The pre-Tertiary
rocks have been intruded by early Tertiary
stocks, sills, and dikes. The geology of Battle
Mountain is further complicated by its posi-

460

Great Basin Naturalist

Vol. 43, No. 3

tion along the Roberts Mountain and Gol-
conda thrust zones.

Alteration at Battle Mountain took place
at various times throughout the Paleozoic
and Mesozoic, and most recently during the
Tertiary. Potassic, argillic, and silicic altera-
tion are present but are difficult to dis-
tinguish from each other in the field; for this
report, rocks are categorized only as unal-
tered or altered (Roberts and Arnold 1965,
Roberts et al. 1971, Theodore and Roberts
1971, Shawe and Stewart 1976, Silberman et
al. 1976).

Three formations were chosen for the
study. The Devonian Scott Canyon Forma-
tion is composed predominantly of chert, ar-
gillite, and greenstone with minor shale,
limestone, and orthoquartzite. As the clastic
content increases, the chert grades into argil-
lite. Recrystallization of Scott Canyon chert
by hydrothermal alteration resulted in an al-
tered chert difficult to distinguish in the field
from unaltered chert except for the presence
of small amounts of altered argillite (Theo-
dore and Blake 1975).

The Late Cambrian Harmony Formation is
composed of feldspathic sandstone inter-
bedded with shale and limestone. The sand-
stone is medium grained, subangular to sub-
rounded, and poorly sorted. The formation is
highly susceptible to alteration due to its tex-
ture and fine fracture patterns. Many of the
mineralized areas were also enriched by su-
pergene alteration (Theodore and Blake
1975, Suczek 1977).

The Pumpernickel Formation is Early
Pennsylvanian to Early Permian in age. It
consists of chert and argillite with minor
shale, greenstone, limestone, sandstone, and
conglomerate. Alteration resulted in recrys-
tallization to quartzose hornfels. In addition,
some silica was added and most oxides were
decreased (Roberts 1964, Theodore and Blake
1975).

Vegetation

The vegetation of the study areas is in-
cluded in the sagebrush and pinyon-juniper
zones described by Billings (1951), Blackburn
et al. (1968, 1969), Cronquist et al. (1972),
Tueller (1975), Young et al. (1976, 1977), and
MacMahon (1979). Artemisia tridentata (big

sagebrush), commonly with Chrysothamnus
nauseosus (rabbitbmsh), Purshia tridentata
(antelopebrush), grasses, and forbs, inhabits
wide valleys and lower slopes, and occurs
more sparsely within the pinyon-juniper
woodland. Juniperus osteosperma (Utah juni-
per) is the most prevalent tree, found in
stands having little undergrowth or scattered
among the shrub communities. Pinus mon-
ophylla (single-leaf pinyon pine) is a common
associate. Stream valleys and moist north-
facing slopes contain a dense growth of Acer
grandidentatum (bigtooth maple), Prunus vir-
giniana (choke cherry), Symphoricarpos oreo-
philus (snowberry), and Amelanchier tita-
hensis (shadbush). Many altered areas support
a sparse and low flora, often including Arte-
misia nova (black sagebrush), Petradoria pu-
mila (rock goldenrod), and other low matted
shrubs and herbs. In addition, several halo-
phytic species from the surrounding bajada
slopes are found in altered areas in Battle
Mountain.

Nomenclature for Utah follows Welsh and
Moore (1973); that for Nevada follows Munz
(1968).

Methods

Sites for sampling of vegetation and soils
were chosen from geologic and alteration
maps. To minimize variables, slopes chosen
ranged from 1,500 to 2,100 m in altitude,
were south facing (azimuth 135° to 225°),
and inclined between 12° and 20°. Side
slopes and spurs were chosen rather than
coves to minimize drainage-catchment differ-
ences, and slopes containing springs were
avoided. In the field, the sampling sites were
further limited to those near roads and rela-
tively undisturbed (e.g., not recently chained
or burned).

Vegetation Sampling

A floristic list for each site was made, in-
cluding forbs and grasses. Unfortunately, the
lists are not complete, because each study
area could not be visited during each part of
the growing season, and the vegetation data
presented here do not include forbs. How-
ever, shrubs and trees can be used to indicate
soil and water conditions, as well as lithologic
variations (e.g., Chikishev 1965).

July 1983

Milton, Purdy: Plant-Soil Relationships

461

Areal cover of vegetation was measured by
using a modification of the line-interception
method (Canfield 1941). Two 15 m tapes
were stretched at right angles, one along the
contour. The intercept of each species along
the tapes was measured to the nearest cen-
timeter, and the percent cover was calcu-
lated. Dead organic matter (standing dead
and litter) was measured as "mulch." For
each slope, fifteen 30-meter transects were
measured and averaged to give a representa-
tive vegetation sample for each site. Total
vegetation cover is the percentage of ground
covered by trees, shrubs, grasses, or mulch.

Soil Sampling

Soil samples were taken from the same
slopes on which vegetation was measured.
Four sites on each alteration type in each
geologic formation were sampled at depths
of 20 to 60 cm. Rocks larger than about 1.0
cm were removed from samples at the time
of collection. Because of the difficulty of an-
gering such dry, rocky soils, holes were dug
with a small shovel as deeply as possible. As a
result, measurements of pore space or bulk
density were not possible.

Silicification of flow and tuff units ob-
scures the original texture so that these units
are difficult to distinguish from one another
in the field; also they would be expected to
form identical soils because of their similar
initial chemical composition. For this reason,
the silicified Laguna Springs latite soil sam-
ples used in the summaries are the same for
flow and tuff examples.

Soil pH was measured by a glass electrode
pH meter on a 1:1 soil-water suspension. Ca-
tion exchange capacity (CEC) was measured
by the modified barium chloride-triethanola-
mine procedure (Chapman and Pratt 1961),
using a flame photometer.

Clay minerals were identified by X-ray dif-
fraction analysis. Oriented slides were run
imtreated, after heating to 350 C and 500 C,
and after treatment with ethylene glycol.
Minerals having (001) peaks at 7.1 A that dis-
appeared after hearing to 500 C were identi-
fied as kaolinite-group. Illite-group minerals
were identified as those having (001) peaks at
10.2 A. Minerals having (001) reflections that
expanded from 14 A on untreated samples to

15 A to 17 A on samples treated with eth-
ylene glycol were called mixed-layer illite-
montmorillonites, and the percentage of ex-
pandable layers was estimated from the rela-
tive peak intensities. In the absence of other
important peaks, relative proportions of the
three clay groups could be estimated from
the peak intensities (M. Hess, oral comm.,
1977).

Particle size analysis was done by the hy-
drometer method and a series of sieves, using
methods adapted from ASTM (1978) and
Lambe (1951). Organic matter was not re-
moved. Weighed samples were soaked over-
night in sodium metaphosphate and sonified
to aid dispersion. Soils were separated into
the following fractions: clay (<2 jn), silt
(2-50 ju), fine sand (50-250 ju) and coarse sand
(250-1,000/1).

Statistical Analysis

The Kruskal-Wallis test for central ten-
dency (Gibbons 1976) was used to test the
null hypothesis that there were no significant
differences in the areal cover of vegetation in
the East Tintic Mountains. Areal cover data
for Battle Mountain were analyzed by using
the Mann-Whitney-Wilcoxon test (Gibbons
1976). Different tests were used because of
the different numbers of independent varia-
bles in the two study areas. Vegetation and
soil differences were tested only within, and
not between, the two study areas. Non-
parametric tests were chosen in preference to
the corresponding parametric tests because
of small sample sizes and unknown
distributions.

A binary discriminant analysis (Strahler
1978a, b) was used to compare affinity of
plant species for rock formation and altera-
tion type, resulting in a list of plant species
for each type that best describes its differ-
ence from the other types. This is not a floris-
tic list nor a list of dominant species; only
those species strongly correlated with rock
and alteration type (p = 0.01) are listed. In
this test, each vegetation sample is entered
separately rather than averaged to give a
composite sample for each rock and altera-
tion type. Frequency, rather than areal cov-
er, is the variable used in this analysis.

462

Great Basin Naturalist

Vol. 43, No. 3

Results

East Tintic Mountains, Utah

The two most common species on the
Packard Quartz Latite areas, Artemisia tri-
dentata and Purshia tridentata, have higher
areal cover measurements on unaltered and
silicified sites than on argillized sites (Table
1). Chrysotliamnus nauseosus and Ephedra
viridis are found only in imaltered and silici-
fied areas. Grasses and mulch also have high-
er areal cover on unaltered and silicified sites
than on argillized sites. In contrast, the cover
of Juniperns osteosperma is higher on the ar-
gillized sites. This results in a high total vege-
tation measurement, even though there is ac-
tually substantial bare ground under and
between the trees in argillized areas. With
the exception of /. osteosperma, each species
has lower areal cover on argillized than on
unaltered and silicified sites.

The samples from the latite flow sites of
the Laguna Springs Volcanic Group also
show that samples from unaltered and silici-
fied areas have higher shrub and nontree sub-
totals than do the samples from argillized
sites (Table 1). Grasses and mulch have lower
areal cover in argillized areas. An important
difference is the presence of Artemisia nova
only on the argillized sites.

The argillized samples from the Laguna
Springs latite tuff (Table 1) have the lowest
shrub and nontree subtotals and the lowest
Artemisia tridentata, Purshia tridentata,
grasses, and mulch. Artemisia nova occurs
only on argillized sites. A greater variety of
herbaceous species was observed in argillized
areas than in unaltered and silicified areas on
both tuff and flow, although forbs, as stated
above, are not included in the vegetation
summaries.

The statistically significant differences in
shrub cover, nontree cover, and tree cover in

Table 1. Vegetation cover
Silicified.

(in percent) for the East Tintic Mountains study areas. U, Unaltered. A, Argillized. S,

Packard Quartz Latite

Laguna Springs Volcanic Group

Flows
U A

Tuffs
U A

Tuffs and flows

undifferentiated

Artemisia tridentata

8.23

.81

14.68

16.20

9.19

12.86

7.09

9.28

A. nova

1.42

1.71

Purshia tridentata

L61

.49

6.22

.21

.26

5.74

.63

2.64

Chrysothamntis nauseosus

.08

.11

.78

.98

.17

C. viscidiflorus
Ephedra viridis
Opuntia sp.
Gutierrezia sarothrae

L50

.08

.14
.08
.01

.17
.12

.07

.05
.03

.50

Tetradijynia sp.
Arenaria kingii

.02

.04

1.18

.02
.10

Petradoria pumila

.07

1.01

1.25

.05

Cercocarptis montanus
Subtotal

n.42

L45

.,31

22.58

16.70

11.72

19.58

11.98

12.76

Grasses
Mulch

Subtotal

1.96

8.18

10.14

.06
3.30
3.36

1.17

9.58

10.75

2.02
11.74
13.76

1.28
7.85
9.13

4.03
14.62
18.65

1.60
6.85
8.45

3.71
10.85
14.56

Nontree subtotal

21.56

4.81

,33.,33

30.46

20.85

38.23

20.43

27.32

Juniperus osteosperma
Pinus monophylla
Tree subtotal

6.90
2.47
9.37

29.34

.29

29.63

5.18
2.99
8.17

5.76
2.75
8.51

6.25
1.53

7.78

4.32
1.99
6.31

1.64
1.64

Total vegetation

30.93

,34.44

41.50

38.97

28.63

38.23

26.74

28.96

Standard deviation
Number of samples

±10.5
15

±15.9
15

±12.7
15

± 9.1
15

±13.5
15

± 8.4
15

±11.5
15

±11.6
30

July 1983

Milton, Purdy: Plant-Soil Relationships

463

Table 2. Significant differences in vegetation on different alteration types using the Kniskal-Wallis test on the
East Tintic Mountains data.

Shrubs

Total

Shrub

grasses

Tree

vegetation

cover

mulch

cover

Packard Quartz Latite
Unaltered vs. argillized
Unaltered vs. silicified
Argillized vs. silicified

Laguna Springs Volcanic Group: Flows
Unaltered vs. argillized
Unaltered vs. silicified
Argillized vs. silicified

Laguna Springs Volcanic Group: Tuffs
Unaltered vs. argillized
Unaltered vs. silicified
Argillized vs. silicified

"Significantly different at p < 0.15

samples from the Packard Quartz Latite sites
occur between unaltered and argillized sam-
ples and between silicified and argillized
samples (Table 2). That is, the argillized sam-
ples are significantly different from both
unaltered and silicified samples, whereas
unaltered and silicified samples differ from
each other only in total vegetation cover.

The Lagvina Springs flow data show signifi-
cant differences between unaltered and argil-
lized samples in total vegetation, shrub cov-
er, and nontree cover (Table 2). The
argillized and silicified samples are signifi-
cantly different only in the nontree
measurements.

In the Laguna Springs tuff results, the
unaltered and argillized samples differ signif-
icantly in total cover, shrub cover, and non-
tree cover, and the argillized and silicified
samples differ in nontree cover (Table 2). In

addition unaltered and silicified samples
differ significantly in all but total vegetation
cover.

Thus, the vegetation is similar in unaltered
and silicified areas but different in argillized
areas; on the Laguna Springs Volcanic
Group, silicified areas are more like argil-
lized than like unaltered areas.

The binary discriminant analysis results for
the Packard Quartz Latite data (Table 3)
show that unaltered areas are characterized
by Artemisia tridentata and Ephedra viridis,
argillized areas by Jtiniperus osteosperma,
and silicified areas by Purshia tridentata. The
correlation of A. nova with argillized flow
and tuff is notable in the Laguna Springs
data.

Soils in unaltered areas were dug to depths
of 25-35 cm. A thin organic layer alternates
with pebble- to cobble-sized angular float.

Table .3. Binary discriminant analysis results for East Tintic Mountains. Species listed are significantly correlated
with rock and alteration type at p = 0.01. U, Unaltered. A, Argillized. S, Silicified. + , d > 2.0. -, d < -2.0.

Packard Quartz Latite
U A 5

Laguna Springs Volcanic Group

A S

Artemisia tridentata
Ephedra viridis
Juniperus osteosperma
A. nova

Purshia tridentata
Chrysothamnus nauseosus
C. viscidiflorus
Gutierrezia sarothrae
Arenaria kingii
Petradoria pumila
Haplopappus acauUs

464

Great Basin Naturalist

Vol. 43, No. 3

The rock fragments increase in size as depth
increases. The soil is light brown (Munsell
color 7.5YR5/4) (Munsell Color Company
1969) to grayish brown (7.5YR4/2). Roots de-
crease in frequency as depth increases and
penetrate crevices in the rock beyond the
depth possible to dig by hand.

Soils in argillized areas are shallower
(about 15 cm) and range in color from mod-
erate brown (7 5YR4/4) to light yellowish or
rusty brown (10YR7/4). The organic layer is
absent or thinner than in unaltered areas.
Pebble- to boulder-sized bleached and limo-
nite coated rock fragments cover the top lay-
er of soil and are profuse at all levels. At the
deepest level dug, rocks are friable and roots
are few.

Soils in silicified areas are nearly as deep as
those in unaltered areas, and color is similar.
Bleached rock fragments occur at the surface.
Finer particles are found deeper than in soils
on unaltered areas, and roots are profuse in
lower parts of the soil profile.

Soil pH ranges from 6.6 to 7.7 (Table 4).
No significant differences were found be-
tween soils on different rock or alteration
type. Cation exchange capacity (CEC)
ranged from 17 to 43 Me/lOOg (Table 4). In
general, soils on Packard Quartz Latite areas
have higher CEC than soils on the Laguna
Springs Volcanic Group, except in silicified
areas. Using the Spearman coefficient of cor-
relation (Gibbons 1976), the CEC was found
to be weakly positively correlated with the
total clay content (r = 0.413). CEC is strong-
ly negatively correlated (r = -0.780) with

the amount of kaolinite in the soil and posi-
tively correlated (r = 0.630) with the
amount of montmorillonite. However, the
lack of significant differences between soils
of different alteration types seems to indicate
that, although the tests were done with rea-
sonable accuracy, the CEC and clay content
are too variable within soils of a single altera-
tion type to be diagnostic of it.

Physical analysis of the soil samples in-
cludes particle size analysis for texture and
X-ray diffraction analysis for identification of
clay minerals. In the Packard Quartz Latite
samples, clay content is about the same on all
three alteration types (Table 4). Silt content,
however, is higher and coarse sand content is
lower in silicified areas than in unaltered and
argillized areas.

In the samples from the latite flow in the
Laguna Springs Volcanic Group, the unal-
tered soils have the most silt and the least
coarse sand, argillized soils have the least silt
and the most coarse sand, and silicified soils
have intermediate amounts of each. In the
sites on the tuffs, the argillized and silicified
samples are similar, and the unaltered soils
have less silt and more coarse sand.

Because clay is formed in the alteration
process, a higher clay content would be ex-
pected in altered, particularly in argillized,
areas. The small clay differences recorded in
the samples suggest that dispersion was not
complete in all tests.

The predominant types of clay minerals in
the samples vary (Table 4). The standard de-

Table 4. pH, cation exchange capacity (CEC), particle size distribution and relative clay content for East Tintic
Mountains soils. U, Unaltered. A, Argillized. S, Silicified.

Packard Quartz

Latite

Laguna Springs Volcanic Group

6.8-7.7

6.9-7.6

6.7-7.6

6.7-

7.2

6.6-7.5

6.7-7.4

CEC (Me/lOOg)

36 ±5

37 ±6

25 ±8

±4
Flows

27 ±3

Tuffs

31 ±3

Particle size (percent)

30 ±12

29 ±3

Coarse sand

8 ±6

9 ±1

18 ±3

23 ±3

12 ±4

11 ±6

Fine sand

27 ±5

24 ±3

28 ±5

31 ±5

27 ±8

27 ±3

31 ±6

30 ±4

Silt

16 ±5

19 ±6

36 ±5

,35 ±3

27 ±3

23 ±7

31 ±3

30 ±6

Clay

27 ±5

28 ±3

28 ±2

25 ±3

28 ±7

27 ±2

26 ±4

29 ±5

Relative clay content

(percent)

Kaolinite

Ulite

Mixed-la ver

Montmorillonite

July 1983

Milton, Purdy: Plant-Soil Relationships

465

viations of the X-ray data averages are large,
so that these data can only be used to make
rough comparisons from one alteration type
to another. In the samples from the Packard
Quartz Latite, the unaltered samples have
the least kaolinite and the most mixed-layer
clay and montmorillonite. Soils of silicified
areas have the most kaolinite and the least
mixed-layer clay and montmorillonite. Soils
of argillized areas are intermediate between
the two but have the greatest amount of
illite.

The Laguna Springs latite samples show
fewer differences in clay type (Table 4). All
the soils contain large amounts of illite, less
kaolinite, and small amounts of
montmorillonite.

The soil characteristics of silicified areas
resemble those of argillized latite areas in the
Lagima Springs Volcanic Group. Following a
suggestion of R. P. Ashley (oral comm.,
1978), the silicified areas were examined and
found to contain large amounts of argillized
float around the silicified outcrops. This in-
dicates that the soils from silicified areas are
mixed with argillized material, resulting in
smaller differences in soils and vegetation on
the two alteration types than would be ex-
pected. The amount of argillized float on sili-
cified areas of the Packard Quartz Latite is
small, so that soils and vegetation differences
are large.

Battle Mountain, Nevada

The vegetation patterns in the Battle
Mountain study area show differences in
composition and areal cover from unaltered
to altered sites (Table 5). The altered areas
have lower total vegetation cover on all
three formations, and, except on the Harmo-
ny Formation, more variety of species is
found in altered areas. Shrub cover is lower
on altered sites of the Pumpernickel and
Scott Canyon formations than on unaltered
sites but remains nearly the same on the Har-
mony Formation; areal cover of grasses and
mulch is higher on imaltered than on altered
sites. Areal cover of Artemisia tridentata is
higher on unaltered than on altered sites on
all three formations and is absent on altered
Harmony Formation. Chrysothamnus nau-
seosus is more likely to be found on unaltered
sites, and C. viscidiflorus on altered sites. Ar-
temisia nova ra'ther than A. tridentata is
found on the altered Harmony Formation.

Shrub cover and total vegetation cover are
significantly different on unaltered and al-
tered areas of the Pumpernickel and Scott
Canyon formations (Table 6). On the Harmo-
ny Formation, the shrub cover is similar on
unaltered and altered sites, but the greater
amount of mulch on unaltered sites makes
the total vegetation cover significantly differ-
ent. Differences in vegetation cover between

Table 5. Vegetation cover (in percent) for the Battle Mountain study areas. U, Unaltered. A, Altered.

Pumpernickel
Formation

Scott Canyon
Formation

Harmony
Formation

Artemisia tridentata
A. nova

Purshia tridentata
Chrysothamnus nauseosus
C. viscidiflorus
Tetradymia sp.
Atriplex confertifolia
Ephedra nevadensis
PeucephyUum schottii

Subtotal

15.9

4.4
.4

3.5

1.9

1.8

1.6

trace

15.2

1.5

trace

11.1

.5
1.4

.4
0.9

19.1

18.8
1.0

20.7

9.1

17.1

14.6

19.9

Grasses
Mulch
Subtotal

2.6
10.9
13.5

4.5
3.8
8.3

2.0

9.1

11.1

1.7

9.1

10.8

2.2

1.1

10.0

2.6

12.2

3.7

Total Vegetation

34.2

17.4

28.2

25.4

32.1

23.6

Standard deviation

±7.8

±4.4

±6.9

± 4.3

±4.6

±6.7

Number of samples

466

Great Basin Naturalist

Vol. 43, No. 3

Table 6. Significant differences in vegetation on dif-
ferent alteration types using the Mann-Whitney-
Wilcoxon test on the Battle Mountain data.

Total

Shrub

vegetation

cover

Pumpernickel Formation

Unaltered vs. altered

•

Scott Canyon Formation

Unaltered vs. altered

•

Harmony Formation

Unaltered vs. altered

'Significantly different at p < 0.15

unaltered and altered sites are greater than
those on different rock types.

Binary discriminant analysis results (Table
7) show that unaltered Pumpernickel Forma-
tion sites are characterized by Artemisia tri-
dentata and Purshia tridentata, altered sites
by Atriplex confertifolia, Chrysothamnus vis-
cidiflorus, and Tetradymia sp. (Table 3). The
altered Scott Canyon Formation sites contain
Peucephylhini schottii, Chrysothamnus vis-
cidiflorus, and Ephedra nevadensis, whereas
only Artemisia tridentata is significantly cor-
related with the unaltered sites. Arteynisia tri-
dentata is characteristic of unaltered Harmo-
ny Formation and A. nova, of altered sites.

The pH of unaltered Battle Mountain soils
was lower than that of altered soils in all
three formations (Table 8). Soil depths were
greatest (35 cm) in unaltered Pumpernickel
chert and Scott Canyon chert soils. Depths of
altered soils and unaltered Harmony sand-
stone soil averaged 20-25 cm.

The particle size distribution analysis re-
sults show similar patterns for all three for-
mations (Table 8). Soils of unaltered areas are
coarser than those of altered areas. The silt
difference is greatest in the Harmony Forma-
tion, and the coarse sand difference is great-

est in the Scott Canyon Formation. As in the
Utah study area, the absence of the expected
higher clay content in altered areas suggests
that the clays may not have dispersed
completely.

The soils in altered areas of all three for-
mations have more kaolinite and less illite
than those of unaltered areas (Table 8). In ad-
dition, the unaltered Scott Canyon soils con-
tain montmorillonite. Again, differences are
not large, particularly in the Pumpernickel
soils, and the standard deviations are high.
Consequently, only general comparisons can
be made between soils of unaltered and al-
tered areas.

The altered sites on all three formations
are lower in altitude than the corresponding
unaltered sites. This factor, plus the presence
of Atriplex confertifolia and the higher pH on
the altered sites, suggests that the vegetation
may be influenced by increased salinity as
well as by a factor in the alteration process.

Discussion

In the East Tintic Mountains study area,
total vegetation cover is lower in argillized
areas than in unaltered and silicified areas,
except on the argillized Packard Quartz La-
tite, where large numbers of Junipenis os-
teosperma are found. All other species have
lower areal cover on the argillized Packard
areas, and the ground is relatively bare under
and around the trees. In addition, the compo-
sition of the plant communities varies with
rock type and alteration history. On unal-
tered and silicified areas are large amounts of
mulch, Artemisia tridentata, Purshia triden-
tata, and Ephedra viridis. Argillized areas, in
contrast, contain mostly Juniperus os-
teosperma and a very few shrubs.

Table 7. Binary discriminant analysis results for Battle Mountain vegetation data. Species listed are significantly
correlated with rock and alteration type at p = 0.01. U, Unaltered. A, Altered. +, d > 2.0. -, d < -2.0.

Pumpernickel

Scott

Canyon

Harmony
U

Artemisia tridentata

-1-

A. nova

Purshia tridentata

Ch njsotham mis natiseosus

-1-

C. viscidiflorus

-t-

-1-

Atriplex confertifolia

Tetradymia sp.

Peucephyllum schottii

-1-

Ephedra nevadensis

-t-

July 1983

Milton, Purdy: Plant-Soil Relationships

467

Table 8. pH, particle size distribution and relative clay content for Battle Mountain soils. U, Unaltered. A,
Altered.

Pumpemi(

ckel Formation
A

Scott Canyon

Formation

Harmony

Formation

pH 6.8-7.3

7.6-7.9

6.8-7.3

7.2-7.9

6.7-7.3

6.9-7.8

Particle size (percent)

Coarse sand 7 ± 1

4 ±2

15 ±2

5 ±3

11 ±3

8 ±2

Fine sand 17 ±2

18 ±1

24 ±2

24 ±4

23 ±4

sat 51 ±2

54 ±3

34 ±2

38 ±6

38 ±5

49 ±3

Clay 25 ±3

24 ±3

27 ±6

31 ±1

28 ±6

20 ±6

Relative clay content (percent)
Kaolinite 2.3

Illite 77

Montmorillonite

On the latite of the Laguna Springs Vol-
canic Group, vegetation differences are large
between unaltered and argillized sites, but
small between silicified sites and argillized
sites. The vegetation on unaltered sites is
composed largely of Arteinisia tridentata and
Purshia tridentata, whereas the argillized
areas contain A. nova and a wide variety of
minor shrubs.

In areal cover and in composition of vege-
tation the silicified areas of the Laguna
Springs Volcanic Group fall between argil-
lized areas and unaltered areas. The areas la-
beled 'silicified' contain a mixture of silicified
and argillized float, which accounts for the
similarity of soils and vegetation on argillized
and silicified sites.

In the Battle Mountain study area, shrub
cover and total vegetation cover are greater
in unaltered areas than in altered areas; Arte-
misia tridentata is the most common shrub in
all areas except on altered Harmony Forma-
tion, where it is replaced by A. nova. Atriplex
conferiifolia occurs on altered sites.

Several soil parameters were measured in
an attempt to explain the causes of the vege-
tation patterns. The hypothesis of low nutri-
ent levels to explain the low vegetation cover
in some altered areas was discarded because
of high vegetation cover measurements in the
most highly altered, leached areas. Physical
and chemical analyses of the soils include
measurements of pH, cation exchange capac-
ity, particle size distribution, and X-ray dif-
fraction for identification of clay minerals.
Although not quantitatively significant, some
relationships seem to exist between areal cov-
er of vegetation and soil characteristics. The

hypothesis that water is the most important
limiting factor in plant growth is supported
by the following reasons and comparisons:

1. Junipertis osteospemia and Ari:emisia
nova, which are found on argillized Packard
Quartz Latite and on the argillized Laguna
Springs Volcanic Group and altered Har-
mony Formation, respectively, are known to
occur in the drier habitats in the Great Basin
(Blackburn et al. 1968, 1969, Cronquist et al.
1972, Zamora and Tueller 1973, Vasek and
Thorne 1977).

2. The high percentage of bare ground,
around and under the trees on the argillized
Packard Quartz Latite and between the low
shrubs on the argillized Laguna Springs Vol-
canic Group and altered Scott Canyon and
Pumpernickel Formations, results in high
runoff and low infiltration of rainfall.

3. Although soil was not dug to bedrock
due to the difficulty of digging, the soil ap-
peared to be shallower in argillized areas and
mixed argillized and silicified areas than in
unaltered and silicified areas in the East
Tintic Mountains, and shallower in the al-
tered Battle Mountain areas than in the unal-
tered areas, except on the Harmony
Formation.

4. Fractured unaltered and silicified bed-
rock allows greater infiltration of rainfall
than does the highly compacted argillized
bedrock in the East Tintic Mountains.

5. The argillized areas are low in mixed-
layer clays and montmorillonite, which
would retain more moisture. The presence of
montmorillonite on the unaltered Scott Can-
yon Formation, Packard Quartz Latite, and
Laguna Springs Volcanic Group may in-

468

Great Basin Naturalist

Vol. 43, No. 3

crease water capacity. However, total clay
content is probably more important than
type of clay in areas that have a mixture of
clay types.

6. The other major difference between
unaltered and altered soils in Battle Moun-
tain, the higher pH of altered soils, can ac-
coimt for the presence of salt-loving plants
such as Atriplex confertifolia but does not ex-
plain the decrease in vegetation cover on al-
tered sites, because nearby areas that contain
halophytic communities have quite dense
vegetation.

Conclusions

Vegetation patterns of areal cover and dis-
tribution of species are related to the distri-
bution of hydrothermally altered and unal-
tered rocks in two areas within the Great
Basin. Several factors, including bare ground,
shallow soil, impermeable rock, soil texture,
and, possibly, clay composition in some areas
appear related to low vegetation cover in ar-
gillized areas, and suggest that water may be
limiting in these areas. The results are con-
sistent with those of other workers in the
Great Basin. This type of information is
needed for the development of techniques for
using vegetation as an aid to prospecting in
vegetated regions.

Acknowledgments

The authors are grateful to Drs. J. T. Hack
and F. A. Branson of the U.S. Geological Sur-
vey for critical reviews of the manuscript.

Literature Cited

American Society for Testing and Materials. 1978.
Standard method for particle-size analysis of
soils. Pages 71-81 in ASTM Standards, Part 19.

Billings, W. D. 1950. Vegetation and plant growth as
affected by chemically altered rocks in the west-
em Great Basin. Ecology 31:62-74.

1951. Vegetational zonation in the Great Basin of

western North America. Pages 101-122 in Les
bases ecologiques de la regeneration de la vegeta-
tion des zones arides. International Union of Biol.
Sci., Series B, No. 9.

Black, C. A. 1968. Soil-plant relationships. Wiley &
Sons, New York. 792 pp.

Blackburn, W. H. 1973. Infiltration rate and sediment
production of selected plant communities and
soils in five rangelands in Nevada. College of Ag-
riculture, Univ. of Nevada, Reno, Rept. No. R-92.
99 pp.

Blackburn, W. H., R. E. Eckert, Jr., and P. T.
Tueller. 1969. Vegetation and soils of the Cow
Creek Watershed. College of Agriculture, Univ.
of Nevada, Reno, Rept. No. R-49. 77 pp.

Blackburn, W. H., P. T. Tueller, and R. E. Eckert,
Jr. 1968. Vegetation and soils of the Duckwater
watershed. College of Agriculture, Univ. of Ne-
vada, Reno, Rept. No. R-40. 81 pp.

Brooks, R. R. 1972. Geobotany and biogeochemistry in
mineral exploration. Harper & Row, New York.
290 pp.

Canfield, R. H. 1941. Application of the line inter-
ception method in sampling range vegetation. J.
of Forestry 39:388-394.

Cannon, H. L. 1971. The use of plant indicators in
ground water surveys, geologic mapping, and
mineral prospecting. Taxon 20:227-256.

Chapman, H. D., and P. F. Pratt. 1961. Methods of
analysis for soils, plants, and waters. Univ. of Cal-
ifornia, Div. of Agr. Sci., Santa Barbara, Califor-
nia. 309 pp.

Chikishev, a. G. 1965. Plant indicators of soils, rocks
and subsurface waters. New York Consultants
Bureau, trans.. New York. 210 pp.

Cronquist, a., a. H. Holmgren, N. H. Holmgren, and
J. L. Reveal. 1972. Intermountain flora. Vol. I.
Hafner, New York. 270 pp.

FoTH, H. D., and L. M. Turk. 1972. Fundamentals of
soil science. Wiley & Sons, New York. 454 pp.

Gibbo.ns, J. D. 1976. Nonparametric methods for quan-
titative analysis. Holt, Rinehart & Winston, New
York. 463 pp.

Krohn, M. D., M. J. Abrams, and L. C. Rowan. 1978.
Discrimination of hydrothermally altered rocks
along the Battle Mountain-Eureka, Nevada, Min-
eral Belt using Landsat images. U.S. Geol. Surv.
Open-File Rept. 78-585. 66 pp.

Lambe, T. W. 1951. Soil testing for engineers. Wiley &
Sons, New York. 165 pp.

Lovering, T. S. 1949. Rock alteration as a guide to ore-
East Tintic district, Utali. Econ. Geol., Monogr.
1. 65 pp.

1960. Geologic and alteration maps of the East

Tintic district, Utah. U.S. Geol. Surv. Mineral In-
vestigation Field Studies Map MF-230. Scale
1:9600.

Lovering, T. S., and A. O. Shepard. 1959. Hydro-
thermal argillic alteration on the Helen claim.
East Tintic district, Utah. Pages 193-220 in A.
Swineford, ed., Clays and clay minerals. Vol. 8,
8th National Conference on Clays and Clay Min-
erals, Proc.

MacMahon, J. A. 1979. North American deserts: their
floral and faunal components. Pages 21-82 in D.
W. Goodall and R. A. Perry, eds.. Arid-land eco-
systems: structure, hmctioning and management.
Cambridge Univ. Press, New York. 881 pp.

Malyuga, D. p. 1964. Biogeochemical methods of pros-
pecting. New York Consultants Bureau, trans..
New York. 205 p.

Morris, H. T. 1957. General geology of the East Tintic
Moimtains, Utah. Pages 1-56 in Geology of the
East Tintic Mountains and Ore Deposits of the
Tintic Mining Districts. Utah Geological Society
Guidebook to the Geology of Utah, No. 12.

July 1983

Milton, Purdy: Plant-Soil Relationships

469

1964a. Geology of the Eureka quadrangle, Utah

and Juab counties, Utah. U.S. Geol. Surv. Bull.
1142-K. 29 pp.

1964b. Geology of the Tintic Junction quad-
rangle, Tooele, Juab and Utah Counties, Utah.
U.S. Geol. Surv. Bull. 1142-L. 23 pp.

Morris, H. T., and T. S. Lovering 1961. Stratigraphy of
the East Tintic Mountains, Utah, with a section
on Quaternary deposits by H. D. Goode. U.S.
Geol. Surv. Prof. Paper 361. 145 pp.

1979. General geology and mines of the East

Tintic Mining District, Utah and Juab counties,
Utah. U.S. Geol. Surv. Prof. Paper 1024. 203 pp.

MuNSELL Color Company. 1969. Munsell book of color.
Neighboring hues edition, Matte finish collection,
(looseleaf, unnumbered pages). Baltimore,
Maryland.

MuNZ, P. A. 1968. A California flora. Univ. of California
Press, Berkeley. 1681 pp.

Nesvetailova, N. G. 1970. Botanical prospecting for
ores. Office of Secretary of State of Canada,
trans., Ottawa, Canada. 97 pp.

OsBORN, B. 1955. How rainfall and nmoff erode soil.
Pages 126-135 in Water, USD A Yearbook of Ag-
riculture. U.S. Govt. Print. Off., Washington,
D.C.

Roberts, R. J. 1964. Stratigraphy and structure of the
Antler Peak quadrangle, Humboldt and Lander
counties, Nevada. U.S. Geol. Surv. Prof. Paper
459-A. 93 pp.

Roberts, R. J., and D. C. Arnold. 1965. Ore deposits of
the Antler Peak Quadrangle, Humboldt and
Lander counties, Nevada. U.S. Geol. Surv. Prof.
Paper 459-B. 94 pp.

Roberts, R. J., A. S. Radtke, and R. R. Coats. 1971.
Gold-bearing deposits in north-central Nevada
and southwestern Idaho. Econ. Geol. 66:14-33.

Rommel, M. A. 1968. Application of biogeochemistry to
mineral prospecting, a survey. NASA SP-5056.
134 pp.

Rowan, L. C, and M. J. Abrams. 1978a. Evaluation of
Landsat multispectral scanner images for map-
ping altered rocks in the East Tintic Mountains,
Utah. U.S. Geol. Surv. Open-File Rept. 78-736.
81pp.

1978b. Mapping hydrothermally altered rocks in

the East Tintic Mountains using 0.4-2.38 jum
multispectral scanner aircraft images, [abs]. Page
156 in International Association on the Genesis of
Ore Deposits, Snowbird, Alta, Utah. Program
with abstracts.

Rowan, L. C, and A. B. Kahle. 1982. Evaluation of
0.46 to 2.36 jum multispectral scanner images of
the East Tintic Mining District, Utah, for map-
ping hydrothermally altered rocks. Econ. Geol.
77:441-452.

Salisbury, F. B. 1954. Some chemical and biological in-
vestigations of materials derived from hydro-
thermally altered rock in Utah. Soil Sci.
78:277-294.

1964. Soil formation and vegetation on hydro-
thermally altered rock material in Utah. Ecology
45:1-9.

Sayers, R. W., M. C. Tippett, and E. D. Fields. 1968.
Duval's new copper mines show complex geolog-
ic history. Pages 55-62 in Mining Engineering,
March 1968.

Shawe, D. R., and J. H. Stewart. 1976. Ore deposits as
related to tectonics and magnetism, Nevada and
Utah. American Institute of Mining, Metallur-
gical and Petroleum Engineers, Society of Mining
Engineers, Trans. 260:225-231.

Silberman, M. L., J. H. Stewart, and E. H. McKee.
1976 Igneous activity, tectonics and hydro-
thermal precious-metal mineralization in the
Great Basin during Cenozoic Time. American In-
stitute of Mining, Metallurgical and Petroleum
Engineers, Society of Mining Engineers, Trans.
260:253-263.

Str.\hler, a. H. 1978a. Binary discriminant analysis: a
new method for investigating species-
environment relationships. Ecology 59:108-116.

1978b. Response of woody species to site factors

of slope angle, rock type and topographic posi-
tion in Maryland as evaluated by binary discrimi-
nant analysis. J. of Biogeography 5:403-423.

SuczEK, C. A. 1977. Tectonic relations of the Harmony
Formation, northern Nevada. Unpubli.shed dis-
sertation. Stanford Univ., Stanford, California.

Theodore, T. G., and D. W. Blake. 1975. Geology and
geochemistry of the Copper Canyon porphyry
copper deposit and surrounding area. Lander
County, Nevada: U.S. Geol. Surv. Prof. Paper
798-B,'86pp.

Theodore, T. G., and R. J. Roberts. 1971. Geochem-
istry and geology of deep drill holes at Iron Can-
von. Lander County, Nevada. U.S. Geol. Surv.
Bull. 1318. 32 pp.

TuELLER, P. T. 1975. The natural vegetation of Nevada.
Mentzelia 1:3-28.

Vasek, F. C, and R. F. Thorne. 1977. Transmontane
coniferous vegetation. Pages 797-832 in M. G.
Barbour and J. Major, eds.. Terrestrial vegetation
of California. Wiley & Sons, New York.

Welsh, S. L., and G. Moore. 1973. Utah plants:
tracheophyta. Brigham Young Univ. Press, Provo,
Utah. 474 pp.

Young, J. A., R. A. Evans, and P. T. Tueller. 1976.
Great Basin plant communities— pristine and
grazed. Nevada Archaeological Surv. 6: 186-214.

Young, J. A., R. A. Evans, and J. Major. 1977. Sage-
brush steppe. Pages 763-796 in M. G. Barbour
and J. Major, eds.. Terrestrial vegetation of Cali-
fornia. Wiley & Sons, New York.

Zamora, B., and P. T. Tueller. 1973. Artemisia arbtis-
cida, A. longiloba, and A. nova habitat types in
northern Nevada. Great Basin Nat. 33:225-242.

PLASTICITY AND POLYMORPHISM IN SEED GERMINATION
OF MIMULUS GUTTATUS (SCROPHULARIACEAE)

Robert K. Vickery, Jr.'

Abstract.— Seeds of 12 populations of Mimulus guttatus representative of the Wasatch Mountain ecotype were
incubated for 17 months (one natural season plus a year) in five artificial climates found in phytotron studies to be
important to the growth of the plants of that form of monkey flower. In all but the coldest climate, germination oc-
curred promptly (3-8 days, on average), peaked during the first three weeks, and then tapered off gradually well into
the second season. Generally, the amount and timing of germination was plastic, showing much the same range of
responses in widely different climates both overall and for individual populations. However, in some cases, there
were significant differences between populations indicative of polymorphism within the species. For example, ger-
mination was significantly slower, more variable, and less in amount the higher the elevation of origin of the popu-
lations. The responses of the population suggest the presence of both nuich plasticity and much polymorphism for
germination characteristics in this form of M. guttatus.

The purpose of this investigation is to
study seed germination in the yellow Monkey
flower, Mimulus guttatus Fischer ex DC, in
greater depth than was possible in the earlier
surveys (Vickery 1963, 1967). In those sur-
veys small samples of a series of species and
varieties of Mimulus were studied in a broad
range of artificial climates. The present study
concentrates on the Wasatch ecotype of
Mimulus guttatus (Vickery 1978) and on five
artificial climates found in phytotron studies
to be important for the growth of M. guttatus
(Vickery 1972, 1974).

Materials and Methods

Seeds from 12 populations of M. guttatus
were collected for the study from the
Wasatch mountain area of northern Utah and
southern Idaho (Table 1). The experiments
were carried out in four laboratory artificial
climates and one greenhouse climate (Table
2). The seeds were germinated on moist blot-
ters in petri dishes. Samples of 500 seeds, 125
per petri dish, were used for each population
in each climate. The climates included ex-
tremes of the earlier studies (1 and 5), optim-
al and suboptimal growth conditions (3 and
4, respectively), and the contrasts of fluctuat-
ing and steady temperatures (1 and 2 vs. 3, 4,
and 5). Germination was scored for 17
months, that is, through the 5-month germi-

nation season normal for M. guttatus in the
Wasatch Mountains plus an additional year.

Results and Discussion

Overall, germination in the four warmer
climates (2-5), started as early as the third

Table 1. Origins of the populations of Mimulus gut-
tatus studied, arranged by culture number, locality, and
elevation.

M. guttatus Fischer ex DC, n = 14

5839 Spruces, Big Cottonwood Canyon, Salt Lake
Co., Utah, 2350 m.

7273 Draper, Salt Lake Valley, Salt Lake Co., Utah,
1390 m.

7274 Gorgoza Ranch, Parley's Summit, Summit Co.,
Utah, 1910 m.

7311 Fish Haven, Bear Lake, Bear Lake Co., Idaho,
2030 m.

7312 Rick's Springs, Logan Canyon, Cache Co.,
Utah, 2000 m.

7314- East Canyon, Salt Lake Co., Utah, 2060 m.

7315 Thousand Springs, Mill Creek Canyon, Salt
Lake Co., Utah, 2215 m.

7316 Mill F East, Big Cottonwood Canyon, Salt
Lake Co., Utah, 2670 m.

7317 Brighton, Big Cottonwood Canyon, Salt Lake
Co., Utah, 2645 m.

7318 Homestead, Heber Valley, Wasatch Co., Utah,
1570 m.

7319 Snow Pine, Alta, Little Cottonwood Canyon,
Salt Lake Co., Utah, 2710 m.

11157 Mill D North, Big Cottonwood Canyon, Salt
Lake Co., Utah, 2520 m.

Note: The experiments were carried out at the University of Utah, elev.
1500 m, near the center of the study area.

'Department of Biology, University of Utah, Salt Lake City, Utah 84112.

470

July 1983

Vickery: Mimulus Germination

471

Table 2. Experimental climates used for the seed ger-
mination study.

1 Steady 4 C day and night, no photoperiod

2 Steady 25 C day and night, no photoperiod

3 Gradually changing from 4 C night to 17 C day, 16-
hour photoperiod

4 Gradually changing from 14 C night to 17C day, 16-
hour photoperiod

5 Standard greenhouse, changing from 10 G nights on
average to 30 G days on average, 16-hour
photoperiod.

NoTE:Artificial climates 1 and 2 employed incubators, whereas climates 3
and 4 employed growth chambers programmed to rise and fall like natural
July climates in the Wasatch Mountains (Dept. of Commerce, Climatolo-
gical Data, 1971-1980).

day, peaked during the following week, ta-
pered off to a low level by the end of the
third week, but continued to occur occasion-
ally well into the next year, forming a typical
(Went 1957, Vegis 1963) leptokurtic curve
(Fig. 1). Despite the overall pattern, germina-
tion varied noticeably from climate to cli-
mate in both speed and amount (Table 3) as
Stakanov (1976) observed in similar studies
on beans. For example, in the 17/14 C cli-
mate (4), the monkey flower seeds were sig-
nificantly slower than in the other cliamtes
both in starting to germinate and in achiev-
ing 50 percent of the ultimate, total germina-
tion for the 17-month trial period (Table 4).
The slowing effect on germination of the sub-
optimal, 17/14 C climate parallels the strik-
ing reduction in plant growth observed in
that climate in the phytotron (Vickery 1972,
1974) and suggests that the posited cause, too
similar day and night temperatures, acts on
speed of germination as well as plant growth.
In fact, if the temperature is constant as in
climate 2, total germination is significantly
less than in the fluctuating climates, 3, 4, 5
(Table 4). Overall, the variable, but generally
similar ranges of germination results in the
four diverse, warmer climates suggest wide
plasticity of response in M. guttatus.

In the cold, steady 4 C climate (1), in sharp
contrast to tiie pattern of early germination
in the four warmer climates, no germination
occurred at all during the first four weeks
(Fig. 1). After that, apparently the cumula-
tive effect of the time spent at room temper-
ature while the seeds were being watered and
scored triggered a little germination— 2 or 3
seedlings per petri dish— followed by a spurt
of germination when the watering and scor-
ing time was inadvertently prolonged. Thus,

4^ -U

Days

Fig. 1. The average number of seeds of all 12 popu-
lations of M. guttatus that germinated per day (Table 3)
for each of the five climates studied (full data available
on request). Peak germination occurred on day 7 in cli-
mate 3 with an average of close to 50 of the 500 seeds
studied per population per climate germinating that
day. The germination rate dropped off to approximately
one seed germinating per 25,000 seeds per day by day
255 and to one-tenth of that rate by day 512, the end of
the experiment.

the latter data for climate 1 are ambiguous
and were not analyzed. However, the early
results are clear and are consonant with the
extremely slow growth of the young plants in
the steady 4 C climate of the phytotron
(Vickery 1972, 1974).

In general, the 12 individual populations of
M. guttatus exhibit statistically similar ranges
of germination responses to the test climates
(Table 5) with some apparent differences
(Table 2, 5), much as Wright (1978, 1980) ob-
served in Panicwn. The differences range
from slight and insignificant to moderate to
three cases in which they are so pronounced
as to be statistically significant (Table 5). On
one hand, the overall similarities suggest a
wide plasticity of the populations of re-
sponse. On the other hand, the differences
appear to reflect underlying genetic differ-
ences—polymorphisms—of the populations.

The germination results of the 12 popu-
lations correlate with the elevation of origin
of the populations (Table 6). The populations
show significantly longer times to first germi-
nation and to 50 percent germination as well
as less total germination with increasing ele-
vation. This is true not only overall but in
most of the individual climates as well (Table
6). The variances of the overall times to first
germination are also significantly greater the
higher the elevation of origin (Table 6). The

472

Great Basin Naturalist

Vol. 43, No. 3

Table 3. Germination results for four replicates of each of the 12 populations of M. guttatus in each of the four
warmer climates (Table 2). Climate 1, steady 4 C, was omitted due to the ambiguity of the later results.

Population

Climate

Days to first
germination

Days to 50%
germination

Total
germination

5839
Spruces

7273
Drape

7274
Gorgoza

7311
Bear Lake

7312

Logan Canyon

7314

East Canyon

7315

1000 Springs

7316

Mill Creek
East

7317
Brighton

7318
Homestead

7319
Alta

11,157
MillD

North

All
populations

2
3
4
5
Average

5
Average

2
3
4
5
Average

5
Average

2
3
4
5

5.25 ±

2.50

5.00 ±

1.41

8.00 ±

1.41

5.00 ±

1.82

5.81 ±

2.10

4.00 ±

0.00

5.00 ±

0.00

6.00 ±

0.82

4.50 ±

0.58

4.88 ±

0.89

4.00 ±

0.00

4.25 ±

0.50

5.75 ±

0.96

4.00 ±

0.00

4.50 ±

0.89

6.33 ±

0.00

5.25 ±

0.00

7.75 ±

0.58

5.00 ±

0.50

6.08 ±

0.73

6.33 ±

2.31

5.25 ±

0.50

7.75 ±

0.96

5.00 ±

1.41

6.08 ±

1.67

4.00 ±

0.00

6.00 ±

1.41

5.25 ±

0.50

4.75 ±

0.50

5.00 ±

1.03

4.25 ±

0.50

6.00 ±

0.00

7.00 ±

0.82

5.00 ±

0.00

5.56 ±

1.15

8.00 ±

2.16

6.00 ±

0.82

9.50 ±

1.73

8.50 ±

2.52

8.00 ±

2.16

7.25 ±

2.63

5.25 ±

0.50

7.50 ±

0.58

6.50 ±

1.00

6.63 ±

1.59

3.25 ±

0.50

4.25 ±

0.50

6.75 ±

0.50

5.00 ±

0.00

4.81 ±

1..38

16.00 ±

10.74

7.25 ±

0.96

11.75 ±

1.26

8.50 ±

1.91

10.88 ±

6.04

3.75 ±

0.50

5.25 ±

0.50

7.50 ±

1.73

4.75 ±

0.50

5.31 ±

1.66

5.71 ±

4.60

5.29 ±

1.11

7.35 ±

2.02

5.52 ±

1.80

15.00 ±

9.90

34.75 ±

16.38

8.00 ±

1.83

81.50 ± 51.45

13.50 ±

2.08

73.25 ±

11.38

11.00 ±

0.82

52.25 ±

15.84

11.90 ±

5.37

60.44 ±

31.86

5.75 ±

0.96

56.25 ±

13.77

6.50 ±

0.58

63.50 ±

16.09

8.50 ±

0.58

68.00 ±

41.29

6.50 ±

1.00

86.25 ±

34.60

6.81 ±

1.28

68.50 ±

28.30

14.75 ±

13.10

24.75 ±

6.95

6.50 ±

1.00

109.00 ±

36.21

10.25 ±

2.50

98.75 ±

20.37

6.00 ±

1.15

95.75 ±

23.16

9.38 ±

7.03

82.06 ±

40.69

6.25 ±

0.50

65.50 ±

15.20

6.25 ±

0.50

102.75 ±

22.88

8.75 ±

1.26

78.00 ±

23.85

6.50 ±

1.00

90.25 ±

9.18

6.94 ±

1.34

84.13 ±

22.06

6.50 ±

3.69

2.67 ±

1.41

10.00 ±

0.82

79.25 ±

6.45

12.75 ±

0.50

21.25 ±

16.19

14.00 ±

4.24

67.25 ±

6.85

10.81 ±

4.47

55.11 ±

32.97

10.50 ±

2.65

31.50 ±

7.85

8.00 ±

2.71

88.75 ±

35.88

9.50 ±

1.29

98.00 ±

10.86

8.00 ±

2.16

97.75 ±

11.35

9.00 ±

2.31

79.00 ±

33.21

8.25 ±

2.22

15.25 ±

2.36

9.50 ±

0.58

68.72 ±

7.50

16.50 ±

7.59

77.75 ±

23.21

10.50 ±

1.00

59.00 ±

17.15

11.19 ±

4.85

55.19 ±

28.24

17.25 ±

8.34

17.50 ±

9.61

22.50 ±

12.40

60.50 ±

28.18

32.00 ±

4.32

49.25 ±

7.89

13.50 ±

2.38

34.00 ±

10.95

21.31 ±

10.10

40.31 ±

22.19

10.50 ±

2..38

8.75 ±

7.59

10.50 ±

1.73

47.75 ±

8.66

12.50 ±

1.00

61.50 ±

6.66

9.25 ±

2.06

35.75 ±

9.54

10.69 ±

2.06

38.43 ±

21.34

5.00 ±

0.00

65.25 ±

9.43

6.25 ±

0.50

85.00 ±

7.44

9.75 ±

0.50

74.50 ±

19.16

5.50 ±

1.00

80.25 ±

14.22

6.63 ±

2.00

76.25 ±

14.16

16.75 ±

10.20

3.25 ±

1.21

15.75 ±

12.30

8.50 ±

4.65

15.00 ±

5.35

8.50 ±

3.70

11.50 ±

0.58

9.00 ±

3.74

14.75 ±

7.82

7.31 ±

4.05

5.00 ±

0.00

82.00 ±

26.99

6.00 ±

0.00

97.25 ±

10.34

11.75 ±

1.71

75.25 ±

9.00

5.00 ±

0.00

77.00 ±

22.69

6.94 ±

3.00

83.00 ±

19.09

9.98 ±

7.25

33.90 ±

28.32

9.65 ±

6.56

74.38 ±

34.14

13.40 ±

6.77

69.54 ±

28.18

8.94 ±

3.42

65.38 ±

30.94

July 1983

Vickery: Mimulus Germination

473

Table 4. Comparison of germination in the different
artificial climates. Climates that are underlined together
are statistically inseparable according to the Student,
Newman, Keuls Multiple Range Test (Woolf 1968). aver-
age values are given below the lines.

Days to 1st germination

Days to 50% germination

Total germination
(Av. per petri dish)

5.3

5.5

5.8

7.4

8.9 9.6 10.2 13.4

2 5 4 3

34.6 65.4 69.5 74.4

slower and more varied germination re-
sponses seem a reasonable adaptation to the
increasing unpredictability of the climate
with increasing elevation in the Wasatch
area (Department of Commerce, Climatolo-
gical Data, 1971-1980). This finding of de-
creased germination is compatible, also, with
my field observations that the populations
appear to rely more on rhizomes than on
seeds at higher elevations, i.e., there appears

to be a cline of decreasing r-selection and in-
creasing K-selection with increasing
elevation.

Although the great majority of seeds ger-
minated promptly during the first season in
the four warmer artificial climates, a few,
about 1 percent on average, in each of the
populations did not germinate until the fol-
lowing season (Fig. 1). This result parallels
the germination response of M. guttatus in
the Wasatch Mountains. There, most of the
seeds germinate soon after shedding, i.e., dur-
ing the later part of the summer. They do not
require an after-ripening period as many spe-
cies do (Mayer and Poljakoff-Mayber 1975).
They continue to germinate in decreasing
numbers well into the fall. The seedlings
overwinter as small (1-2 cm) rosette stage
plants. A few new seedlings appear in the
spring along the streams as new habitats are
exposed with receding water levels. The late
germinating seeds appear to constitute small
but important seed banks (Harper 1977), both
for springtime germination and for the survi-
val of the populations in unfavorable years.

Table 5. Comparison of germination of the various populations. Populations within underlined groups are statis-
tically inseparable, whereas populations in nonoverlapping groups are statistically distinct according to the Student,
Newman, Keuls Multiple Range Test (Woolf 1968). Population culture numbers (Table 1) are given above the lines
and average values for each population below the line.

Days to first germination 7274 7311 7318 7273 7314 11157 7315 5839 73127317

4.5 4.6 4.8 4.9

5.0

5.3 5.6 5.8 6.1

6.6

7317 7316

6.6

8.0

7319
10.9

Days to 50% germination 7318 7273 7311 11157 7314 7274 7317 7312 7315

6.6

6.8 6.9 6.9 9.0

9.4 10.7 10.8 11.2
7274 7317 7312 7315 5839

7319

9.4 10.7 10.8 11.2 11.9

14.8

7316

21.3

Average total germination 7319
7.3

7317 7316 7315 7312

5839

38.4 40.3 55.2 .58.6 60.4

7315 7312 5839 7273 7318 7314 7274 11157 7311
55.2 58.6 60.4 68.5 76.3 79.0 82.1 83.0 84.1

474

Great Basin Naturalist

Vol. 43, No. 3

Table 6. Regression analysis of seed germination in the artificial climates versus elevation of origin of the popu-
lation, given as F-ratio and p value. Significant values are underlined.

Climates

First germination
F= p =

50% germination
F= p =

Total germination
F= p =

All together

Standard deviation (all)

11.462, .007

7.667,

.019

16.586,

.002

16.599,

.002

11.096,

.007

6.168,

.032

5.520, .041
7.942, .018
4.434, .061
2.767, .127
7.649, .019
4.265, .066

2.444, .149
4.946, .050
6.894, .025
19.650, .001
7.112, .024
2.578, .139

The Wasatch ecotype of Mimulus guttatus
exhibits both much plasticity in the overall
similarity of its range of seed germination re-
sponses to widely different climates and some
apparent genetic polymorphisms for speed
and amount of germination during the first
season, for the speed and amount of germina-
tion in populations with different elevations
of origin and for the ability to delay germina-
tion until the second season. Thus, the eco-
type appear to be well adapted for survival
in its climatically unpredictable area.

Acknowledgments

I thank Michael Nellestein for his technical
assistance with the seeds and Harold Hurst
and Dennis Phillips for their assistance with
the statistical checks.

Literature Cited

Harper, J. L. 1977. Population biology of plants. Aca-
demic Press, London. 892 pp.

Mayer, A. M., and A. Poljakoff-Mayber. 1975. The
germination of seeds. 2d ed. Pergamon Press, Ox-
ford, 178 pp.

Stakanov, F. S. 1976. Changes in the rate and simulta-
neity of bean seed germination in relation to ger-

mination temperature. Krishnev Inst. Transac-
tions 153:27-29.

U.S. Department of Commerce. 1971-1980. Climatolo-
gical data. Utah. Vols. 73-82.

Vegis, a. 1963. Climatic control of germination, bud
break, and dormancy. In L. T. Evans, ed.. Envi-
ronmental control of plant growth. Academic
Press, New York. 449 pp.

VicKERY, R. K., Jr. 1963. The evolutionary potential, as
measured by seed germination, of chromosome
races of Mimulus (Scrophulariaceae). Proc. XI In-
ter. Cong. Cenet. 1:46.

1967. Ranges of temperature tolerance for germi-
nation of Mimulus seeds from diverse popu-
lations. Ecology 48:647-651.

1972. Range of climatic tolerance as an in-
dication of evolutionary potential in Mimulus
(Scrophulariaceae). Symp. Biol. Hung. 12:31-42.

1974. Growth in artificial climates— an indication

of Mijuulus' ability to invade new habitats. Ecol-
ogy 55(4):796-807.'

1978. Case studies in the evolution of species

complexes in Mimulus. Evol. Biol. 11:405-506.

Went, F. W. 1957. The experimental control of plant
growth. Chronica Botanica, Waltham, Massachu-
setts. 343 pp.

WooLF, C. M. 1968. Principles of biometry: statistics for
biologists. Van Nostrand, Princeton, New Jersey.
359 pp.

Wright, L. N. 1978. Recurrent selection for changing
gene frequency of germination rate in Blue Panic
grass. Crop Sci. 18(5):789-791.

Wright, L. N. 1980. Germination rate and growth char-
acteristics of Blue Panic grass, Panicum anti-
dotale. Crop Sci. 20(l):42-44.

PREDATORY BEHAVIOR OF LARVAL AMBYSTOMA TIGRINUM NEBULOSUM
ON LIMNEPHILUS (TRICHOPTERA) LARVAE

Joseph R. Holomuzki'

Abstract.— Examination of stomach contents indicated that second-year Ambystoma tigrintim nebulostiin larvae
consumed Limneptnlus sp. larvae but rarely ingested the case. Feeding observations of captive salamanders on cad-
disfly larvae supported this finding. Extraction of caddisfly larvae from their case was accomplished only when larval
.salamanders quickly seized the anterior portion of ambulatorv Limnephihis sp. extended from their case and vigor-
ously shook the trichopteran from side to side.

Interest in the predatory behavior of Am-
hystoina tigrinum nebtiloswn on Limnephihis
sp. was prompted by the examination of
stomach contents of 29 second-year larvae
(^-13 mos. old) from east central Arizona.
Fourteen salamanders had eaten a total of 71
Limnephilus sp. larvae, yet remains of only 7
cases were evident. This indicated that A.
tigrinum were extracting caddisfly larvae
from their case. Such feeding behavior appar-
ently contrasts with some A. tigrinum in Utah
where individuals ingested caddisfly larvae
with cases (Tanner 1931). Moreover, extrac-
tion of larva from the case is discordant with
the notion that larval A. tigrinum exploit
aquatic resources in a manner nearly identi-
cal to freshwater fish (Zaret 1980), since fish
typically consume both caddisfly case and
larva (Elliot 1967, Tippets and Moyle 1978,
W. L. Minckley, pers. comm.). This paper
describes the ability of captive A. tigrinum
nebulosum larvae to extract Limnephilus sp.
larvae from their cases.

Salamanders used for feeding observations
and stomach analyses were collected in June
1981 from Big Meadows Tank 1, a per-
manent pond located 1.0 km NNW of the
western edge of Sunrise Lake, Apache Co.,
Arizona (elev. 2,774 m). Eight second-year
larvae varying from 78 to 98 mm from tip of
snout to posterior margin of vent were indi-
vidually kept in 36 X 22 X 26 cm aquaria
partially filled with 50 percent Holtfreter's
solution. Animals were acclimated for 24
hours before feeding observations were in-
itiated. Limnephilus sp. collected from Big
Meadows Tank 1 also were kept in 50 per-

cent Holtfreter's solution. Each salamander
was provided six caddisflies during feeding
runs. The number of strikes and successful
captures were counted in each 1-3 hour run.
Salamanders were not fed between
observations.

Movement by Limnephilus sp. seemed to
provide a visual stimulus for a strike response
by these salamanders. Previous studies also
noted A. tigrinum larvae typically striking
moving prey (Dodson and Dodson 1971, Rose
and Armentrout 1976). My observations,
however, suggest tactility may also play a
role in stimulating an attack on prey. Attrac-
tion of a salamander to a caddisfly case was
apparently frequently caused by any move-
ment of the case. The salamander usually
halted and placed its snout or chin against a
case that had moved. Further movement by
the caddisfly stimulated a strike. Salamanders
withdrew from the case if cessation of move-
ment was protracted.

During 26 hours of observations, only 2
(3.4 percent) Limnephilus sp. larvae were
eaten in 58 strikes. Unsuccessful attempts at
prey capture consisted of a salamander tak-
ing the entire case into its mouth. The animal
then manipulated the case and discarded it
after about 14 seconds (N = 12, range: 3-85
seconds). On no occasion was the case con-
sumed. Caddisfly larvae were successfully at-
tacked and eaten only when a salamander
slowly approached an ambulatory Limne-
philus sp. extended from its case and quickly
seized the anterior portion of the larva. The
salamander then vigorously shook the tri-
chopteran from side to side until extracted.

'Department of Zoology, Arizona State University, Tempe, Arizona 85287.

475

476

Great Basin Naturalist

Vol. 43, No. 3

Similar head-whipping behavior by metamor-
phosed A. trigrinum on elongate prey was de-
scribed by Larsen and Guthrie (1975) and
Lindquist and Bachmann (1980). This method
of Limnephilus sp. capture seemed to be sup-
ported by the stomach analyses, in which 10
of 71 consumed larvae were severed 0-2 mm
behind the metanotum.

The relatively poor capture success by
salamanders in the laboratory may also be
characteristic of the natural habitat. For ex-
ample, Dodson and Dodson (1971) found rel-
atively few tichopteran larvae in the diet of
A. tigrinum larvae from Colorado, even
though the insects were abundant in the sam-
led pond. Limnephilus sp., however, com-
prised about 16 percent of the total volume
of prey in the diet of larvae in June from Big
Meadows Tank 1. This suggests salamander
larvae of this population frequently attacked
trichopterans.

In sum, stomach contents showed that A.
tigrinum larvae from this population in-
frequently ingested caddisfly cases. Absence
of case consumption in the laboratory sup-
ported the finding. Successful attacks on Lim-
nephilus sp. were few and occurred only
when A. tigrinum quickly seized the anterior
portion of a caddisfly extended from its case.

These observations suggest caddisfly cases are
an effective means of deterring predation by
larval salamanders of this population.

Acknowledgments

Thanks are extended to J. P. Collins and
W. L. Minckley for commenting on an ear-
lier draft of the manuscript.

Literature Cited

Dodson, S. I., and V. E. Dodson. 1971. The diet of Ajji-
bijstoma tigrinum larvae from western Colorado.
Copeia 1971:614-624.

Elliot, J. M. 1967. The food of trout (Salmo tnitta) in a
Dartmoor stream. J. Appl. Ecol. 4:59-71.

Larsen, J. H., Jr., and D. J. Guthrie. 1975. The feeding
system of terrestrial tiger salamanders {Ambtjs-
toma tigrinum mekmostictum Baird). J. Morph.
147:137-154.

Lindquist, S. B., and M. D. Bachmann. 1980. Feeding
behavior of the tiger salamander, Ambystoma
tigrinum. Herpetologica .36:144-158.

Rose, F. L., and D. Armentrout. 1976. Adaptive strate-
gies oi Ambystoma tigrinum Green inhabiting the
Llano Estacado of west Texas. J. Anim. Ecol.
45:71.3-729.

Tanner, V. M. 1931. A synoptical study of Utali am-
phibia. Utah Acad. Sci. 8:159-198.'

Tippets, W. E., and P. B. Moyle. 1978. Epibenthic
feeding by rainbow trout. J. Anim. Ecol.
47:549-559.

Zaret, T. M. 1980. Predation and freshwater commu-
nities. Yale Univ. Press, New Haven.

NOTES ON REPRODUCTION OF THE SIDE-BLOTCHED LIZARD
UTA STANSBURIANA STANSBURIANA IN SOUTHWEST IDAHO

George Bakewell', Joseph M. Chopek', and Gary L. Burkholder'

Abstract.— In 1968 and 1973 in southwestern Idaho, 275 Ufa stanshuriana were collected. Uta occupies low hills
with rocky outcrops and flat sparse vegetation consisting of the following shrubs: Arternisia tridentata, Grcnjla spin-
osa, Atriplex confertifolia, Chrysothamnus nauseosus, and Atriplex canescens. Emergence from hibernation occurs
from mid-March to early April and the first yolked follicles appear in early April, with oviducal eggs present in late
April. Testicular cycle begins with the emergence of males, and spermatozoa are produced from April through July.
Uta reaches sexual maturity in one year at a SVL of 40.0 mm in males and 43 mm in females. Overall clutch size is
3.75 eggs (1-2 clutches per year). Fat body size at emergence is not known, but what is present shows a decline in
males and females until July, when a substantial increase occurs. Sex ratios are about 1:1 in all months and seasons
except June 1968, according to chi-square analysis.

This study has hmited scope for two rea-
sons: (1) the years of study are five years
apart; (2) in 1968 there were only six trips to
the field (which was about 10 miles from the
1973 area) in May and June, whereas in 1973
field trips were made once each week from 4
April through 25 July. In view of this, the
1968 data are minimized to some extent in
drawing our conclusion and emphasis is on
the 1973 data. Despite these limitations, the
study sheds some light on the reproductive
cycle of Uta stanshuriana in the northern
portion of its range.

Uta has been studied in considerable detail
in Texas (Tinkle 1961, 1967a, Hahn and
Tinkle 1965), Colorado (Tinkle 1967b), south-
em Nevada (Hoddenbach and Turner 1968,
Turner, Hoddenbach, Medica, and Lannom
1970, Medica and Turner 1976, Tanner
1972), and Oregon (Nussbaum and Diller
1976). Our results show that latitudinal (and
therefore climatic) differences exert observ-
able modifications of the reproductive cycle
as compared to that of southern populations
in Colorado, Nevada, and Texas, which are
800-1200 km south of our study, respective-
ly. In the case of the northern population
studied by Nussbaum and Diller (1976), our
results are similar in some aspects, as would
be expected, since their study area is only
225 km further north.

Physical and Biotic Environment

These counties included in the study area
are part of the Snake River Valley (Fig. 1).
Elevations of study sites and collection areas
range from 530 to 750 m. Dominant topo-
graphical features consist of small canyons,
which are a part of the Snake River Drain-
age, low rolling hills, boulder-strewn areas,
rocky outcroppings, and intermittent streams
that form sandy washes and/ or alluvial fans

Fig. 1. Geographic location for capture sites (hollow
circles) of Uta stanshuriana in southwestern Idaho. In-
sert shows the portion of Idaho where collections were
made.

'Mount Vemon Nazarene College, Martinsburg Road, Mount Vemon, Ohio 43050.

477

478

Great Basin Naturalist

Vol. 43, No. 3

where they reach the desert floor. Annual
rainfall varies from 10.1 to 30.4 mm; most of
this occurs as snow in winter (Shreve 1942).
Summers are typically hot and dry, with few
overcast days. Frost free days average 150
per year from April to September.

The dominant floral description for the
study areas is that of "cold desert formation"
(Shreve 1942). The dominant floral associ-
ations consist of several different com-
binations of the following desert shrubs:
Grayia spinosa (Hook.), Atriplex canescens
(Pursh, Ex. Nutt.), Atriplex confertifolia
(Torr.), Artemisia tridentata (Nutt.), and
Chrysothamniis nauseosus (Poll.). These
shrubs reach heights of 1 to 1.8 m and grow
in a clumped pattern that creates open areas
(Burkholder and Walker 1973). Many species
of annuals are present in the open areas be-
tween the shrubs, the majority of which flow-
er during the early and midspring rainy peri-
od. Others flower during the occasional rainy
periods of late spring and summer.

Methods and Materials

Specimens for this study were collected in
1968 and 1973 in Canyon, Ada, and Owyhee
counties of southwestern Idaho. All lizards
were weighed, measured (SVL), and autop-
sied (except hatchlings). The ovaries and ovi-
ducts of the females were removed and
counts were made of yolked ovarian follicles,
oviducal eggs, and corpora lutea. Yolked fol-
licles and oviducal eggs were measured to the

Table 1. Sex ratios for six samples of Vta stansbu-
ruina in 1968 and 1973. Each year total was tested for
goodness of fit by the chi-square statistic as well as the
month of Jime, where the largest differences occurred.
All fit the 95 percent confidence limit except June 1968.

nearest 0.1 mm and weighed to the nearest
.01 g. In males, one testis was removed and
used to determine sexual maturity by a
squash preparation using aceto-orcein stain.
Clutch size data followed the procedure out-
lined by Tinkle (1961). Fat bodies were also
removed and weighed to the nearest .01 g.

Results

Sex ratio.— Sex was determined in 275
specimens by the examination of gonads. In
May and June 1968, 34 females and 50 males
were collected. The April through July 1973
sample consisted of 90 females and 101
males. Both years combined resulted in 124
(45.1 percent) females and 151 (54.9 percent)
males. Data for sex ratios are summarized in
Table 1. Statistical analysis for fitting the
expected ratio of 1:1 are also included in
Table 1.

Size at maturity and at hatching.— Size
of individuals when reaching sexual maturity
was determined by analyzing the gonads. In
females, yolked ovarian follicles, oviducal
eggs, and corpora lutea were used as criteria
for maturity. From this analysis, it was deter-
mined that the smallest sexually mature fe-
male was 43 mm SVL for the 1973 sample
and 44 mm SVL for the 1968 sample.

Size at sexual maturity in males was deter-
mined by analysis of a single testis removed
from each male. An aceto-orcein squash
preparation was performed to check for the
presence of mature spermatozoa. All male
specimens, even the smallest at 40 mm SVL,
yielded a positive test for spermatozoa. The

Table 2. Data for hatchling Vta stansbiiriana cap-
tured in July 1973. Two lizards not listed were observed
on 26 June but not captured.

Month Year

Females
14

Males
13

SVL (mm)

Date Captured

May 1968

26.0

3 July

June 1968

25.5

18 July

Total

25.0

23 July

Chi-square test .95 = 5.06 for

Tune

25.0

23 July

Chi-square test .95 = 3.04 overall

26.0
31.0

23 July
23 July

April 1973

31.0

23 July

May 1973

31.5

23 July

June 1973

36.0

23 July

July 1973

28.0

25 July

Total

101

29.0

25 July

Chi-square test .95 = 1.472 for June

31.0

25 July

Chi-square test .95 = .632 overall

33.0

25 July

July 1983

Bakewell et al.: Side-Blotched Lizard

479

6 -

X=6.00

5 -
4 -

5a?. Q?

X=4.13

11 II

c cc

Y— •3 £1 3 •» ^^

VO 1
i-l C\J

11 II
c OS

A-J.O 1

A-J.O/

un 1

CJ CO

II 11

^ .

X=3.00

c<^ 1
II 11

in
O 1

11 1!

2 -

1 -
0

11 II

C PC

43-44

45-46

47-48

49-50

51-52

53-54

Size class of females (SVL)

Fig. 2. The mean number of yolked ovarian follicles, oviducal eggs, and corpora lutea for six arbitrarily sized class-
es of females. N = number of individuals, R = range, and X = mean 1968 and 1973 females combined.

12 hatchlings were not sexed but were as-
sumed not to be reproductively mature.

The first hatchlings were observed, but not
collected, on 26 June 1973. Twelve speci-
mens were collected from 3 July 1973 to 25
July 1973. The data for hatchlings is pre-
sented in Table 2.

Mean size of mature individuals.— The
largest male specimen was 57.5 mm SVL and
the smallest 40.0 mm SVL. The largest fe-
male was 53.0 mm SVL and the smallest was
43 mm SVL. Table 3 summarizes the remain-
ing data related to mean size of mature
individuals.

Reproduction in females.— Counts were
made of oviducal eggs, corpora lutea, and
yolked ovarian follicles in the ovaries of 124
females. Figure 2 shows a comparison of six
arbitrary size classes that indicates that
clutch size increases with size of the female,
with the two smallest categories (43-44 and
45-46 SVL) presumably being first-year re-
producive females, and those 47 and over
second-year females. Using 1973 female data
alone, because it covers the entire reproduc-

tive season, there was an attempt to deter-
mine the number of clutches per year, which
is difficult in multiple clutch species.

The 1973 data appears to indicate a two-
clutch ability by some individuals. The first
clutch group is clustered between 23 April
and 22 May (based on shelled oviducal eggs).
The second clutch group is clustered between
2 June and 10 July. Between the period of 22
May and 2 June, there are females with ovi-
ducal eggs but these are late individuals re-
productively, which will probably produce
only one clutch because of time constraints.
To substantiate the double clutch, we began
with the correlation of the first appearance
of hatchlings, assuming a 45-50 day in-
cubation period, which is based on Bur-
kholder and Tanner's (1974) work on Scelo-
porus graciosus, which has a mean incubation
period of 49 days (from laboratory and field
incubation data). If the females of the first
group laid eggs from 23 April through 22
May, with a 50-day incubation period, the
hatchlings would appear as early as 11 June

and as late as 10 July. The first hatchlings
Table 3. The mean SVL in millimeters of sexually mature males and females for 1968 and 1973, based on sperma-
tozoa in testis and epididymus and yolked ovarian follicles, oviducal eggs, or corpora lutea, respectively.

Males

Females

Year

1968
1973

49.27
49.66

51
100

44-57
40-55

47.70
48.06

34
99

44.5-52
43-53

480

Great Basin Naturalist

Vol. 43, No. 3

were observed on 26 June and 3 July (Table
2), which falls within the predicted time. The
second clutch hatchlings would appear 21
July through 28 August. The smallest hatch-
lings captured between 23 July and 25 July
(Table 2) fit into this category.

The second point that supports two
clutches is that of the time span between the
first and second onset of vitellogenesis. Turn-
er, et al. (1970) state that 31 days are suf-
ficient for production of a second clutch. If
the days are counted between the proposed
first and second clutch (clustering of females
with shelled oviducal eggs), there are 42
days, which would appear to be ample time.

The only nonsupportive evidence is the ab-
sence of corpora lutea when the second pro-
duction of yolked ovarian follicles occurs.
However, it is our opinion, based on observa-
tions of autopsied individuals, that corpora
lutea in Uta disappear very quickly (1-5
days), which would explain the lack of
overlap.

Along with the double clutch phenomena
is the feature of clutch size fluctuation. The
mean size of the first clutch (based on ovi-
ducal eggs only) was 4.40 (N = 14) and the
second was 3.81 (N = 16). This corresponds to
that reported by Tinkle (1967b) for Texas
populations, Turner et al. (1970), and Medica
and Turner (1976) for Nevada, and Nussbaum
and Diller (1976) for Oregon, though the dif-
ference is not as significant.

The earliest onset of vitellogenesis is 14
April (1973). This is based on yolked ovarian
follicles of 2 mm diameter or larger and a
definite yellow color. Table 4 summarizes the
numbers of yolked ovarian follicles, oviducal
eggs, and corpora lutea for the determination
of clutch size (1968 and 1973 data were

Table 4. Data for yolked ovarian follicles, oviducal
eggs, corpora lutea, and mean clutch size per female for
the years 1968 and 1973.

Ovarian Mean

yolked Oviducal Corpora clutch

Year follicles eggs lutea size

1968 45(N=12) 65(N=17) 0 3.79

1973 112(N = 33) 135(N = 33) 7(N = 2) 3.74

Total 157(N = 45) 200(N = 50) 7(N = 2) 3.64

Mean 3.49 4.00 3.50 3.75

Clutch size
Clutches per year

pooled). Reproductive potential was not cal-
culated; the actual numbers of females laying
a second clutch was not determined.

Fat bodies.— The fat bodies decrease in
weight from the time of emergence of adults
through the end of the reproductive period in
both males and females (Table 5)

Discussion

Although the volume of data and length of
time are much less than studies which we
will use for comparison and contrast, this
study provides some additional information
concerning the total knowledge of natural
history of Uta stansburiana. In essence, the
comparisons made with regard to Uta con-
cern that of northern vs. southern popu-
lations and the changes that occur along that
continuum.

The first feature is that of sex-ratio. On a
yearly basis, all (1968 and 1973) fit the

Table 5. Fat body weight changes in 1973 male and
female Uta stansburiana April-July in grams. Single as-
terisk means that all had yolked follicles or oviducal
eggs. Double asterisk means the two at < .01 still had
oviducal eggs and corpora lutea and the one at .08 cor-
pora lutea only, and the rest had no reproductive activi-
ty. Triple asterisk means the .25 specimen had just be-
giui vitellogenesis; the two at < .01, two at .02, and two
at .03 had not started vitellogenesis, whereas all others
had.

April

May

July

Males 16=<.01 32=<.01 26=<.01 3=<.01

1 = .08 1 = .01 2 = .01
1 = .03
2=.04

1 = .05

2 = .06
2 =.07

1 = .08

2 =.09
1 = .10
1 = .15

Females 11 = < .01 18= < .01 14= < .01 1 = <.01

4 =.03
1 = .05
3 =.02
1 = .04
1 = .25

2 =.04
2 =.03
2 = .02

1 = .05

2 =.03
1 = .02

3.75
1-2

1 = .02

2 = .05
1 = .06
1 = .07
1 = .08

1 = .09
2=.10

2 = .ll
1 = .12
1 = .15

21.0.

24°

18°

15°

July 1983

Bakewell et al.: Side-Blotched Lizard

481

expected ratio of 1:1. Monthly fluctuations
are not as consistent, especially in June for
both years. These data indicate a 3:2 ratio of
males to females, which is similar to what
Tinkle (1961) recorded in Texas. Tinkle
(1961) attributed this to territorial vigilance
in males, and this appears to be true for those
in Idaho. In addition to this, the difference
may be further enhanced by female oviposi-
tion in June. By July these activities (male
and female) cease, and the ratio returns to
1:1.

Idaho Uta attain sexual maturity in one
year; they hatch, over winter, and emerge as
adults (Tinkle 1961, Tanner 1972, Medica
and Turner 1976). Nussbaum and Diller
(1976), however, observed that in north cen-
tral Oregon some Uta that hatched late in
the year (late August) would not have suf-
ficient time for growth and therefore would
not be sexually mature by the time the next

Table 6. Comparison of data for six Uta populations.

reproductive season arrived. We think that
this is rare in Idaho, based on size of earliest
individuals collected, because they had
yolked ovarian follicles, spermatozoa, and
larger SVL than the minimum SVL as deter-
mined for sexual maturity of Oregon Uta.
The longer growth period of Idaho vs. Ore-
gon is most likely the basis for the differences
in the two populations.

The remainder of our findings concerning
Idaho Uta have been added to Table 6,
which is a modification from Nussbaum and
Diller (1976). Table 6 calls attention to sever-
al areas of interest. The elevation is as low or
lower than all the previous studies, though
further north than all but the one in Oregon.
Length of growing season and therefore
length of reproductive season are longer than
that found in Oregon but less than the four
southern studies. These physical aspects put
the Idaho population in a somewhat inter-
mediate position between that of Oregon and

Texas

Colorado

Nevada

(Rock

Valley)

Nevada
(Rainier

Mesa)

Oregon

Idaho

(1973)

Elevation (ft)

Length of growing season (days)

Length of reproductive season

(davs)
Habit'

Sexual dichromatism
Aggressiveness
Social stnicture
Density (individuals /acre)
Male home range (acres)
Female home range (acres)
Hatchling size (nun(SVL)
Size range mature males (mm)
Size range mature females

(mm)
Average size adult males (mm)
Average size adult females

(mm)
Clutch size
Clutch frequency
Percent males fail to breed first

season
Percent females fail to breed

first season
Percent males two years old

and older
Percent females two years old

and older
Date hatchlings appear

2900

^4250

3400

7840

2400

1722-2437

215

175

225

200

110-140

150-180

121-141

120

135

90-100

arenicolous

saxicolous

—

saxicolous

high

103

—

high

103

—

low

territorial

dominance

—

dominance (?)

36.109

17.5

0.50

0.27

—

0.54

—

0.17

0.23

—

0.43

—

^22

s22

^22

26'

40-60

42-

40-56

40-53

40-55

40-60

37-

40 +

41-49

43-53

48.4

49.6

48.9

42.8

45.4

48.0

^4.0

3.20

.6- .4

4.85

3.33

3.75

3.5

.3-5

1-2

0.0

<25

= 19.0

0.0

<25

= 47.0

18-28

36-65

57.6

18-28

36-65

69.4

20 June

25 June

17 July

26 June

Texas and Colorado (Tinkle 1961, 1967a, b, 1969a, Tinkle and Woodward 1967)

Oregon (Nussbaum and Diller 1976)

Nevada-Rock Valley (Turner et al., 1970, Medica and Turner 1976)

Nevada— Rainier Mesa (Tanner 1972)

Idaho— (this study)

"Smallest individual caught 3 July 1973

482

Great Basin Naturalist

Vol. 43, No. 3

the four southern populations, even though it
is not located halfway between from the
standpoint of miles. Because of these factors
and others (i.e., precipitation and primary
production), there should be some influence
on reproductive cycle, average adult size,
date of hatching, appearance, etc.

In reference to the above-mentioned pa-
rameters and the information in Table 6, it
becomes apparent that Idaho Uta do lay
more than one clutch per year. This is based
on the broad period of time when females
have oviducal eggs present (the same is true
for yolked ovarian follicles) and the appear-
ance and size (SVL) of hatchlings from 26
June through late July. The percentage of fe-
males laying a second clutch is speculative
because of incomplete data, but it would ap-
pear to occur in the majority of the popu-
lation as compared to a small percentage in
the Oregon population as reported by Nuss-
baum and Diller (1976). It is doubtful if any
could produce a third clutch, as is the case in
Colorado Uta (Tinkle 1976), due to length of
reproductive season, though the possibility
does exist if favorable conditions prevail.

Coupled with this is the fairly high overall
clutch size of 3.75 or, if based on oviducal
eggs only (first and second clutch), 4.40 and
3.81/female, which approaches the size if not
equals that of the Texas, Colorado, and Ne-
vada (Rock Valley) populations. The only dif-
ference is the number of clutches per year.
The size is significantly higher than that of
Oregon. The explanation for the similarity to
populations much farther south and the dif-
ference from the more northern Oregon pop-
ulation is complicated but again probably is
within the realm of the response of Uta to
changes in the elevation and/or latitude,
length of growing and reproductive seasons,
annual precipitation as it affects primary
production, and general habitat. Because of
the limitation of this study, we feel in-
adequate to speculate on these inter-
relationships at this time.

Average size of males and females (SVL),
as well as size range for mature males and fe-
males, does not appear to deviate dramatical-
ly from other populations. Fat body cycle
follows that reported first by Hahn and
Tinkle (1965) and by many subsequent au-
thors with regard to various lizard species.

Literature Cited

BuRKHOLDER, G. L., AND W. W. Tanner. 1974. Life his-
tory and ecology of the Great Basin sagebrush
swift, Sceloporus graciosits graciosus Baird & Gi-
rard, 1972. BYU Sci. Bull., Biol. Ser. 19(5): 1-42.

BuRKHOLDER, G. L., AND J. M Walker. 1973. Habitat
and reproduction of the desert whiptail lizard,
Cnemidophorns tigris Baird and Girard at the
northern part of its range. Herpetologica
29:76-83.

Hahn, W. E., and D. W. Tinkle. 1965. Fat body cycling
and experimental evidence for its adaptive signif-
icance to ovarian follicle development in the liz-
ard Uta stansbuhana. J. Exp. Zool. 158(l):79-86.

Hoddenbach, G. a., and F. B. Turner. 1968. Clutch
size of the lizard Uta stanshuriana in southern
Nevada. Amer. Midi. Nat. 80(l):262-265.

Medica, p. a., and F. B. Turner. 1976. Reproduction
by Uta stanshuriana (Reptilia, Lacertilia, Igua-
nidae) in southern Nevada. J. of Herpetology
10(2): 123-128.

NussBAUM, R. a., and L. v. Diller. 1976. The life his-
tory of the side-blotched lizard, Uta stanshuriana
Baird and Girard, in north-central Oregon.
Northwest Sci. 50(4):243-260.

Shreve, F. C. 1942. The desert vegetation of North
America. Bot. Rev. 8:195-246.

Tanner, W. W. 1972. Notes on the life history of Uta
stanshuriana. BYU Sci. Bull., Biol. Ser.
15(4):31-39.

Tinkle, D. W. 1961. Population structure and reproduc-
tion in the lizard Uta stanshuriana stejnegeri.
Amer. Midi. Nat. 66:206-234.

1967a. Home range, density, dynamics, and struc-
ture of a Texas population of lizard Uta stanshu-
riana. Pages 5-59 in W. W. Milstead, ed.. Lizard
ecology: a symposium. Univ. of Missouri Press.

1967b. Life and demongraphy of the side-
blotched lizard. Univ. Michigan Misc. Publ. Mus.
Zool. 1.32:1-182.

Turner, F. B., G. D. Hoddenbach, P. A. Medica, and
J. R. Lannon. 1970. The demography of the liz-
ard Uta stanshuriana Baird and Girard in south-
ern Nevada. J. Anim. Ecol. 39:505-519.

OBSERVATIONS ON ALPINE VEGETATION NEAR SCHOOLROOM GLACIER,

TETON RANGE, WYOMING

John R. Spence' and Richard J. Shaw'

Abstract.— Quadrat and propagiile trapping studies were made on the moraine of the Schoolroom Glacier and in
adjacent dry alpine meadow vegetation in the Teton Range in 1978-1979. Forty-six species of vascular plants were
collected. Distributionally, three groups of species exist. One of these is concentrated primarily on the moraine, a
second in the meadow, and the third in a narrow ecotonal band at the base of the distal slope of the moraine. The
moraine slopes are steep and unstable, with vegetation cover ranging from 1 to 9 percent, dominated by Cirsium
tweedyi (Rydb.) Petr. Along the more stable moraine crest the vegetation cover is heavier, and is similar to that in
the meadow. The meadow vegetation cover is about 50 percent, dominated by Astragalus kentrophyta Gray. Using a
combination of cover and frequency as a measure of importance, dominance-diversity curves were constructed for
the moraine and meadow. Both approach geometric series, which are suggested as indicating harsh environments.
Abiotically pollinated species are more successful on the moraine than biotically pollinated species, but the reverse is
true for the meadow. Propagule trapping studies suggest that dispersal of anemochorous propagules onto the mo-
raine is very low compared with dispersal in the meadow.

Relatively little is known about the struc-
ture and dynamics of alpine vegetation in the
Teton Range, Grand Teton National Park,
Wyoming. In this paper we report some pre-
liminary quadrat and propagule trapping
studies from alpine vegetation in the south-
em fork of Cascade Canyon in the center of
the range.

The selected study area is a complex of al-
pine meadow and morainal deposits at the
head of the south fork of Cascade Canyon,
about 4 km southwest of the Grand Teton
(Fig. 1). An east-facing cliff called The Wall
bounds the study area on the west; this for-
mation forms part of the hydrographic divide
of the Teton Range. A shallow cirque has
been carved into this cliff by the Schoolroom
Glacier, so called because of the almost per-
fect end moraine fronting it (Fig. 2). The age
of this moraine is unknown, but similar de-
posits elsewhere in the range are of Neo-
glacial age, which places the moraine age
from about 100 to 3,000 years (Mahaney
1975, Mahaney and Spence 1983). Between
this moraine and the glacier lies a small melt-
water lake about 50 m across, which is
drained by a stream that has cut through the
center of the moraine. The two ends of the
moraine merge into extensive talus derived
from The Wall to either side of the glacier.

Northeast of the glacier is a large outcrop of
gneiss and schist of Precambrian age, which
is vegetated by a mosaic of fen and tim-
berline krummholz stands. To the east and
southeast lies an extensive, slightly undulat-
ing dry alpine meadow underlaid by glacial
and talus deposits of Pinedale or older age.
The Wall and Schoolroom Glacier moraine
are composed primarily of Death Canyon
Limestone of the Gros Ventre Formation
(Cambrian), with some debris of the Wolsey
Shale member of the Gros Ventre Formation,
in addition to Flathead Sandstone (Cambrian,
Love, and Reed 1968, Reed 1973).

Climate data from the Teton Range and
park are summarized elsewhere (Spence
1981). Mean annual temperature from Jack-
son Hole to the east of the range, at an eleva-
tion of 2,040 m, is about 1.3 C. Using the ele-
vation of the study area, 3,060 m, and the
adiabatic lapse rate (Cole 1975), average an-
nual temperature in front of the Schoolroom
Glacier would be about -4.8 C. Snow depth
during the winter is unknown, but it is usual-
ly gone from the area by late June to early
July during an average year. Common ani-
mals at the site include marmots {Marmota
flaviventris), pikas {Ochotona princeps), and
Rosy Finches {Leucosticte atrata).

The purposes of this paper are (1) to char-
acterize and contrast the vegetation on the

'Department of Botany, University of British Columbia, Vancouver, B.C., Canada V6T 1W5.
'Department of Biology, Utah State University, UMC45, Logan, Utah 84322.

483

484

Great Basin Naturalist

Vol. 43, No. 3

3 N 42'

rpn glacial moraine

I'^I^I glacier/snowbank

lake ''
fen

it^

Legend

stream
geologic unit

major topographic
barrier

g-mo gneiss-quartz monzonite

qt glacial/periglacial
deposits

qa alluvial deposits

ewf Wolsey Shale

edc Death Canyon Limestone

A peak

Fig. 1. A sketch map of the study area at the head of the south fork of Cascade Canyon. Only major features are
shown on the map. The boundaries for the geologic units are only approximate. The map is derived from 7..5 minute
uses topographic maps. Love and Reed (1968), Reed (1973), and personal observations of the senior author (JRS).
See the legend for map details.

Schoolroom Glacier moraine and in the dry
alpine meadow adjacent to it, and (2) to sug-
gest possible dynamics between the two
communities.

Methods and Materials

Initial site observations and collections
were made in the summer of 1978. In 1979,

July 1983

Spence, Shaw: Schoolroom Glacier Plants

485

Fig. 2. Schoolroom Glacier at the head of the south fork of Cascade Canyon, t.rand ieton i\ational Park,
Wyoming.

the quadrat data were gathered on 19 Sep-
tember. Two transects, each 90 m long, were
run from the base of the proximal slope of
the moraine adjacent to the meltwater lake,
up to and over the moraine crest, down the
distal slope, and out into the meadow. At 3 m
intervals, a 0.5 X 0.5 m quadrat was placed
down on alternating sides of the transect line.
Cover and presence of all species was noted
visually, using a modified Braun-Blanquet
scale, as follows: + = 0-1 percent, 1 = 1-5
percent, 2 = 5-25 percent, 3 = 25-50 per-
cent, 4 = 50-75 percent, 5 = 75-100 per-
cent. For calculating the total and average
cover values, the midpoints of the ranges
were used. Prominence Values (PV) were cal-
culated using the formula PV = percent cov-
er X the square root of percent frequency.
Along the transect lines, slope was measured
using an Abney level.

Propagule trapping was studied in two
ways, by water-filled plastic trays 40 X 15
X 5 cm in size, and 15 X 15 cm wooden
plates coated with petroleum jelly. The wa-
ter trays were used in 1978, with six of them

placed in a line from the edge of the melt-
water lake up and over the moraine to the
meadow. In 1979, nine wooden plates were
used, and were placed in a line 10 m apart
from the edge of the lake up and over the
moraine and out into the meadow. Once in
July and once in September the wooden
plates were in operation, for a total of 38
hours in July and 24 hours in September. This
amounts to a total of 558 trap-hours in oper-
ation. Trapped propagules were placed in
glass vials for later identification. A reference
set of propagules from species at the site was
made to aid in identification. Specimens col-
lected are on deposit at the Moose Herba-
rium in Grand Teton National Park and the
Intermountain Herbarium at Utah State Uni-
versity (UTC). Nomenclature follows Shaw
(1976).

Results and Discussion

Forty-six species of vascular plants were
collected from the moraine and adjacent
meadow in 1978 and 1979. Details on the

Great Basin Naturalist

Vol. 43, No. 3

floristics and comparisons with other glacial
moraine sites in the Tetons will be published
elsewhere (Spence 1983). One species, Ta-
raxacum lyratum Ledeb., is a new report for
the park. The distribution of life forms of the
species is: 1 shrub, 1 fern ally, 1 annual dicot,
11 graminoids, and 32 biennial/perennial
forbs. Of these species, 31 were encountered
along the two transects (Table 1). The re-
maining species are quite rare at the site,
many of them consisting of only a few indi-
vidual plants.

Along the first transect, average cover was
0.6 percent for the moraine proximal slope,
8.9 percent for the distal slope, and 47.4 per-
cent for the meadow. For the second tran-
sect, the respective values were 2.8 percent,
9.2 percent, and 63.3 percent. Cover values

per quadrat ranged from 0 percent (7 times;
6 on the moraine proximal slope, 1 on the
moraine distal slope, and 0 in the meadow) to
106.5 percent (once, in the meadow). Pooling
the results of the two transects, total cover on
the moraine is 1.7 percent for the proximal
slope and 9.1 percent for the distal slope. Av-
erage cover for the meadow is 54.6 percent.
These values are all significantly different
from one another at a = 0.05. Data from the
two transects are summarized in Table 1 for
the moraine and the meadow.

Although most of the species encountered
in the transects are found on both the mo-
raine and in the meadow, they tend to be
much more common on one or the other (see
Table 1). General observations elsewhere on
the moraine and in the meadow tend to sup-

Table 1. The quadrat data from the two Hne transects in summarized form. Each transect was 90 m long, with a
0.5 X 0.5 m quadrat placed at every 3 m interval, for a total of 60 quadrats and 15 m^. The species are arranged
alphabetically, and three numbers are listed for each species; percent frequency, average percent cover, and promi-
nence value, which is calculated as average percent cover X square root of percent frequency. For details on tran-
sect placement and method of measuring species, see Methods and Materials.

Moraine (N = 38)

Meadow (N = 22)

Species

(percent frequency/average percent cover/prominence value)

Achillea millefolium
Agmpyron caninuvi
A. scribneri
Antennaria umbrinella
Arabis h/allii
Arenaria nutiallii
Arnica longifolia
Astragalus kentrophyta
Carex species
Cirsium tweedyi
Cymoptertts hendersonii
Epilobium alpiniim
Erigeron compositus
E. leiomerus
Erysimum asperum
Festuca ovina
Hymenoxys grandiflora
Oxyria digymt
Phacelia sericea
Poa alpina
P. pattersonii
Polemonitim viscosurn
Salix arctica
Selaginelki densa
Senecio fremontii
Silene acaulis
Solidago multiradiata
Taraxacum lyratum
T. officinale
Townsendia montana
Trisetum spicatum
Unknown grasses
Unknown herbs

10.5/0.54/1.75
2.6/0.07/0.11
7.9/0.09/0.25

2.6/0.01/0.02
2.6/0.45/0.73

26.3/0.92/4.72
7.9/0.04/0.11

18.4/0.63/2.70
7.9/0.14/0.39

2.6/0.01/0.02
2.6/0.01/0.02

7.9/0.14/0.39
15.8/0.24/0.95
18.4/0.58/2.49

23.7/0.12/0.58
2.6/0.39/0.63
5.3/0.08/0.18
5.3/0.03/0.07

10.5/0.05/0.16
7.9/0.14/0.39
7.9/0.14/0.39

42.1/0.70/4.54

18.4/0.09/0.39

4.5/0.68/1.44
22.7/4.25/20.25

45.5/4.82/32.51

13.6/0.07/0.26

68.2/28.20/232.S
9.1/0.14/0.42

4.5/0.68/1.44

4.5/0.02/0.04
13.6/0.34/1.25

4.5/0.68/1.44
18.2/0.09/0.38

4.5/0.02/0.04
72.7/4.20/35.81

9.1/0.14/0.42

40.9/3.59/22.96
4.5/0.68/1.44
9.1/1.82/5.49
4.5/0.68/1.44

4.5/1.70/3.61
31.8/0.43/2.42

22.7/0.11/0.52

9.1/0.14/0.42
27.3/1.64/8.57
50.0/4.36/30.83

July 1983

Spence, Shaw: Schoolroom Glacier Plants

487

port the conclusions drawn from the transect
data. Only a few species, such as Poa patter-
sonii, Agropyron caninum, and Taraxacum
officinale, appear to be equally common in
both areas. Furthermore, a group of species
appears to be restricted to an area at the base
of the distal moraine slope. They are found in
a band ranging from 1 m up to 10 m wide be-
tween the moraine slope and the meadow
proper. This band is formed primarily of
debris derived from slumping and sliding off
the distal moraine slope. Some of the species
that were found in this ecotonal region in-
clude Anemone multifida, Draba loncho-
carpa, Oxytropis deflexa. Taraxacum lyratum,
Androsace septentrionalis, Castilleja sul-
phurea, and Eritrichium nanum. Few of these
species were found on the moraine, and they
were all either rare or absent from the mead-
ow. The transect data show that this ecotonal
region is richer in species than either the mo-
raine or meadow. Average number of species
per quadrat ranged from 4 on the moraine, to
5.1 in the meadow, to 7.3 in the ecotone be-
tween the two.

Using prominence values (PV), the most
important species on the moraine are Cirsium
tweedyi, Epilobium alpinum, Polemonium
viscosum, Agropyron caninum, and Poa pat-
tersonii. In the meadow the most important
species are Astragalus kentrophyta, Hyme-
noxys grandiflora, Antennaria umbrinella,
Poa pattersonii, and Agropyron caninum. Us-
ing the contribution of each species PV to
the total summed PV for the moraine and the
meadow, dominance diversity curves can be
constructed (Fig. 3). Both curves approach
geometric series, which have been suggested
to indicate harsh environments in which do-
minance by one or a few species is strong
(Whittaker 1975). On the moraine, several
species share dominance; these are Cirsium
tweedyi (28 percent of the total summed
PV's), Epilobium alpinum (16 percent), Pole-
monium viscosum (15 percent), and Agropy-
ron caninum (10 percent). In the meadow.
Astragalus kentrophyta dominates (63 per-
cent). Other species include Hymenoxys
grandiflora (10 percent), Antennaria umbri-
nella (9 percent), and Poa pattersonii (6 per-
cent). Of the eight species listed above, the
majority are either western North American
alpine species {P. viscosum, A. kentrophyta.

Moraine
Meadow

0.1%

Species Rank

Fig. 3. Dominance-diversity curves for the moraine
and the meadow. The measure of importance used was
each species contribution to the total summed promi-
nence values in percent. There are 21 species in the mo-
raine curve and 23 species in the meadow curve. All
species with values less than 0.1 percent had 0.1 percent
added to their value to include them on the graph. This
tended to slightly flatten the two curves at the bottom.

488

Great Basin Naturalist

Vol. 43, No. 3

P. pattersonii, and A. umbrinella) or
south/central Rocky Mountain alpine species
(C. tweedyi and H. grandi flora). Of the other
two, Epilobium alpinum (sensu lato) is a
widespread circumpolar arctic-alpine species
and Agropyron caninum is a North American
boreal-montane species.

Using information derived from Fryxell
(1957), Swales (1979), and Ostler and Harper
(1978), the species from the transects can be
divided into two groups, biotically and
abiotically pollinated species. Wind-polli-
nated, suspected autogamous and apomictic
species (i.e., Arabis lyallii. Taraxacum offici-
nale, and Epilobium alpinum), and the fern
ally Selaginella densa, which requires water
for fertilization, are included in the abioti-
cally pollinated group. Biotically pollinated
(primarily entomophilous) dicot species com-
prise the biotically pollinated group. Average
number of abiotically pollinated species per
quadrat on the moraine is 1.0, in the meadow
1.4. These are not significantly different at a
= 0.05. Average number of biotically polli-
nated species per quadrat on the moraine is
1.2, in the meadow 3.1. These are significant-
ly different at a = 0.05 (using a t-test for un-
equal sample sizes). These and other com-
parisons between these two groups of species
are summarized in Table 2. The total

Table 2. Comparisons between the moraine and the
meadow using the quadrat data derived from the hne
transects. Two groups of species, abiotically pollinated
(the abiotic group) and biotically pollinated (biotic
group) are compared for the two areas. See the text for
details on the two groups of species. The sample size (N)
is the number of quadrats.

Moraine (N = 38) Meadow (N = 22)

Average percent cover
Abiotic

2.5

10.7

Biotic

3.0

44.5

Average number of species
per quadrat
Abiotic

1.0

1.4

Biotic

1.2

3.1

2) Prominance values (PV)

Abiotic

11.0

54.9

Biotic

6.4

343.1

Number of species

Abiotic

Biotic

Total number of species

Abiotic

Biotic

summed PV for all the moraine species in the
abiotically and biotically pollinated groups
are 11.0 and 6.4, respectively. For the mead-
ow they are 54.9 and 343.9, respectively. The
ratio of PV for the abiotically pollinated spe-
cies in the meadow and the moraine is 5:1,
and that for the biotically pollinated species
is 53:1. The total number of species on the
moraine and in the meadow for the abioti-
cally pollinated group are 8 and 9, respec-
tively. For the biotically pollinated group the
values for the moraine and meadow are 13
and 14, respectively. All these comparisons
suggest that the species in the abiotically pol-
linated group are relatively more succesful at
colonizing and establishing on the moraine
deposits than the species in the biotically pol-
linated group. The reverse is true for the
meadow, where the biotically pollinated spe-
cies dominate. There could be several reasons
for these differences. Perhaps the open na-
ture of the vegetation on the moraine makes
it more difficult for biotically pollinated spe-
cies to attract pollinators. Thus species that
are autogamous, apomictic, or anemophilous
may be at a reproductive advantage. It is also
possible that such breeding systems are
linked with other traits that confer greater
colonizing abilities than is found in the bioti-
cally pollinated species (Jain 1976). Differen-
tial dispersion of propagules onto the mo-
raine by species in the two categories does
not appear to be the reason (see below).

Turning to the propagule trapping results,
the water trays caught only five propagules
during several weeks of operation. The water
in the trays tended to evaporate quickly, and
many propagules were probably blown out of
the trays once they had dried out. These five
propagules and those trapped by the petro-
leum-jelly-coated wooden plates used in 1979
are listed in Table 3. No propagules were
trapped by the wooden plates in July, pre-
sumably because the plants were still flower-
ing and had yet to set seed. In September,
the trap furthest out into the meadow (30 m)
trapped 23 propagules over a 24-hour period.
The only other trap that caught anything was
placed on the distal moraine slope in about
midslope position. This trap caught a single
unidentifiable composite achene with a pap-
pus. Extrapolating from the data on number
of propagules trapped and the size of the

July 1983

Spence, Shaw: Schoolroom Glacier Plants

489

plates, the 23 propagules trapped in the
meadow represent about 1022 propagules
dispersing into and through every 1 m^ every
24 hours, at a time in September when most
species had finished flowering and were dis-
persing propagules. The calculated value for
the trap on the moraine is 45 propagules per
1 m^ per 24 hours, assuming that the single
propagule trapped is representative of the
propagule rain on the deposits. Most of the
propagules (90 percent) have some mor-
phological feature that might aid in more ef-
ficient wind dispersal. Such features include
the coma of the Salix and Epilobium seeds
and the pappus of the composite achenes.
Two propagules, the Carex achene and the
Astragalus kentrophyta legume, have no ap-
parent features that could enhance wind dis-
persal ability. The propagule of Geum rossii
represents a special case. This species is the
only one that did not occur in the vicinity of
the moraine or meadow. The propagule con-
sists of a persistent calyx with numerous stiff
hairs, enclosing many small achene fruits. It
was found in one of the water trays in 1978,
which raises the possibility that the prop-
agule could have accidently dropped into the
water from the fur of an investigating pika.
Pikas are known to collect plants of Geum
rossii for their hay piles (Johnson 1967). All
the propagules trapped consist of dicot spe-
cies except for the achene of the Carex
species.

The moraine deposits appear to be in an
active state of collapse. Fresh slumps and old
slump scars can be found on both slopes, and
are especially common on the proximal
slope. The deposits are very loose and tend to
shift easily underfoot. Numerous small ero-
sion channels (rills) exist, attesting to the ef-
fects of snow meltwater erosion. The steep-
ness of the slopes, which are from 36° to 38°
on the distal slope and 35° to 41° on the pro-
ximal, contributes to the instability of the
moraine. The combination of continual dis-
turbance and apparently low rates of dis-
persal of propagules from the meadow is the
probable explanation for the low average
vegetation cover on the moraine slopes. The
only portions of the moraine where the plant
cover is as dense as in the meadow is along
the crest, which in places is flat and presum-
ably more stable than the steep slopes. The

patches of vegetation on the flat portions of
the crest are very similar to the meadow veg-
etation, including the presence of the three
most common meadow species. Astragalus
kentrophyta, Hymenoxys grandiflora, and An-
tennaria umbrinella. This suggests that, as the
moraine deposits stabilize, they will become
vegetated by the species that dominate and
characterize the meadow.

A perusal of alpine vegetation literature
from the south central Rocky Mountains
failed to reveal any reports of vegetation sim-
ilar to that found in the meadow (Rydberg
1914, Cox 1933, Cain 1943, Hayward 1952,
Ellison 1954, Langenheim 1962, Johnson and
Billings 1962, Bamberg and Major 1968, Ha-
beck 1969, Bonham and Ward 1970, Lewis
1970, Anderson et al. 1979, Komarkova
1979), although many of the species in the
meadow are common and widespread in the
Rocky Mountains. On the other hand, several
reports list vegetation that is strongly similar
to that of the moraine (Buttars 1914, Ma-
haney 1974, Given and Soper 1975). In a de-
tailed study from the Colorado Front Range,
Komarkova (1979) listed several species that
are characteristic of scree, talus, and glacial
deposits (her Order Aquilegio-Cirsietalia sco-
pulorum). Many of these species are also
found at the Schoolroom Glacier moraine.
These include Senecio fremontii, Poa patter-
sonii, Oxyria digyna, Draba lonchocarpa,
Epilobiujn alpinum. Taraxacum officinale,
and Trisetum spicatum. The thistle Cirsium

Table 3. The identity of the propagules trapped dur-
ing 1978 and 1979 are hsted, along with the number
caught, and the presence of any morphological feature
that might aid in more efficient wind dispersal.

Number

Morphological

Species

trapped

feature

Epilobium alpinum

(seeds)

coma

Salix arctica (seeds)

coma

Arnica longifolia

(achenes)

pappus

Geum rossii (achenes

enclosed in calyx)

hairy calyx

Solidago multiradiata

(achene)

pappus

Astragalus kentrophyta

(legume)

—

Carex species (achene)

—

Unknown composites

(achenes)

pappus

490

Great Basin Naturalist

Vol. 43, No. 3

scopulonim appears to play an ecological
role similar to Cirsium tweedyi in the Tetons.
In summary, the vegetation on the School-
room Glacier moraine is very open, with av-
erage cover ranging from less than 1 percent
up to 9 percent, compared with over 50 per-
cent in the alpine dry meadow adjacent to it.
Dominance-diversity curves using promi-
nence values as the measure of importance
were constructed for the meadow and the
moraine. Both curves approach geometric
series, suggesting harsh environments in
which dominance by one or a few species is
strong. The legume Astragalus kentrophyta
dominates the meadow, and the thistle Cir-
sium tweedyi dominates the moraine. Using
cover and frequency data, abiotically polli-
nated species are relatively much more suc-
cessful on the moraine than biotically polli-
nated species. The reverse is true in the
meadow. Propagule trapping studies suggest
that wind dispersal onto the moraine is very
low compared with wind dispersal within the
meadow. Distributionally, three groups of
species can be discerned. One group is con-
centrated primarily on the moraine, only
rarely straying into the meadow. The second
group is found primarily in the meadow. The
third group consists of species that are found
in a narrow band between the base of the dis-
tal moraine slope and the meadow. This band
of vegetation has many of the characteristic
of an ecotone. The moraine deposits are
highly unstable, and, in the few places along
the crest of the moraine that tend to be the
most stable, the vegetation is strongly similar
to the vegetation in the meadow.

Acknowledgments

This paper represents partial requirements
for a M.Sc. thesis by JRS, under the super-
vision of RJS. We thank the National Park
Service, particularly Linda Olson and Bob
Wood, for collecting permits and for other
help.

Literature Cited

Anderson, D. C, R. S. Hoffman, and K. B. Armitage.
1979. Aboveground productivity and floristic
structure of a high subalpine herbaceous mead-
ow. Arctic and Alpine Res. 11:467-476.

Bamberg, S. A., and J. Major. 1968. Ecology of the veg-
etation and soils associated with calcareous par-
ent materials in three alpine regions of Montana.
Ecol. Monogr. .38:127-167.

BoNHAM, C. D., AND R. T. Ward. 1970. Phytosociolog-
ical relationships in alpine tufted hairgrass (Des-
champsia caespitosa [L.] Beauv.) meadows. Arctic
and Alpine Res. 2:267-275.

BuTTARs, F. K. 1914. Some peculiar cases of plant distri-
bution in the Selkirk Mountains, British Colum-
bia. Minnesota Bot. Studies 4:313-331.

Cain, S. A. 1943. Sample-plot technique applied to al-
pine vegetation in Wyoming. Amer. J. Bot.
30:240-247.

Cole, F. 1975. Introduction to meterology. 2d ed. Wiley
& Sons, New York.

Cox, C. F. 1933. Alpine plant succession on James Peak,
Colorado. Ecol. Monogr. 3:300-372.

Ellison, L. 1954. Subalpine vegetation of the Wasatch
Plateau, Utah. Ecol. Monogr. 24:89-184.

Fryxell, p. a. 1957. Mode of reproduction of higher
plants. Botanical Rev. 23:135-233.

Given, D. R., and J. H. Soper. 1975. Pioneer vegetation
on moraines near Clachnacudainn Snowfield,
British Columbia. Syesis 8:349-354.

Habeck, J. R. 1969. A gradient analysis of a timberline
zone at Logan Pass, Glacier Park, Montana.
Northwest Sci. 43:65-73.

Hayward, C. L. 1952. Alpine biotic communities of the
Uinta Mountains, Utah. Ecol. Monogr.
22:93-120.

Jain, S. K. 1976. The evolution of inbreeding in plants.
Ann. Rev. Ecol. Syst. 7:469-495.

Johnson, D. R. 1967. Diet and reproduction of Colorado
pikas. J. Mammol. 48:311-315.

Johnson, P. L., and W. D. Billings. 1962. The alpine
vegetation of the Beartooth Plateau in relation to
cryopedogenic processes and patterns. Ecol.
Monogr. 32:105-135.

Komarkova, v. 1979. Alpine vegetation of the Indian
Peaks Area, Front Range, Colorado Rocky Moun-
tains. Flora et Vegetatio Mundi, 7. R. Tiixen, ed.
Cramer, Vaduz.

Langenheim, J. H. 1962. Vegetation and environmental
patterns in the Crested Butte area, Gunnison
County, Colorado. Ecol. Monogr. 32:249-285.

Lewis, M. E. 1970. Alpine rangelands of the Uinta
Mountains. U.S. Forest Service— Region 4.

Love, J. D., and J. C. Reed. 1968. Creation of the Teton
Landscape. Grand Teton Natural Hist. Assoc.
Press.

Mahaney, W. C. 1974. Soil stratigraphy and genesis of
Neoglacial deposits in the Arapaho and Hender-
son cirques, central Colorado Front Range.
Geogr. Monogr. 5:197-240.

1975. Soils of post-Audubon age, Teton Glacier

area, Wyoming. Arctic and Alpine Res.
7:141-153.

Mahaney, W. C, and J. R. Spence. 1983. Late Qua-
ternary deposits, soils, chronology, and floristics.
Jaw Cirque area, Central Teton Range, Wyom-
ing. In preparation.

Ostler, W. K., and K. T. Harper. 1978. Floral ecology
in relation to plant species diversity in the
Wasatch Mountains of Utah. Ecology
59:848-861.

July 1983

Spence, Shaw: Schoolroom Glacier Plants

491

Reed, J. C. 1973. Geological Map of Grand Teton Na-
tional Park. U.S. Geol. Surv.

Rydberg, p. a. 1914. Phytogeographical notes on the
Rocky Mountain region III. Formations in the al-
pine zone. Bull. Torrey Bot. Club 41:459-474.

Shaw, R. J. 1976. Field guide to the vascular plants of
Grand Teton National Park and Teton County.
Utah State Univ. Press, Logan.

Spence, J. R. 1981. Comments on the cryptogam vegeta-
tion in front of glaciers in the Teton Range. Bryo-
logist 84:564-568.

1983. A floristic analysis of the vegetation on

Neoglacial deposits in the Teton Range, Wyom-
ing. Submitted to Arctic and Alpine Res.

Swales, D. E. 1979. Nectaries of certain arctic and sub-
arctic plants with notes on pollination. Rhodora
81:367-407.

Whittaker, R. H. 1975. Communities and Ecosystems.
2d ed. Macmillan, New York.

WINTER STOMACH CONTENTS OF SOUTH DAKOTA BADGERS

E. Blake Hart' and Michael Truinbo-

Abstract.— Stomach contents of 33 hunter and trapper killed badgers, Taxidea taxtis, from northeastern South
Dakota were examined during the 1980-81 fur season. Mammal prey species made up the bulk of the ingested food,
followed by plant materials. Approximately 40 percent of stomach contents were mammals, 35 percent were of plant
origin, 10 percent were birds, 10 percent were insects, while the remainder was mostly inorganic materials. Little
significant difference was found in variety of food items consumed by each sex. South Dakota badgers are opportu-
nistic foragers in the wintertime when food is scarce. Individual food items are usually taken in quantity when en-
countered by badgers.

The severity of typical northern plains
winters fixes food as a priority item in the
lives of nonhibernating carnivorous mam-
mals. Answers to the question as to varieties
of prey consumed was sought by examination
of a ubiquitous carnivore, the American
badger, Taxidea taxus.

Stomachs of 33 badgers (15 male, 18 fe-
male) from the northeastern South Dakota
counties of Faulk, Brown, Spink, and Ed-
munds were obtained from a local fur broker
during the 1980-1981 season. Several keys
were used to identify prey animal hairs, par-
ticularly Moore, Spence, and Dugnole (1974).
Cuticular scale patterns of hairs were pressed
into a film of nail polish and observed (Wein-
gart 1973).

A variety of materials were found in stom-
achs, including plants, inorganic materials
(soil, stones), insects, and prey animals. Stom-
achs contained an average of 98 grams of
food materials each. From the standpoint of
relative volume and variety of identifiable re-
mains, 40 percent were mammal prey ani-
mals, 35 percent were of plant origin, 10 per-
cent were birds, 10 percent were insects, and
the remainder were mostly undigested in-
organic materials. Similar foods were found
in stomachs of both sexes with the exception
that straw and small stones were not found in
female stomachs.

At least one-half of all badger carcasses
purchased by the fur dealer had empty stom-
achs. This suggests that either stomach con-

tents had largely been digested during long
trap stays or that many badgers spend consid-
erable time with empty stomachs.

Most of the food matter contained within
the stomachs had been thoroughly crushed
and mascerated; the largest bone fragment
was 1.7 X 0.8 cm. In fact, skull remnants of
mammal prey were difficult to identify. In
several instances portions of hair-covered
epidermis of larger prey were present; these
were the largest pieces of food material seen,
although occasional intact mouse viscera and
other assorted prey extremities were
observed.

Sunflower seeds, corn, and millet were
present in several stomachs in considerable
quantities, though these too had mostly been
chewed into a paste, with few kernels intact.
We observed what possibly may have been
several masses of feces. The majority of stom-
achs contained from 5 to 20 Ascaris-\ike
parasites.

We found, as did Jense (1968), who studied
badger food habits and energy utilization in
east central South Dakota, that badgers are
opportunistic foragers of edible plant and an-
imal materials. Inasmuch as they feed on
whatever is at hand, variety is often lacking,
especially if the prey is a large animal or is
plant material. One markedly distended
stomach contained 375 grams of plant and
animal materials. Full stomachs contained
little variety; rather, they were often replete
with a single food type.

'Department of Mathematics, Natural Sciences and Health Professions, Northern State College, Aberdeen, South Dakota 57401.
'Tulare, South Dakota 57476.

492

July 1983

Hart, Trumbo: Badger Diet

493

Jense (1968) suggested that grains were im-
portant fall foods, that birds and eggs were
eaten only during the spring and summer.
We found that squirrels, mice, rabbits, and
other small mammals formed much of the
typical winter badger diet, but also evidence
was foimd of birds, insects, and grains. Snead
and Hendrickson (1942) found many of the
same food items in diets of Iowa badgers;
they found that percentages of kinds of foods
taken changed as availability changed with
the seasons.

Hibernating prey animals appear to be at-
tractive food sources throughout the winter,
especially ground squirrels. During the mid-
winter 1978-1979 fur season, 17 toads were
found in a single badger stomach. Rosen-
weig's (1966) statement that there is a clear
trend for larger predators to seek larger prey
appears true enough, although, judging from
the quantities of mice in stomachs, badgers
obviously do not ignore such small mammals
as important auxiliary food sources.

In summary, winter diets of South Dakota
badgers vary with the prey species, which
represent most nonaquatic vertebrate groups,
as well as insects. Opportunistic feeding ap-
pears to be common among badgers, which is
not unexpected considering sparse food re-
sources during the severe cold of typical
northern plains winters.

Literature Cited

Jense, G. K. 1968. Food habits and energy utilization of

badgers. Unpublished thesis. South Dakota State

Univ., Brookings. 39 pp.
Moore, T. D., L. E. Spence, and C. E. Dugnole. 1974.

Identification of the dorsal gviard hairs of some

mammals of Wyoming. Wyoming Fish and Game

Dept. Bull. 14. 177 pp.
Rosenweig, M. L. 1966. Community structure in sym-

patric carnivora. J. Mammal. 47:602-612.
Snead, F., and G. O. Hendrickson. 1942. Food habits of

the badgers fn Iowa. J. Mammal. 23:380-391.
Weingart, E. L. 1973. A simple technique for revealing

hair scale patterns. Amer. Midi. Nat.

90(2):508-509.

A LIST OF UTAH SPIDERS, WITH THEIR LOCALITIES

Dorald M. Allred' and B. J. Kaston-

Abstract. — The 621 species of spiders known to occnr in Utah as recorded in the Hterature or Utah universities'
collections are listed with their junior synonyms and collection localities. Two-fifths (265 species) are known from
onlv one locality each, and only one-fifth (123 species) from five or more localities in the state.

Little is known of the distribution or eco-
logical relationships of Utah spiders. Each of
265 species of the 621 recorded for the State
is known from only one locality. Even the
ubiquitous black widow, Latrodectus hes-
perus, has been recorded from only 24 sites in
Utah. Most collections from the 249 local-
ities, listed primarily from the literature and
some unpublished data, are from a few places
that seem to have been favorite or conven-
ient collecting areas for early naturalists and
students of arachnology. For example, 166
species are recorded from Box Elder County
in the northwest comer of Utah, principally
in the Raft River Mountains, which apparent-
ly was a favorite place of Wilton Ivie, a con-
temporary and associate of Ralph Chamber-
lin and Willis Gertsch. All three men were
associated with the University of Utah in Salt
Lake City, and 166 species are listed from
that environs. Two naturalists and avid col-
lectors who were contemporaries and associ-
ates of Chamberlin were Vasco Tanner and
Angus Woodbury. Both had family ties in St.
George in Washington County in the south-
west comer of the state and lived and fre-
quently revisited there for many years. One
hundred species are listed for St. George, and
80 from nearby Zion National Park, where
Woodbury was employed as a naturalist for
several years. Chamberlin frequently visited
Tanner in St. George, and collected in that
area. Such large numbers of species recorded
from these localities is indicative of the lack
of study done in other areas of the state, fur-
ther exemplified by the fact that each of only
123 species is known from five or more
localities.

Much of our knowledge of Utah spiders
was contributed by Ralph Chamberlin, who
authored or coauthored the naming of 220 of
the species listed for Utah. Wilton Ivie and
Willis Gertsch authored 106 and 73 species,
respectively, some in coauthorship with
Chamberlin. Stanley Mulaik, arthropodolo-
gist, naturalist, and avid collector, was also
contemporary with these men and coau-
thored several species with Gertsch.

We are indebted to Willis Gertsch, who
provided some unpublished records and valu-
able criticism of the manuscript. James Mac-
Mahon of Utah State University also provid-
ed unpublished records of specimens
collected by him and his students. Anne
Bond, research assistant, initially helped with
much of the literature search.

Sources of collection records are desig-
nated by initials and dates in parentheses im-
mediately following specific, or a group of,
localities. A key to these is given below.
Where more than one publication is repre-
sented by an author's initials, the specific
source is indicated by a date.

Key to Locality Sources

(Published articles except as indicated)

= Allred

= Archer

= Allred & Gertsch

= Allred (unpublished)

= Beck (unpublished)

= Beedlow & Abraham

= Brady

= Bowling & Sauer

= Chamberlin

= Chamberlin & Gertsch

= Chamberlin & Ivie

'Life Science Museum and Department of Zoology, Brigham Young University, Provo, Utah 84602.
'Department of Zoology, San Diego State University, San Diego, California 92182.

494

April 1983

Allred, Kaston: Utah Spiders

495

= Chamberlin & Woodbury

= Dondale

= Dondale & Redner

= Exline

= Gertsch

= Gertsch & Ivie

= Gertsch & Russell

= Gertsch (unpublished)

= Gertsch & Wallace

= Hatley

= Ivie

= Ivie (unpublished)

= Knowlton

= Kaston

KsU

= Kaston (unpublished)

= Keyserling

= Levi

= Lowrie & Dondale

= Leech

= Levi & Levi

= Levi & Randolph

= Millidge

McU

= MacMahon (unpublished)

= Munia & Gertsch

= Platnick

= Platnick & Shadab

= Roewer

= Schick

= Sauer & Platnick

= Van Helsdingen

= Waagen

= Wallace & Exline

List of Utah Collection Localities
with County Designation

Allen Cvn (Rich)

Alta (Salt Lake)

American Fork Cyn (Utah)

Aquarius Plateau (Garfield)

Aspen Grove (Utah)

Bear Lake (Rich)

Beaver (Beaver)

Beaver Crk (in 12 counties)

Beaver Cyn (Beaver)

Beaver Dam Mts (Washington)

Beaver Dam Wash (Washington)

Benson (Cache)

Bicknell (Wayne)

Big Cottonwood Cyn (Salt Lake)

Big Indian Rock (San Juan)

Bills Cyn (Salt Lake)

Black Rock (Salt Lake)

Blanding (San Juan)

Blue Sprvice Cyn (Garfield)

Bluff (San Juan)

Boulder Mt (Garfield)

Bountiful (Davis)

Brigham (Box Elder)

Brigham Cyn (Box Elder)

Brigham Plains Flat (Kane)

Bryce Cyn Nat Park (Garfield)

Butterfield Cyn (Salt Lake)

Caineville (Wayne)

Calf Crk (Garfield)

Carter Crk (Daggett)

Carter & Deep Crk Jet (Daggett)

Castle Cliffs (Washington)

Castle Dale (Emery)

Castle Park (?)

Cedar Cyn (Iron)

Cedar Hills (Box Elder)

Cedar Jet (Box Elder)

Cedar Mt (Juab)

Cedar Mts (in Uinta Mts — Summit?)

Chalk Crk (Summit)

City Crk Cyn (Salt Lake)

Clarkston (Cache)

Clear Crk (Box Elder)

Clear Crk Cyn (Box Elder)

Clear Lake (Millard)

Cobble Rest (Wasatch)

Cove Fort (Millard)

Coyote Gulch (Garfield)

Curlew Valley (Box Elder)

Cyclone Lake (Garfield)

Daniels (Wasatch)

Deep Crk (Daggett)

Delta (Millard) *

Devils Cyn (San Juan)

Diamond Valley (Washington)

Dinosaur Nat Mon (Uintah)

Dixie Nat Forest (Iron)

Dove Crk (Box Elder)

Dry Cyn (Salt Lake)

Duchesne (Duchesne)

East Cyn (Salt Lake)

Egg Island (Tooele)

Elk Ridge (San Juan)

Elsinore (Sevier)

Emery (Emery)

Emigration Cyn (Salt Lake)

Emory (Summit)

Ensign Peak (Salt Lake)

Escalante (Garfield)

Eureka Cyn (Utah)

Farmington (Davis)

Farmington Cyn (Davis)

Ferron (Emery)

Ferron Res (Sanpete)

Fillmore (Millard)

Fillmore Cyn (Millard)

Fish Lake (Sevier)

Fisher Pass (Tooele)

Four-mile Bench (Kane)

Franklin Basin (Cache)

Fremont River (Wayne)

Fruita (Wayne)

Ft Douglas (Salt Lake)

Garden City (Rich)

Glen Cyn City (Kane)

Glenwood (Sevier)

Government Crk (Tooele)

Grand Gulch (San Juan)

Grandaddy Lake (Duchesne)

Granite (Salt Lake)

Grantsville (Tooele)

Green Cyn (Cache)

Green Lake (Daggett)

496

Great Basin Naturalist

Vol. 43, No. 3

Greenriver (Emery)

Grouse Crk (Box Elder)

Gunnison Butte (Emery)

Hanksville (Wayne)

Hat Island (Tooele)

Hatch (Garfield)

Heber (Wasatch)

Helper (Carbon)

Henry Mts (Garfield)

Hidden Lake (Kane & Summit)

Holliday (Salt Lake)

Horse Valley (Wayne)

Hughes Cyn (Salt Lake)

Hurricane (Washington)

Junction (Piute)

Kaibab Forest (Kane)

Kanab (Kane)

Kems (Salt Lake)

Kelton (Box Elder)

Lake Powell (Kane)

Laketown (Rich)

Lambs Cyn (Salt Lake)

La Sal Jet (San Juan)

La Sal Mts (Grand)

La Sal Pass (San Juan)

Layton (Davis)

Lehi (Utah)

Leidy Peak (Uintah)

Levan (Juab)

Liberty (Weber)

Little Cottonwood Cyn (Salt Lake)

Loa (Wayne)

Locomotive Spngs (Box Elder)

Logan (Cache)

Logan Cyn (Cache)

Lynn (Box Elder)

Lynndyl (Millard)

Manila (Daggett)

Marysvale (Piute)

Marysvale Cyn (Piute)

Mill Crk (Summit)

Mill Crk Cyn (Salt Lake)

Mirror Lake (Duchesne)

Moab (Grand)

Monroe Cyn (Sevier)

Monticello (San Juan)

Moroni (Sanpete)

Motauqua (Washington)

Mounds (Emery)

Mt Agassiz (Summit)

Mt Ellen (Garfield)

Mt Nebo (Juab)

Mud Spngs (Emery)

Navajo Mt (San Juan)

Nipple Bench (Kane)

Noton (Wayne)

Ogden (Weber)

Ogden Cyn (Weber)

Ophir (Tooele)

Oquirrh Mts (Tooele)

Orton (Garfield)

Ouray (Uintah)

Pangiiitch (Garfield)

Paradise (Cache)

Park Valley (Box Elder)

Parleys Cyn (Salt Lake)

Parowan (Iron)

Payson (Utah)

Pickleville (Rich)

Pinecrest (Salt Lake)

Pine Cyn (Millard)

Pine Spngs (Garfield)

Pine Valley (Washington)

Pine Valley Mts (Washington)

Pink Sand Dunes (Kane)

Pintura (Washington)

Plain City (Weber)

Posey Lake (Garfield)

Price (Carbon)

Promontory Point (Box Elder)

Provo (Utaii)

Provo River, Upper (Duchesne)

Puffer Lake (Beaver)

Raft River (Box Elder)

Raft River Mts (Box Elder)

Red Butte Cyn (Salt lake)

Red Cyn Camp (Garfield)

Richardson (Grand)

Richfield (Sevier)

Rock Island (Utah)

Rotary Park (Salt Lake)

Salina (Sevier)

Saltair Beach (Salt Lake)

Salt Lake Airport (Salt Lake)

Salt Lake City (Salt Lake)

San Rafael (Emery)

San Rafael River (Emery)

Santa Clara (Washington)

Santaquin (Utah)

Santaquin Res (Utah)

Scipio (Millard)

Silver Lake (Salt Lake & Utah)

Smith & Morehouse Cyn (Summit)

Smokey Mt (Kane)

Snow Crk (Sevier)

Snow Crk Cyn (Sevier)

Snowville (Box Elder)

Spring Crk (Carbon)

Spring Cyn (Carbon)

Spring Lake (Utah)

St John (Tooele)

St George (Washington)

Standardville (Carbon)

Steep Crk (?)

Straight Wash (Emery)

Stockton (Tooele)

Strawberry (Wasatch)

Swan Lake (Rich)

Table Cliff Pass (Garfield)

Table Cliff Plateau (Garfield)

Teapot Lake (Summit)

Terrys Ranch (Washington)

Thompsons (Grand)

Three Lakes (Garfield & Kane)

Tibbet Spng (Kane)

Timpanogos Cave Nat Mon (Utah)

Timpanogos Park (Utali)

Tooele Cyn (Tooele)

Torrey (Wayne)

Tremonton (Box Elder)

April 1983

Allred, Kaston: Utah Spiders

497

Tropic (Garfield)

Tropic Res (Garfield)

Trout Crk City (Juab)

Uinta Mts (in 5 counties)

Utah Lake (Utah)

Utah State Univ School Forest (Rich)

Valley City (San Juan)

Verdure (San Juan)

Vermillion Castle (Iron)

Vernal (Uintah)

Vernon (Tooele)

Wah Wah Mts (Beaver)

Wanship (Summit)

Wasatch (Salt Lake & Summit)

Wasatch Mts (Salt Lake)

Wasatch Plateau (in 6 counties)

White River (Uintah)

Widtsoe (Garfield)

WiUard (Box Elder)

Willow Crk (Tooele)

Willow Tank Spngs (Kane)

Yost (Box Elder)

Zion Nat Park (Washington)

Species and Localities

If the name associated with a specimen in
a collection or a record in the literature is
not foimd in this list of species, consult the
list of "Synonymies of Utah Records" in the
latter part of this report.

Adiaearanea ambera Levi 1963 (Bull. Mus.
Comp. Zool. 128:204). Mill Crk Cyn (L63).

Achaearanea canionis (Chamberlin &
Gertsch) 1928 (J. Ent. Zool. Pomona Coll.
21:103). American Fk Cyn, Pinecrest (lU),
Beaver Cyn, Cobble Rest, Dinosaur Nat Mon,
Dry Cyn, Richfield, Salt Lake City, St
George (L55), Zion Nat Park (CG28).

Achaearanea tepidarioritm (C.L. Koch)
1841 (Die Arachn. 8:75). HoUiday, Provo,
Salt Lake City (L55).

Actinoxia sp. Glen Cyn City (AU).

Aculepeira carbonaria (L. Koch) 1869
(Zeits. Ferd. Tirol. Voral. 15:58). St George
(CW).

Aculepeira packardi (Thorell) 1875 (Kongl.
Svenska Vet. Akad. Handl. 13:3-203). Green
Cyn (H), Kelton Pass (K), USU School Forest
(W).

Agelenopsis aperta (Gertsch) 1934 (Amer.
Mus. Novitates 726:24). Aspen Grove (BU),
Motauqua (CI41), Salt Lake City, Trout Crk,
Zion Nat Park (G34c), St George (CW).

Agelenopsis californica (Banks) 1896 (J.
New York Ent. Soc. 4:88). Dove Crk, Grouse
Crk (City), Lynn, Raft River S fk, Yost
(CI33), St George (CW).

Agelenopsis Oklahoma (Gertsch) 1936
(Amer. Mus. Novitates 852:12). Duchesne
(CI41).

Agelenopsis iitahana (Chamberlin & Ivie)
1933 (Bull. Univ. Utah Biol. Ser. 2(2):43).
Clear Crk (CI33a), Fish Lake, La Sal Mts,
Wasatch Mts (CI41).

Agelenopsis sp. Nipple Bench (AU).

Agroeca ornata Banks 1892 (Proc. Acad.
Nat. Sci. Philadelphia 44:23). Clear Crk, Raft
River S fk (CI33).

Agroeca pratensis Emerton 1890 (Trans.
Conn. Acad. Sci. 8:155). Raft River S
fk(CI33).

Agroeca trivittata (Keyserling) 1887 (Verb,
zool. bot. Ges. Wien 37:444). Fillmore (C19),
Moab (CG28), St George (CW).

Alopecosa gulosa (Walckenaer) 1837 (Hist.
Nat. Ins. Apt., 1:38). Clear Crk, Yost (CI33),
La Sal Jet (CG28), St George, Zion Nat Park
(CW).

Alopecosa kochi (Keyserling) 1877 (Verb,
zool. bot. Ges. Wien 26:636). Green Cyn (H),
Posey Lake, Steep Crk, Table Cliff Pass,
Three Lakes (N Kanab) (BU), USU School
Forest (W).

Amaurobius americanus (Emerton) 1888
(Trans. Conn. Acad. Sci. 7:443). Bluff, Fruita,
Moab, Monticello, Mounds, San Rafael River,
Verdure (CI28), Clear Crk (in Raft River
Mts), Dove Crk, Grouse Crk, Lynn (CI33),
Lake Powell (CW).

Anacornia microps Chamberlin & Ivie
1933 (Bull. Univ. Utah Biol. Ser. 2(2):29).
Clear Crk, Raft River S fk, Uinta Mts (CI33).

Anacornia proceps Chamberlin 1948 (Ann.
Ent. Soc. Amer. 41:483). Chalk Crk, Cobble
Rest, Mirror Lake (C48a), USU School Forest
(W).

Antrodiaetns hageni (Chamberlin) 1917
(Bull. Mus. Comp. Zool. 61:74). Raft River S
fk (CI33).

Antrodiaetns montanus (Chamberlin &
Ivie) 1935 (Bull. Univ. Utah Biol. Ser. 2(8):4).
Green Cyn (H), USU School Forest (W).

Anyphaena californica (Banks) 1904 (Proc.
Calif. Acad. Sci. 3:338). Mill Crk (C20b).

Anyphaena pacifica (Banks) 1896 (Trans.
Amer. Ent. Soc. 23:63). Green Cyn (H), Ver-
dure (CG28).

Anyphaena sp. Clear Crk, Dove Crk, Raft
River S fk (CI33), Escalante, Three Lakes (N
Kanab) (BU).

498

Great Basin Naturalist

Vol. 43, No. 3

Aphonopelma angusi Chamberlin 1940
(Bull. Univ. Utah Biol. Ser. 5(8):21). W of
Beaver Dam Mts (C40).

Aplwnopelma simulatiim (Chamberlin &
Ivie) 1939 (Bull. Univ. Utah Biol. Ser. 5(1):8).
Fruita (CI39a).

Aphonopelma zionis Chamberlin 1940
(Bull. Univ. Utah Biol. Ser. 5(8) :24). Zion Nat
Park (C40).

ApollopJianes texanus Banks 1904 (J. New
York Ent. Soc. 12:113). Blanding, Bluff,
Moab, Verdure (CG28), Glen Cyn City 5 km
W (AG).

Aranetis gemma (MacCook) 1888 (Proc.
Acad. Nat. Sci. Philadelphia 50:193). Clear
Crk, Lynn, Park Valley (in Raft River Mts)
(CI33), Ferron, Salt Lake City (CI35b),
Green Cyn (H), Locomotive Spngs (K), Pro-
vo, Three Lakes (nr Escalante) (BU), Zion
Nat Park (CW).

Araneus mammatus (Archer) 1951 (Amer.
Mus. Novitates 1487:17). Mill Crk Cyn
(L81a).

Araneus marmoreus Clerck 1757 (Aranei
Suecici, p. 29). Chalk Crk (in Uinta Mts)
(C19).

Araneus nordmanni (Thorell) 1870 (Rem.
Syn. Europ. Spid., p. 4). USU School Forest
(W).

Araneus pima Levi 1971 (Bull. Mus.
Comp. Zool. 141:176). "Utah" (L71).

Araneus saevus (L. Koch) 1872 (Zeits.
Ferd. Tirol Voralberg (3) 17:323). Salt Lake
City (Ac).

Araneus trifolium (Hentz) 1847 (J. Boston
Soc. Nat. Hist. 5:471). Provo (BU).

Araneus varians Thorell 1899 (Bih.
Svenska Vet. Akad. Handl. 25:49). Zion Nat
Park (CW).

Araneus sp. Coyote Gulch (BU).

Araniella displicata (Hentz) 1847 (J. Bos-
ton Soc. Nat. Hist. 5:476). Aspen Grove (BU),
Clear Crk (CI33), Green Cyn (H), USU
School Forest (W).

Araniella octopunctata Chamberlin & Ivie
1942 (Bull. Univ. Utah Biol. Ser. 7(1) :76).
Emigration Cyn, Pine Vallev, Zion Nat Park
(CI42a), Steep Crk (BU).

Arctosa alpigena (Doleschall) 1852 (Sitz.-
ber. Akad. Wiss. Wien, 9:643). USU School
Forest (W).

Arctosa chamberlini Gertsch 1934 (Amer.
Mus. Novitates 693:4). Vernal (G34).

Arctosa littoralis (Hentz) 1844 (J. Boston
Soc. Nat. Hist. 4:388). Calf Crk, Coyote
Gulch (in Escalante Basin), La Sal Mts, Rock
Island (in Utah Lake), Steep Crk, Three
Lakes (nr Kanab), Willow Tank Spngs (BU),
Grand Gulch, Utah Lake (G34), Greenriver,
Moab (CG28), St George (CW).

Arctosa parva (Banks) 1894 (J. New York
Ent. Soc. 2:52). "Utah" (C8).

Arctosa rubicunda (Keyserling) 1877 (Verh.
zool. bot. Ges. Wien 26:663). Raft River S fk
(CI33).

Arcuphantes decoratus Chamberlin & Ivie
1943 (Bull. Univ. Utah Biol. Ser. 7(6): 17).
Zion Cyn (Nat Park) (CI43).

Arcuphantes fragilis Chamberlin & Ivie
1943 (Bull. Univ. Utah Biol. Ser. 7(6): 17).
American Fork Cyn, Timpanogos Nat Mon
(CI43).

Argenna obesa Emerton 1911 (Trans.
Conn. Acad. Sci. 16:399). Utah Lake W shore
(CI35b), Zion Nat Park (CG58).

Argenna saphes Chamberlin 1948 (Bull.
Univ. Utah Biol. Ser. 10(6):6). Mirror Lake
(C48).

Argenna sp. Raft River S fk (CI33).

Argennina reclusa Gertsch & Ivie 1936
(Amer. Mus. Novitates 858:1). Cove Fort 10
mi N (GI36).

Argiope trifasciata (Forskal) 1775 (Descr.
Anim., p. 86). Green Cyn (H), Kelton, Kelton
Pass, Snowville (K), Salt Lake City (BU), St
George (CW).

Ariadna bicolor (Hentz) 1842 (J. Boston
Soc. Nat. Hist. 4:225). Lake Powell (CW),
Verdure (CG28).

Aysha gracilis (Hentz) 1847 (J. Boston Soc.
Nat. Hist. 5:452). Zion Nat Park (CW).

Aysha incursa (Chamberlin) 1919 (Ann.
Ent. Soc. Amer. 12:239). St. George (CW).

Bathyphantes concolor (Wider) 1834 (In:
Reuss, Zool. Misc. Mus. Senck. 1:267). Provo,
Salt Lake City (169).

Bathyphantes latescens (Chamberlin) 1919
(Ann. Ent. Soc. Amer. 12:248). Aspen Grove,
Logan Cyn, Mill Crk Cyn, Ogden River Cyn
(169), Chalk Crk (in Uinta Mts) (C19), Clear
Crk, Grouse Crk, Raft River S fk (CI33).

Bathyphantes pullatus (O. Pick. -Cam-
bridge) 1863 (Zoologist 21:8580) Cobble Rest
Camp, Uinta Mts, Salt Lake City (169).

Bathyphantes sp. USU School Forest (W).

Batroceps sp. Kelton (K).

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Allred, Kaston: Utah Spiders

499

Brachybothritim montaniim Chamberlin &
Ivie 1935 (Bull. Univ. Utah Biol. Ser. 2(8):4).
Provo River N fk (in Uinta Mts), Raft River
Mts, Salt Lake City (CI35).

Calilena gertschi Chamberlin & Ivie 1941
(Ann. Ent. Soc. Amer. 34:612). Monticello
(CI41).

Calilena multiformis dixiana Chamberlin
& Ivie 1941 (Ann. Ent. Soc. Amer. 34:608).
Diamond Valley, Pintura, St George (CI41).

Calilena restricta Chamberlin & Ivie 1941
(Ann. Ent. Soc. Amer. 34:606). Aquarius
Plateau, Cedar Mt, Escalante, Eureka Cyn,
Ferron, Loa, Mt Ellen, Noton, Panguitch,
Price, Raft River Mts, Richfield, Standard-
ville, Utah Lake W side, Zion Nat Park
(CI41), Locomotive Spngs (K), Tibbet Spng 2
km NE (AG), Widtsoe (BU).

Calilena sp. Calf Crk, Cyclone Lake (nr
Escalante), Steep Crk, Three Lakes (nr Ka-
nab) (BU), Smokey Mt (AU).

Callilepis eremellus Chamberlin 1928
(Proc. Biol. Soc. Washington 41:77). Caine-
viUe (CG28).

Callilepis zionis Chamberlin & Woodbury
1929 (Proc. Biol. Soc. Washington 43:133).
Zion Nat Park (CW).

Callilepis sp. Kelton, Kelton Pass (K), Four-
mile Bench, Smokey Mt, Tibbet Spng 2 km
NE (AU).

Callobiiis nevadensis (Simon) 1884 (Bull.
Soc. Zool. France 9:318). American Fk Cyn,
City Crk Cyn, Dry Cyn, Logan, Logan Cyn,
Mill Crk Cyn, Timpanogos Park (Le), Fill-
more (C19), Green Cyn (McU), Hughes Cyn,
Ogden, Ogden Cyn, Wasatch Mts (C47).

Callobius nomeus (Chamberlin) 1919 (Ann.
Ent. Soc. Amer. 12:240). Cedar Mts, Chalk
Crk (in Uinta Mts), Ferron Res, Mirror Lake,
Upper Provo River (C47), Cobble Rest,
Grand Daddy Lake, La Sal Pass, Mill Crk
Cyn, Ouray (Le), USU School Forest (W),
Wasatch Plateau (CI47a).

Castianeira aurata (Hentz) 1847 (J. Boston
Soc. Nat. Hist. 5:459). Raft River S fk (CI33).

Castianeira descripta (Hentz) 1847 (J. Bos-
ton Soc. Nat. Hist. 5:456). Zion Nat Park
(CW).

Castianeira longipalpa (Hentz) 1847 (J.
Boston Soc. Nat. Hist. 5:457). "Utah"(DR82).

Castianeira occidens Reiskind 1969 (Bull.
Mus. Comp. Zool. 138:211). Brigham Plains
Flat, Four-mile Bench (8 km SE cow camp at

head Wesses Cyn) (AG), Green Cyn (H), USU
School Forest (W).

Castianeira zionis Chamberlin & Wood-
bury 1929 (Proc. Biol. Soc. Washington
42:139). Zion Nat Park (CW).

Catabrithorax clypiellus Chamberlin 1920
(Canad. Ent. 52:199). Bear Lake, Logan Cyn
(C20), Clear Crk, Dove Crk, Raft River S fk.
Raft River Mts (CI33).

Catabrithorax plwnosus (Emerton) 1882
(Trans. Conn. Acad. Sci. 6:53). Fish Lake,
Moab, Price (C48).

Catabrithorax stylifer Chamberlin 1948
(Ann. Ent. Soc. Amer. 41:486). Clear Crk,
Dove Crk, Lynn 8 mi S, Raft River S fk
(C48).

Catabrithorax iitus (Chamberlin) 1919
(Ann. Ent. Soc. Amer. 12:253). Clear Lake
(C19), Cyclone Lake (nr Escalante), Posey
Lake (BU), Dove Crk, Raft River S fk, Yost
(CI33).

Ceraticelus crassiceps Chamberlin & Ivie
1939 (Verb. 7 intern. Kongr. Ent. 1:68). Mir-
ror Lake (CI39).

Ceraticelus subniger Chamberlin 1948
(Ann. Ent. Soc. Amer. 41:491). Salt Lake City
10 mi W (C48).

Ceraticelus sp. Curlew Valley (K).

Ceratinella acerea Chamberlin & Ivie 1933
(Bull. Univ. Utah Biol. Ser. 2(2):26). Raft Riv-
er S fk, Wasatch Mts (CI33).

Ceratinella brunnea Emerton 1882 (Trans.
Conn. Acad. Sci. 6:36). Clear Crk, Dove Crk
(CI33).

Ceratinops uintana Chamberlin 1948 (Ann.
Ent. Soc. Amer. 41:508). Cobble Rest, Mirror
Lake (C48).

Ceratinops watsinga Chamberlin 1948
(Ann. Ent. Soc. Amer. 41:516). Salt Lake City
(C48).

Ceratinops sp. Clear Crk (CI33).

Cesonia gertschi Platnick & Shadab 1980
(Bull. Amer. Mus. Nat. Hist. 165:352). Zion
Nat Park (PS80).

Cesonia sincera Gertsch & Mulaik 1936
(Amer. Mus. Novitates 851:10). Smokey Mt
(AG).

Cheiracanthium inclusum (Hentz) 1847 (J.
Boston Soc. Nat. Hist. 5:451). Calf Crk, Ka-
nab, Kanab Cyn, Three Lakes (N Kanab)
(BU), Green Cyn (H), St George, Zion Nat
Park (CW).

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Great Basin Naturalist

Vol. 43, No. 3

Cheiracanthitim rnildei L. Koch 1864 (Abh.
naturh. Ges. Nurnberg 1864:144). Provo
(A80), Brigham (McU).

Chrosiothes chirica (Levi) 1954 (Trans.
Amer. Microscop. Soc. 73:184). Utah (LR).

Chrysso nordica (ChamberHn & Ivie) 1947
(Ann. Ent. Soc. Amer. 40:29). Utah (LR).

Chrysso pelyx (Levi) 1958 (Psyche 64:104).
Salt Lake City (L58).

Cicurina deserticola Chamberlin & Ivie
1940 (Bull. Univ. Utah Biol. Ser. 5(8) :65).
Bicknell, Bluff, Perron, Henry Mts, Pintura, S
Price, Santa Clara, Thompsons, Tropic
(CI40), Glen Cyn City 5 km SW, Nipple
Bench, Smokey Mt (14 km from Last Chance
Jet) (AG).

Cicurina intermedia Chamberlin & Ivie
1933 (Bull. Univ. Utah Biol. Ser. 2(2) :46).
Clear Crk, Henry Mts, Raft River S fk, Uinta
Mts, Wasatch Mts (CI33), Fish Lake, Salt
Lake City (E), Green Cyn (H).

Cicurina parma Chamberlin & Ivie 1940
(Bull. Univ. Utah Biol. Ser. 5(9):67). Bryce
Cyn Nat Park (CI40).

Cicurina rohusta Simon 1886 (Ann. Ent.
Soc. Belg. 30:40). Bluff (CG28), Chalk Crk (in
Uinta Mts) (C19), Clear Crk, Henry Mts, Raft
River S fk (CI33), Perron, Pish Lake, La Sal
Mts, Liberty, Mirror Lake, Provo River N fk.
Raft River Mts, Smith & Morehouse Cyn,
Verdure, Wasatch Mts (CI40), Pine Cyn
(C13), Steep Crk, Widtsoe (BU), USU School
Porest (W).

Cluhiona abbottii Koch 1866 (Arach. Fam.
Drassiden., p. 303). City Crk Cyn, Utah Lake
W side (G41), Clear Crk (CI33), Moab
(CG28).

Cluhiona canadensis Emerton 1890 (Trans.
Conn. Acad. Sci. 8:181). Logan (McU).

Cluhiona mimula Chamberlin 1928 (Proc.
Biol. Soc. Washington 41:184). Clear Crk
(CI33), Pruita (CG28).

Cluhiona maesta Banks 1896 (Trans. Amer.
Ent. Soc. 23:64). Chalk Crk (in Uinta Mts)
(C19), Clear Crk, Dove Crk, Park Valley
(City), Raft River S fk. Raft River Mts (CI33).

Cluhiona mutata Gertsch 1941 (Amer.
Mus. Novitates 1184:14). Salt Lake City
(G41).

Cluhiona norvegica Strand 1900 (Kong.
Norske Vidensk. Selsk. Skr., p. 30). "Utah"
(DR82).

Cluhiona pacifica Banks 1896 (Trans.
Amer. Ent. Soc. 23:65). Zion Nat Park (CW).

Cochlemholus provo Chamberlin 1948
(Ann. Ent. Soc. Amer. 41:522). Cobble Rest
(C48).

Coreogonal hicornis (Emerton) 1923 (Ca-
nad. Ent. 55:242). USU School Porest (W).

Coriarachne hrunneipes Banks 1893 (J.
New York Ent. Soc. 1:133). Raft River S fk
(CI33), NE Utah (BS).

Coriarachne versicolor Keyserling 1880
(Spinn. Amer. Lat. 1:53). Benson (G53), Bluff,
Salt Lake City, Zion Nat Park (G32), Clear
Crk (CI33), Greenriver, Moab (CG28), Heber
(McU).

Crustulina sticta (O. Pick. -Cambridge)
1861 (Ann. Mag. Nat. Hist., ser. 3, 7:432).
Moab, Price, San Rafael River (CG28), Salt
Lake City, St George (lU), Uinta Mts (L57).

Ctenium eremophilus (Chamberlin) 1928
(Proc. Biol. Soc. Washington 41:180). Devils
Cyn (CG28), Verdure (Ks46).

Ctenium fusca (Emerton) 1894 (Trans.
Conn. Acad. Sci. 9:407). Locomotive Spngs
(K).

Ctenium vigerens (Chamberlin & Ivie)
1933 (Bull. Univ. Utah Biol. Ser. 2(2):9).
Dove Crk, Raft River S fk, Yost (CI33), Raft
River Mts, Salt Lake City, Verdure (Ks46).

Cybaeota concolor Chamberlin & Ivie 1937
(Ann. Ent. Soc. Amer. 30:211). Mill Crk Cyn
(CI37).

Cybaeota wasatchensis Chamberlin & Ivie
1937 (Ann. Ent. Soc. Amer. 30:211). Hughes
Cyn, Mill Crk Cyn (CI37).

Cyclosa conica (Pallas) 1772 (Spicil. Zool.
1(9):48). Clear Crk (CI33), Logan SE (McU),
Zion Nat Park (CW).

Delopelma iodius Chamberlin & Ivie 1939
(Bull. Univ. Utah Biol. Ser. 5(1):6). Castle
Cliffs 2 mi W, Zion Nat Park (CI39a).

Delopelma marxi (Simon) 1891 (Acta Bull.
Soc. Linn. Bordeaux 44:321). Pruita (G35).

Delopelma melanius Chamberlin & Ivie
1939 (Bull. Univ. Utah Biol. Ser. 5(1):5). Brig-
ham, Salt Lake City, Stockton, and Davis and
Utah counties (CI38a).

Delopelma steindachneri (Ausserer) 1875
(Verb. zool. bot. Ges. Wien 25:199). St
George (C21).

Dendryphantes (prob is Metaphidippus)
mylothrus Chamberlin 1925 (Proc. Calif.
Acad. Sci. (4)14:134). Mill Crk Cyn (C25).

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Allred, Kaston: Utah Spiders

501

DendrypJiantes (prob is Metaphidippus)
uteanus Chamberlin & Gertsch 1929 (J. Ent.
Zool. Pomona Coll. 21:110). Clear Crk, Raft
River S fk (CI33), Lambs Cyn, Zion Nat Park
(CG29).

Dictyna abundans Chamberlin & Ivie
1941 (Bull. Univ. Utah Biol. Ser. 6(3):5). St
George (CI41a), Zion Nat Park (CG58).

Dictyna apacheca Chamberlin & Ivie 1935
(Bull. Univ. Utah Biol. Ser. 2(8):28). Ameri-
can Fk Cyn (CG58).

Dictyna artemisia Ivie 1947 (New York:
priv. publ.). Provo River N fk, Scipio,
Wasatch Mts (nr Salt Lake City) (CG58).

Dictyna bellans Chamberlin 1919 (Ann.
Ent. Soc. Amer. 12:242). Moab (CG58).

Dictyna bicornis Emerton 1915 (Trans.
Conn. Acad. Sci. 20:141). Salt Lake City
(G46).

Dictyna borealis cavernosa Jones 1947
(Field & Lab. 15:12). Salt Lake City (CG58).

Dictyna brevitarsus Emerton 1915 (Trans.
Conn. Acad. Sci. 20:140). USU School Forest
(W).

Dictyna calcarata Banks 1904 (Proc. Calif.
Acad. Sci. 3:342). Calf Crk (BU), St George,
Zion Nat Park (CG58).

Dictyna cholla Gertsch & Davis 1942
(Amer. Mus. Novitates 1158:12). Salt Lake
City (CG58).

Dictyna completa Chamberlin & Gertsch
1929 (J. Ent. Zool. Pomona Coll. 21:101).
Green Cyn (H), Moab (CG28).

Dictyna coloradensis Chamberlin 1919
(Ann. Ent. Soc. Amer. 12:241). Utah (C58).

Dictyna cornupeta Bishop & Ruderman
1946 (Proc. Biol. Soc. Washington 59:1). Salt
Lake City (CG58).

Dictyna horta Gertsch & Ivie 1936 (Amer.
Mus. Novitates 858:4). City Crk Cyn, Layton
(CG58).

Dictyna idahoana Chamberlin & Ivie 1933
(Bull. Univ. Utah Biol. Ser. 2(2) :4). Cedar
Hills, Kelton, Kelton Pass, Snowville (K),
Green Cyn (H), Grouse Crk (CI33), Mt Nebo
(CG58).

Dictyna littoricolens Chamberlin & Ivie
1935 (Bull. Univ. Utah Biol. Ser. 2(8):30).
Black Rock, Salt Lake City (CG58), Locomo-
tive Spngs 3 mi E (K), Utah Lake W side
(CI35).

Dictyna major Menge 1869 (Schr. naturf.
Ges. Danzig 2:247). Farmington Cyn (LL).

Dictyna minuta Emerton 1888 (Trans.
Conn. Acad. Sci. 7:447). Price (CG58), Salt
Lake City (GI36).

Dictyna moaba Ivie 1947 (New York, priv.
publ.). Moab (C48).

Dictyna pallida Keyserling 1887 (Verb,
zool. bot. Ges. Wien 37:472). Richfield
(CG58).

Dictyna personata Gertsch & Mulaik 1936
(Amer. Mus. Novitates 851:9). East Cyn, Hur-
ricane (CG58), Glen Cyn City (AG), Glen-
wood 2 mi E (G46).

Dictyna pictella Chamberlin & Gertsch
1958 (Bull. Amer. Mus. Nat. Hist. 116:97).
Pintura, Scipio (CG58).

Dictyna piratica Ivie 1947 (New York,
priv. publ.). Dry Cyn (CG58), East Cyn, Pro-
vo River N fk, Wanship (C48).

Dictyna reticulata Gertsch & Ivie 1936
(Amer. Mus. JVovitates 858:7). Curlew Valley
(K), Richfield, Santa Clara 4 mi SW (GI36).

Dictyna secuta Chamberlin 1924 (Proc.
Calif. Adad. Sci. 12:583). St George (CG58).

Dictyna stulta Gertsch & Mulaik 1936
(Amer. Mus. Novitates 851:7). Aspen Grove,
Ogden Cyn (G46).

Dictyna terrestris Emerton 1911 (Trans.
Conn. Acad. Sci. 16:399). Bill's Cyn, Delta,
Dry Cyn, Mill Crk Cyn (CG58).

Dictyna tertanea Ivie 1947 (New York,
priv. publ.). American Fork Cyn, Bluff,
Castle Park, Dinosaur Nat Mon, Greenriver,
Henry Mts, Richfield, Salt Lake City (CG58).

Dictyna tridentata Bishop & Ruderman
1946 (Proc. Biol. Soc. Washington 58:2). Elk
Ridge (CG58).

Dictyna tucsona Chamberlin 1948 ( Bull.
Univ. Utah Biol. Ser. 10(6):8). Kanab (CG58).

Dictyna uintana Chamberlin 1919 (Ann.
Ent. Soc. Amer. 12:240). Bluff (CG28), Chalk
Crk (in Uinta Mts) (C19), City Crk Cyn
(C48), Clear Crk, Grouse Crk, Raft River
Mts, Raft River S fk, Yost (CI33), Elsinore,
Fish Lake, Richfield, Salt Lake City (G46).

Dictyna variana Chamberlin & Gertsch
1958 (Bull. Amer. Mus. Nat. Hist. 116:57). St
George (CG58).

Dictyna vincens Chamberlin 1919 (Ann.
Ent. Soc. Amer. 12:243). Clear Crk, Raft Riv-
er S fk. Raft River Mts (CI33).

Dictyna volucripes Keyserling 1882 (Verb,
zool. bot. Ges. Wien 31:286). Central &
northern Utah, St George (C21), La Sal Mts
(C28).

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Great Basin Naturalist

Vol. 43, No. 3

Dictyna zaha Barrows & Ivie 1942 (Ohio J.
Sci. 42:21). City Crk Cyn, Layton, Richfield
(CG58).

Dictyna sp. Coyote Gulch, Posey Lake,
Steep Crk, Table Cliff Plateau, Willow Tank
Spngs (BU).

Dictynina eutypa (Chamberlin & Ivie)
1929 (J. Ent. Zool. Pomona Coll. 21:101).
Bluff (CG28).

Dictynoides sp. Zion Nat Park (CW).

Digiietia canities (McCook) 1890 (Amer.
Spiders 2:136). St George (G35), Zion Nat
Park (CW).

Dipoena atopa (Chamberlin) 1948 (Ann.
Ent. Soc. Amer. 41:541). City Crk Cyn, Dry
Cyn, Timpanogos Nat Mon (L53), Grantsville
(L63), Red Butte Cyn (C48a).

Dipoena malkini Levi 1953 (Amer. Mus.
Novitates 1647:33). Government Crk, Mill
Crk Cyn (L53).

Dipoena nigra (Emerton) 1882 (Trans.
Conn. Acad. Sci. 6:21). Beaver Cyn, City Crk
Cyn, Government Crk, Wanship (L53), Pick-
leville (W).

Dipoena prona Menge 1868 (Schr. naturf.
Ges. Danzig 2:177). Moab (L53).

Dipoena provalis Levi 1953 (Amer. Mus.
Novitates 1639:34). Cobble Rest, Hughes Cyn
(L53), Provo River N fk (GU).

Dipoena tibialis Banks 1906 (Proc. Ent.
Soc. Washington 7:96). Green Cyn (H), USU
School Forest (W).

Dipoena sp. Clear Crk, Raft River Mts
(CI53).

Disembolus alpha (Chamberlin) 1948 (Ann.
Ent. Soc. Amer. 41:550). Dry Crk Cyn
(C48a).

Disemholus anguineus Millidge 1981 (J.
Arach. 9:276). Utah (M81a).

Disemholus beta Millidge 1981 (J. Arach.
9:274). Dry Cyn (M81a).

Disemholus galeatus Millidge 1981 (J.
Arach. 9:268). Utah (M81a).

Disemholus implexus Millidge 1981 (J.
Arach. 9:279). Fish Lake (M81a).

Disemholus implicatus Millidge 1981 (J.
Arach. 9:277). Cobble Rest, Upper Provo
River (M81a).

Disemholus kesimbus (Chamberlin) 1948
(Ann. Ent. Soc. Amer. 41:552). Fish Lake,
Henry Mts, Strawberry Res (C48a).

Disemholus stridulans Chamberlin & Ivie
1933 (Bull. Univ. Utah Biol. Ser. 2(2) :20).

Fillmore Cyn, Lynn 8 mi S, Mirror lake,
Snow Crk Cyn, Wasatch Mts (nr Salt Lake
City) (CI45), Henry Mts, Raft River S fk,
(CI33).

Disemholus vicinus Millidge 1981 (J.
Arach. 9:281). Grantsville (M81a).

Dolomedes triton (Walckenaer) 1837 (Hist.
Nat. Ins. Apt. 1:340). Raft River Mts (CI33).

Drassodes gosiutus Chamberlin 1919 (Ann.
Ent. Soc. Amer. 12:245). City Crk Cyn, Dry
Cyn, Ensign Peak, Greenriver, Helper, E
Monticello, Mud Spngs, Rotary Park, Salt
Lake Airport 5 mi SW, Spring Crk, Utah
Lake W side. Willow Crk (PS76), Fillmore
(C19), Nipple Bench (AG), Zion Nat Park
(CW).

Drassodes neglectus (Keyserling) 1887
(Verb. zool. bot. Ges. Wien 37:434). Bluff,
Marysvale, Verdure (CG28), Fish Lake (C36),
La Sal Mts (C28), also counties of Garfield,
Iron, Salt Lake, Summit, Tooele, and Utah
(PS76).

Drassodes saccatus (Emerton) 1890 (Trans.
Conn. Acad. Sci. 8:178). Fillmore (C19), Glen
Cyn City 5 km SW, Nipple Bench (AG),
Green Cyn (H), Raft River Mts (CI33), also
counties of Carbon, Emery, Garfield, Grand,
Salt Lake, San Juan, Sevier, and Washington
(PS76).

Drassodes sp. Curlew Valley (K), Glen Cyn
City (AU), Steep Crk, Widtsoe (BU).

Drassyllus conformans Chamberlin 1936
(Amer. Mus. Novitates 841:22). Richfield,
Salt Lake City (C36).

Drassyllus dromeus Chamberlin 1922
(Proc. Biol. Soc. Washington 35:169). Dry
Cyn, Pinecrest (C36).

Drasyllus inanus Chamberlin & Gertsch
1940 (Amer. Mus. Novitates 1068:17). Bluff
(CG40).

Drassyllus insularis (Banks) 1900 (Canad.
Ent. 32:97). Bluff, Salt Lake County, Straight
Wash, Valley City (CG28), Green Cyn (H),
Grouse Crk (CI33), Lake Powell (C58), St
George (CW).

Drassyllus lamprus (Chamberlin) 1920 (Ca-
nad. Ent. 52:193). Mill Crk (C22), USU
School Forest (W).

Drassyllus lepidus (Banks) 1899 (Proc. Ent.
Soc. Washington 4:190). Pine Valley Mts
(C36).

Drassyllus mexicanus (Banks) 1898 (Proc.
Calif. Acad. Sci (3)1:217). La Sal Mts (CG40).

April 1983

Allred, Kaston: Utah Spiders

503

Drassyllus mormon Chamberlin 1936
(Amer. Mus. Novitates 841:27). St George
(C36).

Drassyllus nannelliis Chamberlin &
Gertsch 1940 (Amer. Mus. Novitates
1068:11). Tremonton 10 mi W (CG40),
Green Cyn (H).

Drassyllus notonus Chamberlin 1928
(Proc. Biol. Soc. Washington 41:179). Noton
(CG28).

Drassyllus tonaquintus Chamberlin &
Gertsch 1940 (Amer. Mus. Novitates 1068:1).
Farmington, St George (CG40).

Drassyllus sp. Curlew Valley (K).

Ebo evansae Sauer & Platnick 1972 (Ca-
nad. Ent. 104:41). Butler, Logan, Salt Lake
City (SP72), Green Cyn (H).

Ebo sp. Locomotive Spngs (K), Raft River
S fk (CI33).

Enoplognatlm Joshua Chamberlin & Ivie
1942 (Bull. Univ. Utah Biol. Ser. 7(1) :44).
Brigham Plains Flat (AG), Hurricane, Pin-
tura, Helper, Spring Cyn (CI42a), counties of
Emery, Grand, Salt Lake, and Sevier (L57).

EnoplognatJia marmorata (Hentz) 1850 (J.
Boston Soc. Nat. Hist. 6:273). Dry Cyn, Emi-
gration Cyn (lU), Oquirrh Mts, Santaquin
(CI42), Park Valley, Raft River Mts, Raft Riv-
er S fk (CI33), Sevier County (L57).

Enoplognatha ovata (Clerck) 1757 (Aranei
Suecici, p. 58). Green Cyn (H).

Enoplognatha wyuta Chamberlin & Ivie
1942 (Bull. Univ. Utah Biol. Ser. 7(1):43).
American Fork Cyn, Hughes Cyn (CI42a),
and Emery County (L57).

Erigone canthognatha Chamberlin & Ivie
1935 (Bull. Univ. Utah Biol. Ser. 2(8): 13).
Colorado River 5 mi up from Moab (CI35).

Erigone denticulata Chamberlin & Ivie
1939 (Verh. 7 intern. Kongr. Ent. Berlin
1:57). Mirror Lake (CI39).

Erigone dentosa O. Pick. -Cambridge 1894
(Biol. Centr. Amer. Arach. Ar. 1:128). Clear
Crk, Grouse Crk, Raft River Mts, Raft River
S fk, Yost (CI33), Green Cyn (H), Posey Lake
(BU), USU School Forest (W).

Erigone uintana Chamberlin & Ivie 1935
(Bull. Univ. Utah Biol. Ser. 2(8): 14). Mirror
Lake (CI35).

Erigone viabilis Chamberlin & Ivie 1933
(Bull. Univ. Utah Biol. Ser. 2(2): 11). Mirror
Lake, Raft River S fk (CI33).

Eris marginata (Walckenaer) 1837 (Hist.
Nat. Ins. Apt. 1:466) Mill Crk Cyn (C25), St
George (CW).

Eris nigromaculatus (Keyserling) 1884
(Verh. zool. bot. Ges. Wein 34:500) Lambs
Cyn (CG29).

Ero canionis Chamberlin & Ivie 1935 (Bull.
Univ. Utah Biol. Ser. 2(8):23 ). City Crk Cyn
(CI35).

Eularia chelata Chamberlin & Ivie 1939
(Verh. 7 intern. Kongr. Ent. Berlin 1:61). Mir-
ror Lake (CI45).

Eularia dela Chamberlin & Ivie 1933 (Bull.
Univ. Utah Biol. Ser. 2(2): 15). Clear Crk, Raft
River S fk, Raft River Mts (CI33), Lynn 8 mi

5 (CI45).

Eularia kaiba Chamberlin 1948 (Ann. Ent.
Soc. Amer. 41:530). Mirror Lake (C48).

Eularia mana Chamberlin & Ivie 1935
(Bull. Univ. Ufah Biol. Ser. 2(8): 16). Mill Crk
Cyn (C45).

Eularia schediana Chamberlin & Ivie 1933
(Bull. Univ. Utah Biol. Ser. 2(2): 17). Cedar
Mt, City Crk Cyn, Ferron, Fish lake, Henry
Mts, Horse Valley, Kaibab Forest, Mirror
Lake, Upper Provo River (CI45), Dove Crk
(CI33), Raft River Mts (CI45).

Euryopis coki Levi 1954 (Amer. Mus. Novi-
tates 1666:33). Salt Lake City (L54).

Euryopis formosa Banks 1908 (Canad. Ent.
40:206). American Fork, Salt Lake City
(CI41), Dry Cyn, Smith & Morehouse Cyn
(L54).

Euryopis scriptipes Banks 1908 (Canad.
Ent. 40:206). Clear Crk, Grouse Crk, Raft
River Mts (CI33), Glen Cyn City (AG), Mt
Agassiz, Bryce Cyn Nat Park, Jet Deep Crk

6 Carter Crk (in Uinta Mts), Fish Lake,
Fruita, Hughes Cyn, Mill Crk Cyn, Trout
Crk, Vermillion Castle (nr Parowan), Zion
Nat Park (L54), Green Cyn (H).

Euryopis spinigera O. Pick. -Cambridge
1895 (Biol. Centr. Amer. Arachn. Ar. 1:146).
City Crk Cyn, Junction, Parleys Cyn, Pintura
10 mi N (L54).

Euryopis taczanowskii Keyserling 1886
(Spinn. Amer., Theridiidae 2:47). Orton
(L54).

Euryopis texana Banks 1908 (Canad. Ent.
40:207). Hurricane (L54).

Euryopis sp. Brigham Plains (AU), Snow-
ville (K), Coyote Gulch, Escalante, Steep Crk
(BU).

504

Great Basin Naturalist

Vol. 43, No. 3

Evarcha leucophaea (Koch) 1846 (Die
Arachn. 13:216). Dove Crk, Raft River S fk
(CI33).

Filistata hurca Chamberlin & Ivie 1942
(Bull. Univ. Utah Biol. Ser. 7(1):3). Hurricane
(CI42a).

Filistata utahana Chamberlin & Ivie 1935
(Bull. Univ. Utah Biol. Ser. 2(8) :5). Brigham
Plains Flat, Glen Cyn City 7 km SW, Tibbit
Spng 2 km NE (AG), Marysvale, St George
(CI35b).

Floricottius littoralis Chamberlin & Ivie
1935 (Bull. Univ. Utah Biol. Ser. 2(8): 16).
Utah Lake W shore (CI35b).

Frontinella pyramitela (Walckenaer) 1841
(Hist. Nat. Ins. Apt. 2:261). Green Cyn (H),
Washington County, Willow Tank Spngs
(BU).

Gayenna dixiana Chamberlin & Wood-
bury 1929 (Proc. Biol. Soc. Washington
42:138). St George (CW).

Geolycosa fatifera (Hentz) 1842 (J. Boston
Soc. Nat. His. 4:229). Utah (C9).

Geolycosa rafaelana (Chamberlin) 1928
(Proc. Biol. Soc' Washington 41:186). Big In-
dian Rock, San Rafael Desert (CG28), Glen
Cyn City 5 km SW (AG).

Gnaphosa antipola Chamberlin 1933
(Amer. Mus. Novitates 631:4). Farmington,
Moab, Salt Lake City (PS75), Utah Lake W
shore (C33).

Gnapliosa brwnalis Thorell 1875 (J. Boston
Soc. Nat. Hist. 17:497). La Sal Mts, Verdure
(CG28), Salt Lake City (PS75).

Gnapliosa calif ornica Banks 1904 (Proc.
Calif. Acad. Sci. (Zool.) 3:335). Four-mile
Bench (5 km SW cow camp at head Wesses
Cyn) (AG), Ft Douglas, Glenwood, Grant-
sville. Hat Island, Manilla, Pintura, Raft Riv-
er Mts, Richfield, Salt Lake City, St George,
Stockton, Straight Wash, Tooele Cyn, Tre-
monton (PS75), Grouse Crk (CI33).

Gnaphosa clara (Keyserling) 1887 (Verh.
zool. bot. Ges. Wien 37:429). Bridger Basin
(C22), Greenriver, Grouse Crk, Moab, Mud
Spngs, Wah Wah Mts (PS75).

Gnaphosa dentata Platnick & Shadab 1975
(Bull. Amer. Mus. Nat. Hist. 155:18). Utah
Lake and Washington County (PS75).

Gnapliosa gosoga Chamberlin 1928 (Proc.
Biol. Soc. Washington 41:178). Marysvale,
Straight Wash (CG28), Tooele Cyn (C36).

Gnaphosa hirsutipes Banks 1901 (Proc.
Acad. Nat. Sci. Philadelphia 53:573). Fruita,
Salt Lake City, San Rafael River, Verdure
(CG28), Grouse Crk (CI33), St George (CW).

Gnaphosa musco^'uni (L. Koch) 1866
(Arachn. Fam. Drassiden., p. 14). Blue Spruce
Camp, Castle Dale, Emigration Cyn, Fish
Lake, Fillmore, Hatch, Leidy Peak, Mill Crk,
Provo, Richfield, Salt Lake City (PS75),
Clear Crk, Raft River Mts, Raft River S fk,
Yost (CI33), Curlew Valley, Kosmo (K), La
Sal Mts (C28a), Monroe Cyn (C36), Oquirrh
Mts (CG28), Rock Island (in Utah Lake) (BU),
St George (CW).

Gnaphosa salsa Platnick & Shadab 1975
(Bull. Amer. Mus. Nat. Hist. 155:22). Terrys
Ranch (Beaver Dam Wash) (PS75).

Gnaphosa saxosa Platnick & Shadab 1975
(Bull. Amer. Mus. Nat. Hist. 155:17). Salt
Lake City (PS75).

Gnaphosa sericata (L. Koch) 1866 (Arach.
Fam. Drassiden., p. 31). Green Cyn (H), Lo-
comotive Spngs (K), Moab, Price, Salt Lake
City (PS75).

Gnaphosa synthetica Chamberlin 1924
(Proc. Calif. Acad. Sci. 12:620). St George
(PS75).

Gnaphosa utahana Banks 1904 (J. New
York Ent. Soc. 12:110). Egg Island, Farm-
ington NW, Hat Island, Plain City, San Ra-
fael River, Silver Lake (PS75).

Gnaphosa sp. Brigham Plains (AU), Posey
Lake, Provo, Widtsoe (BU).

Gnathantes ferosa Chamberlin & Ivie 1943
(Bull. Univ. Utah Biol. Ser. 7(6):5). City Crk
Cyn (CI43).

Gosiphrurus unicolor Chamberlin & Ivie
1935 (Bull. Univ. Utah Biol. Ser. 2(8):40).
Ferron (CI35b).

Grammonota salicicola Chamberlin 1948
(Ann. Ent. Soc. Amer. 41:532). Salt Lake City
(C48a).

Habrocestrum sp. Raft River S fk (CI33).

Habronattus sp. Calf Crk, Kanab, Steep
Crk (BU).

Hahnia cinerea Emerton 1890 (Trans.
Conn. Acad. Sci. 8:197). Butterfield Cyn, Mill
Crk, Logan Cyn (G34b), Zion Nat Park
(CW).

Hahnia ononidiwn Simon 1875 (Arach. de
France 2:135). Fish Lake, La Sal Mts, Mt El-
len, Pine Spngs (in Henry Mts) (CI42a).

April 1983

Allred, Kaston: Utah Spiders

505

Haplodrassus hicornis (Emerton) 1909
(Trans. Conn. Acad. Sci. 14:218). Blanding,
City Crk Cyn (CG28), Marysvale Cyn (C36a).

Haplodrassus dixiensis Chamberlin &
Woodbury 1929 (Proc. Biol. Soc. Washington
42:134). St George (CW).

Haplodrassus eunis Chamberlin 1922
(Proc. Biol. Soc. Washington 35:162). Cedar
Cyn, East Cyn, Emory, Provo River N fk,
Smith & Morehouse Cyn, Tropic Res (CI46),
Nipple Bench, Smokey Mt (14 & 23 km from
Last Chance Jet) (AG), USU School Forest
(W).

Haplodrassus signifer (Koch) 1839 (Die
Arachn. 6:31). Blanding, Marysvale, Moab,
Noton, Salt Lake County, Straight Wash
(CG28), Cyclone Lake (nr Escalante) (BU),
Green Cyn (H), Gunnison Butte (nr Green-
river), Moab, Richardson (CG40), La Sal Mts
(C28a).

Haplodrassus sp. Posey Lake (BU).

Helopfiora orinoma (Chamberlin) 1920 (Ca-
nad. Ent. 52:195). Bear Lake (C20).

Helophora reducta (Keyserling) 1886
(Spinn. Amer. Theridiidae 2:54). Chalk Crk
(in Uinta Mts), Clear Lake (C19), Clear Crk,
Raft River S fk, Yost (CI33).

Helopliora hinagijna Chamberlin & Ivie
1943 (Bull. Univ. Utah Biol. Ser. 7(6):6). USU
School Forest (W).

Herpyllus cockerelli (Banks) 1901 (Proc.
Acad. Nat. Sci. Philadelphia 53:571). Red
Cyn Camp (11 mi SE Panguitch) (P).

Herpyllus convallis Chamberlin 1936
(Amer. Mus. Novitates 841:2). St George,
Zion Nat Park (P).

Herpyllus ecclesiastica Hentz 1832 (Amer.
J. Sci. 21:102). Bluff, HanksviUe, San Rafael
River, Valley City (CG28).

Herpyllus hesperolus Chamb^lin 1928
(Proc. Biol. Soc. Washington 42:176). Big In-
dian Rock, Bluff, Fruita, HanksviUe, Mary-
svale, Moab, San Rafael River, Valley City
(CG28), Locomotive Spngs (K), Richfield
(C36a), St George, Zion Nat Park (CW), and
counties of Carbon, Davis, Duchesne, Mor-
gan, Salt Lake, San Juan, Tooele, Utah, and
Wayne (P).

Herpyllus propinquus (Keyserling) 1887
(Verb. zool. bot. Ges. Wien 37:430). Four-
mile Bench (8 km SE cow camp at head
Wesses Cyn) (AG), La Sal Mts (CG28), Lynn,
Raft River Mts (CI33), St George (CW) and

counties of Beaver, Carbon, Duchesne, Mil-
lard, Uintah, Utah, and Wayne (P) and San
Juan (PS77).

Herpyllus sp. Curlew Valley, Kosmo (K),
Glen Cyn City (AU), Green Cyn (H).

Hillhousia microtarsus (Emerton) 1882
(Trans. Conn. Acad. Sci. 6:57). Mirror Lake,
upper Provo River (CI45).

Hololena hola (Chamberlin & Gertsch)

1928 (Proc. Biol. Soc. Washington 41:183).
Blanding, Devils Cyn, Moab, Noton, Verdure
(CG28), Monticello (CI42).

Hololena mimoides (Chamberlin) 1919
(Ann. Ent. Soc. Amer. 12:256). Devils Cyn,
Moab, Pine Valley, Pintura, St George, Zion
Nat Park (CI42), Fillmore Cyn (C19), Posey
Lake (BU).

Hololena nevada (Chamberlin & Gertsch)

1929 (J. Ent. Zool. Pomona Coll. 21:107).
Beaver Cyn, Moab 5 mi NE, Price, Richfield
(CI42).

Hololena oquirrhensis (Chamberlin &
Gertsch) 1930 (Proc. Biol. Soc. Washington
43:142). Butterfield Cyn (CG30), Oquirrh
Mts, Wasatch Mts (CI42).

Hybocoptus dentipalpis (Emerton) 1915
(Trans. Conn. Acad. Sci. 20:149). Chalk Crk
(in Uinta Mts) (C19).

Hypselistes florens (O. Pick. -Cambridge)
1875 (Proc. Zool. Soc. London, p. 403). Smith
& Morehouse Cyn (CI35b).

Hypselistes reducens Chamberlin & Ivie
1935 (Bull. Univ. Utah Biol. Ser. 2(8): 17).
Mirror Lake (CI35b).

Hypsosinga funebris (Keyserling) 1893
(Spinn. Amer. Epeiridae 4:37). Alta (L75).

Hypsosinga groenlandica (Thorell) 1872
(Ofvers Kongl. Vet. Akad. Forh. 29:157).
Franklin Basin (L75).

Hypsosinga pygmaea (Sundevall) 1831
(Kongl. Svenska Vet. Akad. Handl., p. 121).
Kanab Cyn (BU), Salt Lake City (L75).

Hypsosinga singaefomiis (Scheffer) 1904
(Ent. News 15:259). Green Cyn (H).

Hijptiotes gertschi Chamberlin & Ivie 1935
(Bull. Univ. Utah Biol. Ser. 2(8): 12). Cedar
Cyn (nr Cedar City), Henry Mts, Salt Lake
City (CI35b).

Hyptiotes puebla Muma & Gertsch 1964
(Amer. Mus. Novitates 2196:14). N central
Utah (MG).

Icius annectans Chamberlin & Gertsch
1929 (J. Ent. Zool. Pomona Coll. 21:110).
Zion Nat Park (CG29).

506

Great Basin Naturalist

Vol. 43, No. 3

Idionella anomala (Chamberlin & Ivie)
1939 (Verb. 7 intern. Kongr. Ent. Berlin
1:69). Dry Cyn(CI39).

Idionella tuganus (Chamberlin) 1948 (Ann.
Ent. Soc. Amer. 41:494). Salt Lake City 10
mi W (C48).

Lahuella prosaica Chamberlin & Ivie 1943
(Bull. Univ. Utah Biol. Ser. 7(6): 10). Smith &
Morehouse Cyn (CI43).

Larinia borealis Banks 1894 (Ent. News
5:8). Green Cyn (H), nr Salt Lake City
(L75a).

Larinia famitlatoria (Keyserling) 1883
(Ver. zool. bot. Ges. Wien 32:201). Central
and southern Utah (L75a).

Latrodectus hesperus Chamberlin & Ivie
1935 (Bull. Univ. Utah Biol. Ser. 3(1): 15).
Ahlstrom Point, Glen Cyn City 6 km SW,
Tibbet Spng 2 km NE (AG), Bluff, Moab,
Price (CG28), Clear Crk, Grouse Crk, Raft
River Mts (CI33), Curlew Valley, Kosmo (K),
Dry Cyn, Mill Crk, Logan Cyn, Weber
Coimty, Millard County (GU), Green Cyn
(H), Provo (A74), St George, Zion Nat Park
(CW), Salt Lake City (CI35a), Tooele (L59),
Utah Lake, Three Lakes (N Kanab) (BU).

Lathys deliculata Gertsch 1946 (Amer.
Mus. Novitates 1319:3). Moab, Pintura
(CG58).

Lathys hesperus Chamberlin 1948 (Bull.
Univ. Utah Biol. Ser. 10(6): 14). Utah (C48).

Lepthypliantes agressus Chamberlin & Ivie
1943 (Bull. Univ. Utah Biol. Ser. 7(6): 15).
Lambs Cyn head. Mirror Lake, Provo River
(in Uinta Mts) (CI43).

Lepthypliantes arhorea (Emerton) 1915
(Trans. Conn. Acad. Sci. 20:151). Posey Lake
(BU), USU School Forest (W).

Lepthypliantes calcarata (Emerton) 1909
(Trans. Comi. Acad. Sci. 14:197). USU School
Forest (W).

Lepthypliantes concolor (Wider) 1834 (In:
Reuss, A. Zool. Misc. Arachn., Mus. Senck.
1:267). Provo, Salt Lake City (169).

Lepthypliantes furcillifer Chamberlin &
Ivie 1933 (Amer. Mus. Novitates 631:32).
Clear Crk, Raft River S fk (CI33).

Lepthypluintes lamprus Chamberlin 1920
(Canad. Ent. 52:195). Clear Crk, Raft River S
fk (CI33), Lake Powell (C58), La Sal Mts
(C28), Logan Cyn (C20b), Posey Lake (BU).

Lepthypliantes nehulosus (Sundevall) 1829
(Kongl. Svenska Vet. Akad. Handl, p. 218).
Lynn (CI33).

Lepthypliantes pollicaris Zorsch 1937
(Amer. Midi. Nat. 18:897). USU School For-
est (W).

Lepthijphantes ranieri Emerton 1926 (Ca-
nad. Ent. 58:118). USU School Forest (W).

Lepthypliantes sp. Cyclone Lake (nr Esca-
lante) (BU).

Linyphantes ephedrus (Chamberlin & Ivie)
1933 (Bull. Univ. Utah Biol. Ser. 2(2):31). Dry
Cyn, Fish Lake, St George, Yost, Zion Nat
Park (CI42a).

Linyphia tattphora Chamberlin 1928 (Proc.
Biol. Soc. Washington 41:180). Bluff, Zion
Nat Park (CG28).

Loxosceles deserta Gertsch 1975 (Toxicon
13:203). Southern Utah (GR).

Loxosceles rufipes (Lucas) 1834 (Mag. Zool.
Guerin 4:354). Clear Crk, Raft River Mts
(CI33), San Juan County (CG28), Zion Nat
Park (CW).

Loxosceles sp. Lake Powell (C58).

Lycosa antelucana Montgomery 1904
(Proc. Acad. Nat. Sci. Philadelphia 56:282).
St George (C21a).

Lycosa carolinensis Hentz 1842 (J. Boston
Soc. Nat. Hist. 4:230). Utah (COS).

Lycosa frondicola Emerton 1885 (Trans.
Conn. Acad. Sci. 6:484). Clear Crk (CI33).

Lycosa helluo Walckenaer 1837 (Hist. Nat.
Ins. Apt. 1:337). Utah (C8).

Lycosa uinticolens Chamberlin 1936 (Proc.
Biol. Soc. Washington 49:15). Green Lake (in
Uinta Mts) (C36).

Lycosa sp. Green Cyn (H), Provo (A78).

Mallos alpheus Chamberlin 1948 (Bull.
Univ. Utah Biol. Ser. 10(6): 14). American
Fork Cyn (C48).

Mallos eutyptis (Chamberlin & Gertsch)
1929 (J. Ent. Zool. Pomona Coll. 21:101).
Bluff, Monroe Cyn, Richfield, St George
(G46).

Mallos niveus O. Pick.-Cambridge 1902
(Biol. Centr. Amer., Arachn. Ar. 1:308). Calf
Crk, Posey Lake, Three Lakes (nr Kanab),
Willow Tank Spngs (BU), Hughes Cyn, Oph-
ir, Richfield, Salt Lake City, Zion Nat Park
(G46), Hurricane (C48).

Mallos trivittatiis (Banks) 1901 (Proc. Acad.
Nat. Sci. Philadelphia 53:577). Calf Crk, Ka-
nab, Posey Lake, Steep Crk, Three Lakes (N
Kanab) (BU), Richfield (G46), USU School
Forest (W), Zion Nat Park (C48).

April 1983

Allred, Kaston: Utah Spiders

507

Marpissa californica (Peckham) 1888
(Trans. Wisconsin Acad. Sci. 7:81). San Ra-
fael (CG28).

Mmo perplexa Chamberlin & Ivie 1939
(Verb. 7 intern. Kongr. Ent. Berlin 1:64).
Cobble Rest (€145), Provo River N fk (in
Uinta Mts) (C139).

Meioneta fiUmomna (Chamberlin) 1919
(Ann. Ent. Soc. Amer. 12:249). Fillmore Cyn
(C19).

Meioneta sp. Green Cyn (H).

Metacyrba taeniola (Hentz) 1846 (J. Boston
Soc. Nat. Hist. 5:353). Moab (CG28), Nipple
Bench (AG).

Metaphidippus aeneolus (Curtis) 1892 (Zoe
3:332). Kelton Pass, Snowville (K), USU
School Forest (W).

Metaphidippus diplacis Chamberlin 1924
(Proc. Calif. Acad. Sci. (4)12:616). Zion Nat
Park (CW).

Metaphidippus galathea (Walckenaer)
1837 (Hist. Nat. Ins. Apt. 1:456). La Sal Mts
(C28), Raft River S fk (CI33), Zion Nat Park
(CW).

Metaphidippus helenae (Banks) 1921 (Proc.
Calif. Acad. Sci. 11:101). Fish Lake, Richfield
(G34c).

Metaphidippus paiutus Gertsch 1934
(Amer. Mus. Novitates 726:1). Richfield, St
George (G34c).

Metaphidippus unicus (Chamberlin &
Gertsch) 1930 (Proc. Biol. Soc. Washington
43:143). Uintah County (CG30).

Metaphidippus verecundus (Chamberlin &
Gertsch) 1930 (Proc. Biol. Soc. Washington
43:144). Clear Crk, Raft River S fk (CI33),
Dry Cyn (CG30), Green Cyn (H).

Metaphidippus vitis (Cockerell) 1894 (En-
tomologist 27:207). Kanab Cyn, Three Lakes
(N Kanab) (BU).

Metaphidippus sp. Coyote Gulch, Esca-
lante, Steep Crk, Table Cliff Plateau, Three
Lakes (N Kanab), Widtsoe, Willow Tank
Spngs (BU), Green Cyn (H), Nipple Bench
(AU).

MeteUina mimetoides Chamberlin & Ivie
1941 (Bull. Univ. Utah Biol. Ser. 6(3): 15).
Utah (L80).

Metepeira foxi Gertsch & Ivie 1936 (Amer.
Mus. Novitates 858:20). Curlew Valley (K),
Fish Lake, Richfield (GI36), Green Cyn (H).

Metepeira grandiose alpina Chamberlin &
Ivie 1942 (Bull. Univ. Utah Biol. Ser. 7(1):74).
Fish Lake (CI42a).

Metepeira gosoga Chamerlin & Ivie 1935
(Bull. Univ. Utah Biol. Ser. 2(8):21). Esca-
lante, Steep Crk (BU).

Metepeira labyrinthea (Hentz) 1847 (J. Bos-
ton Soc. Nat. Hist. 5:471). Clear Crk, Dove
Crk, Raft River S fk (CI33).

Metepeira nanella Chamberlin & Ivie 1942
(Bull. Univ. Utah Biol. Ser. 7(1):71). Fillmore
(CI42a).

Metepeira sp. Smokey Mt (AU).

Micaria ahana Gertsch 1933 (Amer. Mus.
Novitates 637:6). Utah (G35).

Micaria formicoides Chamberlin & Wood-
bury 1929 (Proc. Biol. Soc. Washington
42:139). St George (CW).

Micaria gosiuta Gertsch 1942 (Amer. Mus.
Novitates 1195:1). City Crk Cyn (G42).

Micaria jeanae Gertsch 1942 (Amer. Mus.
Novitates 1195:4). Glenwood (G42).

Micaria rjiontana Emerton 1890 (Trans.
Conn. Acad. Sci. 8:168). Clear Crk, Dove
Crk, Lynn (CI33).

Micaria mormon Gertsch 1935 (Amer. Mus.
Novitates 805:17). City Crk Cyn (G35).

Micaria saUna Gertsch 1942 (Amer. Mus.
Novitates 1195:5). Salina (G42).

Micaria sp. Ahlstrom Point, Four-mile
Bench (AU), Green Cyn (H), Kelton, Lo-
comotive Spngs (K), USU School Forest (W).

MicroHnyphia mandibulata (Chamberlin &
Ivie) 1943 (Bull. Univ. Utah Biol. Ser.
7(6):24). Emery, Fillmore, Provo River mth,
Scipio, Verdure, Wasatch Mts, Zion Nat Park
(CI43).

Microneta anopla Chamberlin & Ivie 1933
(Bull. Univ. Utah Biol. Ser. 2(2) :35). Clear
Crk, Dove Crk, Grouse Crk, Raft River S fk
(CI33).

Microneta cornupalpis (O. Pick. -Cam-
bridge) 1875 (Proc. Zool. Soc. London, p.
401). Clear Crk (CI33).

Microneta fratrella (Chamberlin) 1919
(Ann. Ent. Soc. Amer. 12:250). Clear Crk
(CI33), Uinta Mts (C19).

Microneta lophophor Chamberlin & Ivie
1933 (Bull. Univ. Utah Biol. Ser. 2(2) :35).
Raft River S fk (CI33).

Microneta orines Chamberlin & Ivie 1933
(Bull. Univ. Utah Biol. Ser. 2(2):35). Clear
Crk, Raft River S fk (CI33).

Microneta protrudens Chamberlin & Ivie
1933 (Bull. Univ. Utah Biol. Ser. 2(2) :37).
Clear Crk, Uinta Mts (CI33).

508

Great Basin Naturalist

Vol. 43, No. 3

Microneta tumoa Chamberlin & Ivie 1933
(Bull. Univ. Utah Biol. Ser. 2(2):37). Clear
Crk (CI33).

Microneta uta Chamberlin 1920 (Canad.
Ent. 52:196). Logan Cyn (C20b).

Microneta viaria (Blackwall) 1841 (Trans.
Linn. Soc. London 18:645). Chalk Crk (in
Uinta Mts) (C19), Clear Crk, Dove Crk
(CI33).

Microneta sp. Zion Nat Park (CW).

Minietus aktius Chamberlin & Ivie 1935
(Bull. Univ. Utah Biol. Ser. 2(8) :23). Green
Cyn (H), Utah Lake W side (CI35b).

Mimetus hesperus Chamberlin 1923 (J.
Ent. Zool. Pomona Coll. 15:5). Grouse Crk
(CI33).

Minyriolus plesius Chamberlin 1948 (Ann.
Ent. Soc. Amer. 41:538). Cobble Rest (C48a).

Misumena vatia (Clerck) 1757 (Aranei
Suecici, p. 128). Aspen Grove (BU), Clear
Crk (CI33), St George, Zion Nat Park (CW).

Misumenoides aleatoria (Hentz) 1847 (J.
Boston Soc. Nat. Hist. 6:444). La Sal Mts
(C28a).

Misumenops asperatus (Hentz) 1847 (J.
Boston Soc. Nat. Hist. 5:447). Green Cyn (H).

Misumenops californicus (Banks) 1896 (J.
New York Ent. Soc. 4:91). Coyote Gulch
(BU).

Misumenops celer (Hentz) 1847 (J. Boston
Soc. Nat. Hist. 5:446). Clear Crk, Raft River
S fk (CI33), Curlew Valley (K), Green Cyn
(H).

Misumenops coloradensis Gertsch 1933
(Amer. Mus. Novitates 635:17). Upper Esca-
lante Basin at 10-mile Crk (BU).

Misumenops importunus (Keyserling) 1881
(Verb. zool. bot. Ges. Wien 31:307). Cedar
Jet, Kelton Pass (K).

Misumenops oblongus (Keyserling) 1880
(Spirm. Amer. Lat. 1:79). Three Lakes (N Ka-
nab) (BU).

Misumenops utanus Chamberlin 1929
(Proc. Biol. Soc. Washington 2:137). Zion Nat
Park (CW).

Misumenops varia (Keyserling) 1880
(Spinn. Amer. Lat. 1:94). USU School Forest
(W).

Misumenops sp. Calf Crk, Coyote Gulch,
Escalante, Kanab Cyn, Steep Crk, 10-mile
Crk (in Escalante Basin), Three Lakes (N Ka-
nab), Widtsoe, Willow Tank Spngs (BU).

Nanavia monticola Chamberlin & Ivie
1933 (Bull. Univ. Utah Biol. Ser. 2(2):27).
Clear Crk (CI33).

Neoanagraphis pearci Gertsch 1941 (Amer.
Mus. Novitates 1147:19). Ahlstrom Point,
Brigham Plains Flat, Four-Mile Bench (5 km
SE cow camp head Wesses Cyn), Nipple
Bench, Smokey Mt (14 & 23 km from Last
Chance Jet) (AG).

Neoantistea gosiuta Gertsch 1934 (Amer.
Mus. Novitates 712:24). Lynn 8 mi S, Raft
River S fk, Yost (G34b), Table Cliff Pass (BU).

Neoantistea magna (Keyserling) 1887
(Verh. zool. bot. Ges. Wien 37:463). Raft Riv-
er S fk, Yost (CI33a), Spring Lake (G34b).

Neoantistea sp. Glen Cyn City, Nipple
Bench (AU), USU School Forest (W).

Neon nellii G. & E. Peckham 1888 (Trans.
Wisconsin Acad. Sci. 7:88). City Crk, Jordan
River (nr Salt Lake City) (GI55), Clear Crk
(CI33), Verdure (CG28).

Neon pixii Gertsch & Ivie 1955 (Amer.
Mus. Novitates 1743:15). Salt Lake City
(GI55).

Neoscona arabesca (Walckenaer) 1841
(Hist. Nat. Ins. Apt. 2:74). Curlew Valley (K),
Green Cyn (H), St George (CW).

Neoscona benjamina (Walckenaer) 1841
(Hist. Nat. Ins. Apt. 2:42). St George (CW).

Neoscona oaxacensis (Keyserling) 1864
(Sitz.-ber. naturw. Ges. Isis, Dresden., p. 121).
Saltair Beach (C20a), Washington County
(CW), West Mt (A73).

Neoscona utahana (Chamberlin) 1919
(Ann. Ent. Soc. Amer. 12:254). Fillmore
(C19).

Neriene montana (Clerck) 1757 (Aranei
Suecici, p. 64). Clear Crk (CI33), St George,
Zion Nat Park (CW).

Nodocion eclecticus Chamberlin 1924
(Proc. Calif Acad. Sci. 12:613). Ophir
(CG40), Price, Promontory Point, Utah Lake
W side, Zion Nat Park (PS80a), Valley City
(CG28).

Nodocion moronius Chamberlin 1936
(Amer. Mus. Novitates 853:5). Moroni
(C36b).

Nodocion rufithoracicus Worley 1928
(Ann. Ent. Soc. Amer. 21:620). American
Fork Cyn mth, Mill Crk Cyn, Willard
(PS80a), Green Cyn (H).

Nodocion utus (Chamberlin) 1936 (Amer.
Mus. Novitates 841:7). Glen Cyn City 7 km

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Allred, Kaston: Utah Spiders

509

SW (AG), Richfield (C36a), Santaquin 3 mi
E, Utah Lake W side (PS80a).

Nodocion voluntarius (Chamberlin) 1919
(J. Ent. Zool. Pomona Coll. 12:5). Panguitch
11 mi SE, Red Cyn Camp (PSSOa).

Novalena idahoana (Gertsch) 1934 (Amer.
Mus. ovitates 726:25). Price, Wasatch (CI42).

Novalena lutzi (Gertsch) 1933 (Amer. Mus.
Novitates 637:12). Cedar Cyn, Horse Valley
(in Henry Mts) (CI42).

Nuctenea patagiata (Clerck) 1757 (Aranei
Suecici, p. 38). Aspen Grove, St George (BU),
Benson (McU), Dove Crk, Grouse Crk, Raft
River S fk (CI33), Lake Powell (C58), La Sal
Mts (C28a).

Oedothorax lasalanus Chamberlin & Ivie
1935 (Bull. Univ. Utah Biol. Ser. 2(8): 18). La
Sal Mts (CI35b).

Oedothorax sp. Dove Crk (CI33).

Olios fasiculatiis Simon 1880 (Acta Soc.
Linn. Bordeaux 34:307). St George (C21),
Zion Nat Park (CW).

Orchestina moaba Chamberlin & Ivie 1935
(Bull. Univ. Utah Biol. Ser. 2(8): 10). Bluff,
Moab (CI35b).

Orchestina utaliana Chamberlin & Ivie
1935 (Bull. Univ. Utah Biol. Ser. 2(8): 10).
Utah Lake W shore (CI35b).

Oreonetides vaginatus (Thorell) 1872 (Of-
vers. Kongl. Vet. Akad. Forh. 29:153). Utah
(V).

Orodrassus coloradensis (Emerton) 1877
(Bull. U.S. Geol. Surv. 3:528). Posey Lake,
Steep Crk, Widtsoe (BU), Raft River S fk
(CI33), USU School Forest (W), Uinta Mts
(C19).

Orthonops gertschi Chamberlin 1928
(Psyche 35:235). Straight Wash (in San Rafael
Desert) (C28).

Oxyopes rufipes Banks 1893 (J. New York
Ent. Soc. 1:133). lear Crk, Raft River S fk
(CI33), Kanab, Kanab Cyn, Escalante (BU), St
George (CW).

Oxyopes salticus (Hentz) 1845 (J. Boston
Soc. Nat. Hist. 5:196). St George (C21).

Oxyopes scalaris (Hentz) 1845 (J. Boston
Soc. Nat. Hist. 5:196). Green Cyn (H), Lo-
comotive Spngs (K).

Oxyopes tanneri Chamberlin 1928 (Canad.
Ent. 60:95). La Sal Mts (C28a).

Oxyopes tridens Brady 1964 (Bull. Mus.
Comp. Zool. 131:472). Glen Cyn City 5 km
W (AG).

Oxyopes sp. Escalante (BU), USU School
Forest (W).

Ozyptila heaufortensis Strand 1916 (Arch.
Naturgesch. 81A(9):124). Smith & Morehouse
Cyn (DR74).

Ozyptila conspurcata Thorell 1877 (Bull.
U.S. Geol. Surv. 3:496). Dry Cyn, Fish Lake,
Fruita, Salt Lake City, White River, Zion
Nat Park (DR74).

Ozyptila sp. Cyclone Lake (nr Escalante)
(BU).

Pachygnatha dorothea McCook 1894
(Amer. Spiders 3:270). nr Salt Lake City
(L80).

Pachygnatha mccooki Banks 1916 (Proc.
Acad. Nat. Sci. Philadelphia 68:79). Kanab
(BU).

Pachygnatha xanthostoma C. L. Koch 1845
(Die Arachn. 12:148). Grouse Crk (CI33), &
other localities not designated (L80).

Pagomys monticola (Gertsch & Mulaik)
1936 (Amer. Mus. Novitates 851:2). Mirror
Lake (CG58).

Pagomys uinta Chamberlin 1948 (Bull.
Univ. Utah Biol. Ser. 10(6): 15). Mirror Lake
(C48).

Paidisca camano Levi 1957 (Bull. Amer.
Mus. Nat. Hist. 112:105). Mill Crk Cyn (L57).

Paidisca pallida (Emerton) 1913 (Trans.
Conn. Acad. Sci. 18:213). Counties of Salt
Lake & Utah (L57).

Pardosa altamontis Chamberlin & Ivie
1946 (Bull. Univ. Utah Biol. Ser. 9(5):7). East
Cyn (CI46).

Pardosa atra Banks 1894 (J. New York Ent.
Soc. 2:52). Blanding, Devils Cyn (CG28),
Clear Crk (CI33), Salt Lake City, Zion Nat
Park (G34), Green Cyn (H), Steep Crk (BU),
USU School Forest (W).

Pardosa bellona Banks 1898 (Proc. Calif.
Acad. Sci. 1:275). Monroe Cyn (G34).

Pardosa californica Keyserling 1887 (Verh.
zool. bot. Ges. Wien 37:483). Utah (C8).

Pardosa coloradensis Banks 1894 (J. New
York Ent. Soc. 2:51). Raft River S fk (CI33).

Pardosa concinna (Thorell) 1877 (Bull. U.S.
Geol. Surv. 3:506). Sevier County (G34).

Pardosa distincta (Blackwall) 1846 (Ann.
Mag. Nat. Hist. London (1)17:32). Aspen
Grove (BU), Beaver Crk (G34), Clear Crk,
Dove Crk, Lynn, Raft River S fk, Yost (CI33).

Pardosa dorsalis Banks 1894 (J. New York
Ent. Soc. 2:51). Several localities not desig-
nated (LD81).

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Great Basin Naturalist

Vol. 43, No. 3

Pardosa dorsuncata Lowrie & Dondale
1981 (Bull. Amer. Mus. Nat. Hist. 170:130).
Several localities not designated (LD81).

Pardosa falcifera F. Pick.-Cambridge 1902
(Biol. Centr. Amer., Arachn. Aran. 2:318).
Hurricane, Salt Lake City, Zion Nat Park
(G34).

Pardosa giebeli (Pavesi) 1873 (Atti Soc.
Ital. Sci. Nat. 16:30). Navajo Mt (AG), Steep
Crk (BU), Uinta Mts (G34).

Pardosa groenlandica (Thorell) 1872 (Oefv.
Vet. Akad. Forh. 29:157). Devils Cyn, Green-
river, Moab, Monticello, Price, San Rafael
River, Valley City, Verdure (CG28), Dove
Crk, Grouse Crk, Raft River S fk, Yost
(CI33).

Pardosa lapidicina Emerton 1885 (Trans.
Conn. Acad. Sci. 6:494). Blanding, Bluff,
Fniita, Greenriver, Hanksville, Moab, San
Rafael River, Straight Wash (CG28), La Sal
Mts (C28a), St George, Zion Nat Park (CW).

Pardosa mackenziana (Keyserling) 1877
(Verb. zool. bot. Ges. Wien 26:621). Aspen
Grove, Washington County, Steep Crk (BU),
Clear Crk, Raft River S fk (CI33), La Sal Mts
(C28a).

Pardosa modica (Blackwall) 1846 (Ann.
Mag. Nat. Hist. London (1)17:33). Fish Lake
(G34), Utah Lake (C8).

Pardosa saxatilis (Hentz) 1844 (J. Boston
Soc. Nat. Hist. 4:392). Marysvale Cyn (G34).

Pardosa sierra Banks 1898 (Proc. Calif.
Acad. Sci. 1:274). Coyote Gulch (in Escalante
Basin) (BU).

Pardosa solituda Levi & Levi 1951 (Zoo-
logica 36:225). Hidden Lake, Mt Timpanogos
(LL).

Pardosa sternalis (Thorell) 1877 (Bull. U.S.
Geol. Surv. 3:504). Blanding, Fruita, Moab,
Price, Valley City, Verdure (CW), Clear Crk,
Dove Crk, Grouse Crk, Raft River S fk, Yost
(CI33), Rock Island (in Utah Lake), Steep
Crk, Three Lakes (N Kanab), Kanab (BU), St
George (C21), USU School Forest (W).

Pardosa tristis (Thorell) 1877 (Bull. U.S.
Geol. Surv. 3:510). Cyclone Lake (nr Esca-
lante) (BU).

Pardosa iiintana Gertsch 1933 (Amer. Mus.
Novitates 636:27). Uinta Mts (G33), SW Utah
(LD81).

Pardosa uncata (Thorell) 1877 (Bull. U.S.
Geol. Surv. 3:508). Undesignated localities in
Utah (LD).

Pardosa titahensis Chamberlin 1919 (Ann.
Ent. Soc. Amer. 12:258). Chalk Crk (in Uinta
Mts) (C19), Navajo Mt (AG), Steep Crk (BU).

Pardosa wasatchensis Gertsch 1933 (Amer.
Mus. Novitates 636:25). Utah (R).

Pardosa xerampelina (Keyserling) 1876
(Verb. zool. bot. Ges. Wien 26:622). Utah
(C8).

Pardosa yavapa Chamberlin 1925 (Bull.
Mus. Comp. Zool. 68:231). Fish Lake, Henry
Mts, Richfield, Weber Cyn (G34), La Sal Mts
(C28a), Boulder Mt, Steep Crk, Table Cliff
Pass (BU).

Pardosa zionis Chamberlin & Ivie 1942
(Bull. Univ. Utah Biol. Ser. 7(1):33). Coyote
Gulch (BU), Zion Nat Park (CI42a).

Pardosa sp. Calf Crk Crossing (nr Esca-
lante), Kanab Cyn, Steep Crk (BU), Four-
mile Bench (AU).

Pelecopsis digna Chamberlin & Ivie 1939
(Verb. 7 Kongr. intern. Ent. Berlin, p. 70).
Clear Crk, Uinta Mts (CI33), Provo River (in
Uinta Mts), Raft River Mts (CI39).

Pelecopsis sculptum (Emerton) 1917 (Ca-
nad. Ent. 49:261). Clear Crk, Uinta Mts
(CI33).

Pellenes americanus (Keyserling) 1884
(Verb. zool. bot. Ges. Wien 34:506). USU
School Forest (W).

Pellenes candidus G. & E. Peckham 1901
(Bull. Wisconsin Nat. Hist Soc. (N.S.) 1:206).
Rock Island (in Utah Lake) (BU), Verdure
(CG28).

Pellenes clypeatus (Banks) 1895 (Canad.
Ent. 27:102). Moab, Verdure (CG28).

Pellenes decorus (Blackwall) 1846 (Ann.
Mag. Nat. Hist. (1)17:34). Utah (C58).

Pellenes griseus G. & E. Peckham 1901
(Bull. Wisconsin Nat. Hist. Soc. (N.S.) 1:206).
Price (CG28).

Pellenes hirsutus (G. & E. Peckham) 1888
(Trans. Wisconsin Acad. Sci. 7:64). Green
Cyn (H).

Pellenes laggani G. & E. Peckham 1909
(Trans. Wisconsin Acad. Sci. 16:532). USU
School Forest (W).

Pellenes mimtda Chamberlin & Gertsch
1929 (J. Ent. Zool. Pomona Coll. 21:111). St
George (CG29).

Pellenes oregonense (G. & E. Peckham)
1888 (Trans. Wisconsin Acad. Sci. 7:66). Val-
ley City CG28).

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Allred, Kaston: Utah Spiders

511

Pellenes viridipes (Hentz) 1846 (J. Boston
See. Nat. Hist. 5:362). Verdure (CG28).

Pellenes sp. Curlew Valley (K), Four-mile
Bench, Glen Cyn City, Nipple Bench (AU).

Phidipptis apacheanns Chamberlin &
Gertsch 1929 (J. Ent. Zool. Pomona Coll.
21:109). Black Rock (W Salt Lake City), City
Crk Cyn, Fisher Pass, Strawberry (CG29),
Green Cyii (McU), Locomotive Spngs (K).

Phidippus asottis Chamberlin & Ivie 1933
(Amer. Mus. Novitates 631:50). Grouse Crk
(CI33).

Phidippus johnsoni (G. & E. Peckham)
1883 (Desc. Spiders Fam. Attidae, p. 22). As-
pen Grove, Provo, Rock Island (in Utah
Lake) (BU), Clear Crk, Dove Crk, Raft River
S fk (CI33), Green Cyn (H), Mounds, San Ra-
fael, Straight Wash (CG28), St George (C21).

Phidippus octopunctatus (G. & E. Peck-
ham) 1883 (Descr. Spiders Fam. Attidae, p.
6). Green Cyn (H).

Phidippus pogonopus Chamberlin 1925
(Proc. Calif. Acad. Sci. 14:132). Greenriver
(C25).

Phidippus pruinosus G. & E. Peckham
1909 (Trans. Wisconsin Acad. Sci. 16:388).
Blanding, Bluff (CG38).

Phidippus purpuratus Keyserling 1884
(Verh. zool. bot. Ges. Wien 34:489). USU
School Forest (W).

Phidippus workmanii (G. & E. Peckham)
1901 (Trans. Wisconsin Acad. Sci. 13:287). St
George (C21).

Phidippus sp. Glen Cyn City (AU), Posey
Lake, Provo (BU).

Philodromus alascensis Keyserling 1884
(Verh. zool. bot. Ges. Wien 33:674). Farm-
ington Cyn (LL), Salina (CG28), USU School
Forest (W).

Philodronius californicus Keyserling 1884
(Verh. zool. bot. Ges. Wien 33:676). Big Cot-
tonwood Cyn, Fruita, Mill Crk, Ogden River
Cyn, Payson, Salt Lake City (D), Bluff,
Moab, San Rafael River, Zion Nat Park
(CG28), Green Cyn (H), St George (CW).

Philodromus cespitum (Walckenaer) 1802
(Faune Parisienne, Insecta, 2:230). Curlew
Valley (K), Richfield, W Utah Lake (D).

Philodromus histrio (Latreille) 1819 (N.
Die. hist. nat. Nouv. edit., Paris, p. 36). Green
Cyn (H).

Philodromus imbecillus Keyserling 1880
(Spinn. Amer., Lat., p. 224). Paradise (DR).

Philodromus infuscatus Keyserling 1880
(Spinn. Amer., Lat., p. 222). St George (C21).

Philodromus insperatus Schick 1965 (Bull.
Amer. Mus. Nat. Hist. 129:66). East Cyn, Salt
Lake City, Silver Lake (DR68).

Philodromus pernix Blackwall 1846 (Ann.
Mag. Nat. Hist (1)17:38). Clear Crk, Raft
River S fk, Yost (CI33), Mill Crk (D).

Philodromus praelustris Keyserling 1880
(Spinn. Amer., Lat., p. 209). Allen Cyn (D).

Philodromus quercicola Schick 1965 (Bull.
Amer. Mus. Nat. Hist. 129:56). Kelton Pass
(K).

Philodromus rodecki Gertsch & Jellison
1939 (Amer. Mus. Novitates 1032:27). Red
Cyn Camp (DR68).

Philodromus rufus Walckenaer 1826
(Faune frangaise, Paris, p. 91). Green Cyn
(H), USU School Forest (W).

Phildromits satullus Keyserling 1880
(Spinn. Amer., Lat., p. 211). Green Cyn (H),
Little Cottonwood Cyn, Salt Lake City, Zion
Nat Park (DR68).

Philodromus speciosus Gertsch 1934
(Amer. Mus. Novitates 707:22). Green Cyn
(H), Salt Lake City (G34a), Vermillion Castle
(nr Parowan) (LL).

Philodromus spectabilis Keyserling 1884
(Verh. zool. bot. Ges. Wien 33:676). SW Utah
(DR76).

Philodromus sp. Escalante, Kanab Cyn,
Posey Lake, Provo, Rock Island (in Utah
Lake), Three Lakes (N Kanab), Widtsoe, Wil-
low Tank Spngs (BU).

Pholcophora americana Banks 1893 (Trans.
Amer. Ent. Soc. 23:57). Four-mile Bench (5
km SE cow camp at head Wesses Cyn) (AG),
Lehi (G35), Lynn, Raft River Mts (CI33).

Phruronellus pictus Chamberlin & Gertsch
1930 (Proc. Biol. Soc. Washington 43:138).
Bountiful (CG30).

Phruronellus similis (Banks) 1895 (J. New
York Ent. Soc. 3:81). Utah (R).

Phrurotimpus alarius (Hentz) 1847 (J. Bos-
ton Soc. Nat. Hist. 5:461). Caineville, Moab,
San Rafael River (CG28), Four-mile Bench (5
km SE cow camp at head Wesses Cyn) (AG),
Green Cyn (H).

Phrurotimpus borealis (Emerton) 1911
(Trans. Conn. Acad. Sci. 16:404). Clear Crk
(CI33).

Phrurotimpus mormon (Chamberlin &
Gertsch) 1930 (Proc. Biol. Soc. Washington

512

Great Basin Naturalist

Vol. 43, No. 3

43:140). Ferron (CI35b), Salt Lake City
(CG30).

Phrurotimpus woodburyi (Chamberlin &
Woodbury) 1929 (J. Ent. Zool. Pomona Coll.
21:18). Dry Cyn, Smith and Morehouse Cyn
(CI35b).

Physocyclus tanneri Chamberlin 1921 (Ca-
nad. Ent. 53:245). Lake Powell (C58), St
George (C21), Zion Nat Park (CW)

Piabuna nanna Chamberlin & Ivie 1933
(Bull. Univ. Utah Biol. Ser. 2(2) :41). Dove
Crk, Grouse Crk (CI33).

Piabuna xerophila Chamberlin & Ivie 1935
(Bull. Univ. Utah Biol. Ser. 2(8):38). Moab
(CI35b).

Pimoa utahana (Gertsch & Ivie) 1936
(Amer. Mus. Novitates 858:16). Salt Lake
City (CI43).

Pirata minutus Emerton 1885 (Trans.
Conn. Acad. Sci. 6:493). Ferron, Logan 1 mi
S (WE).

Pirata montanus Emerton 1885 (Trans.
Conn. Acad. Sci. 6:493). Fruita (CG28).

Pirata piraticus (Clerck) 1757 (Aranei
Suecici, p. 102). Coyote Gulch (in Escalante
Basin), Rock Island (in Utah Lake) (BU),
counties of Davis, Emery, Grand, Millard,
Rich, and Salt Lake (WE).

Pirata sedentarius Montgomery 1904
(Proc. Acad. Nat. Sci. Philadelphia 56:312).
Counties of Cache, Grand, Millard, Salt
Lake, and Utah (WE).

Pirata sp. Kanab (BU).

Pityohypliantes costatus (Hentz) 1850 (J.
Boston Soc. Nat. Hist. 6:31). Bear Lake,
Chalk Crk (in Uinta Mts), Logan Cyn (C20b),
Clear Crk, Raft river S fk (CI33).

Pityohypliantes cristatus Chamberlin &
Ivie 1942 (Bull. Univ. Utah Biol. Ser. 7(1):38)
USU School Forest (W).

Pityohypthantes navajo Chamberlin & Ivie

1942 (Bull. Univ. Utah Biol. Ser. 7(1):60).
Henry Mts (CI42a).

Pityohyphantes sp. Posey Lake (BU).

Plectreurys tristis Simon 1893 (Ann. Ent.
Soc. France 62:300). Blanding, Bluff, Devils
Cyn, Straight Wash (CG28), Nipple Bench
(AG), Zion Nat Park (CW).

Pocadicnemis pumila (Blackwall) 1841
(Trans. Linn. Soc. London 18:639). Dove
Crk, Raft River S fk (CI33), St John, Verdure
(CG28), Zion Nat Park (CW).

Poeciloneta bellona Chamberlin & Ivie

1943 (Bull. Univ. Utah Biol. Ser. 7(6): 13).
Mirror Lake (CI43).

Poeciloneta canionis Chamberlin & Ivie
1943 (Bull. Univ. Utah Biol. Ser. 7(6): 13).
Cobble Rest (CI43).

Porrhomma ocella Chamberlin & Ivie 1943
(Bull. Univ. Utah Biol. Ser. 7(6):4). City Crk
Cyn, Smith & Morehouse Cyn (CI43).

Prolinyphia litigiosa (Keyserling) 1886
(Spinn. Amer., Theridiidae 2:62). Aspen
Grove (BU), Clear Crk (CI33), Zion Nat Park
(CW).

Psilochorus imitatus Gertsch & Mulaik
1940 (Bull. Amer. Mus. Nat. Hist. 77:321).
Four-mile Bench (8 km SE cow camp at head
Wesses Cyn), Nipple Bench (AG), Salt Lake
City (GU).

Psilochorus utahensis Chamberlin 1919
(Ann. Ent. Soc. Amer. 12:247). Clear Crk,
Fillmore, Fillmore Cyn, Pine Cyn (C19),
Coyote Gulch, Cyclone Lake (nr Escalante),
Steep Crk (BU), Devils Cyn, Moab, Straight
Wash, Verdure (CG28), Ahlstrom Pt, Brig-
ham Plains Flat, Four-mile Bench (5 km SE
cow camp at head Wesses Cyn), Glen Cyn
City 5 km W, Nipple Bench, Smokey Mt (14
and 23 km from Last Chance Jet), Tibbet
Spng 2 km NE (AG), Green Cyn (H), Lake
Powell (C58), Grouse Crk, Lynn, Raft River
Mts (CI33), St George (C21).

Rhysodromus alascensis dondalei Schick
1965 (Bull. Amer. Mus. Nat. Hist. 129:73).
Utah (S).

Rhysodromus virescens (Thorell) 1877
(Bull. U.S. Geol. Surv. 3:500). Calf Crk, Pink
Dunes (N Kanab) (BU), Clear Crk, Raft River
S fk (CI33), Curlew Valley (K), Lake Powell
(C58), St George (C21).

Salticus austinensis Gertsch 1936 (Amer.
Mus. Novitates 852:20). St George (C21).

Salticus scenicus (Clerck) 1757 (Aranei
Suecici, p. 117). Benson (McU).

Salticus peckhamae (Cockerell) 1897 (Ca-
nad. Ent. 29:223). Lynn (CI33), Snowville
(K).

Sassacus papenhoei G. & E. Peckham 1895
(Occ. Pap. Nat. Hist. Soc. Wisconsin 2:177).
City Crk Cyn (CG29), Curlew Valley (K),
Green Cyn (H), Steep Crk (BU).

Satilatlas gentilis Millidge 1981 (Bull.
Amer. Mus. Nat. Hist. 170:251). Smith &
Morehouse Cyn (M81b).

Schizocosa avida (Walckenaer) 1837 (Hist.
Nat. Inst. Apt. 1:322). Bluff, Caineville, Fre-
mont River, Fruita, Moab, Straight Wash,

April 1983

Allred, Kaston: Utah Spiders

513

Valley City (CG28), Clear Crk, Raft River S
fk (CI33), Four-mile Bench (8 km SE cow
camp at head Wesses Cyn), Glen Cyn City 5
km W & 6.5 km S, Smokey Mt (23 km from
Last Chance Jet) (AG), Widtsoe (BU).

Schizocosa (?) celerior Chamberlin 1910
(Ent. News 21:2). (Not a Schizocosa accord-
ing to Dondale & Redner 1978). St. George
(GW).

Schizocosa crassipalpata Roewer
1951(Abh. naturw. Ver. Bremen 32:440). nr
Salt Lake City (DR78).

Schizocosa mccooki (Montgomery) 1904
(Proc. Acad. Nat. Sci. Philadelphia 56:283).
Green Cyn (H), Hughes Cyn (CI42a), USU
School Forest (W), other localities not desig-
nated (DR78).

Schizocosa mimula (Gertsch) 1934 (Amer.
Mus. Novitates 726:5). Salt Lake City
(DR78).

Schizocosa minnesotensis (Gertsch) 1934
(Amer. Mus. Novitates 726:4). Utah (DR78).

Schizocosa saltatrix (Hentz) 1844 (J. Boston
Soc. Nat. Hist. 4:387). Fish Lake (GW).

Sciastes simplex (Chamberlin) 1919 (Ann.
Ent. Soc. Amer. 12:250). Bear Lake, Logan
Cyn, Mirror Lake, upper Provo River, Swan
Crk Cyn, Teapot Lake (in Uinta Mts),
Wasatch Mts (CI45), Chalk Crk (in Uinta
Mts) (C19).

Scotinella formidabilis (Chamberlin &
Gertsch) 1930 (Proc. Biol. Soc. Washington
43:137). Fish Lake (CG30), Posey Lake (BU),
Raft River S fk (CI33).

Scotinella pelvicolens (Chamberlin &
Gertsch) 1930 (Proc. Biol. Soc. Washington
43:138). Clear Crk (CI33), Mt Ellen (in Hen-
ry Mts) (CG30), USU School Forest (W).

Scotinella pugnata (Emerton) 1890 (Trans.
Conn. Acad. Sci. 8:188). "Utah" (DR82).

Scotinotylus castorus (Chamberlin) 1948
(Ann. Ent. Soc. Amer. 41:520). Beaver Cyn
(C48).

Scotinotylus diibiosus Millidge 1981 (J.
Arach. 9:205). Logan (M81).

Scotinotylus kenus (Chamberlin) 1948
(Ann. Ent. Soc. Amer. 41:519). Ferron Res,
Mirror Lake (C48).

Scotinotylus pallidus (Emerton) 1882
(Trans. Conn. Acad. Sci. 6:1). Utah (M81).

Scotinotylus sacratus Millidge 1981 (J.
Arach. 9:181). Mirror Lake (M81).

Scotinotylus sanctus (Crosby) 1929 (Ent.
News 40:81). Clear Crk, Raft River S fk
(CI33).

Scotoussa bidentata (Emerton) 1882
(Trans. Conn. Acad. Sci. 6:56). Carter Crk &
Deep Crk jet, Smith & Morehouse Cyn
(CI45).

Scylaceus sp. Four-mile Bench (AU).

Sergiolus angustus (Banks) 1904 (Proc. Ca-
lif. Acad. Sci. 3:337). City Crk Cyn, Richfield
(C36b), Clear Crk Cyn, Hughes Cyn, Salt
Lake City (PS81), Fruita (CG28).

Sergiolus columbianus (Emerton) 1917
(Canad. Ent. 49:269). Salt Lake County
(PS81).

Sergiolus iviei Platnick & Shadab 1981
(Amer. Mus. Novitates 2717:34). Lynn,
Moab, Salt Lake City, West Jordan (PS81).

Sergiolus lowelli Chamberlin & Woodbury
1929 (Procf Biol. Soc. Washington 41:177).
Moab (PS81), St George (CW).

Sergiolus montanus (Emerton) 1890 (Trans.
Conn. Acad. Sci. 8:175). Clear Crk, Lynn,
Raft River Mts (CI33), Green Cyn (H), Rich-
field, Salt Lake City (C36b), Zion Nat Park
(CW), and counties of Garfield, Rich, San
Juan, Uintah, Utah, Wayne and Weber
(PS81).

Sisicottus montanus (Emerton) 1882
(Trans. Conn. Acad. Sci. 6:55). Chalk Crk (in
Uinta Mts) (C19), Clear Crk, Raft River S fk
(CI33).

Sisicottus uintanus Chamberlin & Ivie
1939 (Verb. 7 intern. Kongr. Ent. Berlin
1:65). Mirror Lake (CI39).

Sitticus finschi (L. Koch) 187,9 (Verb, zool
bot. Ges. Wien 28:483). USU School Forest
(W).

Sosticus loricatus (L. Koch) 1866 (Arach.
Earn. Drassiden, p. 131). City Crk Cyn,
Green River, Price (PS76a), Logan (CG40).

Spirembolus humilis Millidge 1980 (J.
Arach. 8:12). Mirror Lake, Tooele County
(M80).

Spirembolus monticolens (Chamberlin)
1919 (Ann. Ent. Soc. Amer. 12:251). Chalk
Crk (in Uinta Mts) (C19), Grouse Crk, Raft
River S fk (CI33), Logan Cyn, Lynn 8 mi S,
Smith & Morehouse Cyn (CI45a).

Spirembolus mundus Chamberlin & Ivie
1933 (Bull. Univ. Utah Biol. Ser. 2(2): 18).
City Crk Cyn, Raft River Mts (CI45a), Clear
Crk (CI33), Green Cyn (H).

514

Great Basin Naturalist

Vol. 43, No. 3

Spirembolus pachygnathus Chamberlin &
Ivie 1935 (Bull. Univ. Utah Biol. Ser. 2(8): 18).
City Crk Cyn, Fish Lake, Mill Crk Cyn,
Smith & Morehouse Cyn (CI45a).

Spirembolus pallidus Chamberlin & Ivie
1935 (Bull. Univ. Utah Biol. Ser 2(8): 19). Pine
Spngs (on Mt Ellen in Henry Mts) (CI45a).

Spirembolus spirotubus (Banks) 1895 (Ann.
New York Acad. Sci. 8:424). Mt Ellen (in
Henry Mts), and other localities not desig-
nated (M80).

Spirembolus vallicolens (Chamberlin) 1920
(Canad. Ent. 52:198). Fillmore, Fish Lake,
Mt Ellen (in Henry Mts), Provo River mth,
Richfield, Salt Lake City 10 mi W, Santaquin
Res (CI45a), Mill Crk (C20b), Verdure
(CG28a). (Millidge 1980 states that many of
Chamberlin and Ivies 1945a records of this
species actually are S. spirotubus.)

Steatoda albomaculata (DeGeer) 1778
(Mem. pour, servir a I'hist. des Ins. 7:257).
Clear Crk, Grouse Crk, Lynn, Raft River
Mts, Raft River S fk (CI33), La Sal Mts
(C28a), Boulder Mt, Cyclone Lake, Posey
Lake, Steep Crk (BU), Bryce Cyn Nat Park,
comities of Kane, Millard, San Juan (L57),
and Wayne (lU).

Steatoda americona (Emerton) 1882 (Trans.
Comi. Acad. Sci. 6:23). Fort Douglas, Salt
Lake City, Washington County (C28), Fruita,
Verdure (CG28), Green Cyn (H), Oquirrh
Mts (lU).

Steatoda castanea (Clerck) 1757 (Aranei
Suecici, p. 49). Logan (McU).

Steatoda fidva (Keyserling) 1882 (Spinn.
Amer. Theridiidae 2:142). Dry Cyn, Fort
Douglas, Great Salt Lake shore (lU), Ahlst-
rom Point, Brigham Plains, Glen Cyn City,
Nipple Bench, Tibbet Spng 2 km NE (AG),
Spring Lake (Ky), and counties of Carbon,
Sevier, Tooele and Weber (L57).

Steatoda grandis Banks 1901 (Proc. Acad.
Nat. Sci. Philadelphia 53:578). Beaver Dam
Wash, Utah Lake W shore (lU), Cedar Cyn
nr Cedar Breaks Nat Mon, Ferron, Henry
Mts, Levan, Salt Lake City, Zion Nat Park
(CI35b), Escalante, Fish Lake, Fruita, Help-
er, Noton, Parowan, Price, Red Cyn, Rich-
field, Scipio, Vernal, Watson, White River
(nr Evacuation Crk) (L57), St George (CW),
Steep Crk, Torrey 15 mi S (BU). (According
to Gertsh (pers. comm.), this species and S.
mexicana need clarification, and the records

above are tentatively assigned to these two
species. Steatoda mexicana actually is south-
ern Mexico in distribution, and the Utah
specimens need other names, possible only
after revision of the group.)

Steatoda hespera Chamberlin & Ivie 1933
(Bull. Univ. Utah Biol. Ser. 2(2):9). Aspen
Grove, Cyclone Lake (nr Escalante) (BU),
Clear Crk, Raft River Mts, Raft River S fk
(CI33), St George (CW), USU School Forest
(W).

Steatoda medialis (Banks) 1898 (Proc. Ca-
hf. Acad. Sci. (3)1:239). Several localities not
designated (L59).

Steatoda mexicana Levi 1957 (Bull. Mus.
Comp. Zool. 117:415). Beaver Dam Wash,
Boulder 15 mi N, Bryce Cyn Nat Park, Fish
Lake, Henry Mts, Lehi, Panguitch 10 mi SE,
Salt Lake City, St George, Utah Lake W
shore (L57). (See comments under S. grandis).

Steatoda triangulosa (Walckenaer) 1802
(Faune parisienne, Paris 2:207). Utah (LR).

Steatoda variata Gertsch 1960 (Amer. Mus.
Novitates 1982:24). Ferron, Fruita, Green-
river, Grouse Crk, Moab, Raft River Mts, Salt
Lake City, White River (on Evacuation Crk)
(G60), Glen Cyn City (AG).

Steatoda washona Gertsch 1960 (Amer.
Mus. Novitates 1982:21). Ferron, Fish Lake,
Glenwood, Loa, Moab, Richfield, Salt Lake
City (G60).

Steatoda sp. Three Lakes (N Kanab) (BU).

Stemonyphantes blauveltae Gertsch 1951
(Amer. Mus. Novitates 1514:1). Strawberry
Res (G51a).

Stylophora puUata (O. Pick. -Cambridge)
1863 (Zoologist 21:8580). Cobble Rest (169).

Synagales sp. Green Cyn (H), Cedar Hills,
Curlew Valley (K).

Tachygyna haydeni Chamberlin & Ivie
1939 (Verh. 7 intern. Kongr. Ent. Berlin, p.
63). Mirror Lake (CI39).

Tachygyna paita Chamberlin 1948 (Ann.
Ent. Soc. Amer. 41:548). Mirror Lake (C48a).

Tachygyna pallida Chamberlin & Ivie
1939 (Verh. 7 intern. Kongr. Ent. Berlin, p.
63). Provo River N fk (in Uinta Mts) (CI39).

Tachygyna tuoba (Chamberlin & Ivie)
1933 (Bull. Univ. Utah Biol. Ser. 2(2):23).
Raft River S fk (CI33), Raft River Mts (CI39).

Tachygyna watona Chamberlin 1948 (Ann.
Ent. Soc. Amer. 41:549). Mirror Lake (C48a).

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Allred, Kaston: Utah Spiders

515

Talavera mintita (Banks) 1895 (Canad. Ent.
27:99). Green Cyn (H), USU School Forest
(W).

Tapinocyba gamma Chamberlin 1948
(Ann. Ent. Soc. Amer. 41:550). Mirror Lake
(C48a).

Tetragnatha extensa (Linnaeus) 1758 (Syst.
Nat. 10th ed., p. 621). Aspen Grove, Calf Crk
Crossing (nr Escalante), Boulder (BU), Clear
Crk, Dove Crk, Raft River S fk (CI33), Cur-
lew Valley (K), Zion Nat Park (CW).

Tetragnatha laboriosa Hentz 1850 (J. Bos-
ton Soc. Nat. Hist. 6:27). Curlew Valley (K),
Dove Crk, Raft River S fk (CI33), Green Cyn
(H), Moab (CG28), Calf Crk, Kanab Cyn,
Posey Lake, Steep Crk, Three Lakes (N Ka-
nab), Utah Lake (BU), Zion Nat Park (CW).

Tetragnatha marginata (Thorell) 1890
(Ann. Mus. Civ. Stor. Nat. Genova 28:230).
Lake Powell (C58).

Tetragnatha pallescens F. Pick. — Cam-
bridge 1903. (Biol. Centr. Amer., Arachn., Ar.
2:436). NE Utah (L81).

Tetragnatha shoshone Levi 1981 (Bull.
Mus. Comp. Zool. 149:312). Bear Lake SE
shore. Granite, Laketown, Provo River mth,
Richfield, Salt Lake City, Utah Lake W side
(L81).

Tetragnatha straminea Emerton 1884
(Trans. Conn. Acad. Sci. 6:335). N central
Utah (L81).

Tetragnatha versicolor Walckenaer 1841
(Hist. Nat. Ins. Apt. 2:215). Many localities
not designated (L81).

Tetragnatha sp. Willow Tank Spngs (BU).

Thanatus altimontis Gertsch 1933 (Amer.
Mus. Novitates 636:6). Glen Cyn City (AG).

Thanatus coloradensis Keyserling 1880
(Spinn. Amer. Lat. 1:206). Clear Crk, Raft
River S fk (CI33), St George (BU).

Tlianatus formicinus (Clerck) 1757 (Aranei
Suecici, p. 134). Green Cyn (H), USU School
Forest (W).

Theridion albidum Banks 1895 (J. New
York Ent. Soc. 3:84). Green Cyn (H).

TJieridion australe Banks 1899 (Proc. Ent.
Soc. Washington 4:191). Richfield (L57).

Ttieridion berkeleyi Emerton 1924 (Pan Pa-
cif. Ent. 1:30). Counties of Salt Lake & Utah
(L57).

TJieridion denticulatum (Walckenaer) 1802
(Fauna parisienne, Paris 2:208). Utah (LR).

Theridion differens Emerton 1882 (Trans.
Conn. Acad. Sci. 6:9). Counties of Box Elder,
Grand, Rich, Salt Lake and Sevier (L57).

Theridion dihititm Levi 1957 (Bull. Amer.
Mus. Nat. Hist. 112:37). Richfield, Washing-
ton County (L57).

Theridion goodnightorum Levi 1957 (Bull.
Amer. Mus. Nat. Hist. 112:41). Counties of
Sevier and Utah (L57).

TJieridion Jiaivea Levi 1957 (Bull. Amer.
Mus. Nat. Hist. 112:48). Zion Nat Park (L57).

TJieridion leecJii Gertsch & Archer 1942
(Amer. Mus. Novitates 1171:8). Counties of
Morgan, Salt Lake and Utah (L57).

Theridion montanum Emerton 1882
(Trans. Conn. Acad. Sci. 6:10). Pickleville
(Wa), USU School Forest (W), and counties
of Beaver, Emery (L47) and Rich (L57).

TJieridion murarium Emerton 1882 (Trans.
Conn. Acad.* Sci. 6:11). Counties of Davis,
Tooele and Utah (L57).

TJieridion neomexicanum Banks 1901
(Proc. Acad. Nat. Sci. Philadelphia 53:577).
Beaver, Bryce Cyn Nat Park, counties of Car-
bon, Grand, Juab, Kane, Sevier, Wayne and
Weber (L57), Green Cyn (H), Hughes Cyn,
Logan Cyn (lU), Pickleville (Wa), St George
(CW), Straight Wash (CG28), USU School
Forest (W).

TJieridion ohlerti Thorell 1870 (Remarks
Syn. European Spiders, p. 85). Pickleville
(Wa), USU School Forest (W), counties of
Tooele and Utah (L57).

TJieridion ornatum Hahn 1831 (Monogr.
der Spinnen, H.6, T.3, 7.c). Richfield, Salt
Lake County (L57).

TJieridion petraeiim L. Koch 1872 (Zeits.
Ferd. Tirol Voral. 17:246). Green Cyn (H),
counties of Grand, Millard, Morgan, Salt
Lake, Summit, Utah and Washington (L57).

TJieridion pictum (Walckenaer) 1802
(Favine parisienne, Paris 2:207). Utah (LR).

TJieridion rabuni Chamberlin & Ivie 1944
(Bull. Univ. Utah Biol. Ser. 8(5):53). Green
Cyn (BA), Pickleville (Wa), USU School For-
est (W), counties of Morgan, Salt Lake, San
Juan and Utah (L57).

TJieridion sexpunctatum Emerton 1882
(Trans. Conn. Acad. Sci. 6:12). Clear Crk,
Raft River Mts (CI33), Zion Nat Park, coun-
ties of Salt Lake, Summit, Utah and Wasatch
(L57).

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Great Basin Naturalist

Vol. 43, No. 3

Theridion timpanogos Levi 1957 (Bull.
Amer. Mus. Nat. Hist. 112:31). American Fk
Cyn, Mt Timpanogos (L57).

Theridion transgressum Petrunckevitch
1911 (Bull. Amer. Mus. Nat. Hist. 29:208).
Ogden Cyn, Weber River, counties of Gar-
field, Washington and Utah (L57).

Theridion sp. Steep Crk (BU).

Theridula opidenta (Walckenaer) 1837
(Hist. Nat. Ins. Apt. 1:322). Utah (LR).

TInodina sylvana (Hentz) 1846 (J. Boston
Soc. Nat. Hist. 5:364). Utah (C58).

TInodina sp. Willow Tank Spngs (BU).

TJiomisiis sp. La Sal Mts (C28a).

Thymoites camano Levi 1957 (Bull. Amer.
Mus. Nat. Hist. 112:105). Mill Crk Cyn (L57).

Tfiymoites edinburgensis (Gertsch & Mu-
laik) 1936 (Amer. Mus. Novitates 863:9).
Hughes Cyn (L57).

Thymoites palUdns (Emerton) 1913 (Bull.
Amer. Mus. Nat. Hist. 32:255). Utah (LR).

Tibelhis chamberlini Gertsch 1933 (Amer.
Mus. Novitates 593:10). Elsinore, Monroe,
Zion Nat Park (G33), Green Cyn (H), Kelton
Pass (K).

TibeUus diittoni (Hentz) 1847 (J. Boston
Soc. Nat. Hist. 5:448). Lake Powell (C58), St
George (C21).

Tibelhis gertschi Chamberlin & Ivie 1942
(Bull. Univ. Utah Biol. Ser. 7(1):81). Smith &
Morehouse Cyn (CI42a).

TibeUus maritimus (Menge) 1875 (Schr. na-
turf. Ges., Danzig, N.F. 3:398). Zion Nat
Park (G33).

TibeUus oblongus (Walckenaer) 1802
(Faune parisienne, Paris 2:228). Clear Crk,
Raft River S fk (CI33), Curlew Valley (K),
Green Cyn (H), USU School Forest (W), Utah
Lake (BU).

Tibelhis sp. Moab (CG28), Kanab Cyn,
Three Lakes (N Kanab) (BU).

Titanebo magnificus Chamberlin & Ivie
1942 (Bull. Univ. Utah Biol. Ser. 7(1):81). Salt
Lake City (CI42a).

Titanebo sp. Escalante, Posey Lakes, Three
Lakes (N Kanab) (BU).

Titanoeca nigrella (Chamberlin) 1919 (J.
Ent. Zool. Pomona Coll. 12:2). Beaver 10 mi
E, Beaver Cyn, Beaver Mt, Bountiful, City
Crk Cyn, Cobble Rest, Daniel, Dry Cyn, Eu-
reka, Evacuation Crk, Farmington, Fruita,
Garden City, Glenwood, Junction, Moab,
Parleys Cyn, Richfield, Utah Lake (Le),
Green Cyn (H).

Titanoeca silvicola Chamberlin & Ivie
1947 (Bull. Univ. Utah Biol. Ser. 10(3): 15).
City Crk Cyn (C47), Fish Lake, Glendale (at
Virgin River), La Sal Forest, Snow Crk (Le).

Tmarus angulatiis (Walckenaer) 1837
(Hist. Nat. Ins. Apt. 1:537). Zion Nat Park
(CW).

Trachelas deceptus (Banks) 1895 (J. New
York Ent. Soc. 3:81). Moab, Verdure, Wayne
County (CG28), St George (CW).

Trachelas mexicanus Banks 1898 (Proc.
Calif. Acad. Sci. 1:226). St George, Zion Nat
Park (CI35b).

Trachelas tranquillus (Hentz) 1847 (J. Bos-
ton Soc. Nat. Hist 5:450). Bluff (CG28), St
George, Zion Nat Park (CW).

Trachelas sp. Three Lakes (BU).

Tricholathys spiralis Chamberlin & Ivie
1935 (Bull. Univ. Utah Biol. Ser. 2(8):28).
Kelton (K), Salt Lake City, Utah Lake W side
(CI35b).

Trochosa avara (Keyserling) 1877 (Verb,
zool. bot. Ges. Wien 27:661). La Sal Mts
(BU).

Trochosa gosiuta (Chamberlin) 1908 (Proc.
Acad. Nat. Sci. Philadelphia 60:281). Devils
Cyn, Fruita, San Rafael, Verdure (CG28), St
George, Zion Nat Park (CW), Utah Lake
(BU), counties of Carbon, Duchesne, Grand,
Salt Lake, Sevier, Tooele, and Wasatch (Br).

Trochosa terricola Thorell 1856 (Nova Acta
Reg. Soc. Sci. Upsala (3)2:171). Aspen Grove
(BU), City Crk Cyn, Lambs Cyn (CG29),
Clear Crk, Dove Crk, Grouse Crk (CI33),
Salt Lake City, Tooele (G34c), counties of
Daggett, Rich, Sevier and Summit (Br).

Trogloneta paradoxuin Gertsch 1960
(Amer. Mus. Novitates 1981:12). Timpanogos
Cave Nat Mon (G60).

Tutelina similis (Banks) 1895 (Canad. Ent.
27:100). Clear Crk (CI33), Green Canyon
(H).

Uloborus diversus Marx 1898 (Proc. Calif.
Acad. Sci. 1:234). Fillmore (C19), N, S and
central Utah (MG).

Usofila flava Chamberlin & Ivie 1942
(Bull. Univ. Utah Biol. Ser. 7(1):8). Provo Riv-
er (in Uinta Mts), Raft River Mts, Wasatch
Mts (CI42a).

Usofila gracilis Marx 1891 (Proc. Ent. Soc.
Washington 2:9). Mill Crk (G35).

Walckenaeria communis (Emerton) 1882
(Trans. Conn. Acad. Sci. 6:41). Dove Crk
(CI33), Steep Crk (BU).

April 1983

Allred, Kaston: Utah Spiders

517

Walckenaeria perditits (Chamberlin) 1948
(Ann. Ent. Soc. Amer. 41:555). Salt Lake City
(C48a).

Walckenaeria spiralis (Emerton) 1882
(Trans. Conn. Acad. Sci. 6:39). Raft River S
fk (CI33).

Walckenaeria thrinax (Chamberlin & Ivie)
1933 (Bull. Univ. Utah Biol. Ser. 2(2) :24).
Dove Crk (CI33).

Walckenaeria weber (Chamberlin) 1948
(Ann. Ent. Soc. Amer. 41:557). Smith &
Morehouse Cyn (C48a).

Walckenaeria sp. Fniita (CG28).

Wubana drassoides (Emerton) 1882 (Trans.
Conn. Acad. Sci. 6:72). Chalk Crk (in Uinta
Mts), Fillmore, La Sal Mts (CI36).

Wubana reminescens Chamberlin 1948
(Ann. Ent. Soc. Amer. 41:560). USU School
Forest (W).

Wubana utahana Chamberlin & Ivie 1936
(Ann. Ent. Soc. Amer. 29:89). Chalk Crk (in
Uinta Mts), Fish Lake, Mirror Lake, Smith &
Morehouse Cyn (CI36).

Xysticus apachecus Gertsch 1933 (Amer.
Mus. Novitates 593:22). Blanding (G3), Glen-
wood (G53).

Xysticus benefactor Keyserling 1880
(Spirm. Amer., Lat. 1:22). Bridger Basin, Brig-
ham Cyn, Logan (G53).

Xysticus californicus Keyserling 1880
(Spinn. Amer., Lat. 1:37). Brigham, Clarks-
ton, Escalante 22 mi N, Fish Lake, Logan,
Ogden, Parowan, Salina, Willard, Zion Nat
Park (G53), Grouse Crk, Raft River S fk,
Lynn (CI33), St George (CG29), Straight
Wash, Verdure (CG28), Green Cyn (H), Utah
Lake (BU).

Xysticus coloradensis Bryant 1930 (Psyche
37:133). Tooele County (G53).

Xysticus facetus O. Pick. -Cambridge 1896
(Biol. Centr. Amer., Arach. Ar. 1:179). Moab
(G53).

Xysticus ferox (Hentz) 1847 (J. Boston Soc.
Nat. Hist 5:445). Bluff, Fruita, Moab, Price,
San Rafael River (CG28).

Xysticus gertschi Schick 1965 (Bull. Amer.
Mus. Nat. Hist. 129:159). Clear Crk Cyn, El-
sinore, Monroe Cyn (G34a).

Xysticus gosiutus Gertsch 1933 (Amer.
Mus. Novitates 593:20). Little Cottonwood
Cyn, Zion Nat Park (G33).

Xysticus gulosus Keyserling 1880 (Spinn.
Amer., Lat. 1:43). Blanding (CG28), Glen

Cyn City (AG), Kelton Pass (K), Lake Powell
(C58), Provo (BU), St George (CW).

Xysticus imitarius Gertsch 1953 (Bull.
Amer. Mus. Nat. Hist. 102:442). Little Cot-
tonwood Cyn (G53).

Xysticus knowltoni Gertsch 1939 (Bull.
Amer. Mus. Nat. Hist. 76:399). Vernon (G53).

Xysticus lassanus Chamberlin 1925 (Bull.
Mus. Comp. Zool. 67:218). Glen Cyn City,
Smokey Mt (AG), Hanksville, St George
(G53).

Xysticus locuples Keyserling 1880 (Spinn.
Amer., Lat. 1:24). Brigham (G53), Green Cyn
(McU), Grouse Crk (CI33).

Xysticus lutulentus Gertsch 1939 (Bull.
Amer. Mus. Nat. Hist. 76:396). Mill Crk Cyn
(CI42a).

Xysticus lutzi Gertsch 1935 (Amer. Mus.
Novitates 792:27). Glen Cyn City (AG).

Xysticus '^ montanensis Keyserling 1887
(Verb. zool. bot. Ges. Wien 37:479). Clear
Crk (CI33), Ferron, Salt Lake City (G34a),
Green Cyn (H), Kelton (K), USU School For-
est (W).

Xysticus Orizaba Banks 1898 (Proc. Calif.
Acad. Sci. 1:260). Beaver Dam Wash (G53),
Little Cottonwood Cyn, St George (G33).

Xysticus pellax O. Pick.-Cambridge 1894
(Bio. Centr. Amer., Arach. Ar. 1:138). Kearns
(G53).

Xysticus sp. Coyote Gulch, Steep Crk,
Three Lakes (N Kanab), 10-mi Crk (in Esca-
lante Basin) (BU), USU School Forest (W).

Zanomys kaiba Chamberlin 1948 (Bull.
Univ. Utah Biol. Ser. 10(6):18). Dry Cyn
(C48).

Zanomys ochra Leech 1972 (Mem. Canad.
Ent. Soc. 84:90). Lynndyl 10 mi N (Le).

Zelotes fratris Chamberlin 1920 (Canad.
Ent. 2:193). Blanding, Bluff, Grantsville, Ver-
dure (CG28), Clear Crk, Raft River Mts, Raft
River S fk, Yost (CI33), Curlew Valley (K),
Fish Lake, Monroe Cyn, Richfield (C36a),
Green Cyn (H), Logan Cyn (C20b), St
George (CW).

Zelotes latnpra Chamberlin 1920 (Canad.
Ent. 52:193). Mill Crk (C20b).

Zelotes lasalanus Chamberlin 1928 (Ca-
nad. Ent. 60:93). La Sal Mts (C28a).

Zelotes nannodes Chamberlin 1936 (Amer.
Mus. Novitates 853:10). Tremonton 10 mi W
(C36b).

518

Great Basin Naturalist

Vol. 43, No. 3

Zelotes nanniis Chamberlin & Gertsch
1940 (Amer. Mus. Novitates 1068:18). Bluff,
Richardson (CG40).

Zelotes puritanus Chamberlin 1922 (Proc.
Biol. Soc. Washington 35:164). Puffer Lake,
Tooele Cyn (C36a).

Zelotes tuohus Chamberlin 1919 (Ann. Ent.
Soc. Amer. 12:247). Ahlstrom Point, Brigham
Plains (AG), Dove Crk, Raft River Mts
(CI33), Fillmore (C19), USU School Forest
(W).

Zelotes sp. Brighams Plains, Glen Cyn City
(AU), Kelton, Locomotive Spngs (K), St
George, Tliree Lakes (N Kanab) (BU).

Zornella ctiltrigera (L. Koch) 1879 (Kongl.
Svenska Vet. Akad. Handl. 16:11). USU
School Forest (W).

Synonymies of Utah Records

In the list below, the names on the left are
as recorded for some specimens in collections
of Utah universities, or as listed in some pub-
lished articles, and are considered as junior
synonyms of the names on the right, which
are the only ones included in the main body
of this report.

Aculepeira verae = A. packardi
Agelena californica = Agelenopsis

califomica
Agelena hola = Hololena hola
Agelena mimoides = Hololena mimoides
Agelena oquirrhensis = Hololena

oquirrhensis
Agelenopsis mimoides = Hololena mimoides
Agelenopsis naevia = A. aperta
Agroeca oaba = A. trivittata
Alopecosa helluo = Lycosa helluo
Amaurobius nevadensis = Callobius

nevadensis
Amaurobius nomeus = Callobius nomeus
Amaurobius utahensis = Callobius

nevadensis
Aranea carbonaria = Aculepeira carbonaria
Aranea cucurbitina = Araniella displicata
Aranea displicata = Araniella displicata
Aranea ocellatula = Nuctenea patagiata
Areanea ocellatus = Nuctenea patagiata
Aranea solitaria = Araneus saevus
Aranea tusigia = Araneus marmoreus
Aranea utahana = Neoscona utahana
Arctachaea pelyx = Chrysso pelyx
Arctosa cinerea = A. littoralis

Aysha nigrifrons = A. incursa
Bathyphantes fillmoranus = Meioneta

fillmorana
Bathyphantes phylax = Helophora orinoma
Bathyphantes spatulifer = Leptyphantes

lamprus
Ceraticelus guttatus = Idionella anomala
Ceraticelus tuganus = Idionella tugana
Cheraira castoris = Scotinotylus castoris
Cheraira kena = Scotinotylus kenus
Cicurina garrina = C. robusta
Clubiona orinoma = C. moesta
Cochembolus sanctus = Scotinotylus sanctus
Cornicularia communis = Walckenaeria

communis
Cornicularia thrinax = Walckenaeria thrinax
Cylphosa gosoga = Gnaphosa gosoga
Cylphosa sericata = Gnaphosa sericata
Delopelma simulatum = Aphonopelma

Simula tum
Dendryphantes diplacis = Metaphidippus

diplacis
Dendryphantes mylothrus = prob.

Metaphidippus mylothrus
Dendryphantes nigromaculatus = Kris

nigromaculatus
Dendryphantes pruinosus = Phidippus

pniinosus
Dendryphantes unicus = Metaphidippus

unicus
Dendryphantes uteanus = prob.

Metaphidippus uteanus
Dendryphantes verecundus =

Metaphidippus verecundus
Dendryphantes workmanii = Phidippus

workmanii
Dictyna dactylata = D. calcarata
Dictyna eutypa = Mallos eutypus
Dictyna hoples = D. calcarata
Dictyna socarina = D. uintana
Diplocentria bidentata = Scotoussa

bidentata
Diplocentria perplexa = Maso perplexus
Dipoena daltoni = D. atopa
Dipoena hamata = D. prona
Drassodes celes = D. saccatus
Drassodes robinsoni = D. saccatus
Drassyllus apacheus = D. insularis
Drassyllus devexus = D. dromeus
Drassyllus gertschi = D. conformans
Drassyllus lasalus = D. mexicanus
Drassyllus mephisto = D. lepidus
Emblyna completa = Dictyna completa

April 1983

Allred, Kaston: Utah Spiders

519

Emblyna rena = Dictyna completa
Emblyna reticulata = Dictyna reticulata
Emblyna urica = Mallos niveus
Emblyna utesca = Dictyna piratica
Epeira labyrinthea = Metapeira labyrinthea
Eperigone taibo = Tachygyna tuobo
Eularia simplex = Sciastes simplex
Euryopis nigripes = E. taczanowskii
Evarcha leucophaea = E. hoyi
Frontinella communis = F. pyramitela
Fuentes taeniola = Metacyrba taeniola
Garritus vigerens = Ctenium vigerens
Gayenna saniuana = Anyphaena pacifica
Geodrassus gosiutus = Drassodes gosiutus
Geolycosus carolinensis = Lycosa

carolinensis
Gertschia sp. = Synagales sp.
Gnaphosa gigantea = G. muscorum
Hahnia inomata = H. ononidium
Haplodrassus dystactus = H. signifer
Haplodrassus uncifer = H. bicornis
Haplodrassus utus = H. eunis
Herpyllus atopophysis = Nodocion

eclecticus
Herpyllus piedicus = H. propinquus
Herpyllus validus = H. hesperolus
Herpyllus vasifer = H. ecclesiasticus
Icius similis = Tutelina similis
Labuella utahana = Pimoa utahana
Lathys moabana = Dictyna moaba
Latrodectus curacaviensis = L. hesperus
Latrodectus geometricus = L. hesperus
Latrodectus mactans = L. hesperus
Latrodectus variolus = L. hesperus
Linyphia communis = Frontinella communis
Linyphia ephedra = Linyphantes ephedrus
Linyphia litigiosa = Prolinyphia litigiosa
Linyphia montana = Neriene montana
Linyphia phrygianus = Pityohyphantes

costatus
Liodrassus metalleus = Nodocion eclecticus
Liodrassus utus = Nodocion utus
Lithyphantes albomaculatus = Steatoda

albomaculata
Lycosa avara = Trochosa avara
Lycosa gosiuta = Trochosa gosiuta
Lycosa orophila = Trochosa terricola
Lycosa piraticus = Pirata piraticus
Lycosa pratensis = Trochosa terricola
Lycosa rafaelana = Geolycosa rafaelana
Meriola sp. = Trachelas sp.
Metaphidippus nigromaculatus = Eris

nigromaculatus
Metaphidippus verecundus = M. diplacis

Metastinus oblongus = Tibellus oblongus
Metepeira alpina = M. grandiosa alpina
Misumenops lepidus = M. celer
Neoantistea riparia = N. magna
Neoscona naiba = N. arabesca
Paraphidippus marginatus = Eris marginatus
Pardosa atromedia = P. sierra
Pardosa wyuta = P. atra
Peocilochroa montana = Sergiolus montanus
Peponocranium pumila = Pocadicnemis

pumila
Phidippus borealis = P. purpuratus
Phidippus capitatus = Metaphidippus

galathea
Phidippus formosus = P. johnsoni
Philodromus cespiticolis = P. cespitum
Philodromus hoples = P. californicus
Philodromus virescens = Rhysodromus

virescens
Pirata sylvestris = P. piraticus (part) & P.

insularis (part)
Pityohyphantes phrygianus = P. costatus
Platyxysticus utahensis = Coriarachne

versicolor (part) & C. utahensis (part)
Prosopotheca sp. = Walckenaeria sp.
Pselothorax atopus = Dipoena atopus
Pterotrichia clara = Gnaphosa clara
Pusillia mandibulata = Microlinyphia

mandibulata
Robertus fuscus = Ctenium fusca
Robertus eremphilus = Ctenium

eremophilus
Sassacus uteanus = S. papenhoei
Schizocosa wasatchensis = S. mccooki
Sergiolus clarus = S. angustus
Sergiolus fruitanus = S. angustus
Singa variabilis = Hyposinga pygmaea
Sostogeus zygethus = Sosticus loricatus
Spirembolus chera = Scotinotylus sanctus
Steatoda punctulata = S. medialis
Tapinocyba alpha = Disembolus alpha
Tapinocyba kesimba = Disembolus kesimbus
Teutana castanea = Steatoda castanea
Theridion placens = T. neomexicanum
Theridion pygmaea = Hyposinga pygmaea
Theridium canione = Achaearanea canione
Tigellinus weber = Walckenaeria weber
Tigellinus perditus = Walckenaeria perditus
Tosyna calcarata = Dictyna calcarata
Tosyna cholla = Dictyna cholla
Tosyna terrestris = Dictyna terrestris
Trachelas utahanus = T. mexicanus
Tricholathys reclusa = Argennina reclusa
Trochosa frondicola = Lycosa frondicola

520

Great Basin Naturalist

Vol. 43, No. 3

Trochosa pratensis = T. terricola
Uloborus utahensis = U. diversus
Xysticiis ciinctator = X. californicus
Xysticus quinquepunctatus = X. gertschi
Zelotes pananus = Sergiolus angustus
Zelotes subterraneus = Z. fratris

Literature Cited

Allred, D. M. 1973. An unusual population of spiders

in Utah. Great Basin Nat. 33:51-2.
1974. An unusual spider bite. Great Basin Nat.

34:82.
1975. Arachnids as ecological indicators. Great

Basin Nat. .35:405-6.
1978. Bite of wolf spider. Great Basin Nat.

38:478.
1980. A Chiracanthitim spider bite. Great Basin

Nat. 40:116.
Allred, D. M. and W. J. Gertsch. 1976. Spiders and

scorpions from northern Arizona and southern

Utah. J. Arach. 3:87-99.
Archer, A. F. 1951. Studies in the orb-weaving spiders.

Part 1. American Mus. Novitates 1487. 52 pp.
Bowling, T. A., and R. J. Sauer. 1975. A taxononiic re-
vision of the crab spider genus Coriarachne for

North America north of Mexico. J. Arach.

2:183-93.
Brady, A. R. 1979. Nearctic species of the wolf spider

genus Trochosa. Psyche 86:167-212.
Chamberlin, R. V. 1908. Revision of North American

spiders of the family Lvcosidae. Proc. Acad. Nat.

Sci. Phila. 60: 158-318. '
1919. New western spiders. Ann. Ent. Soc. Amer-
ica 12:239-60.
1920. A new American Amaurobius. Ent. News

31:293.
1920a. The spider of Saltair Beach. Ent. News

31:165-9.
1920b. New spiders from Utah. Candadian Ent.

52:193-201.
1921. A new genus and a new species of spiders

in the group Phrurolitheae. Canadian Ent.

53:69-70.
1921a. On some arachnids from southern Utah.

Canadian Ent. 53:245-7.
1922. The North American spiders of the family

Gnaphosidae. Proc. Biol. Soc. Washington

35:145-72.
1925. Diagnosis of new American Arachnida.

Harvard Univ. Bull. Mus. Comp. Zool.

67:221-48.
1925a. New North American spiders. Proc. Calif.

Acad. Sci. 14:105-42.
1928. A two-eyed spider from Utah. Psvche

35:235-6.
1928a. Notes on the spiders from the La Sal

Mountains of Utah. Canadian Ent. 60:93-5.
1933. Four new spiders of the family Gnapho-
sidae. American Mus. Novitates 631. 8 pp.
1936. A new burrowing spider of the genus Ly-

cosa from the Uinta Mountains, Utah. Proc. Biol.

Soc. Wash. 49:15-16.

1936a. Records of North American Gnaphosidae

with descriptions of new species. American .Mus.

Novitates 841. .30 pp.
1936b. Further records and descriptions of North

American Gnaphosidae. American Mus. Novi-
tates 853. 21 pp.
1940. New American tarantulas of the family

Aviculariidae. Bull. Univ. Utah Biol. Ser. 5(8). 39

pp.
1947. A summary of the known North American

Amaurobiidae. Bull. Univ. Utah Biol. Ser. 10(5).

31pp.
1948. The genera of North American Dictynidae.

Bull. Univ. Utah Biol. Ser. 10(6). 31 pp.
1948a. On some spiders of the family Erigonidae.

Ann. Ent. Soc. America 41:483-562.
1958. Araneida. In Preliminary report on biologi-
cal resources of the Glen Canyon Reservoir.

Univ. Utah Anthropological Papers 31:167-71.
Chamberlin, R. V., and W. J. Gertsch. 1928. New spi-
ders from Utah and California. Pomona College

J. Ent. and Zool. 21:101-12.
1928a. Notes on spiders from southeastern Utah.

Proc. Biol. Soc. America 41:175-88.
19.30. On fifteen new North American spiders.

Proc. Biol. Soc. Washington 43:137-44.
1940. Descriptions of new Gnaphosidae from the

United States. American Mus. Novitates 1068. 19

pp.
1958. The spider family Dictvnidae in America

north of Mexico. Bull. American Mus. Nat. Hist.

116:1-152.
Chamberlin, R. V., and W. Ivie. 1933. Spiders of the

Raft River Mountains of Utah. Bull. Univ. Utah

Biol. Ser. 2(2). 79 pp.
1933a. A new genus in the family Agelenidae.

Bull. Univ. Utah Biol. Ser. 2(3). 7 pp.
1935. Nearctic spiders of the family Urocteidae.

Ann. Ent. Soc. America 28:265-79.
1935a. The black widow spider and its varieties

in the United States. Bull. Univ. Utali Biol. Ser.

3(1). 29 pp.
1935b. Miscellaneous new American spiders.

Bull. Univ. Utah Biol. Ser. 2(8). 79 pp.
1936. Nearctic spiders of the genus Wubana.

Ann. Ent. Soc. America 29:85-98.
1937. New spiders of the family Agelenidae from

western North America. Ann. Ent. Soc. America

.30:211-41.
1939. Studies on North American spiders of the

family Micrvphantidae. Verb. VII Int. Kongr.

Ent. 1:56-73.'
1939a. New tarantulas from the southwestern

states. Bull. Univ. Utah Biol. Ser. 5(1). 17 pp.
1940. Agelenid spiders of the genus Ciciirma

Bull. Univ. Utah Biol. Ser. 5(9). 108 pp.
1941. North American Agelenidae of the genera

Agelenopsis, Calilena, Ritalena, and Tortolena.

Ann. Ent. Soc. America 34:585-628.
1941a. Spiders collected by L. W. Saylor and

others, mostly in California. Bull. Univ. Utah

Biol. Ser. 6(3). 49 pp. _
1942. Agelenidae of the genera Hololena, Nova- fl

lena, Rualena, and Melpomene. Ann. Ent. Soc. ^

America 35:203-41.

April 1983

Allred, Kaston: Utah Spiders

521

1942a. A hundred new species of American spi-
ders. Bull. Univ. Utah Biol. Ser. 7(1). 117 pp.

1943. New genera and species of North American

linvphiid spiders. Bull. Univ. Utah Biol. Ser. 7(6).
39 pp.

1945. Some erigonid spiders of the genera Eularki

and Diploccntrki. Bull. Univ. Utah Biol. Ser. 9(4).

19 pp.

1945a. Erigonid spiders of the genera Spirein-

bohis, Disembohis. and Bactroceps. Trans. Conn.
Acad. Arts and Sci. 36:215-35.

1946. On .several new American spiders. Bull.

Univ. Utah Biol. Ser. 9(5). 15 pp.

1947. North American dictynid spiders: the ben-

netti group of Amaurobius. Ann. Ent. Soc. Amer-
ica 40:29-55.

Chamberlin, R. v., and A. M. Woodbury. 1929. Notes
on the spiders of Washington County, Utah.
Proc. Biol. Soc. Washington 42:131-42.

DoNDALE, C. D. 1961. Revision of the aureolus group of
the genus Philodromus in North America. Cana-
dian Ent. 93:199-222.

DoNDALE, C. D., AND J. H. Redner. 1968. The imbe-
cillus and nifus group of the spider genus Philo-
dromus in North America. Ent. Soc. Canada,
Memoir 55. 78 pp.

1975. The genus Ozyptila in North America. J.

Arach. 2:129-81.

1976. A review of the spider genus Philodromus

in the Americas. Canadian Ent. 108:127-57.

1978. Revision of the Nearctic Wolf Spider genus

Schizocosa. Canadian Ent. 110:143-81.

1979. Revision of the wolf spider genus Alopecosa

Simon in North America. Canadian Ent.
111:1033-55.

1982. The sac spiders of Canada and Alaska. Can-
ada Dept. Agr. Bull. 1724. 194 pp.

ExLiNE, H. 1936. Nearctic spiders of the genus Ciciirina
Menge. American Mus. Novitates 850. 25 pp.

Gertsch, W. J. 1932. A new generic name for Coria-
rachne versicolor Keyserling, with new species.
American Mus. Novitates 563. 7 pp.

1933. Notes on American spiders of the family

Thomisidae. American Mus. Novitates .593. 22
pp.

19.34. Notes on American Lycosidae. American

Mus. Novitates 693. 25 pp.

19.34a. Notes on American crab spiders. Ameri-
can Mus. Novitates 707. 25 pp.

19.34b. Some American spiders of the family Hah-

niidae. American Mus. Novitates 712. 32 pp.

1934c. Further notes on American spiders. Amer-
ican Mus. Novitates 726. 26 pp.

1935. Spiders from the southwestern United

States, with descriptions of new species. Ameri-
can Mus. Novitates 792. 29 pp.

1941. New American spiders of the familv Club-

ionidae. Part 1. American Mus. Novitates 1147.

20 pp.

1942. New American spiders of the family Club-

ionidae. Part III. American Mus. Novitates 1195.

18 pp.
1946. Notes on American spiders of the familv

Dictynidae. American Mus. Novitates 1319. 21

pp.

1953. The spider genera Xijsticus, Coriarachne,

and Oxtjptila (Thomisidae, Misumeninae) in
North America. Bull. American Mus. Nat. Hist.
102:417-82.

1960. Descriptions of American spiders of the

family Smyphytognathidae. American Mus. Novi-
tates 198l' 40 pp.

Gertsch, W. J., and W. Ivie. 1936. Descriptions of new
American spiders. American Mus. Novitates 858.
21pp.

1955. The spider genus Neon in North America.

American Mus. Novitates 1743. 17 pp.

Gertsh, W. J., and F. E. Russell. 1975. Loxosceles
deserta Gertsch. Toxicon 13:203-4.

Gertsch, W. J., and H. K. Wallace. 1937. New Ameri-
can Lycosidae with notes on other species. Amer-
ican Mus. Novitates 919. 22 pp.

Hatley, C. L. 1978. The role of vegetation architecture
in determining spider community organization.
Unpubublished thesis, Utah State Univ., Logan.

Helsdingen, p. J. VAN 1981. The Nearctic species of
Oreonetides. Bull. Amer. Mus. Nat. Hist.
170:229-41.

IviE, W. 1967.' Some synonyms in American spiders. J.
New York Ent. Soc. 75:126-31.

1969. North American spiders of the genus Batli-

ijphantes. American Mus. Novitates 2364. 70 pp.

Kaston, B. J. 1946. North American spiders of the genus
Ctenium. American Mus. Novitates 1.306. 19 pp.

Knowlton, G. F. 1971. Some Curlew Valley spiders.
Utah State Univ. Ecol. Center, Terrestrial Ar-
thropods, Mimeo. Ser. 2. 5 pp.

Leech, R. 1972. A revision of the Nearctic Amauro-
biidae. Canadian Ent. Soc. Memoir 84. 182 pp.

Levi, H. W. 1953. New and rare Dipoena from Mexico
and Central America. American Mus. Novitates
1639. 11 pp.

1954. Spiders of the genus Eurijopis from North

and Central America. American Mus. Novitates
1666. 48 pp.

1955. The spider genera Coressa and Achaea-

ranea in America north of Mexico. American
Mus. Novitates 1718. 33 pp.

1957. The spider genera Enoplognatha, Theri-

dion, and Paedisca in America north of Mexico.
Bull. Amer. Mus. Nat. Hist. 112:1-12.

1958. Spiders of the genus Arctachaea. Psyche

64:102-6.

1959. Problems in the spider genus Steatoda. Syst.

Zool. 8:107-16.

1963. American spiders of the genus Achaearanea

and the new genus Echinotheridion. Harvard
Univ. Bull. Mus. Comp. Zool. 129:187-240.

1975. Additional notes on the orb-weaver genera

Araneus, Hypsosinga, and Singa north of Mexico.
Psyche 82:265-74.

1975a. The American orb-weaver genera Larinia,

Cercidia, and Mangora north of Mexico. Harvard
Univ. Bull. Mus. Comp. Zool. 147:101-35.

1980. The orb-weaver genus Mecijnogea, the sub-

familv Metinae and the genera Pachijgmitha, Gle-
nognatlia, and Azilia of the subfamily Tetragna-
thinae north of Mexico. Harvard Univ. Bull. Mus.
Comp. Zool. 149:1-75.

522

Great Basin Naturalist

Vol. 43, No. 3

1981. The American orb-weaver genera Dolichog-

natha and Tetragnatha north of Mexico. Harvard
Univ. Bull. Mus. Comp. Zool. 149:271-318.

. 1981a. More on the genus Araneus from North

America. Bull. Amer. Mus. Nat. Hist. 170:254-6.

Levi, H. W., and L. R. Levi. 1951. Report on a collec-
tion of spiders and harvestmen from Wyoming
and neighboring states. Zoologica .36:219-37.

Levi, H. W., and D. E. Randolph. 1975. A key and
checklist of American spiders of the family Theri-
diidae north of Mexico. J. Arach. 3:31-51.

LowRiE, D. C, and C. D. Dondale. 1981. A revision of
the nigra group of the genus Pardosa in North
America. Bull. Amer. Mus. Nat. Hist. 170:125-39.

MiLLiDGE, A. F. 1980. The erigonine spiders of North
America. Part 2. The genus Spirembolus Cham-
berlin. J. Arach. 8:109-58.

1981. The erigonine spiders of North America.

Part 3. The genus Scotinotyhis Simon. J. Arach.
9:167-213.

1981a. The erigonine spiders of North America.

Part 4. The genus Disembolus Chamberlin and
Ivie. J. Arach. 9:259-84.

1981b. The erigonine spiders of North America.

Part 5. The genus Satilathis. Bull. Amer. Mus.
Nat. Hi.st. 170:243-53.

(In press) The erigonine spiders of North Ameri-
ca. Part 6. The genus Wolckenaeha Blackwall. J.
Arach.

MuMA, M. H., and W. J. Gertsch. 1964. The spider
family Uloboridae in North America north of
Mexico. American Mus. Novitates 2196. 43 pp.

Platnick, N. L and M. U. Shadab. 1975. A revision of
the spider genus Gnaphosa in America. Bull.
Amer. Mus. Nat. Hist. 155:1-66.

1976. A revision of the spider genera Drassodes

and Tivodrassus in North America. Amer. Mus.

Novitates 2593. 29 pp.
1976a. A revision of the spider genera Rach-

odrassus, Sosticiis, and Scopodes. Amer. Mus.

Novitates 2594. 33 pp.
1977. A revision of the spider genera Herpi/Uiis

and Scotophaeits in North America. Bull. Amer.

Mus. Nat. Hist. 159:1-44.
1980. A revision of the spider genus Cesonia.

Bull. Amer. Mus. Nat. Hist. 165:337-85.
1980a. A revision of the North American spider

genera Nodocion Litopyhis, and Synaphosus.

Amer. Mus. Novitates 2691. 26 pp.
1981. A revision of the spider genus Sergiolus.

Amer. Mus. Novitates 2717. 41 pp.
1982. A revision of the American spiders of the

genus Drassyllus. Bull. Amer. Mus. Nat. Hist.

173:1-97.
(In press) A revision of the American species of

the genus Zetotes. Bidl. Amer. Mus. Nat. Hist.
Sauer, R. J., and N. I. Platnick. 1972. The crab spider

genus Ebo in the United States and Canada. Ca-
nadian Ent. 104:.35-60.
Schick, R. X. 1965. The crab spiders of California. Bull.

Amer. Mus. Nat. Hist. 129:1-180.
Waagen, G. N. 1979. Changes in spider community at-
tributes along a subalpine successional gradient.

Unpubublished dissertation, Utah State Univ.^

Logan.
Wallace, H. K., and H. Exline. 1977. Spiders of the

genus Pirata in North America, Central America,

and the West Indies. J. Arach. 5:1-112.

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1-84 650 68551

ISSN 017-3614

The Great Basin Naturalist

Published at Provo, Utah, by
Brigham Young University

ISSN 0017-3614

Volume 43

October 31, 1983

No. 4

ALPINE AND SUBALPINE WETLAND PLANT COMMUNITIES
OF THE UINTA MOUNTAINS, UTAH

George M. Briggs' and James A. MacMahon-

Abstract.— Seven wetland areas in the subalpine and alpine region^ of the Uinta Mountains are described. Most
areas have a mosaic vegetation structure, comprised of several patches that are usually sharply demarcated from one
another. Each patch has a distinct species composition, usually with only one or two species. Water level is associ-
ated with some of the vegetation patterns found in these sites. Standing crop varied from 28 to 360 g/m^. Sites show
considerable variation in standing crop that could not be explained by elevation differences between sites. Sites with
water flowing over them have substantially greater standing crops than sites where water stagnates. Seasonal pat-
terns in shoot density and standing crop indicate one late summer peak in standing crop and little recruitment of
shoots over the summer. On stagnant sites, the average stem weight shows a strong relationship to stem density. This
pattern did not appear to be caused by thinning mortality and did not follow a "3/2 power law" pattern.

A conspicuous aspect of many of the west-
em North American mountain ranges is their
wetlands. The Uinta Mountains of north-
eastern Utah have an abundance of wetland
areas, from the lower elevations (around
streams in sagebrush slopes) to areas near
springs at high elevations in the alpine zone.
The majority of wetlands are found at the
heads of glaciated valleys in a zone just
above and below treeline. The vegetation in
these areas is dominated by members of the
Cyperaceae (sedges), as is much of the upland
region in the alpine zone of the Uintas. Al-
though both wetland and upland regions are
dominated by Carex spp., the structure of the
two regions is markedly different (Briggs and
MacMahon 1982). In this study we describe a
variety of sedge-dominated wetlands, detail
some of their structural attributes, and dis-
cuss some of the factors that we think are im-
portant in determining their patterns of
occurrence.

Study Sites and Methods

All sites are in the Uinta Mountains of
northeastern Utah (40°45'N, 110°-111°W),
the largest east-west trending mountain range
in North America. Bedrock throughout the
upper elevations of the range is quartzite.
The range was heavily glaciated in Pleisto-
cene time (Hansen 1975) and is dominated by
numerous large U-shaped valleys ending
abruptly in steep-walled cirques.

No attempt was made to describe or cata-
log all the wetland areas of the Uintas. In-
stead, we chose to study a variety of wetland
regions that varied in their species composi-
tion and standing crop. Most of these regions
had a patchwork appearance because they
were composed of regions (2-100 m^) of dis-
tinct species composition, often monotypic,
which abruptly gave way to regions of a dif-
ferent species mixture. In sampling these
areas, we first made a rough map that demar-
cated the "patches" found in them. Each

'University of Michigan Biological Station, Pellston, Michigan 49769.
'Department of Biology, Utah State University, Logan, Utah 84322.

523

524

Great Basin Naturalist

Vol. 43, No. 4

patch was then sampled by the methods out-
lined below. We refer to each patch as a site
and present figures representing the species
composition and standing crop of each of
these. The larger units of vegetation (the en-
tire meadow composed of several patches)
are not quantitatively described in this paper.
Each site was given a two-letter code that
represented the area from which it came and
a number that indicated the specific patch
sampled (e.g., SC-1 is patch 1 in area SC).

The boundaries to each site were sub-
jectively demarcated and a point was picked
within the stand. A 20 X 50 cm frame was
placed at that point and all vegetation (ex-
cept bryophytes) was clipped at ground level.
Nine additional samples were taken at regu-
lar intervals from the first point. These sam-
ples were sorted to species in the field, re-
turned to the lab, air dried for at least two
months, and oven dried at 40 C for at least
24 h. Sorting to species was quite easy, even
when based on vegetative characters, since
there were rarely more than two species per
sample. For larger species, the number of
stems was directly counted and the average
weight per stem calculated. For the smaller
species, the average weight per stem was de-
termined by weighing three replicates of 100
stems. The total number of stems was obtain-
ed by dividing the total weight of that spe-
cies in the sample by the weight/ stem. All
sites were sampled in August, when the fruits
of the dominant species were mature. Sea-
sonal changes in aboveground standing crop,
number of shoots, and number of fruiting
shoots were determined in three of the sites
by sampling three or four times during the
summer. In addition, several individual plants
were observed. The length of each leaf and
the length and condition of the fruiting culm
(if present) were measured at each sampling
time.

At each site, soil samples were dug and pH
of the soil was determined. All these sites
were submerged at least part of the summer,
and notes were made of both the length of
time the site was submerged and the degree
of water movement over the site.

Site Descriptions

Seven wetland areas were mapped and 21
sites were sampled within these areas. Each

wetland area contains between one and five
sites. Table 1 lists the dominant species, ele-
vation, and standing crop of all 21 sites. Fig-
ure 1 maps the vegetational patterns found in
the more complex areas.

The highest wetland found was at an ele-
vation of 3,768 m, in an area adjacent to the
steep, rocky talus pile that makes up the
northeast side of King's Peak in the east cen-
tral section of the Uinta range. Springs moist-
en the site all summer and feed a small
stream that traverses the site. Large rocks are
scattered throughout. The wetlands of this re-
gion consist of a series of small irregularly
shaped pools containing 1-10 cm of water at
the end of the summer. The emergent vege-
tation consists solely of Eriophorum scheiich-
zeri Hoppe. These pools were sampled as site
ER-1.

To the north of King's Peak lies Henry's
Fork Basin. The head of this basin has large
expanses of sedge meadows and willow thick-
ets interrupted by upland regions. In spite of
a homogeneity in species composition (much
of the area is dominated by Carex aquatilis),
there is considerable variation in stature and
standing crop of wet meadows within upper
Henry's Fork Basin. We sampled two sites
that represent extremes in Carex aquatilis
meadows. HF-1 is a border to a small (20 m
diameter) pond at an elevation of 3,278 m.
Vegetation consisted of robust individuals of
Carex aquatilis and Caltha leptosepala. Wa-
ter stands at least 10 cm deep throughout the
year, draining off the site to the north. HF-2
is a very different Carex aquatilis region,
with a more dense stand of much smaller
plants. There is no proximate open water and
the ground surface was dry at the end of the
summer although the ground was saturated
with water at a depth of 2-5 cm.

The remaining five areas are within four
miles of Mirror Lake on the west end of the
Uintas. Area TL is part of a circular meadow
approximately 200 m in diameter. The mead-
ow has a small stream meandering through it
and has numerous small "oxbow lakes" and
"kettleholes." The vegetation displays a pat-
tern commonly found in moist subalpine
meadows of the Uintas: monotypic stands of
Carex aquatilis and Eleocharis pauciflora,
areas of Carex aquatilis and Eleocharis pau-
ciflora mixed, and sections of bare ground, all

October 1983

Briggs, MacMahon: Uinta Plant Communities

525

surrounded by an upland that has soil with
less organic matter and a vegetation of Carex
illota L. H. Bailey, Deschampsia cespitosa,
and Ligusticwn filicinum var. tenuifolium (S.
Wats.) Mathias & Constance. A large portion
of this area has standing water year around.
This area was sampled in five sites (Fig. 1).

Areas FT and SE are small meadows in
Picea enge/mannii-dominated forests. The

vegetation in them is similar to that in TL
and is mapped in Figure 1. Both areas were
sampled as four sites. Area SC is a meadow in
a Picea engelmannii-Pinns contorta-domi-
nated forest. The meadow has a stream flow-
ing through it and is kept moist by two large
seepage areas. The vegetation presents a
complex patterning, with part of the area ex-
hibiting the mix described at the TL site.

Table 1. Elevation, standing crop, and species found on the sites.

Site

Elevation

(m)

Standing
crop

(g/m2)

Dominant species
(> 10 percent standing crop)

Other species

3,768

Eriophorum scheuchzeri

Hoppe.

None

HF-1

3,278

360

Carex aquatilis Wahl.
Caltha leptosepala DC.

None

HF-2

3,278

103

Carex aquatilis

Eleoeharis pauciflora (Lightf.) Link,
Pedicularis groenlandica Retz.

SE-1

3,260

172

Carex aquatilis
Carex eanescens L.

None

SE-2

3,260

234

Carex aquatilis

None

SE-3

3,260

264

Carex aquatilis

None

SE-4

3,260

106

Eleoeharis pauciflora
Carex aquatilis

Pedicularis groenlandica

FT-1

3,256

152

Eleoeharis pauciflora
Carex aquatilis

Caltha leptosepala, Pedicularis

■-2

3,256

■-3

3,256

-4

3,256

104

114

Carex aquatilis
Eleoeharis pauciflora
Carex illota L. H. Bailey
Ligusticwn filicinum (S. Wats.)
Mathias and Constance

groenlandica, Dodecatheon pulchellum
(Ref.) Merrill, Epilobium alpinum L.,
Agrostis alba L.
Eleoeharis pauciflora
Carex aquatilis

Carex aquatilis, Eleoeharis pauciflora,
Caltha leptosepala, Pedicularis
groenlandica, Veronica wormskjoldii
Roem. and Schult.

TL-1

3,110

Carex aquatilis

Deschampsia cespitosa (L.) Beauv.

TL-2

3,110

Carex illota
Deschampsia cesp itosa

Eleoeharis pauciflora, Caltha leptosepala,
Ligusticwn filicinum, Pedicularis
groenlandica, Veronica wormskjoldii

TL-3

3,110

139

Eleoeharis pauciflora

None

TL-4

3,110

Carex aquatilis
Eleoeharis pauciflora

None

TL-5

3,110

Carex aquatilis

None

3,085

141

Carex aquatilis

None

SC-1

3,073

194

Scii-pus cespitosus L.
Caltha leptosepala

Carex nigricans A. Meyer, Pedicularis
groenlandica, Veronica wormskjoldii

SC-2

3,073

109

Scirpus cespitosus
Carex aquatilis
Eleoeharis pauciflora

Caltha leptosepala, Ligusticum filicinum,
Pedicularis groenlandica, Veronica
wormskjoldii

SC-3

3,073

Carex aquatilis
Eleoeharis pauciflora

None

SC-4

3,073

251

Carex illota
Carex rostrata Stokes
Carex aquatilis
Caltha leptosepala

Agrostis alba, Carex eanescens,
Ligusticum filicin um

526

Great Basin Naturalist

Vol. 43, No. 4

Other parts of the meadow have a dense turf contains a mix of Carex illota and C aaua-

of Scirpus cespuosus L. associated with Carex tilis. The whole meadow has standing wa"er

aqtuitths Eleochans pauciflora, and Caltha throughout the year. SC was diviL fnto

leptosepala. A final portion of the meadow four sites (Fig. 1).

u^^ CAAO -"r"" °^ '^"r'n^t '''"^ ^' ^'"'^ ^^- ^' ^'^^ ^L. C, Area SC. D, Area FT. ARMO = Arnica
t^ieocnans pauciflora, JUPA - /uncti* parryi, POPA = Po« pra/en^., SCCE = Scirpus cespitosa.

October 1983

Briggs, MacMahon: Uinta Plant Communities

527

Area AQ is on the margin of a small pond
that narrows into a stream at one end. It is
surrounded by a Picea engelmannii-Pinus
contorffl-dominated forest. This area is at
least 30 cm imder water in the spring. Stand-
ing water is present to a depth of 10 cm in
the fall. One site was sampled in this area, a
region of nearly pure Carex aquatilis.

Results and Discussion

Vegetation

The number of species sampled on each of
the 21 sites (Table 1) varied from one to sev-
en. Most of the species present in the wet-
land sites are common species of the western
United States. The Carex species collected
can be separated, phytogeographically, into
two groups: western cordilleran species
{Carex nigricans, C. illota) and circumboreal
species (C. rostrata, C. aquatilis, and C
canescens).

One of the striking features of these wet-
lands is the dominance by plants that are ca-
pable of extensive spread by rhizomes. The
mosaic structure (Fig. 1) of these commu-
nities is a function of the strongly rhizo-
matous nature of the dominant plants and
some, perhaps most, of the "patches" seen
represent single plants (genets. Harper 1977)
formed by growth from a single propagule.
The patterns seen in these wetlands are prob-
ably dependent on both the history of prop-
agule arrival and on variations in environ-
mental conditions affecting the success of
particular species. A complicating factor re-
sults from the "inertia" of these sites— a re-
sistance to vegetational change resulting
from both the harsh nature of the climate and
the vigorous mode of wetland plant growth.
Both these factors could make established
vegetation patterns difficult to disrupt. Al-
though seedling establishment is common in
some wetlands (Lieffers and Shay 1982), both
Costello (1936) and Bernard (1975) noted that
seedling establishment was rare in some
Carex wetlands. Our sites included areas
where the dominant was in very poor condi-
tion (e.g., HF-2) or had died off completely,
leaving areas of bare ground (e.g., part of
TL). Both biotic and abiotic factors at these
sites may change with time. These changes

will sometimes be to the detriment of the
species that has been dominant. But, because
of the difficulty in establishing additional
species, the dominant may continue to exist
in a depauperate condition and may even die
back completely before another species is
able to invade the area. Thus, the species
present today may not reflect present condi-
tions but may represent conditions of an ear-
lier time.

One environmental variable that we found
associated with community structure is water
level. In TL and FT the Carex illota section
was always upland relative to the Carex
aquatilis-Eleocharis pauciflora sections. In
SC the part of the plot containing Scirpus
cespitosus was on higher ground than those
regions containing C. aquatilis and Eleocharis
pauciflora. Some species distributions could
not* be associated consistently with water lev-
el. Carex aquatilis could be found in areas
with both more and less standing water than
the often adjacent E. pauciflora areas (areas
TL, FT). Carex illota, although usually re-
stricted to areas with no standing water (FT-
4, TL-2), sometimes was found in areas 5 cm
deep in running water (SC-4).

It is generally thought that the influence of
water level in wetland regions is based on
variations in waterlogging and aeration. Soil
aeration may influence plants either by af-
fecting root respiration (Mendelssohn et al.
1981), which could affect water and nutrient
flow through roots, or by affecting soil nutri-
ent status. The influence on soil nutrition can
occur directly, by influencing the chemical
forms available and their solubilities (Jones
1971, 1972), or indirectly, by affecting soil
microbial activity.

The importance of aeration on species dis-
tribution is represented in our observations
on Carex illota, which is intolerant of water-
logged soils if the water is stagnant (and pre-
sumably poorly aerated), yet grows quite
well under 5 cm of water when it is flowing
over the site (and probably better aerated).
Further evidence for the importance of aera-
tion in these sites is shown in standing crop
data presented below. If aeration does in-
fluence the structure of these communities,
the complex nature of some of these sub-
alpine sites could be due to underground wa-
ter flow patterns and resultant aeration pat-
terns. These flow patterns may be due to

528

Great Basin Naturalist

Vol. 43, No. 4

300-

200

100-

HF-I

• SE-3

• SC-4

• SE-2

• SC-I

• SE-I

• FT-I

• AQ

SC-2

TL-4
SC-3

TL-3

FT-2
TL-5
FT-3

o^ .• PT-4
SE.4«#HF.2

TL-2: TL-I

• ER
I

Running water
year-round

Running water
in spring,
standing water
all summer

Standing water
all summer

Standing water
half of summer

Standing water only
in early summer

MOISTURE CLASS

Fig. 2. Graph showing the relationship between standing crop on a site and a measure of water movement over
that site.

springs and may not be readily discemable
without measurement of soil conditions.

The peak aboveground standing crop val-
ues (Table 1) varied considerably, both be-
tvi'een areas and also between adjacent sites
in one area (e.g., area SC). The very low
standing crop value for site ER was undoubt-
edly due to the short growing season and low
temperatures associated with this, the highest
site. However, aside from site ER, there is
very little relationship between elevation and
peak aboveground standing crop. Gorham
(1974) derived a regression equation relating
standing crop of Carex meadows to the high-
est monthly mean temperature of the year.
His data on 11 pure Carex stands (on both
rich and poor soils) fit the regression line well
(r = 0.84). The data of Auclair et al. (1976)
on wetland stands that were not pure Carex
also fit this regression. Using weather data for
areas in Colorado that are comparable to
ours, Gorham 's equation predicts a standing

crop of 292 g/m- for a site at 3,109 m
(10,200 ft) and 235 g/m^ for a site at an ele-
vation of 3,566 m (11,200 ft). The standing
crop values that we found are both above
and below these values, with most sites be-
low. Gorham 's equation is most appropriate
for those sites where water is not stagnant
(HF-1, SE-2, SE-3, SC-4). The sites with less
water movement have less standing crop than
predicted by Gorham. The reduced growth
on sites where water stagnates is shown in
Figvire 2, where standing crop is plotted
against a gradient in the amount of water
movement through the site. Reduced growth
on the stagnant sites is probably due to low
oxygen levels that may be disturbing root res-
piration, mineral uptake, or soil nutrient
status.

An oxygen-limited situation in Carex
meadows might explain the timing of nutri-
ent uptake in wetlands. Boyd (1970) and Ber-
nard and Solsky (1977) note considerable

October 1983

Briggs, MacMahon: Uinta Plant Communities

529

300H

Qi
O

I lOOH

— r 1 I'

15 5 2 3

JULY AUG. SEPT. OCT.

Fig. 3. Seasonal trends in aboveground standing crop
for two of the pure Carex aqtiatilis sites, SE-2 and SE-3.

mineral uptake by wetland plants in the early
spring. Boyd hypothesized that this early
mineral uptake (preceding the growth peri-
od) was an adaptation to allow certain spe-
cies to procure nutrients before other species.
Another possible reason for mineral uptake
early in the spring could be related to oxygen
concentrations. In the early spring, oxygen in
wetland soils may be at its highest levels be-
cause of well-oxygenated runoff waters and
because low temperatures increase the
amount of oxygen that water can hold. Thus,
plants may take up nutrients in the early
spring because that is the most favorable
time for root respiration.

In several of the Carex aquatilis-domina.ied
sites, we observed the variation in standing
crop during the summer. The typical pattern
in these subalpine meadows (Fig. 3) includes
a single peak in aboveground standing crop
which occurs in late summer. Gorham and
Somers (1973) and Bernard and MacDonald
(1974) also found such a pattern for wetlands
composed of C. aqimtilis and C. lacustris, re-
spectively. On our sites there was relatively
little new shoot production over the course
of the summer (Fig. 4), and the changes in
aboveground standing crop were being
caused by the growth of individual shoots.
Some Carex species (e.g., C. rostrata) have
biannual periods of shoot production and

Fig,
Carex

4. Seasonal trends in stem density (stems/m^) for
oquatilis on SE-2 and SE-3.

these may be responsible for bimodal pat-
terns in biomass (Gorham and Somers 1973),
although other factors may cause bimodal
patterns. The pattern in shoot production
that we observed for C. aquatilis had been
found previously for this species (Gorham
and Somers 1973), as well as for C. lacustris
(Bernard and MacDonald 1974), and would
be expected in regions with short growing
seasons such as the one we studied.

Based on our observations, the life history
of C. aquatilis is initiated by growth of a new
shoot sometime during the winter or spring.
The shoots grow throughout the summer and
then overwinter. The following spring the
old shoots are distinguishable from shoots of
the current year by their large size and old
leaves. One or two of these old leaves may
undergo additional growth in the spring. At
least some and possibly all of the shoots un-
dergo two or possibly more seasons of growth
before they flower or die. Many shoots may
not flower. Floral initiation is evident very
early in the spring; floral parts are visible less
than two weeks after growth initiation. Dur-
ing the growing season leaves are initiated,
grow quickly to a maximum length, and re-
main green until early September.

There was considerable variation in plant
density in the C. aquatilis stands. On the
stagnant sites (those where water was not
flowing), the variation in density was related
to stem weight. If a plot is made of the natu-
ral logarithm of plant density vs. the natural
logarithm of plant weight (Fig. 5), it can be
seen that the stagnant sites fall on a line

530

Great Basin Naturalist

Vol. 43, No. 4

n (density)

6.3

-0.86

Fig. 5. Graph showing the relationship between stem
density and average stem weight for five Carex aquatilis
stands where water was stagnant.

(r^ = 0.98). Sites with flowing water do not
fall on this line and have substantially greater
plant weights at a particular density than
those on stagnant sites. The regression line of
Figure 5 describes the following equation:

[1] w = 5.53p-o86

where w = individual
weight; p = plant density

plant

The form of this equation is characteristic of
monotypic stands and an exponential value of
-3/2 has been found for a wide variety of
species of differing life forms and habitats
(Gorham 1979, White 1981). The equation
has been labeled the "3/2's thinning law" be-
cause it describes the changes in density and
plant weight that occur in stands during self-
thinning. On our plots it is likely that the
pattern of density and plant weight is not
being caused by mortality (thinning) but
rather by recruitment (see Fig. 4). Gorham
(1979) cites studies with imcrowded condi-
tions, and therefore little self-thinning, where
the exponent value in equation (1) is closer to
-1. Our stagnant sites appear to be similar
areas. With an exponent of -1, standing crop
(w o p) remains constant regardless of density.
Such sites may represent areas where stand-
ing crop is limited by factors (such as fer-
tility?) that prevent densities to reach levels

where self-thinning in the typical 3/2's fash-
ion can occur. The precise factors that dic-
tate the 3/2's thinning law are still not eluci-
dated (White 1981). Perhaps studies on such
areas as these, where the "law" does not
hold, may help to clear up the problem.

Literature Cited

AUCLAIR, A. N. D., A. BuCHARD, AND J. PaJACKOWSKI.

1976. Productivity relations in a Corex-dominated
ecosystem. Oecoiogia 26:9-31.

Bernard, J. M. 1975. The life history of shoots of Carex
lacustris. Canadian J. Bot. 53:256-260.

Bernard, J. M., and J. G. MacDonald. 1974. Primary
production and life history of Carex lacustris. Ca-
nadian J. Bot. 52:117-124.

Bernard, J. M., and B. A. Solsky. 1977. Nutrient cy-
cling in a Carex lacustris wetland. Canadian J.
Bot. 55:630-638.

Boyd, C. E. 1970. Production, mineral accumulation
and pigment concentrations in Typha kitifolia
and Scirpiis americanus. Ecology 51:285-290.

Briggs, G. M., and J. A. MacMahon. 1982. The struc-
ture of alpine plant communities near King's
Peak, Uinta Mountains, Utah. Great Basin Nat.
42:50-59.

CosTELLO, D. F. 1936. Tussock meadows in southeastern
Wisconsin. Bot. Gaz. 97:610-648.

Gorham, E. 1974. The relationship between standing
crop in sedge meadows and summer temper-
ature. J. Ecol. 62:487-491.

1979. Shoot height, weight and standing crop in

relation to density in monotypic stands. Nature
279:148-150.

Gorham, E., and M. G. Somers. 1973. Seasonal changes
in the standing crop of two montane sedges. Ca-
nadian J. Bot. 51:1097-1108.

Hansen, W. R. 1975. The geologic story of the Uinta
Mountains. Geological Survey Bulletin 1291. U.S.
Government Printing Office, Washington, D.C.
144 pp.

Harper, J. L. 1977. The population biology of plants.
Academic Press, London. 891 pp.

Jones, H. E. 1971. Comparative studies of plant growth
and distribution in relation to waterlogging. III.
The response of Erica cinerea L. to waterlogging
in peat soils of differing iron content. J. Ecol. 59:
583-591.

1972. Comparative studies of plant growth and

distribution in relation to waterlogging. VII. The
influence of water table fluctuations on iron and
manganese availability in dune slack soils. J.
Ecol. 61:107-116.

Lieffers, V. J., AND J. M. Shay. 1981. The effects of wa-
ter level on the growth and reproduction of
Scirptis maritiimis var. paludosus. Canadian J.
Bot. 59:118-121.

Mendelssohn, I. A., K. L. McKee, and W. H. Patrick.
1981. Oxygen deficiency in Spartina alterniflora
roots: Metabolic adaption to anoxia. Science
214:439-441.

White, J. 1981. The allometric interpretation of the self-
thinning rule. J. Theor. Biol. 89:475-500.

UTAH FLORA: SALICACEAE

Sherel Goodrich'

Abstract.— A revision of the willow family, Salicaceae, is presented for the state of Utah. Included are 31 species
and 5 subspecific taxa of indigenous and introduced plants. Keys to genera and species are provided, along with de-
tailed descriptions, distributional data, and comments. No new taxa or combinations are proposed.

This paper is another in a series of works
leading to a definitive treatment of the flora
of Utah. The willow family as represented in
Utah is rather small when compared to sever-
al other families, but its taxa cover the state,
and it is complex. Herbarium specimens are
frequently misidentified. Unisexual plants, ex-
treme variation in leaves of fertile and vege-
tative or short and long twigs, and early de-
ciduous flowers all contribute to the
difficulty in identification of taxa. Hybridiza-
tion especially in Populus further complicates
identification.

Several members of the family are culti-
vated for ornamental plants or shade trees.
Not all of these are included in this treat-
ment. Among those not treated are Populus
candicans Ait. (Balm of Gilead), P. simonii
Carr., and Salix viminalis L. (Golden Osier).

Members of the family are important to
many kinds of wildlife. For example, in Utah,
beaver are almost totally dependent on the
family. They utilize aspen, cottonwoods, and
willows and avoid most other woody plants.
Among the very few exceptions are probably
Alnus and Betula.

The arabic numerals following the dis-
cussion of each taxon indicate the number of
specimens examined in the preparation of
this treatment. The roman numerals indicate
the number of specimens collected by me.

Acknowledgments

Appreciation is expressed to Dr. Arthur
Cronquist for permitting me to preview his
Salix manuscript of the intermountain flora.
Appreciation is also expressed to the direc-
tors and curators of the following herbaria of
Utah: Brigham Young University, Provo; For-
est Service Herbarium, Ogden; Garret Her-
barium, University of Utah, Salt Lake City;
Intermountain Herbarium, Utah State Uni-
versity, Logan. I appreciate the loan of speci-
mens from each of these herbaria. These
specimens are the basis of this work.

Salicaceae Mirbel.

Willow Family

Dioecious dwarf shrubs to large trees;
leaves alternate, simple, entire, serrate, cre-
nate, rarely lobed, usually stipulate, but the
stipules often readily deciduous; flowers
borne in aments (catkins), without a perianth,
each subtended by a small, scalelike bract
(commonly referred to as a scale); staminate
flowers of (l)2-many stamens; pistillate flow-
ers of a single pistil with 2-4 carpels and as
many stigmas; placentation parietal or basal;
fruit a sessile or stipitate capsule with 2-4
valves; seeds numerous, small, covered with
long white hairs, dispersed easily by wind.

1. Trees with pendulous aments; leaf buds covered by several, usually resinous
scales; each flower subtended by a cup-shaped disk, without obvious glands;
stamens 6 to many; scalelike bracts subtending the flowers laciniate or
fimbriate (except in P. alba), otherwise glabrous or ciliate Populus

'Forest Service, U. S. Department of Agriculture, Intermountain Forest and Range Experiment Station, Ogden, Utah 84401, stationed in Provo, Utah, at
the Shrub Sciences Laboratory.

531

532 Great Basin Naturalist Vol. 43, No. 4

— Trees, shrubs, or dwarf shrubs with mostly ascending to erect aments; leaf buds
covered by a single nonresinous scale; each flower subtended by 1 or 2 basal
glands, but without a disk; stamens (1)2-8, rarely more; scalelike bracts sub-
tending the flowers entire or occasionally shallowiy toothed,
usually densely pubescent Salix

PopuLus L. valves, glabrous in our taxa except in P.

hahamifera.

Small to large trees; leaf buds covered by
several overlapping scales, resinous in most References

taxa; aments pendulous, mostly appearing be- Eckenwalder, J. E. 1977. North American cottonwoods
fore the leaves, and often soon deciduous, the {Populm, Salicaceae) of sections Abaso and Aieg-

i,.,, . -iij-j J iros. T. Arnold Arboretum 58(3): 194-208.

scalelike bracts very quickly deciduous, deep- hjtchcock, C. L., and a. Cronc^u.st. 1964. Vascular
ly lobed to laciniate, often dilated (entire or plants of the Pacific Northwest. Part 2: Salicaceae to

nearly so and not dilated in P. alha); each Saxifragaceae. Univ. Washington Publ. Biol. Vol. 17.

flower subtended by a cuplike disk; stamens „ ^'^' ,0=1 \n i r u ^ w j u 1

1 f I r Kehder, a. 1951. Manual ot cultivated trees and shrubs

6-60 or more, the filaments free; inserted on hardy in North America. Macmillan Co., New

the disk; capsules pedicellate, with 2-4 York. 996 pp.

1. At least some of the mature leaves deeply 3-5 lobed and aceriform, often
densely tomentose beneath; bracts of flowers entire or shallowiy toothed, long
pilose-ciliate; twigs of the season and winter buds often white-woolly; stigma
lobes slender; plants introduced, cultivated, and escaping P. alha

— Leaves not deeply lobed, not aceriform, merely toothed, glabrous or nearly so;
scales of flowers deeply lobed to lacerate 2

2(1). Bark white and smooth except blackened and rough where scarred, covered
with a whitish powdery bloom; bracts of flowers more or less persistent, deeply
lobed or cleft, ciliate with long white hairs; leaves orbicular to reniform-
cordate; bud scales shiny but hardy resinous; stamens 6-14; capsules 4-6 mm
long, with 2 carpels; stigmas slenderly lobed; plants not confined to water
courses P. tremuloides

— Bark turning gray or brown and roughly furrowed on older trunks; bracts of
flowers laciniate-fringed, otherwise glabrous or inconspicuously short hairy;
stamens 12-60 or more; capsules mostly longer, with 2-4 carpels; stigmas
broadly dilated; plants mostly cultivated or growing along water courses

or edges of lakes 3

3(2). Leaves 0.67-1.3 times longer than wide, deltoid to rhombic or ovate;

petioles compressed laterally 4

— Leaves (1)1.2-7(10) times longer than wide, ovate to lanceolate; petioles terete

or dorsiventrally compressed 6

4(3). Bud scales and twigs of the season pubescent; leaf blades commonly with
4-10(15) fine to coarse teeth on each side; branches widely spreading and the
crown often as broad or broader than the tree is tall; plants native, sometimes
cultivated, most common along the drainages of the Colorado River system,
but sporadic along the Wasatch Front and elsewhere P. fremontii

— Bud scales and twigs mostly glabrous; leaf blades commonly with 15-25(30)
fine teeth on each side; branches ascending to erect and the crown mostly
longer than wide; plants introduced, cultivated, sometimes persisting 5

5(4). Leaf blades rhombic-ovate, cuneate at the base, seldom over 7 cm long, capsules 2
valved; branches often comparatively small, strongly ascending to erect and the
crowna narrow and columnar (in the trees planted in our area) P. nigra

October 1983

Goodrich: Utah Flora, Salicaceae

533

— Leaf blades more or less deltoid or broadly ovate, broadly cuneate at the base,
some regularly over 7 cm long; capsules 2 or more valved; branches
large, spreading-ascending, the crown not columnar P. canadensis

6(3). Leaf blades distinctly darker above than beneath, very strongly resinous espe-
cially when young, the petiole terete or nearly so; ovary and young fruit
hairy or glabrous; stamens 30-60 P. balsamifera

— Leaf blades about equally yellow-green on both sides; ovary and young fruit
glabrous; stamens mostly 12-30 7

7(6). Leaf blades (1.8) 2.5-6 (9.5) times longer than wide; petioles 1/5-1/3 (2/5) as

long as the blades, dorsiventrally compressed; carpels 2 P. angustifolia

— Leaf blades 1-2.4 times as long as wide; petioles 1/5-3/4 as long as the blades,
subterete or somewhat flattened; carpels 2 or 3; plants hybrids, intergrading
into P. angustifolia on one hand and into P. fre7nontii and other broad-leaved
poplars on the other P. acuminata

Popuhis acuminata Rydb. Lanceleaf Cot-
tonwood. A series of hybrids between P. an-
gustifolia and P. fremontii and other taxa
with broad leaves, with features intermediate
between the parents and intergrading into P.
angustifolia on one hand and into the broad-
leaved parent on the other; petioles com-
monly (1.5) 2.5-5.5 (6.5) cm long, 1/5-3/4 as
long as the blade; leaf blades 1-2.4 times
longer than wide. Along streams and rivers,
edges of ponds and lakes, often in mouths of
canyons where the parental types come to-
gether, probably cultivated, from (1370)
1525-1920 m, in Box Elder, Cache, Du-
chesne, Emery, Garfield, Iron, Kane, Salt
Lake, San Juan, Sevier, Uintah, Utah,
Wasatch, Washington, Wayne counties;
throughout the range of P. angustifolia. The
name P. acuminata in the strict sense is ap-
plied to crosses of P. angustifolia and P. del-
toides Marsh, var. occidentalis Rydb. It is
used here in a broad sense to include crosses
with other broad-leaved taxa, including P.
balsamifera and P. fremontii; 29(0).

Populus alba L. White poplar. Trees
spreading by root sprouts, to about 30 m tall,
the tnmk to 1 m or more in diameter, the
branches usually spreading, the crown more
or less roimded; bark gray-green to whitish
and smooth on upper parts of the trunk and
branches, rough and furrowed and turning
blackish on lower parts of old trunks; twigs
tomentose or glabrous; buds tomentose; pet-
ioles terete 1-5 cm long, 0.2-0.6 times as
long as the blade; leaf blades longer than
wide, deltoid-ovate in outline, undulate

toothed to deeply palmately 3-5 lobed and
a-ceriform, the lobes serrate or crenate, the
two primary lateral lobes sometimes has-
tately lobed, dark green above, silvery white-
tomentose beneath or glabrous; aments ap-
pearing before or with the leaves, the rachis
pilose-tomentose, the bracts entire to
toothed, not laciniate, ciliate-fringed with
long-pilose hairs, very quickly deciduous;
staminate aments 8 cm long or more, the
flowers with 6-10 stamens; pistillate aments
4-9 cm long; capsules 2-5 mm long,
glabrous, 2-3 valved, the pedicels about 1(2)
mm long; stigmas 2, each 2 lobed, the lobes
linear, not dilated. Introduced from Eurasia,
cultivated, escaping, and more or less natu-
ralized, in populated areas, along fencelines,
ditchbanks, and abandoned homesteads and
fields, up to about 1980 m, to be expected in
all counties of the state. Trees with leaves
densely white-tomentose beneath are refe-
rable to var. alba. Those with leaves and
twigs glabrous or glabrate and fastigiate
crowns are referable to var. bolleana
Lauche. These may be hybrids between P.
alba and some other species; 21 (ii).

Populus angustifolia James Narrowleaf
Cottonwood. Trees about 7-15(20) m tall, the
trunk 30-60(80) cm in diameter, the branches
erect-ascending, the crown more or less pyra-
midal; bark pale green to whitish when
young, furrowed and grayish on old trunks,
twigs glabrous or pubescent; buds ovoid-con-
ic, pointed, strongly resinous, reddish brown,
glabrous or pubescent; petioles semiterete or
horizontally flattened and channeled above.

534

Great Basin Naturalist

Vol. 43, No. 4

especially near the blade, 3-25 mm long, up
to 0.3 (rarely 0.4) times as long as the blade;
leaf blades 4-14 cm long, 0.7-2.5 (4.0) cm
wide, (1.8) 2.5-6 (9.5) times longer than wide,
lanceolate or occasionally narrow elliptical
or ovate, glabrous or nearly so, usually acute
at the apex, roimded at the base, the margins
finely to coarsely serrate; aments often devel-
oping with the leaves, the rachis glabrous or
nearly so, the bracts broadly obovate, deeply
and irregularly lacrate; staminate aments
2-6 cm long, the flowers with 12-20 stamens;
pistillate aments 6-10 cm long; capsules
3-6(7) mm long, 2 valved, glabrous, the pedi-
cels about 2-10 mm long; stigmas 2, dilated,
irregularly lobed. Along water courses, often
in canyons, from about 1525-2135 (2440) m,
in all counties of the state. Rather freely
crossing with the broad-leaved species of the
genus; 79 (i).

Populus balsamifera L. Balsam poplar,
Black Cottonwood. [P. trichocarpa T. & G.].
Tree 15-30(50) m tall; the trunk mostly 0.6-1
(1.5) m in diameter, bark furrowed and gray-
ish on older trunks; buds large, the scales
very resinous, glabrous or inconspicuously
puberulent; petioles more or less terete,
2-6.2 cm long, 1/4-3/4 as long as the blade;
leaf blades 4.3-11 cm long, 3.2-8 cm wide,
1.3-2.6 times longer than wide, ovate-
accuminate, cuneate to cordate at the base,
the margins crenulate, sometimes short ci-
liate, strongly resinous, glabrous at maturity
on both sides, the upper side dark green, the
lower side distinctly paler and often rufous
tinged in dried specimens; bracts of aments
lacerate-fringed, otherwise glabrous or some-
times with minute hairs, these not over
0.5 mm long; staminate aments 2-3(5) cm
long, readily deciduous; stamens commonly
30-60; pistillate aments 8-20 cm long; cap-
sules 5-8 mm long, glabrous or pubescent,
subsessile; stigmas broadly dilated. Along
streams, mostly in canyons and cultivated,
1370-2350 m, in Cache, Juab, Salt Lake, Se-
vier, Utah, Wasatch, and Wayne counties;
widespread in North America from New-
foundland south to New York and west to
Alaska (ssp. balsamifera), and from Alaska
south to Baja California in the western part
of the continent (ssp. trichocarpa). The native
trees of our area are expected to be ssp.
trichocarpa (T. & G.) Brayshaw with mostly

pubescent and 3 (rarely 2-4) carpellate cap-
sules. Some of the cultivated trees might be
ssp. babamifera with mostly glabrous and 2
(rarely 3-4) carpellate capsules; 9 (0).

Populus X canadensis Moeneh. Carolina
poplar. Gray poplar. Cultivated and per-
sisting, rarely escaping, to 40(50) m tall, the
trunk 0.75-1.5(2) m in diameter; bark deeply
furrowed and grayish on old trunks; buds
large, the scales glabrous, but resinous; pet-
ioles laterally flattened 3.5-8.5 cm long, 1/3
to as long as the blade; leaf blades mostly
3.5-11.5 cm long, 3.5-11 cm wide, or much
larger on stump sprouts, 0.9-1.3 (rarely to
1.5) times as long as wide, deltoid-ovate,
acuminate at the apex, mostly broadly cu-
neate or trvmcate at the base, the margin cre-
nate-serrate; glabrous and equally green on
both sides; staminate aments about 7 cm
long; stamens 15-25; pistillate aments un-
known. Cultivated for shade trees, probably
originated in France as a cross between P.
deltoides Marsh, and P. nigra (Rehder, 1951),
to be expected in nearly all counties of the
state. Populus deltoides might also be ex-
pected in the state as an introduced tree from
the Plains and eastward, but no specimens
were seen that were clearly assignable to that
taxon. The original Carolina poplar was P.
deltoides, but for many years the nursery
stock distributed under that name has been P.
X canadensis (Hitchcock and Cronquist,
1964); 15 (ii).

Populus fremontii Wats. Fremont cotton-
wood. Trees 10-25 m tall with broad
rounded crowns, the crown often as broad or
broader than the tree is high, the trunk 0.5-1
(1.5) m in diameter; bark smooth and whitish
on young trees and on twigs and young
branches, deeply furrowed and grayish or
brownish on old trunks; petioles (0.8)
3-9.5 cm long, one half to as long as the
blade, flattened; rarely with two glands at
the summit; leaf blades (2) 4-10 cm long, (15)
4.5-12.5 cm wide, or much larger on sterile
sprouts, 0.67-1.2 times as long as wide, del-
toid, ovate, rarely nearly rhombic, with trun-
cate, cuneate, or occasionally cordate base,
acuminate at the apex, coarsely to finely cre-
nate or serrate with about 8-11 (15) glandu-
lar teeth, glabrous, greenish or yellow-green
on both sides, turning yellow in autumn;
staminate aments 4-10 cm long, the flowers

October 1983

Goodrich: Utah Flora, Salic ace ae

535

with a broad oblique disk and 50-80 stamens
with dark red anthers; pistillate aments
5-15 cm long, the flowers with a cup-shaped
disk, this to 5 mm wide in fruit; capsules
7-10 (12) mm long, to 8 mm wide, ovoid to
subglobose, 3 to 4 valved, glabrous, the
stipes 2-6 (10) mm long; stigmas strongly di-
lated and irregularly lobed. Along flood
plains of rivers and along washes, irrigation
ditches, and occasionally cultivated, from 762
to about 1860 m, in Cache, Duchesne, Gar-
field, Grand, Iron, Kane, Salt Lake, San Juan,
Sevier, Tooele, Uintah, Utah, Washington,
Wayne, and Weber counties. The Fremont
Cottonwood is abundant along the Colorado,
Green, San Juan, and Virgin rivers and their
tributaries within the Colorado Drainage, to
be expected anywhere in the state as it has
been cultivated for a shade tree. This tree is
part of a transcontinental complex, of which
P. arizonica Sarg., P. deltoides, P. sargentii
Dode, and P. wislizeni (Wats.) Sarg. are a
part. Popuhis arizonica and P. wislizeni have
generally been considered closely allied to P.
fremontii and they have by some authors
been included as varieties of or as synony-
mous with P. fremontii. Specimens that have
capsules with stipes up to 6 or even 10 mm
long are found in Emery County and other
points along the Colorado River system.
These trees have been referred to as P. fre-
montii var. wislizeni Wats. Based on the long
stipes, these trees have recently been as-
signed to P. deltoides var. wislizenii (Wats.)
Eckenwalder (Eckenwalder, 1977). However,
these trees are like P. fremontii in the lack of
glands at the jimction of petiole and blade
and with few, broad, and coarse teeth on leaf
margins. Based on my provincial study, I am
not well prepared to make a judgment as to
the specific assignment of these trees, but I
prefer the traditional approach. If P. fre-
montii is to be kept separate at all from P.
deltoides, I feel these plants are best kept as a
part of P. fremontii., 98 (ii).

Populus nigra L. Black poplar. Tree to
30 m tall; bark deeply furrowed and grayish
on old trunks; bud scales glabrous, resinous;
petioles flattened laterally, slender 1-4.5 cm
long, 0.4-0.8 times as long as the blade; leaf
blades 2.2-6.5 cm long, 1.8-8 cm wide, occa-
sionally larger 0.8-1.2 (rarely 1.4) times as

long as wide, very often as wide or wider
than long, rhombic ovate, or orbicular, usual-
ly strongly accuminate at the apex, cuneate
at the base, glabrous, equally green on both
sides or a little darker above, the margin
crenate-serrate, not ciliate; bracts of aments
laciniate; staminate aments 4-6 cm long; sta-
mens 20-30; pistillate aments not seen. In-
troduced, cultivated for shade and wind
breaks, specimens seen from Beaver, Salt
Lake, and Utah counties, but to be expected
throughout the state. Most of the trees in our
area are from a staminate clone with strongly
ascending branches that produced a narrow,
often nearly cylindrical crown. Trees of this
clone have been assigned to var. italica
Duroi, Lombardy poplar; 6 (0).
. Populus tremuloides Michx. Aspen, quak-
ing aspen, quakey. Colonial tree 10-15
(20) m tall, seldom taller; the trunk seldom
over 40 cm in diameter; bark white and
smooth, covered with a powdery white
bloom, turning black and rough where
scarred and at the base of very old trunks;
branches usually spreading, the crown usually
rounded; bud scales shiny but hardly re-
sinous; petioles laterally flattened, 2-5.5 cm
long (1/2) 3/4 to nearly as long as the blade;
leaf blades 2-6.5 cm long, 1.8-6.5 cm wide,
or much larger on stump sprouts, 3/4-1 1/3
times longer than wide, ovate to reniform-
cordate, the margin subentire to serrate or
undulate, ciliate, glabrous on the surfaces at
maturity; bracts of the aments more or less
persistent, especially the staminate ones, 3-7
lobed or cleft, silky-pilose ciliate, the hairs up
to 2 mm long; staminate aments 2-4 cm long,
readily deciduous; stamens 6-14; pistillate
aments 4-12 cm long, to 13 mm wide; cap-
sules 4-6 mm long, the stipes 1-2 mm long,
subtended by a cuplike disk about 2 mm
across; carpels 2; stigmas 2, each deeply cleft
into 2 or more slender lobes. Along water
courses and forming clones and aggregates of
clones on canyon walls and mountain sides,
from (1400) 1830-3050 (3200) m, in all coun-
ties of the state; widespread in North Ameri-
ca from Labrador to Alaska and south to
Tennessee and northern Mexico. Aspen is cul-
tivated as a shade or ornamental tree. In re-
cent years, nursery stock has become readily
available from commercial nurseries; 96 (i).

536

Great Basin Naturalist

Vol. 43, No. 4

Salix L.

Depressed, mat-forming dwarf shrubs to
large trees; buds covered with one non-
resinous scale; aments erect to spreading,
rarely drooping, developing before (pre-
cocious), with (coetaneous) or after (sero-
tinous) the leaves, the bracts mostly entire,
occasionally with a slightly toothed apex;
flowers with 1, occasionally 2 minute glands
near the base; stamens (1) 2-8 (12), the fila-
ments free or imited toward the base, in-
serted on the base of the bract; capsules ses-
sile or stipitate, glabrous or pubescent.

A large genus of about 300 species, mostly
of the Northern Hemisphere, most common
in arctic and temperate regions.

Identification of the willows is com-
pounded by unisexual plants, aments that are
sometimes precocious and mostly early de-
ciduous, and variation among the usually
smaller leaves of the flowering branches
which often lack or have inconspicuous stip-
ules and the usually much larger leaves and
stipules of vegetative branches and particu-
larly of vigorous young shoots. Thus, her-
barium specimens of each species present
specimens of 3 or 4 phases (pistillate, stami-
nate, flowering twigs with or without the de-
ciduous aments, and vegetative twigs). Vigor-
ous young shoots sometimes add a fifth
dimension. At times whole plants in the field
present only one or two of the various
phases.

To facilitate identification of plants of the
different phases, pistillate, staminate, and
vegetative features have been included in

many of the leads in the key. Thus, some of
the leads are rather long, and features not ap-
plicable to a particular specimen will need to
be skipped. An alternative approach to
lengthy leads is separate keys for the differ-
ent sexual and vegetative phases. Many such
keys have been written, but these sometimes
also contain a mixing of vegetative and sexual
features. To establish an adequate basis for a
staminate key, I feel that many more stami-
nate specimens are needed in the herbaria of
the state.

References

Archer, W. A., and E. E. Little. 1965. "Salicaceae of

Nevada." Contr. Flora Nevada 50: 10-59.
Argus, G. W. 1965. The taxonomy of the Salix glauca

complex in North America. Contr. Gray Herb.

196: 1-142.
197.3. The genus Salix in Alaska and the Yukon.

Nat. Mus. Canad. Publ. Bot. 2: 1-279.
1980. The typification and identity of S. erioce-

phala Michx. (Salicaceae). Brittonia 32(2):

170-177.
Arnow, L., B. Albee, and A. Wycoff. 1980. Flora of

the central Wasatch Front, Utah. Univ. of Utah

Printing Service, Salt Lake City. 663 pp.
DoRN, R. D. 1975. A systematic study of Salix section

Cordatac in North America. Can. J. Bot. 53:

1491-1522.
1976. A svnopsis of American Salix. Can. J. Bot.

54:2769-2789.
1977. Willows of the Rocky .Mountain States.

Rhodora 79:390-429.
Hitchcock, C. L., and A. CRONguisT. 1964. Vascular

plants of the Pacific Northwest. Part 2: Sali-
caceae to Saxifragaceae. Univ. Washington Puhl.

Biol. Vol. 17. ,597 pp.
Rehder, a. 1951. Manual of cultivated trees and shrubs

hardv in North America. Macmillan Co., New

York. 996 pp.

1. Plants shrubs or dwarf shrubs not over 1 (1.5) m tall, subalpine to alpine 2

— Plants shrubs or trees, mostly over 1.5 m tall, of valleys to montane 3

2(1). Plants depressed dwarf shrubs 1-10 (20) cm tall mostly alpine, often forming

mats, the stems creeping on or below the ground surface KEY I

— Plants (10) 20-100 cm tall or taller, subalpine or alpine, not forming mats on

the ground, the stems ascending to erect KEY II

3(1). Leaves (8) 10-20 (32) times longer than wide; plants often strongly colonial,
spreading underground and forming patches and occasionally thickets, our
most common and widespread lowland willow S. exigua

— Leaves less than 8 times as long as wide 4

4(3). Bracts persistent, dark brown to blackish or if pale green or pale brown in age
then silky pilose with the hairs exceeding the bract by 1-2 mm and the cap-
sules pubescent (rarely glabrous in unusual specimens); stamens 2 per flower,

October 1983 Goodrich: Utah Flora, Salicaceae 537

the filaments glabrous or pilose in a few species; plants shrubs or
occasionally treelike, mostly native KEY III

— Bracts of at least the pistillate aments quickly deciduous, pale green or yellow-
ish tan in age, short pubescent, the hairs hardly if at all exceeding the bract by
more than 1 mm; capsules glabrous; stamens more than 2 per flower, or if only
2 then plants introduced trees, the filaments pilose; plants mostly trees

or treehke except in S. lasiandra, mostly of valleys and lower montane 5

5(4). Plants native; stamens 3-9 per flower; stipes of capsules mostly 1-2 mm long,

obviously longer than the gland KEY IV

— Plants introduced trees; stamens 2 except in S. pentandra; capsules sessile or

the stipes mostly less than 1 mm long and hardly longer than the gland KEY V

KEY I.

Depressed, mat-forming dwarf shrubs, 1-10 (20) cm tall, at or above timberline

1. Bracts of aments pale green or yellowish, glabrous dorsally; filaments
1.5-2 mm long; style obsolete or to 0.2 mm long, shorter than the stigma;
leaves elliptic to orbicular, 1.4-2.6 times longer than wide, glaucous and
strongly reticulate-veined beneath, the tips mostly rounded or obtuse S. reticulata

— Bracts of aments blackish, pilose dorsally; filaments over 2 mm long; styles
0.5 mm long or longer, longer than the stigmas; leaves elliptic or narrow ellip-
tic, (1.25)2.3-4.7 times longer than wide, glaucous or not, not strongly reticulate
veined beneath, the tips mostly pointed 2

2(1). Leaves 2-5 (7) mm wide, 2-4.7 times longer than wide, sessile or the petiole to

3 mm long; plants seldom over 3 cm tall, aments 0.5-2.2 cm long S. cascadensis

— Leaves 5-20 mm wide, mostly 2-3 times longer than wide, with petiole
3-13 mm long; plants mostly 5-10 (20) cm tall; aments (1) 2-4 cm long S. arctica

KEY II.

Low shrubs (10) 20-100 (300) cm tall, mat forming, subalpine or alpine

1. Capsules glabrous, the style and stigma together less than 1 mm long; leaves
permanently pubescent on both sides, the lower surface not glaucous but often
more densely pubescent and thus lighter than the upper surface; twigs of the
season glabrous or thinly villous-puberulent S. wolfii

— Capsules pubescent at least until mature or style and stigma together over

1 mm long; leaves often glaucous beneath, glabrous or pubescent 2

2(1). Mature leaves glabrous, dark green and shiny above, strongly glaucous and
glabrous or with a few hairs beneath; twigs of the season glabrous or very scat-
tered pubescent, dark chestnut to lustrous purplish black; aments precocious or
coetaneous, sessile or nearly so or rarely on a stalk to 0.5(1) cm long, this nei-
ther bearing nor subtended by bractlike leaves; style and stigmas collectively
1.5 mm long or longer; filaments of stamens glabrous S. planifolia

— Mature leaves pubescent on both sides, but sometimes glabrate or glabrous in
age; twigs of the current season densely pubescent; aments coetaneous or sub-
serotinous, born on stalks to 2 (4) cm long, these usually bearing and subtended
by bractlike leaves; style and stigmas collectively up to 1.5 mm long; filaments

of stamens sometimes pilose 3

538 Great Basin Naturalist Vol. 43, No. 4

3(2). Bracts of aments pale green when young, tan in age; capsules 3-5 mm long,
pubescent even in age, crowded and nearly sessile so as to mostly conceal the
rachis at the center of the aments, the stipes seldom over 0.5 mm long; pistil-
late aments 0.8-2 (2.5) cm long, 8-10 mm wide; staminate aments about 0.8-1
(1.2) cm long, 5-6 mm wide, the filaments densely pilose at the base and for
1/2 to 3/4 their length, the pilose portion often equaling or exceeding the
scale, the anthers usually less than 0.5 mm long; petioles 1-4 mm long, seldom
exceeding the bud even on vegetative twigs S. brachycarpa

— Bracts of aments brown to blackish, sometimes light brown to whitish tan but
not green even when young; capsules (4) 5-7 (8) mm long, sometimes glabrate
in age, dense but often not so crowded as to conceal the rachis at the center of
the ament, the stipes 0.5-2 mm long; pistillate aments (1.8) 2.5-5 cm long,
11-15 mm wide; staminate aments 0.8-2(4) cm long, sometimes over 6 mm
wide, the filaments glabrous or pilose but usually not so conspicuously pilose as
above, the anthers mostly over 0.5 mm long; petioles (1) 2-6 (10) mm long,
equaling or often exceeding the bud, especially on vegetative twigs S. glauca

KEY III.

Mostly native shrubs or small trees; aments mostly with dark bracts; stamens 2; capsules

glabrous or pubescent

1. Capsules glabrous; leaves not both glaucous and pubescent on the lower sur-
face when fully expanded; hairs of aments mostly crisped-villous and more or
less tangled except in S. wolfii with aments 0.8-2(3) cm long or in S. planifolia
and then plants keyed both ways 2

— Capsules mostly pubescent except in S. lasiolepis; leaves glaucous and pu-
bescent on the lower surface when fully expanded; hairs of aments straight or
slightly wavy but hardly crisped-villous or tangled; aments sometimes longer
than in S. wolfii 6

2(1). Leaves glaucous beneath, not or scarcely pubescent when fully expanded 3

— Leaves not glaucous beneath, although sometimes lighter colored from pu-
bescence; pubescent at least in part on both sides when fully expanded, but
sometimes glabrate in age 5

3(2). Aments sessile or on a stalk, the stalk to 0.5(1) cm long neither bearing nor sub-
tended by bractlike leaves; pubescence of aments straight or nearly so; leaves
mostly entire, often slightly revolute; twigs dark chestnut to lustrous purplish
black, essentially glabrous; plants often less than 1.5 m tall and keyed also in
Key II S. planifolia

— Aments usually stalked, the stalk usually subtended by or bearing 1-4 bractlike
leaves; pubescence of aments crisped-villous; leaves serrate, serrulate, or en-
tire, not at all revolute; twigs variously colored, glabrous or those of the
current season more often pubescent; plants often over 1.5 m tall 4

4(3). Styles 0.7-1.5(1.8) mm long; leaves of fertile and vegetative twigs often less
than 3 times longer than wide, evidently crenulate-serrate or subentire; bark of
older twigs not ashy gray or whitish; plants apparently uncommon, in the
eastern and central part of the state, mostly montane S. monticola

— Styles 0.2-0.7 mm long; leaves of vegetative twigs 2-5 times longer than wide,
serrulate or entire; bark of older twigs usually ashy gray or white; plants
widespread, mostly of valleys and lower montane S. lutea

October 1983 Goodrich: Utah Flora, Salic ace ae 539

5(2). Aments precocious or coetaneous (1.5) 2-5 cm long, with dense crisped-villous,
tangled hairs; leaves subglabrate in age, with inconspicuous hairs, entire or
sometimes serrulate; plants sometimes over 2 m tall S. boothii

— Aments coetaneous, 0.8-1.5 (3) cm long, with hairs straight or nearly so; leaves
permanently pubescent throughout on both sides even in age, the hairs readily
conspicuous with a 10- power lens, entire; plants 0.6-1.5(2)m tall, also keyed in

Key II S. wolfii

6(1). Twigs strongly blue glaucous, the bloom sometimes deciduous, but then the
twigs glabrous or sometimes puberulent; larger leaves mostly 3-5 times longer
than wide, sericeous beneath; capsules densely pubescent 7

— Twigs not glaucous or those of the current season often pubescent, or leaves
not sericeous; the larger leaves various but sometimes wider than above;
capsules pubescent or glabrous 8

7(6). Pistillate aments 2-5 cm long; capsules sessile or the stipes to 1 mm long, the
style and stigmas together 0.8-1.3 mm long; staminate aments about 2 cm
long, the filaments glabrous; aments sessile or nearly so with or more often
without subtending bractlike leaves, precocious or subcoetaneous; bracts of the
aments blackish; leaves permanently silvery, silky-sericeous to subtomentose
beneath, dark green and glabrous above in age S. drummondiana

— Pistillate aments 1-2 cm long; capsules stipitate, the stipes 2-3 mm long, the
style and stigmas together about 0.5 mm long; staminate aments 8-15 mm
long, the filaments pilose on the lower 1/2; aments borne on 2-10 mm long,
bracteate-leafy stalks; coetaneous or subprecocious; bracts of the aments dark
at the tip and pale below; leaves sericeous when unfolding, sparsely or moder-
ately sericeous, especially beneath when fully expanded, glabrate in age
especially above S. geyeriana

8(6). Plants shrubs 0.6-3 m tall, midmontane to above timberline, the stems less than
4 cm thick; leaves mostly less than 2 cm wide, occasionally wider on
vegetative twigs, elliptic to narrowly lanceolate KEY II

— Plants shrubs or small trees, commonly 3-4 m tall or taller, but sometimes
shorter, of valleys or montane, the stems of mature plants often 4-10 cm thick
or thicker; leaves sometimes over 2 cm wide, oblong, obovate, oblanceolate, or
elliptic 9

9(8). Capsules glabrous; filaments about 3-5 mm long, bracts of aments blackish or
purplish black, about as wide as long and rounded at the apex, densely pilose-
tomentose, the hairs exceeding the bracts by about 1 mm; leaves oblong to ob-
lanceolate, less than 15 mm wide except on vigorous young shoots; plants of
Great Basin and Virgin River drainages S. lasiolepis

— Capsules pubescent; filaments longer or bracts not as dark; bracts of aments of
lighter color or if blackish then with hairs exceeding the bracts by about 2 mm,
pointed or somewhat rounded; leaves elliptic, obovate, to oblanceolate,
sometimes over 15 mm wide; plants of various distribution 10

10(9). Twigs of the second and current year and the dark red bud scales velvety vil-
lous; lower surface of leaves densely velvety villous throughout the season,
twigs with longitudinal ridges beneath the bark; aments precocious; plants
introduced, cultivated S. cinerea L.

— Twigs of the second year glabrous, those of the current season villous or with
appressed hairs; lower surface of leaves villous at first but usually rather
scattered-villous to glabrate in age; aments various; plants native 11

540 Great Basin Naturalist Vol. 43, No. 4

11(10). Bracts of aments pale green or tan to very light brown in age, silky pilose, the
hairs exceeding the bract by about 1 mm; aments coetaneous; capsules long
beaked, loosely arranged so as to expose much of the rachis; filaments of sta-
mens 3-6 mm long; leaves mostly elliptic, occasionally lanceolate or obovate;
twigs of the season with mostly appressed or ascending hairs or occasionally
glabrous S. hehbiana

— Bracts of aments black or purplish black, reddish or pale only at the very base,
pilose, the hairs exceeding the bract by about 2 mm; pistillate aments pre-
cocious, or subprecocious, the capsules not long beaked, densely arranged and
mostly concealing the rachis; staminate aments strictly precocious, the fila-
ments about 10 mm long; leaves obovate or oblanceolate; twigs of the season
with mostly widely spreading hairs S. scouleriana

KEY IV.

Plants native, tall shnibs or small trees; bracts of aments pale green or yellow, at least the pis-
tillate ones, deciduous; stamens 3-8(12); capsules glabrous with a 1-2 mm long stipe

1. Bracts of aments 3-4 mm long; bud scales fused, without free overlapping mar-
gins, blunt to rounded at the tip; staminate aments 2-3.5 times longer than
wide, 1-3.5 cm long; styles 0.5-1 mm long; petioles of larger leaves often with
wartlike glands near the base of the blade; plants mostly multistemmed, large
shrubs from large root crowns, rarely trees, widespread in the northern half of
the state S. lasiandra

— Bracts of aments 1-2 mm long; bud scales with free overlapping margins, usu-
ally pointed at the tip; staminate aments often narrower and longer than
above; styles 0.1-0.2 mm long; petioles without wartlike glands or occasionally
with glands in S. nigra; plants mostly trees with solitary or few trunks, of
various distribution 2

2(1). Leaf blades not glaucous beneath, (2.5)4-7 times longer than wide; twigs

whitish or grayish yellow; plants of the southern half of the state S. nigra

— Leaf blades glaucous beneath 3

3(2). Twigs reddish or reddish brown, often pubescent at least near the nodes, hori-
zontal or spreading; some of the leaf blades usually 4-5 times longer than

wide, shiny dark green above; plants of San Juan and Washington counties

S. laevigata

— Twigs ashy gray or yellowish when fresh, glabrous, tending to droop; leaf
blades usually not over 3 times longer than wide, not shiny dark green above;

plants widespread in the state, mostly north of the counties listed above

S. amygdalioides

KEYV.

Plants small or rather large trees, introduced, cultivated, sometimes escaping and persisting;
bracts of aments pale green or yellowish, at least the pistillate ones deciduous; capsules

glabrous, sessile or nearly so
(Note: The cultivated species of this key, except S. fragilis, are not described due to lack of

adequate specimens in herbaria.)

1. Stamens 3-12; leaves with wartlike glands on upper part of petiole and lower
margins of blade, the blade seldom over 3 times longer than wide, usually
glabrous except above along the midrib, lighter beneath than above but not
glaucous (bay willow) S. pentandra L.

October 1983 Goodrich: Utah Flora, Salicaceae 541

— Stamens 2; leaf blades 3-5 times longer than wide, usually glaucous beneath,
glabrous or variously pubescent 2

2(1). Pistillate aments 1-2.5 (3) cm long, the capsules 1-2.5 mm long; staminate
aments to 4 cm long; petioles glandless; trees weeping, with very slender,
greatly elongate, pendulous branches, or if not weeping then the twigs more or
less contorted 3

— Pistillate aments mostly over 3 cm long, the capsules 3-6 mm long; petioles
sometimes with small glands near the base of the blade; trees not weeping,
with upright branches; twigs spreading, not contorted 5

3(2). Trees not weeping; twigs not pendulous, more or less contorted, aments
1-1.5 cm long; (all specimens seen from the state were referrable to f. tortuosa

Rehd. with the branches twisted and contorted — corkscrew willow)

S. matsudana Koidz.

— Trees weeping; twigs pendulous, very straight; aments sometimes longer than
above 4

4(3). Leaves mostly 3-15 mm wide, mostly deciduous in October; twigs often bright

yellow; capsules sessile (weeping willow) S. babylonica L.

— Leaves 15-22 mm wide, often persisting into December; twigs greenish or yel-
low-green; capsules with stipe exceeding the gland; plants hybrids of S.
babylonica x S. fragilis (Niobe or Wisconsin weeping willow) .... S. x blanda Anderss.

5(2). Leaves glabrous when unfolded, the margin of mature leaves usually serrate
with 4-8 teeth per cm; twigs glabrous, or nearly so; stipe of capsules
0.5-0.8(1) mm long; plants common, cultivated, and escaping S. fragilis

— Leaves sericeous, or glabrous when unfolded, the margin of mature leaves fine-
ly serrulate with 9-10 teeth per cm; twigs sometimes pubescent; capsules
sessile or subsessile; plants not known outside of cultivation (white willow) S. alba L.

Salix amygdaloides Anderss. Peach-leaf half; pistillate aments (1.5) 2.5-8 cm long,
willow. Plants mostly small trees, rarely 13-20 mm wide; capsules 4-7 mm long,
shrublike, mostly 4-10 (12) m tall, often with glabrous, the stipe 1.2-3 mm long, the style
2-4 leaning tnmks; twigs whitish, yellowish, about 0.2 mm long, not longer than the
or ashy gray, rarely reddish, glabrous except stigmas. Lake and pond margins and along
when very young; stipules usually minute and ditches, streams, and rivers, and in neglected
soon deciduous; petioles (3) 5-15 (25) mm fields and pastures, from about 1070-1710 m
long; leaf blades, (1.8) 2.3-6 (7.5) cm long, (7) in Box Elder, Davis, Duchesne, Emery, Juab,
12-19 (23) mm wide, or up to 10.5 cm and Salt Lake, Tooele, Uintah, Utah, and Wash-
3.2 cm wide on vigorous young shoots, ellip- ington counties; southern Canada and wide-
tical to lanceolate, entire or serrulate, spread in the United States except the south-
glabrous except when very young, glaucous ern part; 63 (vi).

beneath, green above; aments coetaneous, Salix arctica Pall. Arctic willow. [S. anglo-
rarely subprecocious, on leafy or bracteate rum Cham. var. antiplasta Schneid.]. De-
twigs of the season, 1.5-4 cm long; bracts of pressed shrubs with stems creeping on or un-
the aments 1-2 mm long, at least the pistil- der the ground, seldom rising more than 10
late ones soon deciduous, pale green, orbicu- (20) cm above ground level, tending to form
lar, the dorsal side woolly-pilose below and mats, but not so much as in S. cascadensis or
along the margins, but mostly glabrous to- S. reticulata; stipules minute or lacking; pet-
ward the apex, the ventral surface woolly-vil- ioles 2-12 mm long; leaf blades (5) 11-47 mm
lous throughout, the hairs seldom exceeding long, (4) 6-16 mm wide, elliptical, narrow el-
the bract by more than 0.5 mm; staminate liptical, obovate, or oblanceolate, entire,
aments 2-10 cm long, 7-11 mm wide; sta- slightly paler beneath than above but not
mens 4-7, the filaments pilose on the lower strongly glaucous; pilose-sericeous when

542

Great Basin Naturalist

Vol. 43, No. 4

young, sparingly pubescent or glabrous when
mature; aments coetaneous, borne on
glabrous or pubescent 7-35 mm long leafy-
bracteate or barren twigs of the season;
bracts of the aments persistent, dark brown,
pinkish purple at the base, pilose-sericeus on
both sides, sometimes less so dorsally than
ventrally, the hairs exceeding the bract by
about 1 mm; staminate aments 15-25 mm
long, about 7-9 mm wide; stamens 2, the fila-
ments glabrous, to about 7 mm long; pistil-
late aments 1.5-7 cm long, about 10-12 mm
wide, with 25-75 fruits; capsules 4-7 mm
long, pubescent, the stipe about 1 mm long,
the style and stigmas together about 1-2 mm
long. About snowbanks, meadows, shores of
lakes, and rocky slopes near or a little above
timberline, 2775-3600 m on the Bear River
(Mt. Naomi), Tushar (Delano Peak), western
Uinta, and the Wasatch mountains in Cache,
Piute, Salt Lake, Summit, and Utah counties;
circumboreal and south in mountains of west-
ern North America to California and New
Mexico. Our plants are var. petraea Anderss.
They more or less intergrade into S. casca-
densis in the Uinta Mountains; 14 (0).

Salix bebbiana Sarg. Bebb willow. Plants
shrubs, occasionally treelike, (2) 4-6 (8) m
tall, with 1 to several stems, young twigs
glabrous, puberulent or densely pubescent;
stipules usually inconspicuous and soon de-
ciduous; petioles (2) 3-8 (10) mm long, red-
dish or pale; leaf blades 1-4 cm long,
1.2-2 cm wide or to 7 cm long and 3 cm
wide on vigorous young shoots, 2.2-2.8 times
longer than wide, mostly elliptical, occasion-
ally obovate or oblanceolate, entire to
slightly undulate-crenate, dark green above,
glaucous beneath, pubescent when young on
both sides; fully expanded leaves glabrous
above, usually with a few hairs beneath near
the midrib; aments coetaneous, on a brac-
teate 3-15 mm long peduncle; bracts of the
aments persistent, pale green to very light
brown in age, sometimes reddish at the apex,
particularly in staminate aments, silky pu-
bescent, the hairs exceeding the bract by
about 1 mm; staminate aments 1.5-2 cm
long, to 13 mm wide; stamens 2, the fila-
ments 3-6 mm long, glabrous or sparingly pi-
lose at tlie base; pistillate aments 1.5-4(5) cm
long, to 2 cm wide; capsules 6-8(10) mm
long, rostrate with a rounded basal portion

1-2 mm wide and a long slender beak, pu-
bescent, rather loosely arranged and not con-
cealing the rachis, the stipe 2-3.5 mm long,
the style about 0.1-0.2 mm long; stigmas
0.3-0.5 mm long, bilobed to the base. Ripa-
rian communities on canyon bottoms and
along streams in mountains, occasionally
along irrigation ditches, from (1370)
1830-2710 m in Box Elder, Cache, Daggett,
Davis, Garfield, Grand, Juab, Kane, Rich,
Salt Lake, San Juan, Sevier, Summit, Uintah,
Utah, Wasatch, Washington, and Wayne
counties; across much of Canada and north-
ern United States. Our plants with leaves
sparsely appressed pubescent and soon
glabrous beneath and rather weakly raised
reticulate-veiny are often referred to as var.
perrostrato (Rydb.) Schneid., but the separa-
tion probably merits no recognition. 77(x).

Salix boothii Dorn Booths willow. [S.
pseudocordata (Anderss.) Rydb., misapplied].
Shrubs (1.5) 2-4 m tall; young twigs finely
hairy, stipules small, inconspicuous and soon
deciduous or larger and leaflike on vigorous
young shoots; petioles mostly 2-5 mm long;
leaf blades (0.8) 2.5-6 cm long, (4) 8-22 mm
wide, or to 11.2 cm long and 4 cm wide, with
petiole to 2 cm long on vigorous shoots, ellip-
tical, lanceolate, occasionally nearly linear,
rarely oval, entire or serrulate, not glaucous
beneath, sparingly to moderately pubescent
at least in part on both sides, or glabrate to-
ward the end of the season, about as pu-
bescent at the apex as at the base, coriaceous
in age; aments subprecocious or coetaneous,
sessile or on a barren or 1-3 bracteate pe-
duncle to 8 mm long; bracts of the aments
persistent, dark brown to purplish black at
the apex, often with a lighter base; pu-
bescence of aments sericeus-pilose at first but
soon becoming crisped-villous and somewhat
entangled, the hairs usually exceeding the
bracts by 1-2 mm, sometimes deciduous;
staminate aments 1-2.5 cm long; stamens 2,
the filaments about 5 mm long, glabrous; pis-
tillate aments (1) 2-4 (6) cm long; capsules
3-6 mm long, glabrous, the stipe 1.5-2 mm
long; styles 0.3-1 (1.5) mm long. Riparian
and wet meadow communities from about
2075-3050 m, particularly common on the
plateaus of central Utah, but from all coun-
ties of the state except Millard, Morgan,

October 1983

Goodrich: Utah Flora, Salicaceae

543

Rich, Tooele, Washington, Wayne, and We-
ber and to be expected in some of these; Col-
orado Rockies west to northern California
and north to southern Alberta and British Co-
lumbia. Our plants are closely related to S.
myrtillifolia Anderss. of Alaska and Canada.
They vary from those of Alaska and Canada
by either taller stature or pubescent leaves or
both, and they have longer stipules that are
more sharply acute at the apex. They might
be treated as a variety of S. myrtillifolia, but
no new combination is proposed here. Some-
times referred to as S. pseudocordata, but this
name is synonymous with S. myrtillifolia
(Dom 1975). Occasionally grading toward S.
wolfii in pubescence of leaves and sometimes
difficult to distinquish from that species veg-
etatively. Like S. lutea in color and pu-
bescence of scales and rachis of aments, and
sometimes confused with that species, but
with leaves coriaceous in age and more and
persistently pubescent and not glaucous be-
neath, and generally of higher elevations, but
sometimes nearly impossible to distinguish
from S. lutea in leafless or very young-leaved
specimens with precocious aments. However,
older twigs of S. boothii are not whitish as
they often are in S. lutea, and specimens with
older twigs are more easily distinguished; 139

(xh).

Salix brachycarpa Nutt. Barrenground wil-
low. Short-fruited willow. Shrubs (0.25)
0.6-1.5 m tall, rarely taller; twigs below the
leaves with epidermis breaking in translucent
flakes, twigs of the season dark or reddish un-
der the dense pubescence; stipules inconspic-
uous, deciduous; petioles 1-4 mm long, usual-
ly not longer than the bud, often reddish, the
reddish color sometimes extending up the
midrib of the blade; leaf blades (0.6)
1.5-4 cm long, (3) 5-18 mm wide, or to 7 cm
long and 3 cm wide on sterile branches, 2-4
(5) times longer than wide, elliptical, broadly
lanceolate, occasionally nearly linear, entire,
thinly to moderately sericeous to nearly
glabrous on both sides, strongly glaucous be-
neath; aments coetaneous or serotinous,
nearly sessile or more often on bracteate pe-
dimcles at the ends of leafy twigs; bracts of
the aments pale green, tan, or light brown in
age, rarely pink or pale reddish at apex, scat-
tered to densely pilose on both sides, the
hairs exceeding the bract by about 1 mm or

less; staminate aments (6) 8-10 (12) mm long,
5-6 mm wide; stamens 2, the filaments
2.5-5 mm long, densely pilose at base and
scattered pilose to 1/3 to 3/4 the entire
length, the pubescent portion sometimes ex-
ceeding the scale, anthers 0.3-0.5 (0.6) mm
long, orbicular, yellowish; pistillate aments
8-25 mm long, 3-10 mm wide; capsules
3-5 mm long, densely arranged and mostly
concealing the rachis, sessile or on stipes up
to 0.5 (1) mm long, sometimes persisting over
winter, pubescent, the hairs persistent even
on over-wintering capsules, the style
0.5-1 mm long, the stigmas about 0.5 mm
long, bilobed to the base. Along streams, in
wet meadows, dry rocky and talus slopes, and
rocky, open ground in mountains from
2070-3230 m, mostly on ground with basic
substrate in Cache, Duchesne, Emery, Grand,
Iron, Juab, Kane, Salt Lake, Sanpete, Sevier,
Summit, Utah, and Wasatch counties; wide-
spread in Alaska, Canada, and south in west-
ern United States from Oregon south and east
to Colorado. Our plants are assignable to var.
brachycarpa with bracts greenish at anthesis
and subspherical or short cylindrical, densely
flowered pistillate aments, leaves coarsely
pubescent on both sides and with com-
paratively tall stature. Closely related to and
often confused with S. glauca, but distinct in
the state by small but numerous features. In
addition to the features given in the key, S.
brachycarpa more or less differs from S.
glauca in having twigs with more numerous
aments, distal leaves of fertile twigs often
considerably larger than the 3 or 4 proximal
ones, and reddish as well as yellowish petioles
with the reddish color sometimes extending
up the midrib of the leaf blade. Although
most of our plants seem quite distinct, appar-
ently there is widespread introgression with
S. glauca in the Rocky Mountain Region and
particularly southward in Colorado (Argus
1965); see discussion under S. glauca; 74
(xviii).

Salix cascadensis Cockerell. Cascades wil-
low. Depressed, mat-forming subshrubs,
1-3 cm tall, from tap root and rhizomatously
much-branched caudex; petiole lacking or to
3 mm long; leaf blades 6-18 mm long,
1.5-4 mm wide, 2-4.7 times longer than
wide, linear or narrow elliptical, entire,
pilose-sericeous when young, soon glabrous

544

Great Basin Naturalist

Vol. 43, No. 4

and green on both sides or slightly paler be-
low, some marcescent for 1 or more years;
aments coetaneous, terminal on short leafy
lateral branches, these about 8-22 mm long;
bracts of the aments persistent, black or
purplish black, reddish-purplish at the very
base, about 1-2 mm long, 1 mm wide, pilose
on both sides, but less so to nearly glabrous at
the base ventrally, the hairs about 1 mm
long; staminate aments 3-12 mm long,
5-8 mm wide; stamens 2, separate to the
base, the filaments about 3-4 mm long,
glabrous, the anthers reddish or purplish; pis-
tillate aments 5-22 mm long, 5-11 mm wide;
capsules 3-4 mm long, pubescent, sessile or
the stipe less than 1 mm long, the style and
stigmas together about 1.5 mm long. Alpine
tundra on the Uinta Mountains,
3350-3932 m, in Daggett, Duchesne, Sum-
mit, and Uintah counties; southwestern Brit-
ish Columbia, east to Montana south through
Wyoming to Colorado and Utah; 15 (iv).

Salix drummondiana Barratt in Hook.
Drummond willow. [S. subcoerulea Piper].
Shmbs (1) 2-3(4) m tall; twigs glabrous or
puberulent when very young, heavily
glaucous, the bloom persisting into the sec-
ond year, yellow-brown to blackish purple
beneath the bloom; stipules narrow, small
and deciduous, or larger and more persistent
on vigorous young shoots; petioles 4-12 mm
long; leaf blades 2.2-8 cm long, (5)
13-20 mm wide, or to 14 cm long and 3 cm
wide on vigorous young shoots, lanceolate or
narrowly elliptical, rarely oblanceolate, en-
tire, sometimes with slightly revolute mar-
gins, dark green and glabrous or thinly pu-
bescent above, densely silvery white
pubescence beneath with short appressed or
spreading and slighty tangled hairs, pale
glaucous beneath the pubescence; aments
precocious or sub-coetaneous; bracts of the
aments persistent, purplish black or purplish
brown, pilose on both sides, the longest hairs
exceeding the bract by 1.5-2 mm; staminate
aments 19-22 mm long, 3-10 mm wide, ses-
sile or on a peduncle to 3 mm long; stamens
2, the filaments, 4-9 mm long, glabrous; pis-
tillate aments 2-4.5 cm long, 3-12 mm wide;
capsules 3-6 mm long, pubescent, sessile or
the stipe to 1 mm long, the style 0.5-0.7 mm
long, the stigmas 0.3-0.6 mm long. Along
streams and rivers, wet meadows, and other

wet places from 2135-3140 (3290) m in Bea-
ver, Box Elder, Cache, Daggett, Davis, Du-
chesne, Emery, Grand, Piute, Salt Lake, San-
pete, Sevier, Summit, Uintah, Utah, and
Wasatch counties; British Columbia and Al-
berta south to California and New Mexico;
84 (xxv).

Salix exigua Nutt. Coyote willow. Dusky
willow. Narrow-leaf willow. Colonial shrub
(1) 2-3 m tall or rarely treelike and to 8 m
tall; stems ashy gray, branches often reddish,
twigs of the season greenish, pubescent;
leaves (1) 2-11 cm long, (0.1) 0.2-1 cm wide,
sessile or with 1-3 mm long petiole, or to
17.5 cm long and 1.6 cm wide with petiole
up to 12 mm long on vigorous yoving shoots,
linear, entire or serrulate-dentate with
glandular teeth, glabrate to densely white se-
riceous; aments coetaneous or serotinous on
slender leafy peduncles or twigs of the sea-
son, these 0.5-14 cm long; bracts of the
aments about 2 mm long, about 1 mm wide,
pale green or yellowish, deciduous, pu-
bescent on both sides but often glabrate or
glabrous dorsally especially toward the apex,
occasionally only ciliate ventrally; staminate
aments 1.5-4.5 cm long, 0.5-1 cm wide; sta-
mens 2, the filaments pilose on the lower
half; pistillate aments 1.5-6 cm long,
8-16 mm wide; capsules 4-7 mm long, most-
ly glabrous, sometimes pubescent, sessile or
the stipe up to 0.8 mm long, the style obso-
lete. Along rivers and streams, irrigation dit-
ches, washes, in neglected fields and pastures,
around ponds and reservoirs, tolerant of alka-
line soils, from 825-2315 (2590) m in all
counties of the state. Our plants are part of a
complex that extends from the Atlantic to the
Pacific in the northern United States and
southern Canada and extends from Alaska to
northern Mexico in the western part of North
America. Most of our specimens of ssp.
exigua have glabrous capsules and can be as-
signed to var. stenophyUa (Rydb.) Schneid.
Some specimens from the northern part of
the state have somewhat pubescent capsules
and these may be var. exigua. The closely re-
lated S. melanopsis Nutt. has been reported
for the area, but I have not seen any speci-
men that clearlv belongs to that taxon. 170
(ii).

Salix fragilis L. Crack willow. Large trees
to 20 m tall, the trunks to 1.3 m in diameter,
solitary or few, erect or strongly leaning,

October 1983

Goodrich: Utah Flora, Salicaceae

545

with thick fiirrowed gray or blackish gray
bark; branches ascending, often large; twigs
spreading, not pendulous, very brittle and
easily broken at the base; leaf blades lanceo-
late to narrow elliptic, (2.5)3-17 cm long,
(7)10-32 mm wide, acute or accuminate, ser-
rate, glaucous or glaucescent beneath,
glabrous or sericeous when young, glabrous
when mature; aments coetaneous, on twigs of
the current season, these twigs about
1-2.5 cm long, with (1)2-3(4) reduced leaves,
the leaves like the larger leaves of the non-
floriferous twigs but sometimes oblanceolate;
bracts of the aments pale green, pale yellow-
green, or greenish white, tan or very pale
brown upon drying, the pistillate ones de-
ciduoiLS by the time the capsules mature, se-
riceous with the hairs exceeding the bract by
about 1-1.5 mm; staminate aments
3.5-7(9) cm long, 9-12 mm wide; stamens 2;
filaments about 3-6 mm long, yellow, pilose
toward the base, the pilose portion about
equal or shorter than the subtending bract;
pistillate aments (2.5)4-7 cm long, 10-13 mm
wide; capsules 4-6 mm long, crowded but
usually not so dense as to conceal the rachis,
glabrous, the stipes about 1 mm long, the
styles 0.5-1 mm long, the stigmas 0.2-0.3 mm
long. Introduced from Eurasia, cultivated at
homes and along streets, persisting, escaping,
and naturalized along irrigation and natural
waterways and lake margins, solitary to form-
ing groves, from 1370-2075 m, in Beaver,
Box Elder, Cache, Duchesne, Juab, Rich, Salt
Lake, Sanpete, Sevier, Summit, Tooele, Uin-
tah, Utah, Wasatch, and probably most other
counties of the state. Hybrids of S. fragilis x
S. alba (S. x rubens Schrank.) have been de-
veloped (Rehder 1951). If such hybrids have
been cultivated in the state, they could add
considerable complication to the taxonomic
separation of the two species; 30 (xiv).

Salix geyeriana Anderss. Geyer willow.
Shnibs 1.5-4.5 m tall, twigs glabrous or scat-
tered puberulent, strongly glaucous, the
bloom sometimes deciduous; stipules minute
and deciduous; petioles 3-10 mm long; leaves
(1) 2-4.5 cm long (4) 8-12 mm wide, ellipti-
cal, narrow elliptical to narrow lanceolate,
entire or nearly so, glaucus beneath, sericeous
when unfolding, sparsely to moderately se-
riceous at maturity, especially below, the
hairs white or a few pale reddish; aments
subprecocious to coetaneous; peduncles of

aments leafy or bracteate, the staminate
2-5 mm long, the pistillate 3-10 mm long;
bracts of the aments persistent, sericeous-pi-
lose on both sides, or glabrate or glabrous
ventrally especially in age, the hairs exceed-
ing the bract by 0.5-1 mm, the staminate
ones light brown when very young, turning
reddish to purplish black at the tips, those at
the tips of aments turning first, the pistillate
ones greenish brown to brown; staminate
aments 7-15 mm long, 5-9 mm wide; sta-
mens 2, the filaments about 4 mm long, pi-
lose to about midlength, the pilose portion
about equaling or exceeding the bract; pistil-
late aments 1-2 cm long, 6-15 mm wide;
capsules 4-7 mm long, pubescent, the stipe
(1) 2-3 mm long, the style 0.2-0.3 mm long;
stigmas about 0.2 mm long. Along streams
and rivers and in other wet places from
2195-2895 m, most common in the Uinta
Mountains to Strawberry Valley, occasional
elsewhere in the state, in Beaver, Cache,
Daggett, Duchesne, Emery, Kane, Rich, Salt
Lake, Sevier, Summit, Uintah, Utah,
Wasatch, Washington, and Wayne counties;
southern British Columbia south to California
and east to Montana and Colorado. With
glaucous twigs and whitish pubescence, our
plants are assignable to var. geyeriana; 67
(xix).

Salix glauca L. Glaucous willow, Grayleaf
willow. [S. pseudolapponum Seem, in En-
gler]. Plants mostly low shrubs (0.1) 0.3-1
(3) m tall; twigs sometimes glaucous but
mostly not, those below the leaves with epi-
dermis exfoliating in translucent flakes, those
of the season reddish under whitish pu-
bescence, occasionally glabrate, often with a
tuft of pilose hairs at the node; stipules most-
ly small and soon deciduous; petioles (1) 2-6
(18) mm long, mostly yellowish or greenish,
the color often extending up the midrib of
the blade; leaf blades 2-55 mm long, 7-22
mm wide, or to 9 cm long and to 5 cm wide
on ends of vegetative twigs, elliptical, pu-
bescent when young to glabrate or glabrous
in age, mostly entire or rarely serrate; aments
coetaneous, nearly sessile on old twigs or
more often on bracteate peduncles or leafy,
current twigs; bracts of the aments persistent,
pale brown to blackish, pilose; staminate
aments 1.5-4 cm long; stamens 2, the fila-
ments free or united in imusual specimens,
glabrous or sparsely pilose at the base, the

546

Great Basin Naturalist

Vol. 43, No. 4

anthers 0.5-0.8 mm long; pistillate aments
1.5-5 cm long, 11-15 mm wide; capsules (4)
5-7 (9) mm long, densely pubescent to gla-
brate or glabrous in age, crowded but usually
not so dense as to conceal the rachis, the
stipes 0.5-2 mm long, the style 0.6-1 mm
long, the stigmas about 0.5 mm long. Along
streams, around springs, on talus slopes,
snowflush areas, and dry alpine timdra but
then usually in or near krummholz, from
2775-3660 m, on the Bear River, Uinta, and
Wasatch mountains and Wasatch Plateau in
Cache, Daggett, Duchesne, Salt Lake, San-
pete, Summit, Uintah and Wasatch counties;
circumboreal, south in western North Ameri-
ca in the Rocky Mountains to New Mexico.
Highly variable plants with geographic
phases. The Uinta Mountain plants from high
elevations are more or less comparable to
plants that have passed under the name of S.
pseiidolapponum, and they generally repre-
sent a rather low-statured phase. On wind-
swept alpine summits, these plants approach
the stature of S. arctica, but the stems are still
ascending to erect. These Uinta Mountain
plants tend more toward glabrescence in the
capsules and have darker scales than is typi-
cal of those in the Bear River Range. The
twigs are quite persistently pubescent.
Leaves are seldom over 5 cm long or over
2 cm wide. Plants from the Bear River and
Wasatch ranges have densely and persistently
pubescent capsules, pale brown to dark
brown to occasionally pinkish tan or rarely
whitish tan scales, the twigs are sometimes
early glabrate and some of the leaves are fre-
quently over 5 cm long and over 2 cm wide.
Specimens from Horseshoe Flat area of the
Wasatch Plateau have glabrous or pubescent
capsules, mostly dark scales, and glabrate and
unusual, distinctly serrate leaves. The varia-
bility in S. glauca nearly encompasses S.
brachycarpa. However, I prefer to follow
Argus (1965) and keep the two separate; 57
(xii).

Salix laevigata Bebb. Red willow. Shrub or
tree 2-15 m tall; twigs reddish brown or dull
brown, ashy red or ashy gray during exfolia-
tion, stipules inconspicuous or to 6 mm long
on vigorous vegetative twigs; usually de-
ciduous, petioles stout, 4-14 mm long; leaf
blades (1) 1.8-4 (6) cm long, 5-20 cm wide,
or to 19 cm long and 4 cm wide on vigorous
young shoots, narrowly to broadly lanceolate.

glandular-serrulate, somewhat revolute, usu-
ally thick and firm, dark green and glabrous
above, glaucous, and glabrous or pubescent
toward the base and along the midrib;
aments subprecocious to coetaneous, on leafy
or bracteate twigs of the season; bracts of
aments 1-2 mm long, at least the pistillate
ones deciduous, pale yellow, crinkly pilose on
both sides or often glabrous dorsally, entire
or erose to dentate at apex; staminate aments
3-6 cm long, about 1 cm wide; stamens 3-7,
pilose on lower half; pistillate aments 4-8

(11) cm long, to 1.5 mm wide; capsules 4-5
(6) mm long, glabrous, the stipes 1.5-2.5 mm
long, styles 0.1-0.2 mm long, equaling the bi-
lobed stigmas. Along drainages from
701-1370 m, in Washington, San Juan, and
probably Kane counties; Arizona, California,
Nevada, Utah, and northern Baja California.
Perhaps not distinct from S. bonplandiana
H.B.K., and treated as synonymous with that
taxon by Dorn (1977); 16 (0).

Salix lasiandra Benth. Whiplash willow;
Caudate willow. Shrub or small tree (2) 3-6

(12) m tall; twigs glabrous or finely hairy
when young; stipules often well developed,
broadly rounded, gland toothed, 2-10 mm
long, eventually deciduous; petioles 3-15
(25) mm long, often bearing 2 or more wart-
like glands on the upper side at or near the
base of the blade; leaf blades (2.2)
5.5-11.5 cm long, (5) 12-21 mm wide, or to
26 cm long and 5.5 cm wide on vigorous
young shoots, lanceolate, elliptical or narrow
elliptical, gradually long accuminate, closely
serrulate, glabrous except when very young;
aments coetaneous, on 1-3.5 cm long leafy-
bracteate twigs, the leaves or bracts of the
ament-bearing twigs 3-5 in number to 6.5 cm
long and 1.2 mm wide, deciduous after the
fruit matures; bracts of the aments deciduous
(at least the pistillate ones) by the time the
capsules start to open, 3-4 mm long, glabrous
or nearly on the upper half, pubescent to-
ward the base usually more so ventrally than
dorsally, entire or minutely toothed at the
apex with a few rounded teeth, the staminate
yellow, the pistillate pale greenish; staminate
aments 1.8-4.5 cm long, 3-12 mm wide; sta-
mens 3-8, usually 5, the filaments pilose; pis-
tillate aments 2-7 cm long, 11-18 mm wide;
capsules 4-8 mm long, glabrous, the stipe
1-2 mm long, the style 0.5-1 mm long, the

October 1983

Goodrich: Utah Flora, Salicaceae

547

stigmas to 0.5 mm long. Along streams and
rivers, on flood plains, occasionally along irri-
gation canals, or around ponds, and reser-
voirs, from 1525-2440 (262 l)m in Beaver,
Box Elder, Cache, Carbon, Daggett, Davis,
Duchesne, Emery, Garfield, Juab, Piute,
Rich, Salt Lake, Sanpete, Sevier, Summit,
Uintah, Utah, and Wasatch counties; to be
expected elseu'here; Alaska and Yukon to
California and New Mexico. Our plants are
var. caudata (Nutt.) Sudw, with leaves about
equally colored on both sides. Var. lasiandra
with leaves glaucous beneath has been re-
ported for the state, but I have seen no speci-
men; 98 (xxi).

Salix lasiolepis Benth. Arroyo willow.
Shrubs or small trees mostly 4-6 m tall in our
range; twigs yellowish olive to reddish, usual-
ly soft puberulent when young; stipules mi-
nute, soon deciduous or lacking, occasionally
well developed on vigorous young shoots;
petioles 3-15 mm long; leaf blades 1.5-4.2
cm long, 6-13 mm wide, or to 11 cm long
and 2.5 cm wide on vigorous young shoots,
usually oblanceolate or oblong, occasionally
elliptical, entire, rarely minutely toothed,
somewhat revolute margined, dark green and
glabrous above, at maturity glaucous be-
neath, more or less coriaceous, rather densely
soft pubescent on both sides when unfolding,
less so above than beneath, few to many of
the hairs persisting beneath at maturity;
aments precocious to subcoetaneous on
3-6 mm long bracteate or bare peduncles;
bracts of the aments persistent purple-black,
obovate with broad rounded apex, densely
villous, almost hidden in the hairs; staminate
aments 2.2-4.5 cm long; stamens 2, the fila-
ments glabrous; pistillate aments (1.8)
2.2-4.5 cm long (to 7 cm long outside of our
area), 10-12 mm wide; capsules 3-4 (5) mm
long, glabrous, the stipe 1-2 cm long, the
style about 0.5 mm long, the stigmas 0.2-0.3
mm long. Along streams, ditches, and washes
from about 1463-2328 m, in western Utah,
Great Basin and Virgin River drainages, in
Beaver, Iron, Juab, Millard, Sevier, Tooele,
Utah, and Washington counties; southern
British Columbia south to Baja California and
east to Idaho, Utah, Texas and northern Mex-
ico; 32 (xvii).

Salix lutea Nutt. Yellow willow. [S. I. var.
platyphylla Ball; S. /. var. watsonii (Bebb)

Jeps.]. Shrubs or rarely small trees but then
still generally several stemmed at the base;
(2) 3-5 (9) m tall; young twigs slender, yel-
lowish to reddish at first, often pale on one
side and red-purple on the other, glabrous;
older twigs and smaller branches often gray-
ish white; stipules small and inconspicuous or
to 1 cm long or more and leaflike in texture
on vigorous young shoots, usually deciduous;
petioles 1-11 (20) mm long; leaf blades (1)
2-5.5 cm long, (4) 9-21 mm wide or to
10.7 cm long and 3 cm wide on vigorous
young shoots, elliptical or lanceolate, rarely
linear, entire or occasionally serrulate,
glaucous beneath but hardly so when very
young, usually glabrous at maturity, the
lower surface glabrous from the first or less
pubescent than above, the upper surface
sometimes pubescent toward the base while
th'e leaves are unfolding; aments precocious
or subprecocious, on 1-7 mm long barren or
1-3 bracteate stalks; rachis and usually the
stalk of the aments covered with a tangle of
crisped-villous white hairs; bracts of the
aments persistent, pubescent with crisped-
villous, soon-entangled hairs, sometimes only
moderately pilose-woolly toward the base or
near the apex ventrally, the dorsal side usual-
ly glabrous toward the apex and often
throughout as the crinkly hairs are readily
deciduous; staminate aments 2-5 cm long,
about 1 cm wide; stamens 2, the filaments
glabrous, the anthers yellowish or turning
purple; pistillate aments 2-7 cm long, to
2 cm wide; capsules 3-6 mm long, glabrous,
mostly densely arranged on the rachis, occa-
sionally a little scattered, the stipe (1) 1.3-3
(4) mm long, the style 0.2-0.7 mm long, the
stigmas often scarcely bilobed. Along streams
and ditches in valleys and canyons and occa-
sionally on mountains from 1340-2255
(2350) m, in all counties of the state except
Beaver, Carbon, Davis, Iron, Morgan, and
Rich, and to be expected in some or all of
these; New Mexico to California and north to
Alberta. Our plants are closely related to and
possibly a part of the S. eriocephala Michx.
complex. They have been referred to as
S. rigida Muhl., but Argus (1980) has placed
S. rigida in synonomy under S. eriocephala.
He did not place S. lutea in synonomy, but
suggested that more study is needed. Until
such a study is made, I believe it best to re-
tain the traditional name of S. lutea for our

548

Great Basin Naturalist

Vol. 43, No. 4

plants. Salix ligulifolia Ball has been reported
for southern Utah. Tliis has been separated
from S. lutea by pedicels of capsules 1-2 mm
long versus (1)2-4.5 mm long in S. lutea, and
by having mostly entire rather than mostly
semilate leaves. At the varietal level such
plants are referrable to S. lutea var. liguli-
folia Ball. See discussion under S. boothii; 144
(xxv).

Salix monticola Bebb ex Coult. Shrubs
1.5-4 m tall; twigs yellowish when fresh,
drying blackish, pubenilent at first; stipules
small and inconspicuous or leaflike on vigor-
ous young shoots; petioles 5-10 (15) mm
long; leaf blades 2-5 cm long, 0.7-1.5 mm
wide or up to 1 1 cm long and 4 cm wide on
vigorous young shoots, mostly elliptical or el-
liptic-obovate, crenate-serrate or subentire,
slighty pubescent when very young, more so
above than beneath, usually glabrous when
fully expanded, glaucous beneath when ma-
ture; aments precocious or coetaneous, sub-
sessile or on short stalks to 1 cm long, often
subtended by bractlike leaves; bracts of the
aments persistent, dark brown to blackish, pi-
lose, or soon crisped-villous, the hairs exceed-
ing the bract by about 2 mm, more or less
tangled; staminate aments 2-3.5 cm long,
about 1-1.5 cm wide; filaments 2, glabrous;
pistillate aments 2-6 cm long, 1-1.5 cm
wide; capsules 4-7 mm long, glabrous, sub-
sessile, the stipe less than 1 mm long; style
0.7-1.8 mm long, longer than the stigmas.
Along streams and other wet places from
2195-3200 m, on mountains of eastern and
central Utah in Beaver, Garfield, Piute, San
Juan, Sanpete, Sevier, Uintah, and Wasatch
coimties, and to be expected elsewhere in
eastern Utah, but apparently uncommon in
the state; Rocky Mountains of southern
Wyoming, Colorado, Utah, Arizona, and
New Mexico. Closely allied to S. boothii and
S. lutea and rather easily confused with them.
Separation from S. boothii is often com-
pounded by the lack of glaucescence on
yoimg leaves; I am indebted to Dr. Robert
Dom for his annotations of specimens of this
taxon. More specimens are needed to gain a
better understanding of this plant in the
state; 12 (0).

Salix nigra Marsh. Black willow. [S. good-
dingii Ball]. Trees or occasionally shrubs (2)
6-10 (24) m tall; twigs of the season yellow-

ish, glabrous, or finely pubescent at first; stip-
ules to 8 mm long, more or less glandular,
usually quickly deciduous; petioles 3-7 mm
long; leaf blades 2-7.5 cm long, 6-16 mm
wide or to 10.2 cm long and 18 mm wide
with petiole to 15 mm long on vigorous vege-
tative twigs, narrowly to broadly lanceolate,
apex short to long acuminate, entire or more
often glandular-serrulate, greenish on both
sides, pubescent when unfolding but becom-
ing glabrous or glabrate; aments coetaneous,
on lateral 1-6 cm long twigs of the season
with 3-6 leaves or bracts; bracts of aments
pale green or pale yellow, soon fading to tan,
and at least the pistillate ones deciduous,
pubescent on both sides or glabrous toward
the apex, entire or with 1-3 minute, rounded
teeth; staminate aments 2.5-6.5 cm long,
5-10 mm wide; stamens 3-6, the filaments
pilose to about midlength; pistillate aments
1.5-6 cm long, 10-17 mm wide; capsules
4-7 mm long, glabrous, not so densely ar-
ranged as to conceal the rachis, the stipe
1-2 mm long, the style 0.1-0.3 mm long.
Along the Virgin and San Juan rivers and
other drainages in southern Utah, and up the
Green River to near Moab from 825-1585 m,
in Garfield, Grand, Kane, San Juan, and
Washington counties; widespread in the con-
tinental United States, southern New Bruns-
wick, and southern Quebec and Ontario, and
northeastern Mexico. Utah specimens quite
consistently have light-colored twigs and
have been called S. gooddingii. I feel as did
Archer (1965) that S. gooddingii is not clearly
distinct from S. nigra. Arthur Cronquist (un-
publ. ms.) has placed S. gooddingii in syn-
onomy under S. tiigra var. venulosa (An-
derss.) Bebb, and he recognized our plants as
being different in having smaller stature and
usually having some pubescence on the cap-
sules or stipes, or both, as well as having
light-colored twigs. However, he further
states that these features are not consistent. I
prefer to follow Cronquist's approach and
recognize the differences in our plants at the
varietal rather than at the specific level. Salix
nigra and S. amygaloides come together near
Moab on the Green River, and notes on
specimens from that area by Arthur
Cronquist indicate that the two hybridize at
that location; 23 (0).

October 1983

Goodrich: Utah Flora, Salic ace ae

549

Salix planifolia Pursh. Plainleaf willow. [S.
plnjlicifolia L. ssp. planifolia (Pursh) Hiito-
nen]. Shnibs 0.5-1.5 (4) m tall; twigs below
the leaves often with epidermis exfoliating in
translucent flakes or strips, younger twigs
typically glabrous and lustrous black or
purplish black, rarely glaucous in part; stip-
ules small and asually deciduous; petioles
2-10 mm long; leaf blades 1.2-3.8 (8) cm
long, 4-13 (30) mm wide, or to 5 (13) cm
long and 2 (5) cm wide on vigorous sterile
twigs, elliptical or narrow elliptical, soon
glabrous and dark green above, glaucous and
glabrous to sparingly pubescent below, entire
or rarely with minute teeth; aments pre-
cocious (at least the staminate) to coetaneous,
nearly sessile or rarely on a short, mostly bar-
ren pedvmcle to 0.5-1 cm long; bracts of the
aments persistent, blackish, scattered to
densely villose to pilose, the hairs usually ex-
ceeding the bract by about 2 mm; staminate
aments 10-25 mm long; stamens 2, the fila-
ments glabrous, about 6 mm long; pistillate
aments 2-4 cm long, 1-1.5 cm wide; capsules
3-7 mm long, typically pubescent at least
near the base, occasionally glabrous or nearly
so in age, the stipe mostly less than 1 mm
long, the style and stigmas together mostly
over 1.5 mm long. Streamside meadows,
around lakes and ponds and other wet places,
most abundant and sometimes forming wil-
low fields in the Uinta Mountains, scattered
on high points of the plateaus and mountains
of the central and southern part of the state,
from (2255) 2895-3660 m in Daggett, Du-
chesne, Garfield, Iron, Salt Lake, Sanpete,
Sevier, Summit, and Uintah counties; circum-
boreal, south to California and New England.
I have followed Argus (1973) in listing our
plants under S. planifolia rather than S. phij-
licifolia. Our plants mostly fall well within
the concept of var. monica (Bebb) Jeps.,
though a few taller plants with larger leaves
from moderate elevations of the major drain-
ages in the Uinta Mountains are apparently
var. planifolia. However, the differences are
merely of stature and of leaf size and the two
varieties are hardly worthy of separation; 39
(ix).

Salix reticulata L. [S. nivalis Hook.; S. n.
var. saximontana (Rydb.) Schneid.]. Caespi-
tose dwarf shrubs, stems creeping at or just
below the ground surface, the slender aerial

twigs rarely more than 2-3 cm long, usually
prostrate; stipules minute and deciduous or
none; petioles 1-8 (15) mm long; leaf blades
0.5-3 cm long, 0.3-2 cm wide, ovate, ob-
ovate, orbicular or occasionally broadly ellip-
tical, entire, glabrous, green above, glaucous
beneath, strongly reticulate veined; aments
subcoetaneous, but mostly serotinous on the
ends of shoots of the season; bracts of the
aments persistent, pale green or yellowish,
sometimes with reddish tops, spatulate or
obovate, glabrous or sparsely pubescent ven-
trally, especially toward the margin, with
short hairs that extend less than 1 mm beyond
the bract; staminate aments 0.5-2 cm long,
slender, the flowers loose and not concealing
the puberulent rachis, on a slender glabrous
peduncle about 10-12 mm long; stamens 2;
fikments 1.5-2 mm long, glabrous or pilose
toward the base; anthers soon reddish or
purple; pistillate aments 5-15 mm long, 5-8
mm wide, on a slender 1-2 mm long pe-
duncle; capsules 1.5-3 mm long, pubescent
or glabrous in age, sessile or the stipe to
0.5 mm long, the style obsolete or to 0.2 mm
long, the stigmas about 0.1-0.2 mm long.
Open rocky slopes and ridges and alpine tun-
dra from 2987-3965 m, on the LaSal, Uinta,
and Wasatch Mountains in Duchesne, Grand,
Salt Lake, San Juan, Summit, and Utah coun-
ties; circumboreal, south in the mountains of
western North America to California, New
Mexico, Utah, and Colorado. Most of our
plants are referable to var. saximontana (L.)
Kelso, which may not be distinct from var.
reticulata. A few specimens seem to be like
var. nivalis (Hook.) Anderss. The features
used for separation seem to be poorly corre-
lated in our plants. Some plants with pistil-
late aments less than 1 cm long (that should
be var. nivalis) have leaves well over 15 mm
long, which is indicative of the other variety;
32 (0).

Salix scouleriana Barratt in Hook. Scouler
willow. Shrubs or small trees 3-7 m tall; stip-
ules small and inconspicuous or large and
leaflike on vigorous young shoots, eventually
deciduous; petioles 2-11 mm long; leaf
blades 2-6 cm long, (0.8) 1-3 cm wide or to
11.5 cm long and 4 cm wide on vigorous
young shoots, obovate to oblanceolate,
rounded to acute or occasionally accuminate
at the apex, entire or finely serrate, or occa-

550

Great Basin Naturalist

Vol. 43, No. 4

sionally coarsely crenate or serrate on larger
leaves of vegetative twigs, densely crisp-hairy
or sericeous, especially beneath as they un-
fold, the mature ones dark green and
glabrous above except sometimes puberulent
along the midrib, the lower side strongly
glaucous, sparsely puberulent with trans-
lucent whitish or rusty minute hairs, or occa-
sionally densely felty-villous; bracts of
aments blackish or purplish black nearly
throughout, reddish or pale at the very base,
sericeous-pilose on both sides, the hairs at the
apex usually exceeding the bract by
1.5-2 mm; staminate aments 15-35 mm long,
nearly as wide as long, strictly precocious,
nearly sessile or on thickened bracteate pe-
duncles to 7 mm long, the bracts 3-4 mm
long, about 2 mm wide, pale green to whit-
ish, sericeous; stamens 2, the filaments to
11mm long at maturity, glabrous; pistillate
aments 2-6 cm long, 13-17 mm wide, pre-
cocious or subcoetaneous, nearly sessile or on
thickened bracteate peduncles to 17 mm
long, the bracts to 7 mm long and 2 mm
wide, not at all leaflike; capsules (5) 6-9 mm
long, pubescent, rarely nearly sessile, usually
on a 1-3 mm long stipe, the style 0.3-0.4 mm
long, rarely shorter, the stigmas 0.5-1 mm
long. Around springs, along streams, and on
well-drained slopes in aspen and conifer
woods, from (1400) 2377-2835 (3355) m, in
Box Elder, Cache, Carbon, Daggett, Davis,
Duchesne, Garfield, Grand, Juab, Millard,
Rich, Salt Lake, Sanpete, San Juan, Sevier,
Summit, Tooele, Uintah, Utah, Wasatch,
Washington, and Weber counties; Alaska and
Yukon to California, Arizona, and New Mexi-
co. Salix scouleriana is most closely allied to
S. humilus Marshall and to S. discolor Muhl.
of eastern United States and Canada. Salix
discolor (pussy willow) may be cultivated in
our area. It is generally distinguished from S.
scouleriana by: looser aments with longer
stipes (1.5-3 mm), more elliptic, pointed, and
toothed leaves that are usually more quickly
and fully glabrate, but none of these features
is wholly consistent (Hitchcock and
Cronquist, 1964). Occasionally specimens
have leaves densely pubescent beneath. Ar-
now et al. (1980) attributed this to hybridiza-
tion with S. drummondiana; 69 (xi).

Salix tvolfii Bebb in Rothr. Wolf's willow.
Shrubs 0.6-1.5 (2) m tall; twigs yellow to or-

ange when young, chestnut brown in age,
those of the season thinly villous-puberulent;
stipules 1-5 mm long, often glandular-serru-
late, eventually deciduous; petioles 2-10 mm
long; leaf blades 1.2-4.2 cm long, 5-13 mm
wide or to 5.3 cm long and 16 mm wide to-
ward the ends of vigorous vegetative twigs,
narrow elliptical, linear-lanceolate, or occa-
sionally oblanceolate, entire, sparsely to
densely sericeous-tomentose on both sides
even in age or glabrate beneath very late in
the season; aments coetaneous or sub-
serotinous, nearly sessile or on bracteate pe-
duncles to 1 cm long; bracts of the aments
persistent, blackish or pale at the very base,
pilose-sericeous on both sides, the hairs ex-
ceeding the bract by about 1 mm; staminate
catkins 10-15 mm long, about 8-10 mm
wide; stamens 2, the filaments about 3-4 mm
long, glabrous; pistillate aments 8-20 (30)
mm long, 6-10 mm wide; capsules 3-5 mm
long, glabrous or rarely pubescent, the stipe
less than 1 mm long, the style about 0.5 mm
long, the stigmas about 0.2 mm long. Along
streams and around the margins of lakes and
ponds, occasionally forming willow fields, in
the Bear River and Uinta mountains and
West Tavaputs and Wasatch plateaus from
2470-3290 m, in Cache, Daggett, Duchesne,
Emery, Summit, Uintah, and Wasatch coun-
ties. Oregon to Montana, south to Nevada,
Utah, and Colorado. Our plants are var. tvolf-
ii with mostly glabrous capsules. One speci-
men (B. Maguire, D. Hobson, & R. Maguire
14104) from White Pine Lake, Cache Coun-
ty, has pubescent capsules and leaves that are
larger than others from the state. This speci-
men is like S. wolfii var. idahoensis Ball,
which is known from well north and west of
Utah. Other specimens from the vicinity of
White Pine Lake and other points in the
Bear River Range have glabrous capsules,
and I prefer not to list var. idahoensis for the
state based on this one specimen. The plants
from the Bear River Range with pistillate
aments 15-30 mm long do, however, seem in-
termediate toward var. idahoensis when com-
pared to those of the Uinta Mountains with
pistillate aments 8-15 mm long. The speci-
men with pubescent capsules and somewhat
large leaves is probably the basis of reports of
S. commutata Bebb for Utah; 44 (ix).

A MAMMALIAN HUMERUS FROM THE UPPER JURASSIC OF COLORADO

Donald R. Prothero' and James A. Jensen-

Abstract.— The first reported mammal fossil from Dry Mesa Quarry (Upper Jurassic Morrison Formation, Mesa
Countv, Colorado) is the distal end of a right humerus. It is very similar to humeri described by Jenkins (1973) from
the Morrison Formation at Como Bluff, Wyoming. It has a distinct ulnar condyle and a spiral humero-ulnar joint,
both features found in prototherian mammals but not in therians.

Postcranial remains of Jurassic mammals
are extremely rare. An articulated skeleton of
a dryolestid therian mammal has been report-
ed from the Jurassic of Portugal (Henkel and
Krebs 1977), but is still undescribed. Early
Jurassic mammalian postcranial fossils are
also known from India (Datta et al. 1978),
but are undescribed. A few fragmentary post-
cranial remains of mammals have been de-
scribed from the Upper Jurassic of England
(Seeley 1879, Simpson 1928, Haines 1946)
and from the Upper Jurassic Morrison For-
mation of Wyoming (Jenkins 1973). Of the
five important mammal-producing localities
in the Morrison Formation (listed in Clemens
et al. 1979:23-26), two have produced mam-
malian postcranial fossils prior to this paper:
Como Bluff, Wyoming (Jenkins 1973), and
the Fruita Paleontological Area, Mesa Coun-
ty, Colorado (Rasmussen and Callison 1981).

In 1977, the distal portion of a right hu-
merus of a mammal was found in Dry Mesa
Quarry, Mesa County, Colorado. This speci-
men (BYU 2026) was first mentioned by Cle-
mens et al. (1979:24), and is described below.

Locality and Associated Fauna

Dry Mesa Quarry is located in the lower
section of the Brushy Basin Member of the
Upper Jurassic Morrison Formation, 135 feet
below its contact with the overlying Cre-
taceous Cedar Mountain Formation. The
quarry sediments include very fine to coarse
sands, grits, and fine gravels containing angu-
lar to well-rounded clay and bone pebbles.
Stream gradient was sufficient to move very
large bones, with the long axes of all large

bones usually oriented at right angles to the
stream flow. Sorting was biased by shape
rather than by size.

Sediments overlying the bone layer are
predominantly light-colored, cross-bedded
sands with occasional lenses of clay and fine
gravel, the latter often containing clay peb-
bles. Sediments underlying the bone layer are
principally a light, blue-green clay with oc-
casional traces of bright yellow zones of
oxidation.

The bone layer consists of an imusual vari-
ety of disarticulated bones of all sizes, includ-
ing specimens representing crocodilians, fish,
turtles, pterosaurs, four new theropods, an
unknown variety of sauropods, some ornitho-
pods, and the mammal described herein. Due
to the great variety of disarticulated bones in
the deposit, and the generic novelty of the
fauna, descriptive work has been postponed
until enough material is available. Field work
has been carried out for the last 10 years. The
following have been identified so far:

Lungfish tooth plate (probably Ceratodus—
K. Thomson, pers. comm.)

Pterodactyloid phalanx (Jensen and Ostrom
1977)

Torvosaurus tanneri, a megalosaur (Galton
and Jensen 1979)

Prototherian mammal humerus (this paper)

Description

BYU 2026 (Fig. 1) is the distal portion of a
right humerus of a mammal. It has been bro-
ken at midshaft, but is otherwise well pre-
served. There is relatively little evidence of
crushing or distortion. The shaft cross-section

'Department of Geology, Knox College, Galesburg, Illinois 61401.
■Earth Science Museum, Brigham Young University, Prove, Utah 84602.

551

552

Great Basin Naturalist

Vol. 43, No. 4

is a mediolaterally compressed triangle with
the apex pointing anteriorly. Distally the
shaft expands transversely to form the large
medial and lateral epicondyles. The distal
end of the hiunerus is naturally flattened an-
teroposteriorly. The long axis of the distal
end is not perpendicular to the major axis of
the shaft cross-section, but is rotated about 20
degrees clockwise (viewed from the distal
end). The shaft of the humerus has a strong
anterior crest that is deflected anterolat-
erally. This crest is probably the distal end of
the deltopectoral crest. A faint posterolateral
crest merges with the lateral epicondyle.

The medial epicondyle is considerably
more prominent than the lateral epicondyle.
It flares medially and is anteroposteriorly
compressed. The entepicondylar foramen is
visible in anterior view. It is broken at the
anterior end, where it passes anteromedially.
The lateral epicondyle merges with the radial
condyle. It is connected to the shaft of the
humerus by a thin posterolaterally arched
crest. The radial and olecranon fossae are in-
terconnected, forming an apparent supra-
trochlear foramen. This feature may be an ar-
tifact of breakage, however.

In anterior view, the main body of the
shaft bifurcates to form crests joining the
radial and ulnar condyles. These crests sur-
round the radial fossa. In posterior view, the
olecranon fossa is broadly concave and ex-
tends partially up the shaft. From this view
the apparent supratrochlear foramen has an
irregular margin that is clearly enlarged by
breakage.

In distal view, three main features are
seen: the medial epicondyle, the ulnar con-
dyle, and the radial condyle-lateral epicon-
dyle. The latter two features are confluent
and separated only by a shallow groove. The
ulnar and radial condyles, on the other hand,
are separated by a narrow, deep inter-
condylar groove. The radial condyle is broad
and bulbous in anterior view. The spiral ul-
nar condyle is very similar to that shown by
Jenkins (1973, Fig. 13). It is wrapped around
the distal end of the humerus, with a prox-
imolaterally oriented extensor surface and a
proximodistally oriented flexor surface. How-
ever, the anterior portion of the ulnar con-
dyle is more bulbous than the same feature in
the humerus figured by Jenkins (1973, Fig.

Fig. 1. BYU 2026, distal end of a right humerus. A,
Anterior view. B, Posterior view. C, Distal view. Abbre-
viations: dpc, deltopectoral crest; enf, entepicondylar
foramen; le, lateral epicondyle; me, medial epicondyle;
of, olecranon fossa; re, radial condyle; rf, radial fossa; uc,
ulnar condvle.

18). The axis of the ulnar condyle as it crosses
over the distal end of the humerus is at an
approximately 60 degree angle to the trans-
verse (interepicondylar) axis of the humerus
(seen in distal view). This compares with an-
gles of 58-65 degrees reported by Jenkins
(1973:286) for several humeri from Como
Bluff, Wyoming.

Discussion

The Dry Mesa Quarry mammal very close-
ly resembles the Como Bluff humeri de-
scribed by Jenkins (1973). It differs from
them in having a more bulbous and broader
ulnar condyle. In this respect, it is more like
the humeri referred to the multituberculate
Catopsolis by Jenkins (1973, Fig. 19). The
multituberculates Tugribataar (Kielan-Jawo-
rowska and Dashzeveg 1978) and Ptilodus
(Gidley 1909), the triconodont Eozostrodon
(Jenkins and Parrington 1976), and the mon-
otremes (Howell 1937, Haines 1946) also
have prominent bulbous ulnar condyles. BYU
2026 clearly does not have a trochlear con-
dyle, which Jenkins (1973) considers charac-
teristic of therian mammals.

The only other feature that distinguishes
the Dry Mesa Quarry mammal from the
Como Bluff humeri is the apparent supra-
trochlear foramen. As noted above, this fea-
ture may be an artifact of breakage.

October 1983

Prothero, Jensen: Jurassic Mammalian Humerus

553

The affinities of BYU 2026 are difficult to
assess based on such hmited evidence. The
presence of a distinct ulnar condyle with a
spiral humero-ulnar joint is characteristic of
prototherian mammals (Jenkins 1973). The
advanced therian trochlear condyle is known
from rocks as old as the Lower Cretaceous
(Jenkins 1973, footnote 3). The Dry Mesa
Quarry mammal humerus could have be-
longed to a number of prototherian mammal
taxa presently known from the Morrison For-
mation (Prothero 1981, Clemens et al. 1979).
It could also have come from some of the
primitive Morrison therian mammals that
may or may not have had a trochlear con-
dyle. Until the Portuguese dryolestid therian
skeleton (Henkel and Krebs 1977) is fully de-
scribed, we cannot rule out the possibility
tliat the Dry Mesa Quarry humerus belonged
to a very primitive therian mammal. Of the
possible candidates among nontherian mam-
mals, BYU 2026 resembles the known humeri
of multituberculates and triconodonts. The
skeleton of docodonts is presently unknown.
The systematic affinities of the Dry Mesa
Quarry mammal cannot be determined more
precisely at present than Mammalia incertae
sedis.

Acknowledgments

The Dry Mesa Quarry has been worked for
10 years by the Earth Science Museum of the
Brigham Young University under permits
from the U.S. Forest Service, as authorized
by the Antiquities Act of 1906. Funds for this
work have been provided by research sup-
port grants from Brigham Young University,
with additional support from the National
Geographic Society, Kenneth R. Thomson,
and several other private donors.

Measurements of angles were made with
an E.P.O.I. Shopscope. D.R. Prothero was
supported by a Columbia Faculty Fellowship
diu-ing preparation of this paper. We thank
Henry Galiano for advice on the illustration.

Literature Cited

Clemens, W. A., J. A. Lillec;raven, E. H. Lindsay, and
G. G. Simpson. 1979. Where, when and what— a
survey of known Mesozoic mammal distribution.
Pages 7-58 in J. A. Lillegraven, Z. Kielan-Jawo-
rowska, and W. A. Clemens, eds., Mesozoic
mammals: the first two-thirds of mammalian his-
tory. Univ. of California Press, Berkeley.

Datta, p. M., p. Yadagiri, and B. R. J. Rao. 1978. Dis-
covery of Early Jurassic micromammals from up-
per Gondwana sequence of Pranhita (iodavari
Valley, India. Geol. Soc. India J. 19:64-68.

Galton, p. M., and J. A. Jensen. 1979. A new large
theropod dinosaur from the Upper Jurassic of
Colorado. Brigham Young Univ. Geology Studies.
26(2):1-12.

Gidley, J. W. 1909. Notes on the fossil mammalian
genus Ptilodus, with descriptions of new species:
Proc. U.S. National Museum .36:611-626.

Haines, R. VV. 1946. A revision of the movements of the
forearm in tetrapods. J. Anatomy 80:1-11.

Henkel, S., and B. Krebs. 1977. Der erste Fund eines
Saugetier-Skelettes aus der Jura-Zeit. Umschau in
Wissen.schaft und Technik 77:217-218.

Howell, A. B. 1937. The swimming mechanism of the
platypus. J. Mammal. 18:217-222.

Jenkins, F. A., Jr. 1973. The fimctional anatomy and ev-
olution of the mammalian humero-ulnar articu-
lation. Amer. J. Anat. 137:281-298.

Jenkins, F. A., and F. R. Parrington. 1976. The post-
cranial skeletons of the Triassic mammals Eo-
zostrodon, Megazostrodon, and Enjthrotherium.
Roy. Soc. London Philos. Trans., B (Biol. Sci.)
27.3:387-431.

Jense.n, J. A., and J. H. Ostrom. 1977. A second Jurassic
pterosaur from North .-America. J. Paleont.
51:867-70.

Kielan-Jaworowska, Z., and D. Dashzeveg. 1978. New
Late Cretaceous mammal locality in Mongolia
and a description of a new multituberculate.
Acta Palaeont. Polonica 23:115-130.

Prothero, D. R. 1981. New Jurassic mammals from
Como Bluff, Wyoming, and the interrelationships
of the non-tribosphenic Theria. Bull. Amer. Mus.
Nat. Hist. 167 (art ,5):277-326.

Rasmussen, T. E., and G. Callison. 1981. A new spe-
cies of triconodont mammal from the upper Ju-
rassic of Colorado. J. Paleont. 55:628-6.34.

Seeley, H. G. 1879. Note on a femur and humerus of a
small mammal from the Stonesfield Slate. Quar-
terly Journal (Teological Society of London
35:456-463.

Simpson, G. G. 1928. A catalogue of the Mesozoic mam-
malia in the Geological Department of the Brit-
ish Museum. British Museum (Natural History),
London.

BATS OF THE COLORADO OIL SHALE REGION

Robert B. Finley, Jr.,' William Caire,' and Dallas E. Wilhelm'

Abstract.— New records for Myotis californicus, M. evotis, M. leibii, M. lucifugus, M. thysanodes, M. volans, M.
yunuinensis, Lasionycteris noctivagans, Pipistrelhis hespertis, Eptesicus fiiscus, Lasiuriis cinereus, Plecotus town-
sendii, and Antrozous pallidus and their habitat occurrence in northwestern Colorado are reported. Mortality of 27
bats of six species trapped in an oil sludge pit is described.

In 1974 the National Fish and Wildlife
Laboratory began field work in the Piceance
Basin as part of a survey of the vertebrates of
the coal and oil shale regions of northwestern
Colorado. The information was needed as
baseline data in preparation for energy devel-
opment and to better define the poorly
known distribution of mammals in this area.

From the oil shale region of Colorado,
roughly defined as Rio Blanco and Garfield
counties west of a line between Meeker and
Rifle, records of eight species of bats were
summarized by Armstrong (1972), seven of
which had been reported only from the vi-
cinities of Meeker or Rifle. Since then speci-
mens of five additional species have been col-
lected and numerous locality and habitat
records obtained. This information is report-
ed herein to make it available for manage-
ment decisions, and to facilitate and stimu-
late further work on the bats of northwestern
Colorado.

Study Area and Methods

The elevation of the Oil Shale Region
ranges approximately from 1,585 to 2,805 m,
falling within the Upper Sonoran, Transition,
and Canadian life zones of Gary (1911). The
Roan Plateau extends east-west, roughly
along the Rio Blanco-Garfield county line, di-
viding the two main drainages, the Colorado
River to the south and the White River to the
north. The region is semiarid with pre-
dominantly shaly alkaline soils and has only a
few long permanent creeks, with many short

intermittent tributaries. The few permanent
creek bottoms are occupied by ranches and
irrigated hay meadows. Low cliffs and rock
ledges border some of the creeks, and high
cliffs mark the southern rim of the Roan
Plateau and the western rim of the Cathedral
Bluffs east of Douglas Creek.

Specimens were obtained primarily by
mist-netting and were preserved as study
skins and skulls or in liquid. All specimens ex-
amined are in the Biological Surveys/ Fort
Collins collection (BS/FC) of the U.S. Fish
and Wildlife Service, unless otherwise in-
dicated: (CU) Colorado University Museum
or (KU) Kansas University Museum of Natu-
ral History.

Results

The known kinds of bats from the Colo-
rado oil shale region include seven species of
Myotis and one species in each of six other
genera {Lasionycteris, Pipistrellus, Eptesicus,
Lasiurus, Plecotus, and Antrozous). Species
found to be most common were the hoary bat
{Lasiurus cinereus cinereus), long-legged
myotis {Myotis volans interior), long-eared
myotis {Myotis evotis evotis), and small-foot-
ed myotis {Myotis leibii melanorhinus). The
spotted bat {Euderma maculatum) has been
reported in some environmental impact re-
ports as "potentially" or "possibly" present;
yet to our knowledge no specimen has been
taken in the oil shale region. One was picked
up in Browns Park, 65 km north of the oil
shale region, in 1981 by J. Creasy (Finley and
Creasy 1982).

'U.S. Fish and Wildlife Service, 1300 Blue Spruce Drive, Fort Collins, Colorado 80524-2098.
'Department of Biology, Central State University, 100 North University Drive, Edmond, Oklahoma 73034.
'Department of Biology, Hastings College, P.O. Box 269, Hastings, Nebraska 68901.

554

October 1983

FiNLEY ET AL.: COLORADO BaTS

555

Accounts of Species

Myotis californicus stephensi Dalquest

California Myotis

Specimens examined (5).— Garfield Co.: 5'/2 mi N, 2
mi W Rifle, 5,900 ft, 5.

Five California myotis, four unsexed and
one female, were found partially decomposed
in an oil sludge pit on the east side of the val-
ley of Government Creek north of Rifle.
Four specimens were picked up on 17 Sep-
tember 1974 and one on 17 May 1975. They
are referred to stephensi on geographic
grounds. The circumstances of these and 22
other bat casualties at the same sludge pit are
described in this paper under "Effects of
Energy Development on Bats."

The National Museum of Natural History
has a specimen taken 14 August 1907 by
Merritt Cary (1911) in a house 7 mi W Rifle,
5,300 feet.

Myotis evotis evotis (H. Allen)

Long-eared Myotis

Specimens examined (31).— Garfield Co.: 5'/2 mi N, 2
mi W Rifle, 5,900 ft, 7; Rio Blanco Co.: Cascade Gulch,
T3S, R95W, Sec 8, 1; Duck Cr tank, TIS, R98W, Sec 7,
1; Little Duck Cr, TIS, R98W, Sec 10, 6,200 ft, 10; Ryan
Gulch, 24 mi W, 10 mi S Meeker, 6,500 ft, 5; 25 mi'w,
10 mi S Meeker, 6,400 ft, 2; 1 mi N, 4 mi W Rio Blanco,
6,900 ft, 5.

Seven long-eared myotis carcasses (1 male,
3 females, and 3 unsexed) were picked up on
the bank of the sludge pit north of Rifle on
17 September 1974. This species was the
most abundant bat taken at pools adjacent to
big sage (Artemisia tridentata), greasewood
(Sarcobatus vermiculatus), and pinyon-juni-
per habitats. Sandstone ledges or low cliffs
were also present within a few hundred yards
at all localities.

The most productive mist netting site was
on Little Duck Creek, at 2,000 m elevation.
There on 7 August 1975 Caire and Finley
collected 10 M. evotis, 2 M. leibii, 4 M. vol-
ans, 3 Eptesicus fuscus, and 2 Lasiurus cine-
rus. The valley bottom was covered with tall,
dense big sagebrush and greasewood on deep
alluvial soil. Little Duck Creek was en-
trenched in an arroyo about 3-4V2 m deep.
The steep upper slopes of the valley were of
nearly bare shaly soil. Higher rock outcrops

along the rim supported old mature junipers
(Juniperus osteosperma).

The water in the arroyo was barely enough
to provide a flow, but a deposit of silt at the
mouth of a side gully had dammed the flow
and formed a long pool about 4 m wide and
20 cm deep. Nets were set over this pool and
another about 40 m downstream, where we
dammed the stream with a shovel, creating a
pool.

One long-eared myotis caught at Little
Duck Creek had a large laceration in the left
wing membrane, which had fully healed. The
study skin (BS/FC 2119), pinned with wings
fully extended, has the posterior margin of
the membrane between the tibia and the 5th
metacarpal deeply concave so that the width
(chord) of the left wing measures only 22
njm, compared with 37 mm at the same
place on the right wing. A lobe of smooth
scar tissue extends anteriorly from the de-
formed posterior margin, leaving only an 8
mm band of normal membrane between the
scar tissue and the radius.

In Ryan Gulch Wilhelm netted bats at a
steel stock tank by a windmill on 15 August
1974. He took 5 Af. evotis, 5 M. volans, and 2
M. leihii. The terrain and habitat there were
almost the same as on Little Duck Creek, ex-
cept that Ryan Gulch was dry at the time
and not so deeply gullied. Less than a mile
away in June two macerated skeletons of M.
evotis that had apparently drowned in a
small watering trough were found.

Three of four female M. evotis taken on 4
August 1977 on Piceance Creek, 1 mi N, 4
mi W Rio Blanco, were lactating or recently
lactating. The habitat there is described in
the M. lucifugus account. Of the 26 total
specimens of long-eared myotis that were
sexed, 14 were females and 12 were males.

Myotis leibii melanorhinus (Merriam)

Small-footed Myotis

Specimens examined (14).— Garfield Co.: 3 mi N
Douglas Pass, 7,000 ft, 1; 4.6 mi W Rifle (by rd), T6S,
R94W, Sec 14, 1; 5'/2 mi N, 2 mi W Rifle, 5,900 ft, 5; Rio
Blanco Co.: Cascade Gulch, T3S, R95W, Sec 8, 2; Little
Duck Cr, TIS, R98W, Sec 10, 2; Ryan Gulch stock tank,
24 mi W, 10 mi S Meeker, 6,500 ft, 2; Ryan Gulch, 25
mi W, 10 mi S Meeker, 1.

Five M. leibii were picked up (4 on 17
September 1974 and 1 on 29 April 1975) on

556

Great Basin Naturalist

Vol. 43, No. 4

the bank of the same oil sludge pit north of
Rifle where 5 M. califomicus and 7 M. evotis
were found. There were 2 females, 1 male,
and 2 imsexed carcasses.

This species seems to be found mainly at
lower and intermediate elevations in sage-
bnish, greasewood, and pinyon-juniper habi-
tats, but is less abundant there than M. evotis.
Our highest record of occurrence was at
2,130 m, 3 mi N Douglas Pass, where one
was netted along with Lasiunis cinereus on 2
August 1977 over an earthen stock pond. The
adjacent hillsides supported mixed chaparral,
sagebrush, and grassland with scattered
Douglas-fir (Pseudotsuga menziesii) and juni-
pers (Junipenis).

A female taken on 22 July 1975 at a sheep
stock tank 4.6 mi W Rifle contained an em-
bryo 20 mm long. Of the 12 individuals col-
lected that were sexed, half were females.
The eight skins at hand show a wide varia-
tion in color of the back and sides, from light
buffy or reddish brown to dull medium
brown, but have uniformly blackish ears and
wings.

Myotis lucifugus carissima Thomas

Little Brown Bat

Specitnens examined (3).— Garfield Co.: 4 mi W, 1 mi
S Rifle, 5,300 ft, 1; 1 mi N, 4 mi W Rio Blanco, 6,900 ft
2.

A female was taken on 28 August 1975
over a sheep watering pond in the Colorado
River valley west of Rifle. The pond had
been bulldozed in alkali soil on a greasewood
flat south of cliffs and a steep rock slope with
pinyon-juniper.

Two males were taken in a net across up-
per Piceance Creek west of Rio Blanco on 4
August 1977. At that point the valley is nar-
row between steep canyon sides covered with
pinyon-juniper and mountain shrub, with
Douglas-fir in the tributary gulches. The val-
ley bottom was grazed by sheep, and the
creek was barely flowing between long shal-
low pools. Other bats taken there were four
M. evotis and one Lasionycteris.

Cary (1911:206) collected two M. luci-
fugus on the White River meadows a few
miles east of Meeker in August 1905.

Myotis thysanodes thysanodes Miller

Fringed Mvotis

Specimens examined (1).— Rio Blanco Co.; 11 mi W
Meeker, '/2 mi S Hwy 64, Hav Gulch Rd, 1 (CL').

A male fringed myotis was netted by Jerry
Freeman on 17 August 1978 at a stock tank
where Hay Gulch opens into the White River
Valley.

Myotis volans interior Miller

Long-legged Myotis

Specimens examined (14).— Garfield Co.: 2 mi E Rio
Blanco, 1; Rio Blanco Co.: Little Duck Cr, TIS, R98W,
Sec 10, 4; Little Hills Game Research Station, 15 mi W,
1.9 mi S Meeker, 4 (CU); Ryan Gulch, 24 mi VV, 10 mi S
Meeker, 5,600 ft, 5.

Myotis volans occurs in the sagebrush,
greasewood, and pinyon-juniper habitats
along with M. evotis and M. leihii. Specimens
were taken at sites described under the ac-
counts of these species, and also at a more
wooded site described under Lasiurus
cinereus.

Two males and two females were taken in
the barn at the headquarters of the Little
Hills Game Research Station by Bissell,
Olivas, and Webb of the Colorado Division
of Wildlife on 29 June and 7 July 1977. The
barn is in a gulch flanked by rocky slopes
with pinyon-juniper. One M. ywnanensis and
one Eptesicus fuscus were also obtained
there.

Eight of the 14 M. volans examined were
males and 6 were females.

Myotis yumanensis yumanensis (H. Allen)

Yuma Myotis

Specimens examined (4).— Rio Blanco Co.: Little Hills
Game Research Station, 15 mi W, 1.9 mi S Meeker, 1
(CU); Rio Blanco Lake, 3 (CU).

Three female Yuma myotis were taken on
6 and 7 July 1977 by Ribic and Olivas in the
attic of a deserted house by Rio Blanco Lake,
a small reservoir on the White River near the
mouth of Piceance Creek. One contained
three embryos 4 mm in length.

Lasionycteris noctivagans (Le Conte)

Silver-haired Bat

Specimens examined (8).— Garfield Co.: 5'/2 mi N, 2
mi W Rifle, 5,900 ft, 6; Rio Blanco Co.; 31/2 mi W Rio
Blanco, 1; 1 mi N, 4 mi W Rio Blanco, 6,900 ft, 1.

October 1983

FiNLEY ET AL.: COLORADO BaTS

557

Six silver-haired bats (5 males and 1 fe-
male) were foimd at the sludge pit north of
Rifle. One each day was picked up in a fairly
good state of preservation on 29 April, 14
May, and 17 May, two on 4 June 1975, and
one still alive on 17 September 1974.

Only two Lasionycteris were taken in mist
nets, one each at two sites on Piceance Creek
west of Rio Blanco, where they were associ-
ated with M. lucifugns at one site and La-
siurus cinereus at the other. The sites are de-
scribed under those accounts.

Pipistrelltts hesperus hesperus (H. Allen)

Western Pipistrelle

Specimens examined (2).— Garfield Co.: 5V2 mi N, 2
mi W Rifle. 5,9(X) ft, 1; 4 mi W, 1 mi S Rifle, 5,300 ft, 1.

One P. hesperus was picked up on 17 Sep-
tember 1974 at the sludge pit north of Rifle.
The only pipistrelle netted was in the Colo-
rado River valley at the locality described
under M. lucifugus. These records and the
absence of specimens from our other sites at
higher elevations bear out Cary's (1911:209)
statement that pipistrelles "inhabit the Upper
Sonoran zone in the western and south-
western valleys . . . [and] hve only about cliffs
and in rock- walled canyons."

Eptesicus fuscus pallidus Young

Big Brown Bat

Specimens examined (4).— Rio Blanco Co.: Little
Duck Cr, TIS, R98W, Sec 10, 6,200 ft, 3; Little HilLs
Game Research Station, 15 mi W, 1.9 mi S Meeker, 1

(CU).

One big brown bat was shot flying over
tall sagebrush and two were netted over a
pool in a gully at Little Duck Creek on 7 Au-
gust 1975. The habitat and other captures are
described under the account of Myotis evotis.
Cary (1911:209) obtained a big-brown bat
from the White River, a few kilometers east
of Meeker in August 1905.

The four skins vary in color, as discussed
by Armstrong (1972:70), and fall within the
range of a series of nine pallidus in the
BS/FC collection from 5 mi E, 1 mi N Fort
Collins, except that one is more light reddish
than any in the Fort Collins series.

Lasiurus cinereus cinereus

Palisot de Beauvois

Hoary Bat

Specimens examined (16).— Garfield Co.: 3 mi N
Douglas Pass, 7,000 ft, 1; 3.4 mi N Rifle, on Government
Creek, 1; 5'/2 mi N, 2 mi W Rifle, 5,900 ft, 3; 2 mi E Rio
Blanco, 7,500 ft, 3; Rio Blanco Co.: Little Duck Cr, TIS,
R98W, Sec 10, 2; TIN, R102W, Sec 4, W of Rangely
[5,300 ft], 1; 3'/2 mi W Rio Blanco, 5.

The hoary bat was the most abundant spe-
cies in mist nets set over pools of Piceance
Creek adjacent to cottonwood (Populus) and
Douglas-fir stands between 2,000 and 2,300
m elevation. One was also netted over an
ephemeral pond a few miles west of Rangely
in salt desert shrub at 1,615 m elevation.
Three hoary bats were picked up at the
sludge pit north of Rifle. A female was found
OH 15 July 1974 and a male and a female on
17 September 1979. Two male hoary bats
were netted on Little Duck Creek on 7 Au-
gust 1975 at the site described under the ac-
count of Myotis evotis.

Five male hoary bats were caught in two
nets over Piceance Creek, SVz mi W Rio
Blanco, on the night of 10-11 July 1975. One
net was set over a narrow beaver pond in a
grove of cottonwoods and the other in a
grassy area just below the junction of Cow
Creek and Piceance Creek. The Piceance
valley was narrow between steep hillsides
wooded with small Douglas-fir and mixed
mountain shrubs on the shady side and mostly
pinyon-juniper on the sunny slope.

One female and two male hoary bats, as
well as one female M. volans, were netted on
10-11 July 1975 over a beaver pond on up-
per Piceance Creek, 2 mi E Rio Blanco,
7,300 ft. The site was more wooded than the
one just described (3V2 mi W Rio Blanco).
The north-facing slope above the beaver
pond was covered with Douglas-fir, and the
opposite slope had a heavy stand of pinyon-
juniper and mountain mahogany (Cerco-
carpus montanus). Other localities where
hoary bats were taken are described under
the accounts of Myotis leihii and Antrozous
pallidus.

A pregnant female (BSC/FC 5109, alco-
holic) was taken on 11 July 1975, 2 mi E Rio
Blanco. She weighed 30 grams and carried
two embryos, 22 mm. All 11 of the other

558

Great Basin Naturalist

Vol. 43, No. 4

hoary bats taken by mist nets were males.
The 11 skins available vary considerably in
appearance. The three taken the latest in the
summer (7 August) have longer dorsal fur
with a heavier overlay of silver tips. Some
taken in July have short pelage and some are
moulting.

Plecotus townsendii pallescens (Miller)
Townsend's Big-eared Bat

Speci7nens examined (5).— Rio Blanco Co.: Spring
Cave, 7,850 ft, 9 mi S, 4 mi E Biiford, 4; 5 mi N, 10 mi
W Rangely, 5,800 ft, 1 (KU).

On 19 February 1977 Finley visited Spring
Cave to investigate reports of two kinds of
bats there in winter, one with "long" ears
and one with "short" ears. This limestone
cave is located high on a forested mountain-
side in the White River National Forest. A
detailed description, photos, and maps of
Spring Cave were published by Parris
(1973:221). The mountainside was snow cov-
ered, but temperatures were mild. There
were a few scattered dormant bats on the
wall of the entry passage between the two
main entrances, some even in the twilight
zone. There were about 90 bats in all high on
the wall of the "Long Room" extending
southwest from the entry passage. About 25
were hanging singly and about 65 in 3 clus-
ters, mostly 2 to 3.7 m above the floor. All
appeared to be Plecotus, some with ears ex-
tended and some with ears folded, giving a
first impression of two kinds of bats present.
No bats were seen beyond the bend in the
Long Room or on the walls of the "Tunnel"
or "Pirates' Den." Wilhelm visited Spring
Cave on 15 August 1974 and explored it
without finding any bats or guano deposits.
Armstrong (1972) reported one specimen at
the University of Colorado Museum from
Spring Cave.

A single male big-eared bat was taken by
Finley on 29 August 1948 in a sandstone cave
5 mi N, 10 mi W Rangely, 5,800 ft., Rio
Blanco County. It was found in an overhead
vertical cleft through the roof of an arched
rock shelter that was also occupied by a
bushy-tailed wood rat (Neotoma cinerea
arizonae).

Antrozous pallidus pallidus (Le Conte)
Pallid Bat

Specimens examined (4).— Garfield Co.: .3.4 mi N
Rifle, on Government Cr, 3; Rio Blanco Co.: TIN,
R102W, Sec 4, W of Rangely, 1.

Three pallid bats, lactating females, were
taken by Caire on 25 July 1975 in a net set
under cottonwoods over a trickle of water in
Government Creek north of Rifle. The creek
was nearly dry and lined with large cotton-
woods and a few tamarisks {Tamarix gallica).
On 7 August 1975 another lactating female
was taken west of Rangely on the dry, low
sage-saltbush bench south of the White Riv-
er. The net was set across the lower end of a
shallow, mud-flat pool in a small arroyo. A
single hoary bat was the only bat taken in the
evening, and the pallid bat was found in the
net the following morning.

Gary (1911:206) reported seeing a pallid
bat "among the cliffs along the Grand [Colo-
rado] River, 7 miles west of Rifle, August 14,
1907."

Effects of Energy Development
ON Bats

Man's increasing activities for devel-
opment of energy resources in western Colo-
rado can entail several kinds of adverse ef-
fects on bat populations, such as disturbance
of hibernating bats in caves, mine tunnels,
and other sites; destruction of daytime roost
sites; loss or contamination of watering sites;
and contamination of the food chain by in-
secticides and pollutants.

Bats are extremely vulnerable because of
their specializations as volant, nocturnal in-
sectivores. For protection against ground
predators they require secure daytime roosts
such as cave walls, cliff crevices, or hollow
tree snags. Suitable shelters in the oil shale
region are usually located in narrow canyons
and ravines or on juniper-covered rimrock.
Such terrain features occupy a low percent-
age of total land area and should be avoided
as waste disposal sites for spent shale from
retorts.

Both natural and man-made water surfaces
are widely available to bats for drinking in
northwestern Colorado. The main permanent
streams such as the White River and Pi-
ceance Creek are important and likely to

October 1983

FiNLEY ET AL.: COLORADO BaTS

559

remain dependable drinking sites. But the
small intermittent streams and springs are
very susceptible to loss by lowering water ta-
bles when wells are drilled, and may become
contaminated by waste dumping or spillage.
Loss of springs and small stream pools on the
upland areas may be compensated for by
presence or addition of stock-watering tanks.
As long as cattle raising is maintained as a vi-
able industry in the areas of energy devel-
opment, the adverse effects on bats of spring
and creek flow loss can be at least partially
mitigated. However, production of aquatic
insect food would still be reduced.

Drilling for oil and gas in Colorado results
in a certain amount of drilling fluids and oil
at well sites being spilled and impounded in
pools or sludge pits. These are recognized
hazards to birds and other wildlife, but bats
have not previously been reported as casu-
alties. A source of bat mortality was called to
our attention by Richard E. Fillmore, who
picked up a mummified hoary bat carcass on
15 July 1974 on the bank of a sludge pit 5V2
mi N, 2 mi W Rifle at 1,800 m elevation. Pio-
neer Drilling, Inc., had drilled a test well and
dug two pits for impounding spilled oil. The
sludge pool was situated in the bottom of a
pit with steep banks rising about 3 m above
the surface, which was about 23 by 30 m in
size. Oil in the pool formed a thin slick over
the water, which was a few feet deep. This
lower pool was formed by drainage from an
adjacent higher pit that contained much
heavier oil sludge and was nearly dried up.
The oil well and pits were on a bulldozed
pad of alkaline clay soil on the east side of
the valley. The pad was surrounded by a
gentle slope covered with greasewood. To
the east was a steep rocky slope of pinyon-
juniper, leading up to high cliffs bounding
the east side of the valley of Government
Creek.

The lower of the two sludge pits proved to
be a lucrative source of bats. They were
found 3 to 8 m from the edge of the lower
pool, but none were found around the upper,
nearly dry pit. In the following 16 months re-
peated visits to this site by Finley and others
yielded 27 poor-quality but identifiable spec-
imens. The kinds and numbers of individuals
collected are as follows: hoary bat (3), long-

eared myotis (7), small-footed myotis (5), Cal-
ifornia myotis (5), western pipistrelle (1), sil-
ver-haired bat (6).

A live, oil-soaked Lasionycteris was seen
on 17 September 1974 hanging on the side of
a rock about 1 m from the oil-slick surface of
the pond. It responded to touch but did not
seem to be much affected by the oil.

In comparison with collections from other
localities, there were relatively high numbers
of silver-haired bats, only two of which have
been taken by mist net in the region, and of
California myotis, none of which has been
mist-netted; but the pallid bat, three of
which were netted only 5 km away, is absent
from the sludge pit sample. A more common
species not far away, Myotis volans, was also
absent from the sludge pit. Perhaps various
species of bats differ in their vulnerability to
entrapment.

On 19 September 1974 Finley visited the
Rangely oil field and inspected sludge pits
without finding any bats. The six pits that
contained oil were all much smaller than the
pit north of Rifle and had heavy oil, appar-
ently without water, in the bottom. They had
wires with colored streamers stretched over
them, apparently to keep out birds. It does
not seem likely that bats would mistake such
pits for water ponds. Four sludge pits con-
taining oil on water were inspected on 1 May
1981 between Craig, Colorado, and Rock
Springs, Wyoming, without finding any bats.

Three instances of entrapment of bats in
oil are known to us in the literature:
Krutzsch (1948) described finding 3 bats as
well as insects, lizards, and birds in three
small pools of oil formed by drainage into a
gully from a recently oiled road in the Bo-
rego Desert, California. Barbour and Davis
(1969) cited a report by E. J. Koestner of 15
red bats {Lasiurus borealis) entrapped in oil
on a 500-foot stretch of road in Illinois. Gil-
lette and Kimbrough (1970) reported a bat
seen in a "tar pit" of undescribed origin at
Fort Sill, Lawton, Oklahoma.

We thank Jerry Freeman for permission to
report a specimen of Myotis thysanodes
taken by him, Michael Bogan for identifying
several bats, and the Colorado Division of
Wildlife for generous information and
assistance.

560

Great Basin Naturalist

Vol. 43, No. 4

Literature Cited

Armstrong, D. M. 1972. Distribution of mammals in
Colorado. Mono. Mus. Nat. Hist., Univ. of Kansas
3:1-415.

Barbour, R. W., and W. H. Davis. 1969. Bats of Ameri-
ca. Univ. Press of Kentucky, Lexington. 286 pp.

Gary, M. 1911. A biological survey of Colorado. N.
Amer. Fauna .3.3:1-255.

FiNLEY, R. B., Jr., and J. Creasy. 1982. First specimen

of the spotted bat {Euderma macukitmti) from

Colorado. Great Basin Nat. 42:360.
Gillette, D. D., and J. D. Kimbrough. 1970. Chiropte-

ran mortality. In Slaughter and Walton, eds..

About bats. Southern Methodist Univ. Press,

Dallas.
Krutzsch, p. H. 1948. Water-like surfaces attract volant

desert animals. Ecology 29:391-392.
Parris, L. E. 1973. Caves of Colorado. Pruett Publ. Co.,

Boulder. 247 pp.

NEW GENERIC CONCEPTS IN THE TRITICEAE
OF THE INTERMOUNTAIN REGION: KEY AND COMMENTS

Mary E. Barkvvorth', Douglas R. Dewey% and Riley J. Atkins'

Abstract.— Revision of the perennial genera of North American Triticeae requires recognition of seven new gen-
era in the Intermonntain Region: Eltjtrigia, Letjmiis, Psatlnjwstachys, Pseudoroegneria, Thinopijrum, and the hybrid
genera X Ehjleijmus and X Pseudelymus. One previously recognized genus, Sitanion, is included in Ehjmiis. Several
new combinations are presented to accommodate the taxonomic changes. Ehjmits tmcln/caulus is treated as a wide-
spread, polymorphic species with three subspecies in the region: subsp. trachycauhts, subsecundus, and latigliimis.
Agropyron dd.systuchyum and A. albicans are treated as conspecific subspecies of Elymus lanceolatus. A key to the
genera of the Triticeae occurring in the Intermonntain Region is presented as well as keys and brief descriptions for
those genera not included in, or substantially modified from, other regional treatments.

Agrostologists have been aware for a long
time that traditional North American treat-
ments of the Triticeae (e.g., A. Hitchcock
1951, Gould 1968, C. Hitchcock 1969, Hol-
mgren and Holmgren 1977) do not reflect the
evolutionary relationships within the tribe.
Nevertheless, in the absence of any well-
documented revision that included a high
proportion of the North American species,
most North American taxonomists have
adopted A. Hitchcock's (1951) treatment,
with relatively minor modifications, as the
best available. Recently, however, Dewey
(1982, 1983a, 1983b) has published a revision
of the perennial genera that better reflects
the genomic and ecological data available
and is consistent with the morphological
data. Although written in terms of North
American taxa, Dewey's treatment is based
on data from the full geographic and tax-
onomic range of the tribe.

This paper is designed to assist those who
wish to use Dewey's generic concepts for
plants from the Intermonntain Region. It in-
cludes a key to the genera of the tribe in the
region and, for those genera not included in,
or substantially modified from Holmgren and
Holmgren (1977), brief generic descriptions
and keys to the infrageneric taxa that we rec-
ognize. Readers are referred to the Holm-
grens' article for illustrations, detailed de-
scriptions of the species, and the complete
synonymy. Table 1 summarizes the differ-

ences between the treatment presented here
and that found in their article.

Taxonomic Treatment

The genera that are most affected by the
revised generic boundaries are Agropyron
Gaertn., Elymus L., and Sitanion Raf. Agro-
pyron has been restricted to the crested
wheatgrasses, the remaining species being as-
signed to Elymus, Elytrigia, Pseudoroeg-
neria, or Thinopyrum. Several species of
Elymus have been placed in the segregate
genus Leymus Hochst., but all species of Sita-
nion are now included in Elymus. The rea-
sons for these and other changes are given in
the discussion of individual genera. To assist
those not familiar with the subtribal classifi-
cation of the Triticeae, the genera and spe-
cies within genera are treated alphabetically
after the generic key.

The intergeneric hybrids are treated after
the nonhybrid genera. Readers are advised
that such hybrids are relatively rare in na-
ture. We include them because they do exist
but, in our experience, most plants thought to
be hybrids are aberrant forms of good spe-
cies. Interspecific hybrids are more common,
particularly in disturbed areas. Part of the
problem in identifying hybrids in the Tri-
ticeae, particularly interspecific hybrids, is
that most can backcross to their parents.

'Department of Biology. UMC 45, Utah State University, Logan, Utah 84.322.

-Crops Research Laboratory, UMC 63, U.S. Department of .■\griculture. Logan, Utah 84322.

561

562

Great Basin Naturalist

Vol. 43, No. 4

Table 1. Synopsis of our revised treatment of the Trititeae occurring in the Interniountain Region compared with
that presented in the Intermountain Flora (Hohngren and Hohngren 1977).

Revised treatment

Intermountain Flora

Aegilops cylindrica L.

Agropyron cristatwn L.
Agropyron clesertortim J. A. Shultes
Agrpyron fragile Roth

X Elyhordeiiin macounii (Vasey) Barkworth & D. R.
Dewey

X Ehjleymus aristatits (Merrill) Barkworth & D. R.
Dewey

Elymus canadensis L.
Elymiis elymoides (Raf.) Sweezy
Elymus glaucus Buckley
Elymus X lumsenii Scribner, pro sp.
Elymus lanceolatus (Scribner & J. G. Smith) Gould
subsp. lanceolatus

subsp. albicans (Scribner & J. G. Smith) Barkworth
& D. R. Dewev

Elymus multisetiis (J. G. Smith) M. E. Jones
Elymus X pseudorepens (Scribner & J. G. Smith)

Barkworth & D. R. Dewey
Elymus X saundersii Vasey
Elymus trachycaulus (Link) Gould ex Shinners

subsp. trachycaulus

subsp. subsecundus (Link) Barkworth & D. R.

Dewey, pro parte

subsp. latiglumis (Scribner & J. G. Smith) Barkworth

& D. R. Dewey

subsp. subsecundus (Link) Barkworth & D. R.

Dewey, pro parte
Elymus virginicus var. submuticus Hooker

Elytrigia intermedia (Host) Nevski

subsp. intermedia

subsp. barbulata (Schur) A. Love
Elytrigia repens (L.) Nevski
Elytrigia spicata (Pur.sh) D. R. Dewey

Eremopyrum triticeum (Gaertner) Nevski

Hordeum brachyantherum Nevski

Hordeum depressum (Scribner & Smith) Rydb.

Hordeum marinum Hudson

subsp. gussonianuni (Pari.) Thell.
Hordeum murinum L.

subsp. murinum

subsp. glaucum (Steudel) Tsvelev

.subsp. leporinuni (Link) Arcang.
Hordeum pusillum Nutt.
Hordeum vulgare L.

Leymus cinereus (Scribner & Merrill) A. Love
Leymus flavescens (Scribner & Smith) Pilge
Leymus salinus (M. E. Jones) A. Love

subsp. salinus

subsp. stdnionis (C. L. Hitchc.) R. J. Atkins
Leymus simplex (Scribner & Williams) D. R. Dewey
Leymus triticoides (Buckley) Pilger

Pascopyrum smithii (Rydb.) A. Love

Aegilops cylindrica L.

Agropyron cristatum L.
included in A. cristatum L.
included in A. cristatum L.

X Agrohordeum macounii (Vasey) Lepage

X Elysitanion aristatum (Merril) Bowden

Elymus canadensis L.

Sitanion hystrix (Nutt.) J. G. Smith

Elymus glaucus Buckley

X Elysitanion hansenii (Scribner) Bowden

Agropyron dasystachyum (Hooker) Scribner

var. dasystachyum

Var. riparium (Scribner & J. G. Smith) Bowden
Agropyron albicans Scribner & J. G. Smith

var. albicans

var. griffithsii (Scribner & J. G. Smith) A. A. Beetle
Sitanion jubatum J. G. Smith
Agropyron X pseudorepens Scribner & J. G. Smith

X Agrositanion saundersii (Vasey) Bowden
Agropyron trachycaulum (Link) Malte

var. trachycaulum

var. glaucum (Pease & Moore) Malte

var. latiglumis (Scribner & J. G. Smith) A. A. Beetle
var. unilaterale (Cassidy) Malte

Elymus virginicus var. submuticus Hooker

Agropyron intermedium (Host) Beauv.

var. intermedium

var. trichophora (Link) Halac
Agropyron repens (L.) Beau--'.
Agropyron spicatum Pursh

Eremopyrum triticeum (Gaertner) Nevski

Hordeum brachyantherum Nevski

Hordeum depressum (Scribner & Smith) Rydb.

Hordeum geniculatum All.

Hordeum murinum L.
Hordeum glaucum Steudel
Hordeum leporinum Link
Hordeum pusdlum Nutt.
Hordeum vulgare L.

Elymus cinereus (Scribner & Merrill)
Elymus flavescens Scribner & Smith

Elymus salina M. E. Jones
Elyvtus ambiguus var. salmonis C. L. Hitchc.
Elymus simplex Scribner & Williams
Elymus triticoides Buckley

Agropyron smithii Rydb.

October 1983

Barkworth et al.: Intermountain Triticeae

563

Table 1 continued.

Revised treatment

Intermountain Flora

Psetidoroegneria spicata (Pursh) A. Love

X Psi'udehjnnis saxkolus (Scribner & J. C Smith)
Barkwortli & D. R. Dewey

Psdthtjrostaclnjs jtincea (Fischer) Nevski

Secale cereale L.

Taeniatheriim caput-medusae (L.) Nevski

Thinopyrum ponticum (Podp.) Barkworth
& D. R. Dewey

Triticwn aestivttm L.

Agwpyron spicatum Pursh

X Agrositanion saxicohi (Scribner & ]. G. Smith)
Bowden

Elymus juncetis Fischer

Secale cereale L.

Taeniatheriim capitt-mediisae (L.) Nevski

Agropyron elongatum (Host) Beauv.

Triticwn aestivitm L.

resulting in a morphological continuum as
well as partial restoration of fertility. This
problem is not unique to the Triticeae but it

is exacerbated by the relatively small number
of diagnostic characters available for these
grasses.

Key to the Genera

1. Spikelets three at a node, each with only one floret; only the central floret

fertile (exc. in Hordeum vulgare in which all three florets are fertile) Hordeum

— Spikelets not three at a node OR with more than one floret per spikelet (n.b
watch out for some forms of Elymus elymoides [ = Sitanion hystrix] in which
the second floret of the central spikelet is reduced and the lateral spikelets have
only a single, sterile, floret) 2

2(1). Annual or biennial, introduced cereals or weeds 3

— Perennial; native or introduced 7

3(2). Glumes ovate, with three or more (often many) nerves at midlength 4

— Glumes subulate to lanceolate, only one vein evident at midlength 5

4(3). Spikelets sunk in the rachis, the spike therefore very slender, less than 5 mm in

diameter; rachis disarticulating at maturity Aegilops

— Spikelets not sunk in the rachis, the spike therefore with a larger diameter;
rachis not disarticulating at maturity Triticum

5(4). Spikes less than 2.5 cm long; lemmas 5—7.5 mm long Eremopyrum

— Spikes more than 4 cm long; lemmas more than 8 mm long 6

6(5). Glumes more than 15 mm long, subulate, united at the base and tapering into a

long slender awn; spikelets with only one fertile floret; lemmas glabrous

Taeniatherum

— Glumes 6-15 mm long, narrowly lanceolate to linear, free to the base, gradu-
ally acuminate but not awned; spikelets with two fertile florets; lemmas
conspicuously scabrous on the keel Secale

7(2). Spikelets solitary at each node, closely imbricate, often pectinate; internodes

short, less than one third the length of the spikelets Agropyron

— Spikelets two or more at some nodes OR, if single at all nodes, neither closely
imbricate nor pectinate and with internodes about half as long as the spikelets

or longer 8

8(7). Glumes 3-10 mm long, very narrow, 1-nerved at midbody length, their keels
lying over the sides of the lowest lemmas rather than the midvein; lemmas
awnless or with awns up to 7 mm long 9

564 Great Basin Naturalist Vol. 43, No. 4

— Glumes 5-90 mm long, with 2-5 nerves evident at midbody length; keels of
the glumes lying opposite the midveins of the lowest lemmas; lemmas often
truncate or with awns more than 10 mm long 10

9(8). Rachis disarticulating at maturity; truly cespitose, branching intravaginal,

rhizomes never present; old leaf sheaths becoming fibrous Psathyrostachys

— Rachis not disarticulating at maturity; often rhizomatous, sometimes shortly so,
branching extravaginal; old leaf sheaths not fibrous Leymus

10(8). Plants fertile; anthers well-filled prior to anthesis, dehiscent, usually bleached

and falling off after anthesis 11

— Plants sterile; anthers poorly filled prior to anthesis, nondehiscent, retaining
their color and usually retained on the plant 13

11(10). Glumes 6-12 mm long; linear-lanceolate to lanceolate, tapering from below

midlength into an awn-tip; only one spikelet at most nodes Pascopynim

— Glumes varied but if 6-12 mm long either obtuse or tapering only in the distal
third; number of spikelets per node 1-4, varying between species 12

12(11). Glumes acute to long awned, never truncate or obtuse; anthers 2-3.5 mm long
and plants cespitose or anthers more than 3.5 mm long, plants rhizomatous,
and leaf blades with subequal ribs Elymus

— Glumes varied, often truncate or obtuse; anthers 4-7 mm long, plants rhizo-
matous or cespitose, if both long anthered and rhizomatous [E. repens], leaf
blades with 2-3 minor ribs alternating with the major ribs 13

13(12). Plants cespitose; glumes acute-tipped; spikelets only slightly longer than the

internodes Pseudoroegneria

— Plants rhizomatous or cespitose, if cespitose the glumes truncate to obtuse;
spikelets almost twice as long as the internodes 14

14(13). Plants cespitose Thinopyrum

— Plants rhizomatous Elytrigia

15(10). Lemmas with divergent awns more than 15 mm long; nodes with only one

spikelet X Pseudelymus

— Lemmas awnless or with nondiverging awns; nodes with one or two spikelets 15

16(15). Internodes less than 3 mm long; lowest lemmas usually less than 8.5 mm long ...

X Elyhordeiim

— Internodes more than 3 mm long; lowest lemmas usually 9 mm or longer 16

17(16). Glumes 12-24 mm long X Ehjleymus

— Glumes awnless or 25-85 mm long Elymus hybrids

A „.; T ploid wheats. His treatment has since been

adopted by Morris and Sears (1967) and

Bowden (1959) argued that Aegi/ops should ^°"J*^ ^J??^^', . , ^. ^. ^ .,

be included in Triticum, primarily because The difficulty with this approach is that, if

species of both genera have been involved in ^PP^^^^^ consistently, the tribe has to be re-

the evolution of many of such polyploid ^"^'^^ *« ^ ^^"g^^ g^""^ ^^f^"f '^' members

wheat species at T. durum Desf. and T. aesti- '^'^ connected by a network of introgressants

vum L. He noted also that the International ^nd hybrids. Krause (1898) advocated recog-

Code of Botanical Nomenclature requires i^ition of a single genus, but most taxonomists

that intergeneric hybrids must have a differ- have rejected his position because it ignores

ent generic name from their parents. By in- the differentiation, both morphological and

eluding Aegilops in Triticum he obviated the physiological, that has occurred within the

need for a new generic name for the poly- tribe. MacKey (1975) pointed out that the

October 1983

Barkworth et al.: Intermountain Triticeae

565

combination of annual growth habit and self-
fertihzation, such as occurs in Aegilops and
Triticum, "stimulates morphological and
physiological discontinuity in connection
with ecological specialization without the ne-
cessity for a simultaneous construction of
sterility barriers based on karyological differ-
entiation." Of the two genera in question,
Aegilops has remained a weedy genus with a
relatively narrow ecological amplitude and is
generally restricted to poor soils. Triticum, on
the other hand, has a much wider ecological
amplitude and greater ability to occupy fer-
tile land. This, combined with its tendency to
produce a larger grain, has led to rapid evo-
lution in response to selection pressures
exerted in part by human cultivation. Thus,
we prefer to treat the two as separate genera
both because of their morphological dis-
continuity and their different evolutionary
potentials.

The nomenclatural code requires that a
hybrid genus be given a different name from
any of its parents, but it does not state what
groups of species are to be treated as hybrid
genera. A group that has become sufficiently
well established that its origins are "ancient
history" can be treated as a "normal" genus
even if it is known to have originated
through hybridization. The species of Triti-
cum are such a group.

Agropyron Gaertner

This genus is now restricted to members of
the crested wheatgrass group. Its members
can be recognized by the very short inter-
nodes of the inflorescence and, in most in-
stances, the pectinate arrangement of the
spikelets. All our species are more or less ces-
pitose, although forms that produce short rhi-
zomes exist. Only one genome, the C gen-
ome, has been found in Agropyron s.str. Both
diploids and polyploids are known.

Agropijron s.str. includes about 10 species,
all of which are native to Eurasia. Consid-
erable controversy exists concerning the ap-
propriate taxonomic treatment for the plants
found in North America (cf, e.g., Hitchcock
1951, Sarkar 1956, Schulz-Schaeffer et al.
1963, Dewey 1969a, Taylor and McCoy
1973). The species exhibit considerable mor-
phological intergradation (cf. Tsvelev 1976),
and the problems of identification are exacer-
bated by their ability to hybridize when
brought into contact (Knowles 1955, Dewey
1969a), as has happened in North America.
The genus needs detailed biosystematic
study, based on wild populations, a project
beyond the scope of this paper. The treat-
ment presented here is based in part on Dew-
ey's examination of specimens in the Koma-
rov Institute (the National Herbarium of the
Soviet Union) and discussions with Tsvelev.

2(1).

Key to the Species of Agropyron

Spikelets diverging from the rachis at an angle of more than 40 degrees;
glumes widespread, forming an angle of more than 120 degrees, giving the

spike a bristly appearance; spikes at least 8 mm wide A. cristatum

Spikelets diverging from the rachis at an angle of less than 350 degrees; glumes
appressed; spikes 5-10 mm wide 2

Lemma with an awn 1-2(4) mm long; glumes forming an angle of

approximately 60 degrees A. desertorum

Lemma without an awn, sometimes mucronate; glumes forming an angle of

approximately 45 degrees (not common) A. fragile

(= A. sihiricum)

Elymus L.

Elymus is the largest genus in the Tri-
ticeae, but genomically it is very uniform. All
of its members are allopolyploids in which
two genomes are present, one derived from
Pseudoroegneria spicata or a relative thereof.

and the other from Hordeum. Almost all
plants examined, including all those from the
Intermountain Region, are tetraploids (2n =
28).

Despite their genomic similarity, species of
Elymus fall into two distinct morphological
groups. The largest group consists of self-

566

Great Basin Naturalist

Vol. 43, No. 4

fertilizing, cespitose species with small an-
thers; the other of rhizomatous, outcrossing
species, with long anthers. Dewey (1983a)
earlier included the latter group in Elytrigia
with other rhizomatous, long-anthered, but
genomically distinct species; but he now
(1983b) includes them in Elymiis, a treatment
that better reflects their phylogenetic affi-
nities. These rhizomatous species of Ehjmus
differ from those of Elytrigia in having
glumes that are acute or shortly awned,
rather than truncate or long-awned, and leaf
blades with no evident alteration of major
and minor ribs on the adaxial surface. As in-
terpreted here, there is only one such species
in the Intermountain Region, Elymus lan-
ceolatus [= Agropyron dasystachyum and
Agropyron albicans, cf. Table 1.]. The change
in epithet is necessary because the com-
bination Elymus dasystachys has been used
for a European species.

Elymus includes two other species that
used to be included in Agropyron (£. scribneri
and E. trachycaulus), because they have only
one spikelet per node. We maintain that the
morphological, reproductive, and genomic
similarity of these two species to others with
a similar genomic composition is more signif-
icant than the number of spikelets per node.

Elymus elymoides [ = Sitanion hystrix] and
E. multisetus [ = S. jubatum] have previously
been included in Sitanion (A. Hitchcock
1951, C. Hitchcock 1969, Holmgren and
Holmgren 1977), a genus characterized by a
readily disarticulating rachis and subulate,
long-awned glumes. Genomic studies have
shown, however, that the species included in
Sitanion are just as closely related to the SH
species previously included in Elymus or
Agropyron as these species are to each other
(Stebbins and Snyder 1956, Stebbins et al.
1946, Stebbins and Vaarama 1954, Brown
and Pratt 1960, Dewey 1967, 1969b, Church
1967a, b).

The disarticulating rachis, long subulate
glumes, and reduced sterile florets constitute
a set of adaptations for dispersal in open en-
vironments because the segments of the spike
are easily blown over the ground. Similar fea-
tures are found in one of the forms of the di-
morphic species Aegilops speltoides. The
other form consists of plants with a non-dis-
articulating rachis, short glumes, and more

fertile florets. Zohary and Imber (1963)
showed that the differences between the two
forms are determined by a group of closely
linked genes that are normally inherited as a
block. No studies have been conducted to de-
termine whether the same is true of the char-
acteristics used to delimit Sitanion, but Zo-
hary and Imber's study lends credence to our
conviction that Sitanion does not merit rec-
ognition at the generic level.

Three hybrid species are included in our
interpretation of Elymus, E. X hansenii, E.
X pseudorepens, and E. X saundersii. These
were previously referred to X Elysitanion,
Agropyron, and Agrositanion, respectively.
The change in their generic position results
from changes in the treatment of their paren-
tal taxa.

Our treatment of Elymus trachycaulus
differs somewhat from that endorsed by
Holmgren and Holmgren (1977). The tax-
onomy of the slender wheatgrass complex, of
which Elymus trachycaulus is a part, is ex-
tremely difficult to elucidate. Jozwik (1966)
recognized four groups in North America,
primarily on the basis of field, hybridization,
and herbarium studies. He suggested that
many of the members of two of his groups
may have been derived by hybridization, one
of them comprising plants derived from a va-
riety of different hybrid combinations. He
described the largest of the other two groups
(which corresponds to subsp. trachycaulus in
our treatment) as morphologically diverse,
probably as a result both of innate genetic
plasticity and introgression from other taxa.
It has a wide ecological amplitude, growing
along stream banks and in forests, meadows,
and moist prairies. Geographically it is ex-
tremely widespread, extending from Mexico
to Alaska and to both the west and east coasts
of North America.

Subspecies latiglumis corresponds to Joz-
wik's other, primarily nonhybrid, group. Its
members are more or less restricted to sub-
alpine, alpine, and far northern locations, but
at lower elevations they tend to intergrade
with subsp. trachycaulus, probably in part
because of hybridization.

Our third subspecies, subsp. suhsecundus,
corresponds to Jozwik's second group. This is
the group that he believed consisted almost
entirely of hybrids. His data indicated that

October 1983 Barkworth et al.: Intermountain Triticeae 567

the second parent could be one of several available, it would be impossible to design a

taxa, e.g. E. elymoides, E. multisetus, E. completely satisfying treatment for such a

glaucus, and H. jubatum. Intermediates be- group because the formal requirements of the

tvt'een the members of this subspecies and nomenclatural code cannot perfectly reflect

subsp. trachycaulus were numerous. He also the dynamic interactions occurring in a

found intermediates with subsp. latiglumis, group such as the slender wheatgrass

but these were much less frequent. complex.

We admit that our treatment of this com- Our treatment of E. elymoides also differs

plex is not altogether satisfying, but it seems from that in Holmgren and Holmgren (1977)

the most appropriate treatment considering in that we are not recognizing any in-

the data available. Even if more data were fraspecific taxa.

Key to Species and Hybrids of Elymus

1. Spikelets 2-7 at a node 2

— Spikelets solitary at each node 7

2(1). Glumes subulate, 1-2-nerved at midlength and with awns more than 20 mm

long; rachis disarticulating at maturity 3

— Glumes lanceolate, 2-5-nerved at midlength, if 2-nerved the awns less than 5

mm long; rachis not disarticulating at maturity 5

3(2). Awns not diverging, even at maturity E. X hansenii

— Awns widely divergent at maturity 4

4(3). Glumes longitudinally divided into 3 or more narrow sections E. multisetus

— Glumes entire or bifid E. elym,oides

5(2). Rachis flexible, spike nodding; glumes with an awn 10-30 mm long E. canadensis

— Rachis stiff, spike erect; glumes unawned or short awned 6

6(5). Glumes bowed outward and indurate at the base, the nerves not evident in the

indurate portion E. virginicus var. submuticus

— Glumes not bowed out, membranous at the base, nerves evident throughout

E. glaucus

7(1). Plants rhizomatous; anthers 3-5 mm long 8

— Plants cespitose; anthers 1-3 mm long 10

8(7). Plants sterile, anthers not well filled at anthesis, not dehiscent E. X pseudorepens

— Plants fertile, anthers well filled at anthesis, dehiscent (£. lanceolatus) 9

9(8). Lemmas awnless or with an awn-tip less than 5 mm long

E. lanceolatus subsp. lanceolatus

— Lemmas with a divergent awn 5-12 mm long E. lanceolatus subsp. albicans

10(7). Lemmas awned, the awns widely divergent; culms decumbent, usually less

than 35 cm tall E. scribneri

— Lemmas unawned or, if awned, then erect or only slightly divergent; culms
erect, usually more than 50 cm tall 11

11(10). Glumes l-2(3)-nerved; rachis tending to disarticulate at maturity; plants

sterile E. X saundersii

— Glumes (3) 5-nerved; rachis not disarticulating at maturity; plants fertile

(£. trachycaulus) 12

12(11). Lemma awns 8-24 mm long, erect to divergent . E. trachycaulus subsp. subsecundus

— Lemmas awnless or with short, erect, awns less than 5 mm long 13

568

Great Basin Naturalist

Vol. 43, No. 4

13(12). Culms erect, 30-130 cm tall; glumes with a narrow hyaline margin

E. trachycaulus subsp. trachycaulus

— Culms often geniculate or decumbent, less than 55 cm tall; glumes with a

broad hyaline margin E. trachycaulus subsp. latiglumis

Elymus lanceolatus subsp. albicans (Scribner
& J. G. Smith) Barkworth & D. R.
Dewey, cotnb. nov.— Basionym: Agro-
pyron albicans Scribner & J. G. Smith,
USDA Div. Agrost. Bull. 4:32, 1897.

Elymus X pseudorepens (Scribner & J. G.
Smith) Barkworth & D. R. Dewey,
comb, nov.— Basionym: Agropyron
pseudorepens Scribner & J. G. Smith,
USDA Div. Agrost. Bull. 4:34, 1897,
pro sp.

Elytrigia Desv.

All species of Elytrigia are outcrossing, but
in their other characteristics, including their
genomic composition, they are very diverse.
It is undoubtedly the least satisfactory genus
as presently constituted and the one that
most needs further study. There are only two
species in the Intermountain Region, both of
which are introduced.

2(1).

Key to the Species of Elytrigia

Glumes acute to awn tipped, membranous; rachis only slightly concave

adjacent to spikelet E. repens

Glumes truncate or mucronate, thick; rachis markedly concave adjacent to the
spikelet (£. intermedia) 2

Lemmas glabrous; spikelets 3-8-flowered E. intermedia subsp. intermedia

Lemmas hirsute; spikelets 2-3(-6)-flowered E. intermedia subsp. barbulata

Hordeum L.

The limits of this genus have not been
changed but we have adopted the in-
frageneric treatment recommended by von
Bothmer (pers. comm.), since he has studied
the genus in both North and South America
as well as Europe. This seems particularly
appropriate since the taxa for which von
Bothmer's treatment differs from that in
Holmgren and Holmgren (1977) are all in-
troduced Mediterranean weeds. Moreover,
although Holmgren and Holmgren treated
the subspecies of H. murinum at the specific
level, they noted that the taxa were very
closely related and often difficult to dis-
tinguish. Thus, the differences between the
two treatments are not as great as it may ap-
pear. No key is presented since the Holm-
gren's key can be used, the only changes
needed being nomenclatural. These are in-
dicated in Table 1.

Leymus Hochst.

In our region, the species of this genus can
be recognized by their short, subulate glumes

that lie over the sides rather than the mid-
veins of the lemmas, and by the absence of
long awns. The genus includes both rhizo-
matous and cespitose, but extravaginally
branching, species.

Species of Leymus, both here and else-
where, tend to grow in alkaline or saline
soils. Some are coastal in distribution; others
are inland species. The two groups are mor-
phologically distinct. Our species, not sur-
prisingly, belong to the inland group.

Despite the morphological discontinuity
between its coastal and inland members, spe-
cies of Leymus are genomically similar. They
are all allopolyploids based on the J genome,
from Psathyrostachys, and the X genome
whose origin is unknown. Tetraploids (2n =
28), hexaploids (2n = 42), and octoploids (2n
= 56) are known. Thus species of Leymus
differ from species of Elymus both in their
genomic composition and their tendency to
form higher polyploids.

In traditional treatments of the tribe,
Leymus is included in Elymus since most of
its members have more than one spikelet at a

October 1983

Barkworth et al.: Intermountain Triticeae

569

node. As indicated above, however, the two
genera differ from each other in a number of
other morphological characteristics. More-
over, L. salinits and L. simplex usually have

only one spikelet at most, if not all, nodes.
The treatment of L. salinus presented here is
based on work by Atkins (1983; Atkins et al.,
in press).

Key to Species of Leymiis

1. Plants strongly rhizomatous, the rhizomes long and slender 2

— Plants cespitose, sometimes with short rhizomes 5

2(1). Lemmas conspicuously hirsute, the hairs 1-2 mm long, not closely appressed to

the lemma L. flavescens

— Lemmas glabrous to, at most, inconspicuously hirsute with hairs less than

1 mm long 3

3(2). Leaf blades with more than 7 veins, not densely hirsute above the ligule; most

nodes with two or more spikelets L. triticoides

— Leaf blades with 5-7 prominent veins, densely short-hirsute above the ligule;
most nodes with only one spikelet f. L. simplex

5(1). Leaf blades 4-15 mm wide, flat, many nerved; ligules 2-5 mm long; culms more

than 1 m tall L. cinereus

— Leaf blades 2-4 mm wide when flat, strongly involute, 5-7-nerved; ligules less
than 2 mm long; culms less than 1 m tall (L. salinus) 6

6(5). Basal leaf sheaths glabrous; most nodes with only one spikelet

L. salinus subsp. salinus

— Basal leaf sheaths conspicuously hirsute; most nodes with two or more
spikelets L. salinus subsp. salmonis

Leym.us salinus subsp. salmonis (C. Hitchc.)
Atkins, comb, nov.— Basionym: Elymiis
amhiguus var. salmonis C. Hitchc.
Univ. Wash. Publ. Biol. 17(1):558, 1969.
Holotype: WTU!

Pascopyrum A. Love

Pascopyrum is a monotypic genus com-
prising only P. smithii. This species is an oc-
toploid, its probable parents being Elymus
lanceokitus and Leymus triticoides (Dewey
1975). Morphologically it is intermediate be-
tween its parents. This is particularly evident
in the glumes, which are membranous and
flat at the base, as it typical for Elymus, but
then taper gradually into an acuminate tip
resembling the linear lanceolate glumes char-
acteristic of Leymus. Holmgren and Holm-
gren (1977) recognized two varieties within
the species, but we do not consider either
merits formal recognition.

Psathyrostachys Nevski

This genus is comprised of eight species
that are native to the steppes and arid re-
gions of southeastern Europe. They are all
strictly cespitose and have disarticulating
rachises and two spikelets at a node. All the
species studied so far are diploids based on
the J genome. Psathrostachys juncea (Russian
wild rye) is the only species to have become
established in North America.

Pseudoreogneria A. Love

All species of Pseudoroegneria are based on
a single genome, the S genome. The genus
consists of several Eurasian species but only
one North American species, P. spicata. Most
of its members can be recognized by their
rather slender habit and the single spikelets
that are only slightly longer than the
internodes.

570

Great Basin Naturalist

Vol. 43, No. 4

In previous discussions of the tribe, Dewey
(1982, 1983a, b) has included these species in
Elytrigia in conformity with Tsvelev's (1976)
treatment. It is clear, however, that they are
genomically distinct and consequently, in
keeping with the philosophy guiding this re-
vised treatment, should be recognized at the
generic level. No new combinations are nec-
essary for the Intermountain Region.

Thinopyrum A. Love

This is a Eurasian genus but one of its
members, T. ponticum, has been introduced
into North America and occurs along high-
ways in the Intermountain Region. We have
followed Holub (1973) and Melderis (1980) in
adopting the epithet pontica for the plants
that Holmgren and Holmgren (1977) referred
to Agropyron elongatum. The epithet ehn-
gata is now interpreted as referring to a west-
em Mediterranean species of relatively small,
slender plants, all of which are diploids.
Thinopyrum ponticum (Tall Wheatgrass) con-
sists of robust decaploid plants that are wide-
spread in Eurasia. It has been seeded at scat-
tered locations in the Intermountain Region.
The new combination is presented here:

Thinopyrum, ponticum (Podp.) Barkworth &
D. R. Dewey, comb, nov.— Basionym:
Triticum elongatum (Host), Gram.
Austr. 2:18, 1802).

X Elyhordeum Mansf. ex Zizin & Petr.

One hybrid between Elymus and Hordeum
is established in the Intermoiantain Region,
X E. macounii. Its parents are Elymus
trachycaulus and Hordeum jubatum (Boyle
and Holmgren 1955). In previous treatments
it was included in X Agrohordeum macounii.
The transfer to X Elyhordeum is made neces-
sary by the transfer of Agropyron trachycau-
lum to Elymus.

X Elyhordeum macounii (Vasey) Barkworth
& D. R. Dewey, comb, nov.— Elymus
trachycaulus (Link) Gould ex Shinners
X Hordeum jubatum L.— Basionym:
Elymus macounii Vasey, Grasses U.S.
46, 1883. Macoun, Great Plains of B.C.

X Elyleymus Baum

One hybrid between Elymus and Leymus
occurs in the Intermountain Region, X Ely-
leymus aristatus. Dewey and Holmgren
(1962) have shown that its parents are prob-
ably Elymus elymoides and L. triticoides.
Holmgren and Holmgren (1977) referred it to
X Elysitanion aristatum.

X Elyleymus aristatus (Merrill) Barkworth
& D. R. Dewey, comb, nov.— Elymus
elymoides (Raf.) Barkworth & Dewey
X Leymus triticoides (Buckley) Pil-
ger.— Basionym: Elymus aristatus Mer-
rill, Rhodora 4:147, 1902. Cusick 2712,
"in large clumps, Silver Creek, Harney
Co., Oregon."

X Pseudelymus Barkworth &
D. R. Dewey, gen. hybr. nov.

X Pseudelymus Barkworth & D. R. Dewey,

gen. hybr. nov.— Pseudoroegneria A.
Love X Elymus L.
One hybrid between Pseudoroegneria
and Elymus has become established in west-
ern North America, X P. saxicola. The ge-
neric name X Pseudelymus is presented here
to accommodate it and other such hybrids
that may occur elsewhere.

The parents of X P. saxicola are Pseudo-
roegneria spicata and Elymus elymoides
(Dewey 1964). The plants are usually com-
pletely sterile but, being perennial, once they
are established at a location, they will persist
there. The change in generic name is made
necessary by changes in generic boundaries
affecting its parents.

X Pseudelymus saxicola (Scribner & J. G.
Smith) Barkworth & D. R. Dewey,
comb, nov.— Pseudoroegneria spicata
(Pursh) A. Love X Elymus elymoides
(Raf.) Sweezy.— Basionym: Elymus
saxicolus Scribner & J. G. Smith, USDA
Div. Agrostol. Bull. 18:20, 1899, pro.
sp.

Discussion

Selection of the most appropriate tax-
onomic treatment of a polyploid complex is
always difficult. This is particularly true

October 1983

Barkworth et al.: Intermountain Triticeae

571

when the members of the complex hybridize
as readily as do members of the Triticeae.
The problem is compounded by the great
morphological reduction that characterizes
all grasses and the prevalence of convergent
evolution. The treatment presented here
seeks to reflect as completely as possible all
available data. It is therefore a compromise
between a strictly genomic interpretation
and one based entirely on morphological
data.

The main advantage of this treatment is
that it reflects a higher proportion of the
genomic and morphological information
available than does the traditional treatment.
It is also in closer accord with the systems
adopted by Tsvelev (1976) and Tutin et al.
(1980). Since both the Soviet Union and Eu-
rope have more species of Triticeae than the
United States and Canada combined, it is ap-
propriate to consider seriously the treatments
advocated by taxonomists in those regions.

Some of the new genera are not, perhaps,
as easy to recognize as the old interpretation
of Agropyron and Elymus. On the other hand,
numerous herbarium specimens indicate that
Leymus salinus and Leymus simplex, which
have only one spikelet at most of their nodes,
have often been misidentified as species of
Agropyron rather than Elymus. Thus, even
the traditional treatment was sometimes diffi-
cult to apply. The revised genera can be rec-
ognized on the basis of their gross morpho-
logical characters, although not the same
characters as before. We hope that this treat-
ment will assist those wishing to become fa-
miliar with the new generic concepts.

Acknowledgments

We thank Drs. A. Love and J. McNeill for
their careful review and criticism of the in-
itial draft of this manuscript and for the nu-
merous discussions concerning the taxonomy
of the Triticeae in general.

Literature Cited

Atkins, R. J. 1983. A taxonomic study of Leymus arn-
higuus and L. salinus (Poaceae). Unpublished
thesis. Department of Biology, Utah State Univ.

Atkins, R. J., M. E. Barkworth, and D. R. Dewey. In
press. A taxonomic study of Leymus ambiguus
and L. salinus (Poaceae). Syst. Bot.

BowDEN, W. M. 1959. The taxonomy and nomenclature
of the wheats, barleys, and ryes and their wild
relatives. Canadian J. Bot. 37:657-684.

Boyle, W. S., and A. H. Holmgren. 1955. A cytogenet-
ic study of natural and controlled hybrids be-
tween Agropyron trachycaiilum and Hordeum
jubatum. Genetics 40:539-545.

Brown, W. V., and G. A. Pratt. 1960. Hybridization
and introgression in the grass genus Elyjnus.
Amer. J. Bot. 47:669-676.

Church, G. L. 1967a. Taxonomic and genetic relation-
ships of eastern North American species of
Elymus with setaceous glumes. Rhodora 69:
121-162.

1967b. Artificial hybrids of Elymus virginicus

with E. canadensis, interruptus, riparius, and
weigandii. Amer. J. Bot. 43:410-417.

Dewey, D. R. 1964. Natural and synthetic hybrids of
Agropyron spicatum X Sitanion hystrix. Bull.
Torrey Bot. Club 91:396-405.

1967. Synthetic hybrids of Elymus canadensis X

Sitanion hystrix. Bot. Gaz. 128:11-16.

1969a. Hybrids between tetraploid and hexaploid

crested wheatgrasses. Crop Sci. 9:787-791.

1969b. Synthetic hybrids of Agropyron albicans

X A. dasystachyum, Sitanion hystrix, and
Elijmiis canadensis. Amer. J. Bot. 56:664-670.

1975. The origin of Agropyron smithii. Amer. J.

Bot. 62:524-530.

1982. Genomic and phylogenetic relationships

among North American perennial Triticeae.
Pages 51-88 in J. R. Estes et al., eds.. Grasses and
grasslands, Univ. of Oklahoma Press, Norman.

1983a. New nomenclatural combinations in the

North American perennial Triticeae (Gramineae).
Brittonia 35:30-33.

1983b. Historical and current taxonomical per-
spectives of Agropyron, Elymus, and related gen-
era. Crop Sci. 23:637-642.

Dewey, D. R., and A. H. Holmgren. 1962. Natural
hybrids of Elymus cinereus and Sitanion hystrix.
Bull. Torrey Bot. Club 89:217-228.

Gould, F. W. 1968. Grass Systematics. New York:
McGraw-Hill Book Co.

Hitchcock, A. S. 1951. Manual of the grasses of the
United States, rev. A. Chase. USDA Misc. Publ.
200.

Hitchcock, C. L. 1969. Gramineae. Pages 383-725 in C.
L. Hitchcock, A. Cronquist, and M. Ownbey,
eds.. Vascular plants of the Pacific Northwest Pt.
1. Univ. of Washington Press, Seattle.

Holmgren, A. H., and N. H. Holmgren. 1977. Poaceae.
Pages 175-465 in A. Cronquist et al., eds., Inter-
mountain flora. Columbia Univ. Press, New York.

HoLUB, J. 1973. New names in phanerogamae 2. Folia
Geobot. Phytotax. 8:155-179.

JozwiK, F. X. 1966. A biosystematic study of the slender
wheatgrass complex. Unpublished dissertation.
Univ. of Wyoming.

Knowles, R. p. 1955. A study of variability in crested
wheatgrass. Canadian J. Bot. 33:534-546.

Krause, E. H. L. 1898. Florische Notizen II. Graser.
Bot. Centrabl. 73:332-343.

572

Great Basin Naturalist

Vol. 43, No. 4

Mackey, J. 1975. The boundaries and subdivisions of the
genus Triticum. Paper read at Twelfth Int. Bot.
Congr., Leningrad.

Mackey, J., C. J. Humphries, and T. G. Tutin. 1980.
Triticeae. Pages 185-206 in Flora Europaea Vol.
5. University Press, Cambridge.

Morris, R., and E. R. Sears. 1967. The cytogenetics of
wheat and its relatives. Pages 19-87 in K. S. Qui-
senberry and L. P. Reitz, eds.. Wheat and wheat
improvement, Amer. Soc. Agron. Inc. Madison,
Agron. Ser. 13.

Sarkar, R. 1956. Crested wheatgrass complex. Canadian
J. Bot. 34:328-345.

Schulz-Schaeffer, J., p. W. Allerdice, and G. C.
Creel. 1963. Segmental allopolyploidy in tetra-
ploid and hexaploid Agropyron species of the
crested wheatgrass complex (Section Agropyroii).
Crop Sci. 3:525-530.

Stebbins, G. L., and L. A. Snyder. 1956. Artificial and
natural hybrids in the Gramineae, tribe Hordeae.
V. Amer. J. Bot. 43:305-312.

Stebbins, G. L., and A. Vaarama. 1954. Artificial and
natural hybrids in the Gramineae, tribe Hordeae.
VII. Hybrids and allopollyploids between Ehjmtis
glancus and Sitanion spp. Genetics 39:378-395.

Stebbins, G. L., J. 1. Valencia, and R. M. Valencia.
1946. Artificial and natural hybrids in the Gra-
mineae, tribe Hordeae I. Elymiis, Sitanion, and
Agropyron. Amer. J. Bot. 33:338-351.

Taylor, R. J., and G. A. McCoy. 1973. Proposed origin
of tetraploid species of crested wheatgrass based
on chromatographic and karyotypic analyses.
Amer. J. Bot. 60:576-583.

TsvELEV, N. N. 1976. Zlaki SSSR. Nauka, Leningrad.

Tutin, T. G., V. H. Heywood, D. H. Valentine, S. M.
Walters, and D. H. Webb. 1980. Flora Eu-
ropaea. Vol. 5. Monocotyledones. Cambridge
Univ. Press, Cambridge, U.K.

ZoHARY, D., AND D. Imber. 1963. Genetic dimorphism
in fruit types of Aegilops speltoides. Heredity 18:
223-231.

REPRODUCTIVE ATTRIBUTES OF SOME ROCKY MOUNTAIN SUBALPINE HERBS

IN SUCCESSIONAL CONTEXT

David J. Schiinpfi'2 and Robert L. Bayn, ]i}-^

Abstract.— Selected reproductive attributes of herbaceous plant species were compared among three stages of a
sere in the subalpine zone of Utah's Wasatch Mountains: herbaceous meadow, aspen grove, spruce-fir forest. No suc-
cessional trends in seed size or inferred mode of seed dispersal were detected. We ascribe the deviation of these find-
ings from those of most other studies to differences in climate, life-form composition, or community age between our
sere and those of other studies. A variety of flower colors were found in the meadow stage, grading into a pre-
dominance of white flowers under conifers. Animal vectors of pollen, capable of effecting plant outcrossing, were
most abundant in the meadows and an order of magnitude less abundant under aspen.

Attempts to develop inductive general-
izations about ecological succession have in-
cluded studies of reproductive characteristics
of plants. Among the earliest were those of
Salisbury (1942), who found British woodland
species to have heavier seeds than those of
species in open vegetation. This was largely
due to woody species having heavier seeds
than herbs, but forest herbs were also heavier
seeded than meadow herbs. Salisbury sug-
gested that low light intensity on forest floors
favored individuals with stored materials in
the seed sufficient to fabricate a greater
light-intercepting surface. These results have
been generally corroborated for grassland
(Hayashi 1976, Werner and Piatt 1976) as
well as forest (Opler et al. 1977, Abrahamson
1979) seres. The statistical significance of
these increases in seed size during succession
has not been established. Marino (1980)
found no significant difference in seed size of
foredune herbs and shrubs from those under
dune forest, but those of the intermediate
slack dune stage were significantly smaller.

The mode of seed dispersal, as inferred
from diaspore morphology, has been sug-
gested to change toward a greater frequency
of animal dispersal as forest succession pro-
ceeds (Dansereau 1957, Dansereau and Lems
1957, Harper et al. 1970, Johnston and Odum
1956, Pijl 1972, Opler et al. 1977). Hayashi

(1976) did not detect this trend in a grassland
seje. Several of these authors, as well as
others, suggest that animal dispersal adapta-
tions may be more common among woody
species than among herbaceous species. If so,
the successional changes in dispersal mode
may be due to changes in the proportions of
various life forms. Statistical tests of these
trends are lacking.

Most of the above studies were conducted
in temperate deciduous forest or tropical
moist forest biomes. Our study examined
some plant reproductive attributes in stands
along a sere terminating in subalpine co-
niferous evergreen forest. Seed sizes, dis-
persal modes, flower color frequencies, and
abundance of animal pollen vectors most
likely to effect floral outcrossing were com-
pared among the stages of this sere.

Study Area and Methods

Field work was conducted in the subalpine
zone of the Wasatch Mountains in extreme
northern Utah. The presumed sere involves
vegetative colonization of well-drained, herb-
dominated meadows by clones of quaking as-
pen, Populus tremuloides Michx. The climax
forest is dominated by subalpine fir, Abies
lasiocarpa (Hook.) Nutt., and Engelmann
spruce, Picea engelmannii Parry. No other

'Department of Biology and Ecology Center, Utah State University, Logan, Utah 84322.
'Present address: Department of Biology, University of Minnesota, Duluth, Minnesota 55812.
'Present address: Bio-Resources, Inc., P.O. Box 3447, Logan, Utah 84321.

573

574

Great Basin Naturalist

Vol. 43, No. 4

woody species are significant community
components. For further discussion of com-
munity dynamics and a site description, see
Schimpfetal. (1980).

For each stage of succession, we selected
abimdant herbaceous species producing sub-
stantial numbers of seed, thus excluding vege-
tative remnants of earlier stages. Species
were included if they produced at least 100
seeds from a total of 10 or more mother
plants within a 0.2-ha rectangular plot during
the summers of 1976, 1977, or 1978. There
was one such plot for each stage in the ex-
ample of the sere known as Big Meadow
(Schimpf et al. 1980): meadow, aspen, fir,
spruce-fir. The species inclusion criteria
proved unworkable for the fir and spruce-fir
plots, owing to lower population densities
and seed production per plant; these plots
were treated as a single 0.4-ha conifer plot to
determine species inclusion and for all data
analyses. Species which qualified for in-
clusion were all counted equally in the analy-
ses of reproductive attributes, rather than
being weighted by abundance.

Total weight was determined for a sample
of 100 seeds (or caryopses or achenes) for
each species in a stage. Samples had been
heated overnight at 100 C after collection to
stop respiration. All seeds and caryopses were
inspected microscopically, and all achenes
sectioned transversely to confirm seed devel-
opment before being weighed to the nearest
0.1 mg, in equilibrium with atmospheric hu-
midity. Size was expressed as mean mg per
seed. Baker (1972) reported that mean seed
sizes of a group of plant species are often not
normally distributed. The distribution of
mean sizes in each stage was tested for nor-
mality with the Kolmogorov-Smirnov meth-
od (Sokal and Rohlf 1969), using both un-
transformed and log-transformed variates.
Transformed mean seed sizes were compared
among stages by analysis of variance. Pairs of
stages were compared for their intrinsic vari-
ation in mean sizes by the F-ratio of varian-
ces (log-transformed data) (Lewontin 1966).

Flower color was recorded in the field for
each species. The presumed mode of dis-
persal was classified as animal, wind, or other
from inspection of diaspore structure. The
frequency distributions of flower color and
dispersal mode categories were compared

among stages by chi-square tests. Chi-square
was also employed to test for differences in
frequency of animal dispersal modes among
different plant life forms.

Insects suspected to be capable of effecting
floral outcrossing were sorted from the gen-
eral collections of insects associated with the
herbaceous layer. Suspected pollinators in-
cluded all adults in the following families:
Syrphidae, Bombyliidae, Colletidae, Halic-
tidae, Andrenidae, Megachilidae, Apidae,
Sphingidae, Nymphalidae, Lycaenidae. These
collections were obtained by D-Vac sampling
of from 130 to 310 randomly chosen O.SO-m^
or 0.25-m2 plots that had been quickly cov-
ered with an insect-tight cage (Southwood
1978). Samples were taken at regular inter-
vals throughout the 1977 and 1978 growing
seasons in nearby examples of the succession-
al stages similar to those at Big Meadow. Re-
sults were expressed as m^ sampled per pol-
linator caught.

Results

The 46 herbaceous taxa studied included
26 species in the meadow plot, 22 taxa in the
aspen plot (including two varieties of one
species), and 23 species in the conifer plot
(Table 1). Mean seed sizes fail to exhibit a
normal distribution within a stage, based on
the Kolmogorov-Smirnov test. Following log
transformation, all three stages show a nor-
mal distribution. The mean and standard de-
viation of transformed variates is depicted for
each stage in Figure 1. None of the F values
are significant at the .05 level; the means and
variances of seed sizes in the three stages are
statistically indistinguishable. No successional
trends in community seed size are apparent.
There are also no discernible patterns of size
change within the set of those species that
occupy two or more stages (Table 1), based
on sign tests (Sokal and Rohlf 1969). White-
flowered species become increasingly fre-
quent and the red- and blue-flowered species
less frequent through successional time, al-
though this trend was not significant at the
0.05 level (Table 2). Modes of dispersal have
similar frequencies in all three successional
stages (Table 3), suggesting that there is no
trend in this attribute.

October 1983

ScHiMPF, Bayn: Subalpine Herbs

575

Pollen vector abundance was notably
lower during 1977 than during 1978 (Table
4). In both summers, abundance is an order of
magnitude greater in the meadows than in

the aspen understory, with intermediate val-
ues in conifer understory. The most abundant
pollinator families in the meadows were Ha-
lictidae, Apidae, and Syrphidae, each about

Table 1. Reproductive attributes of the herbaceous taxa.

Taxon

Mean

seed size.

Flower
color

Dispersal

Meadow

Aspen

Conifer

mode

0.166
3.017

0.157
2.513
3.359

0.197
3.324

White
Orange
Green

Other
Wind
Animal

Adnlleu millefolium L. ssp. lanulosa (Nutt.) Piper
Agoseris aurantiaca (Hook.) Greene var. aurantiaca
Agropijron tmdiijcaulum (Link) Malte var. glaiicwn

(Pease & Moore) Malte
Agropijwn tmchijcaulum (Link) Malte var. latiglume

(Scribn. & Smith) A. A. Beetle
Androsace filifomiis Retz.

Aquilegia coerulea James var. ochroleuca Hook.
Arabis drwnmondii Gray
Arnica cordifolia Hook. var. cordi folia
Arnica parnji Gray
Aster engelnumnii (Eat.) Gray
Aster integrifolius Nutt.
Bromiis carinatus Hook. & ,\rn.
Castilleja miniata Dougl. var. miniata
Cknjtonia lanceolata Pursh var. lanceolata
Collomia linearis Nutt.

Delphinium nuttallianum Pritz. var. nuttallianum
Descurainia richardsonii (Sweet) Schulz var. sonnei

(Robins.) C.L. Hitchc.
Draba stenoloba Ledeb. var. nana (Schulz) C.L. Hitchc.
Epilobium brachi/carpnm Presl
Epilobitim lactiflorwn Haussln.
Erigeron speciosus (Lindl.) D.C. var. macranthus (Nutt.)

Cronq.
Eriogoniim heracleoides Nutt.

Erysimum asperum (Nutt.) D.C. var. purshii Durand
Erijthronium grandiflorum Pursh
Galium bifolium Wats.
Geranium viscosissimum Fisch. & C.A. Meyer var.

nervosum C.L. Hitchc.
Gi7ifl aggregata (Pursh) Spreng.
Hackelia micrantha (Eastw.) J.L. Gentry
Hieracium albiflorum Hook.
Hieracium scouleri Hook.

Hijdrophtjllum capitatum Dougl. var. capitatum
Ligusticum filicinum Wats.
Lupinus argenteiis Pursh var. rubricaulis (Greene)

Welsh
Madia glomerata Hook.
Osmorhiza chilensis Hook. & Arn.
Osmorhiza occidentalis (Nutt.) Torr.
Pedicidaris racemosa Dougl. var. alba (Pennell) Cronq.
Poa nervosa (Hook.) Vasev var. wheeleri (Vasey) C.L.

Hitchc.
Polygonum douglasii Green var. douglasii
Potentilla arguta Pursh var. convallaria (Rydb.) Th. Wolf
Rudbeckia occidentalis Nutt. var. occidentalis
Senecio crassidus Gray
Senecio serra Hook.
Stipa lettennanii Vasey
Trisetum spicatutn (L.) Richter
Viola nutiallii Pursh var. major Hook.

2.807

3.125

Green

Other

0..359

White

Other

1.318

White

Other

0.255

Pink

Wind

1.080

Yellow

Wind

1.462

Yellow

Wind

3.091

White

Wind

1.932

Blue

Wind

6.905

Green

Animal

0.383

Red

Wind

0.740

0.823

0.782

White

Other

2.141

Pink

Other

0.601

Purple

Other

0.206

0.205

Yellow

Other

0.084

Yellow

Other

0.115

0.125

White

Wind

0.802

Pink

Wind

0.263

0.294

Blue

Wind

2.786

Y'ellow

Other

0.719

Yellow

Other

5.125

5.992

6.660

Yellow

Other

2.427

3.260

3.338

White

Animal

11.065

Pink

Other

1.369

Red

Other

3.621

4.058

Blue

Animal

0.430

White

Wind

0.814

0.926

Yellow

Wind

4.752

White

Other

6.501

5.932

White

Other

27.706

26.478

Blue

Other

2.664

Yellow

Other

7.746

4.673

White

Animal

12.666

13.953

White

Other

1.976

White

Other

0.470

0.580

Green

Other

2.039

White

Other

0.276

Yellow

Other

2.070

1.657

Yellow

Other

2.633

2.855

Yellow

Wind

0.745

Yellow

Wind

1.105

Green

Animal

0.165

0.172

Green

Animal

3.944

3.478

Yellow

Animal

576

Great Basin Naturalist

Vol. 43, No. 4

"O
QJ
0)
if)

0.5

Oil

Meadow
N=26

Aspen
N=22

F=0.97 (means)

F= 1.23 (variances,A/M)

F=0.87 (variances, C/ A)

Conifer
N = 23

Fig. 1. Seed sizes of species in each stage of succes-
sion, depicted on log-scale. Dots represent means of log-
transformed species means for each stage, with bars ex-
tending one standard deviation above and below these.
N is the number of species in a stage. None of the F val-
ues are statistically significant.

12-15 m^ per individual averaged over the
two years. So few pollinators were caught in
the forest understories that we cannot recog-
nize their numerically dominant families
with confidence.

Discussion

Several explanations can be offered for the
lack of a successional increase in seed size
(Fig. 1). Unlike almost all seres for which
seed size has been studied, ours does not in-
clude a pioneer stage. The meadows we stud-
ied have apparently been unforested for cen-

turies and are being slowly invaded vegeta-
tively by forests due to climatic change
(Schimpf et al. 1980). Thus, we do not expect
the meadows to . be dominated by species
with light, highly vagile diaspores to the ex-
tent that recently deforested sites often are.

Likewise, our study site differs consid-
erably from the deciduous forest and tropical
moist forest sites of other studies with respect
to environmental conditions. Soil at our site
dries quickly after snowmelt, and summer
rain is far less than potential evaporation.
This is especially marked in the meadows,
where evaporative potential is more than
twice that under the conifer canopy (Schimpf
et al. 1980). Because low moisture avail-
ability has been correlated with greater seed
size, both interspecifically (Baker 1972) and
intraspecifically (Schimpf 1977), the dryness
of the meadows may offset the dim illumina-
tion of the spruce-fir understory as a force se-
lecting for larger seeds. Marion (1980) found
equally large seeds in the most xeric and most
shaded stages of a sere.

Some reports of increases in seed size with
succession appear to be equivocal. Werner
and Piatt (1976) found greater seed sizes of
herbaceous Solidago species in climax than in
serai ecosystems, but this is confounded by
the location of the climax stand in a drier cli-
mate than that of the serai stand. The signifi-
cance of an interspecific successional increase
in herb seed size (Abrahamson 1979) is due to
the presence of a single large-seeded climax
species; no significant increase can be shown
if the sizes are first normalized by log trans-
formation. Perhaps the most noteworthy seed
size increases during succession are those as-
sociated with shifts in life form composition.

A number of recent community-level stud-
ies of flower color in western North America
concur that white flowers are relatively more

Table 2. Number of species in each flower color cat-
egory in each successional stage.

Table 3. Number of species in each dispersal mode

Color

Meadow

Aspen

Dnifer

category in

each

successional

stage.

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