/ /? ^
t- o
HARVARD UNIVERSITY
Library of the
Museum of
Comparative Zoology
Tlie Great Basin Naturalist
VOLUME 40, 1980
Editor: Stephen L. Wood
Published at Brigham Young University, by
Brigham Young University
"b
0)
n.
TABLE OF CONTENTS
Volume 40
Number 1 - March 31, 1980
Some aspects of succession in the spnice-fir forest zone of northern Utah. David J.
Schimpf, Jan A. Henderson, and James A. MacMahon 1
Utah flora: Malvaceae. Stanley L. Welsh 27
I'tah flora: miscellaneous families. Stanley L. Welsh 38
The taxononiic status of the rosy boa LicJianura roseofiisca (Serpentes: Boidae). John R.
Ottley, Robert W. Murphy, and Geoffrey V. Smith 59
Hesperoperla hoguei, a new species of stonefly from California (Plecoptera: Perlidae).
Richard W. Baumann and Bill P. Stark ...! 63
Reproduction in three sympatric lizard species from west-central Utah. John B. Andre
and James A. MacMahon 68
Haplopappus aJpinus (Asteraceae): a new species from Nevada. Loran C. Anderson 73
Miscellaneous plant novelties from Alaska, Nevada, and Utah. Stanley L. Welsh and
Sherel Goodrich 78
New genera and new generic synonymy in Scolytidae (Coleoptera). Stephen L. Wood .... 89
The bacterium Thioploca ingrica on wet walls in Zion National Park, Utah. Samuel R.
Rushforth, Sheril D. Burton, Jeffrey R. Johansen, and Judith A. Grimes 98
Number 2 - June 30, 1980
Feeding ecology of Gilu l)or(ixo})itis (Osteichthyes: Cyprinidae) endemic to a thermal
lake in southeastern Oregon. Jack E. Williams and Cynthia D. Williams 101
First record of the pallid bat {Antwzuiis pallidus) from Montana. Jeff Shrver and Dennis
L. Flath ' 115
A CJuracanthiimi spider bite. Dorald M. Allred 116
Identity of narrow-leaved Chnjsothamnus viscidiflorus (Asteraceae). Loran C. Anderson 117
Ribulose diphosphate carboxylase activities in cold-resistant common mallow, Malva ne-
glecta Wallr. and a cold-sensitive tomato, Lycopersicon esculentum L., Ace 55 var.
William R. Andersen and Jack D. Brotherson 121
Recovery of Gambel oak after fire in central Utah. L. M. Kunzler and K. T. Harper 127
Relationships among total dissolved solids, conductivity, and osmosity for five ArtemUi
habitats (Anostraca: Artemiidae). Nicholas C. Collins and Gray Stirling 131
Spawning of the least chub {lotichtlufs pJilegethontis). Thomas M. Baugh 139
Transferrin polymorphism in bighorn sheep, Ovi.s canadensis, in Colorado. Patrick W.
Roberts, Donald J. Nash, and Robert E. Keiss 141
The genus Eriogonum Michx. (Polygonaceae) and Michel Gandoger. James L. Reveal 143
Parasites from two species of suckers (Catostomidae) from southern Utah. J. Craig Brein-
holt and Richard A. Heckmann 149
Soil water withdrawal and root distribution under gnibbed, sprayed, and undisturbed
big sagebnish vegetation. David L. Sturges 157
Swarming of the western harvester ant, Pogononiywiex occidentalis. Dorald M. Allred ... 165
Relationship between environmental and vegetational parameters for understory and
open-area communities. William E. Evenson, Jack D. Brotherson, and Richard B.
Wilcox 167
Seasonal activity pattern ot Columbian ground squirrels in the Idaho primitive area.
Charles L.' Elliott and Jerran T. Flinders I'^S
Habitat and plant distributions in hanging gardens of the Narrows, Zion National Park,
Utah. Ceorge P. Malanson 1^^
Short-term effects of logging on red-backed voles and deer mice. Thomas M. Campbell
III and Tim W. Clark 183
Terminal bud formation in limber pine. Ronald M. Lanner and James A. Bryan 190
Stinger utilization and predation in the scorpion Paniwctonus horeus. Bmce S. Gushing
and Anne Matherne 1^'^
Number 3 - September 30, 1980
Spatiotemporal variation in phenology and abundance of floral resources on shortgrass
prairie. V. J. Tepedino and N. L. Stanton 197
Doe owners and hvdatid disease in Sanpete County, Utah. Peter M. Schantz and Ferron
L. Andersen -i"
New grass distribution records for Arizona, New Mexico, and Texas. Stephan L. Hatch .. 221
A comparison of epiphytic diatom assemblages on living and dead stems of the common
grass Phr(i<fniites austmUs. Judith A. Grimes, Larry L. St. Glair, and Samuel R.
Rushforth ' 223
Poisonous plants of Utah. Jack D. Brotherson, Lee A. Szyska, and William E. Evenson ... 229
The successional status of Cuprcssus arizonica. Albert J. Parker 254
A self-pollination experiment in Pimis echilis. Ronald M. Lanner 265
Comparative floral biology of Penstemon eatonii and Penstenion cyananthus in central
Utah: a preliminary study. Lucinda Bateman 268
Differential habitat utilization by the sexes of mule deer. Michael M. King and H.
Duane Smith 273
Temporal activity patterns of a Dipodomys ordii population. Clive D. Jorgensen, H.
Duane Smith, and James R. Garcia 282
New records of western Trichoptera with notes on their biology. Bernard G. Swegman
and Leonard C. Ferrington, Jr 287
Observations on seasonal variation in desert arthropods in central Nevada. Robert D.
Pietruszka 292
Number 4 - December 31, 1980
Impact of the 1975 Wallsburg fire on antelope bitterbrush [Piiishid tiidcntata). Fred J.
Wagstaff .'. 299
Terrestrial vertebrate fauna of the Kaiparowits Basin. N. Duane Atwood, Clvde L. Prit-
chett, Richard D. Porter, and Benjamin W. Wood 303
A new species of fossil (^hn/sothamnti.s (Asteraceae) from New Mexico. Loran C. Ander-
son 351
New American bark beetles (Coleoptera: Scolytidae), with two recently introduced spe-
cies. Stephen L. Wood 353
Field observations on the response of the Railroad Valley springfish {Crenichtliy.s nc-
vadae) to temperature. Thomas M. Baugh and Bruce G. Brown 359
Woodrat nest flea Anomiopsylhis aniphihohis in southeastern Oregon. Harold J. Ego-
scue 361
Postemergence development and interyear residence of juvenile Columbian ground
.squirrels in the Idaho primitive area. Charles L. Elliott and Jerran T. Flinders 362
Flood frequency and the assenil)latj;e of dispersal types in hanti;in<j; (gardens of the Nar-
rows, Zion National Park, Utah. Geor<j;e P. Malanson and Jeanne Kay 365
Zonation patterns in the potholes of Kalsow Prairie, Iowa. Jack D. Brotherson 372
Plants of Angel Island, Marin County, California. J. D. Kipley 385
Additions to the vascular flora of Teton (bounty, Wyoming. Ronald L. Ilartinan and
Robert W. Lichvar '. '. \ 408
Index to X'oluine 40 414
L^ f<Ll^ ^/oc
rHE GREAT BASIN NATURALIST
Volume 40 No. 1
March 31, 1980
Brigham Young University
-VUS, COMP. ZOOL
LIBRARY
GREAT BASIN NATURALIST
Editor. Stephen L. Wood, Department of Zoology, Brigham Young University, Provo, Utah
84602.
Editorial Board. Kimball T. Harper, Botany; Wilmer W. Tanner, Life Science Museum;
Stanley L. Welsh, Botany; Clayton M. White, Zoology.
Ex Officio Editorial Board Members. A. Lester Allen, Dean, College of Biological and Agricul-
tural Sciences; Ernest L. Olson, Director, Brigham Young University Press, University
Editor.
The Great Basin Naturalist was founded in 1939 by Vasco M. Tanner. It has been published
from one to four times a year since then by Brigham Young University, Provo, Utah. In gener-
al, only previously unpublished manuscripts of less than 100 printed pages in length and per-
taining to the biological and natural history of western North America are accepted. The
Great Basin Naturalist Memoirs was established in 1976 for scholarly works in biological natu-
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Scholarly Exchanges. Libraries or other organizations interested in obtaining either journal
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Great Basin Naturalist Memoirs should be addressed to the editor as instructed on the back
cover.
5-80 650 45971
ISSN 0017-3614
The Great Basin Naturalist
Published at Pkovo, Utah, by
Brigham Young University
ISSN 0017-3614
Volume 40
March 31, 1980
\o. 1
SOME ASPECTS OF SUCCESSION
IN THE SPRUCE-FIR FOREST ZONE OF NORTHERN UTAH
David J. Sthiiiipf'-, Jan A. ilendersou,'^ and James A. MacMahon'
.\bstract.— a site in the Rocky Mountain siibalpine forest zone with which a series of hypotheses concerning
ecosystem succession was tested is characterized. Succession from herh-dominated meadows to climax forests of Eng-
ehnann spruce and suhalpine fir can follow at least four identified pathwa\'s. After fire, spruce and fir may reinvade
a site directly, follow invasion by aspen, or follow invasion by lodgepole pine, the pathway depending on a com-
bination of physical and biotic factors. In other cases, succession begins with long-established meadows which do not
owe their existence to fire. In this latter pathway, aspen invades meadows by suckering and changes the environ-
ment near the soil surface so as to facilitate establishment of the climax tree species. The biota and soils of four
characteristic serai stages (meadow, aspen, tir, spruce-tiri in this latter pathway are described.
Succession may be identified as the change
in ecosystem properties of a specified area
over a time interval of the same magnitude as
the generation time of the conspicuous or-
ganisms in the ecosvstem. Odum (1969) pro-
po,sed 24 trends in ecosystem properties as
successional changes take place. From 1976
through 1978 we and our colleagues .studied
many of these properties in a successional se-
quence of ecosystems in the subalpine zone
of the Wasatch Mountains of northern Utah.
In this paper we review patterns of succes-
sion i- the subalpine central Rockv Moun-
tains, describe our intensively studied se-
quence, and characterize the environment of
the study area. Subsequent papers will report
results of tests for trends in specific ecosys-
tem properties along the successional
gradient in the context of Odum's hypoth-
eses.
Materials and Methods
Rocky Mountain Subalpine Succession
The subalpine zone of the Rocky Moun-
tains is the uppermost forested part of the
Cordillera, characterized by climatic climax
ecosystems dominated by Engelmann spruce
{Picea engebnannii Parry) and subalpine fir
{Abies kisiocarpa [Hook.] Nutt.) (Daubenmire
1943, 1978, Oosting and Reed 1952), here-
after referred to as spruce and fir. Some
workers term this the upper montane zone
(Love 1970). Although long-term climatolog-
ical data from tliis zone are comparatively
scant, the occurrence of these subalpine for-
ests is apparently more closely correlated
with summer air temperatures than with pre-
cipitation patterns (Daubenmire 1956).
A variety of types of ecosystems may oc-
'Department of Biology and Ecology Center, Utah State University, Logan, Utah 84322.
'Present address: Department of Biology, University of Minnesota, Duliith, Minnesota 5.5812.
'Department of Forestry and Outdoor Recreation and Erology Center. Utah State University. Logan. Utah 84,322. Present address: U.S. Forest Service,
Federal Building, P.O. Box 2288. Olympia, Washington 98507.
Great Basin Naturalist
Vol. 40, No. 1
cupv a subalpine site from the time it is her-
baceous in character until its full devel-
opment as a spruce-fir forest. In the lower
range of the subalpine zone, preclimax forests
are sometimes dominated by tree species
which form climatic climax stands in lower
zones; the particular tree species involved
depend on the geographic location in the
Cordillera (Daubenmire 1943). Throughout
the subalpine zone the spruce-fir climax is of-
ten preceded by tree species which form cli-
max stands only under restricted topograph-
ic/edaphic conditions. The predominant
species of this type are quaking aspen {Popii-
lus treiniiloides Michx.) and lodgepole pine
{Pinus contorta Dougl. var. hitifolia Engelm.),
hereafter referred to as aspen and lodgepole.
While lodgepole is usually an invader of
burned areas, it also may invade unforested
sites which have not been binned recently
(Patten 1969).
Few detailed studies of successional path-
ways in the subalpine of the Rocky Moun-
tains have been reported. In Colorado, the lo-
cal presence of lodgepole seeds versus aspen
roots tends to determine the composition of
the preclimax forest which follows fire
(Stahelin 1943), though exposure and soils
also play a role (Langenheim 1962, Feet
1978).
Direct invasion of unforested areas by
spmce and fir can be extremely slow, even
when high numbers of their seeds reach the
site (Noble and Ronco 1978). Competition
with vigorous herbs or shrubs in the open
stands is undoubtedly a factor (Alexander
1974, Dunwiddie 1977), but the direct expo-
sure of spruce and fir seedlings to the sun and
night sky may be more detrimental. Deadfall
remaining after logging or blowdown in-
creases the rate of reestablishment of spruce
and fir via shading (Alexander 1974, Noble
and Alexander 1977). Even greater recruit-
ment of spruce and fir populations occurs in
the .shade of lodgepole or aspen if understory
vegetation is not too dense.
Several mechanisms for the shading ben-
efaction of Engelmann spruce establishment
have been identified. Shade obviously reduc-
es evaporative and heat .stress in the seed-
lings' environment. Even under favorable wa-
ter balance, spruce seedlings may be
intolerant of full intensity sunlight (Ronco
1970), though this was concluded from obser-
vation of seedlings planted at higher eleva-
tions than those of their parents. Shade-cast-
ing objects also lessen the nocturnal radiative
cooling of seedlings; spruce seedling growth
Is enhanced by higher night temperatures
(Hellmers et al. 1970). Under high soil water
potential, spruce seedlings emerge somewhat
faster than lodgepole at 16 C, but distinctly
slower than lodgepole at 35 C (Kaufmann
and Eckard 1977). The mechanisms for
shade-enhanced subalpine fir establishment
are less well understood.
Subalpine succession involving lodgepole
or aspen preclimax ecosystems seems to qual-
ify as a "facilitation" (Council and Slatyer
1977) type of serai sequence, at least with re-
gard to trees. Under certain circumstances
the preclimax .species invade the site more
readily than do the climax species, and great-
ly facilitate the later invasion of climax trees.
Aspen and lodgepole are unable to persist as
more than isolated individuals following
spruce-fir canopy development, possibly due
in part to their shoot geometry (Horn 1971)
and high light requirements in the face of
dense shade cast by spruce and fir.
Adaptive features conferring this greater
colonizing ability on lodgepole and aspen ap-
pear to center on the establi.shment phase of
the life cycle. Aspen bypasses the seedling es-
tablishment barrier of harsh environments by
vegetative reproduction, producing sucker
shoots from the roots. Establishment from
seed is thought to be extremely uncommon in
the central Rocky Mountain portion of the
range of this species (Cottam 1954). Thus as-
pen commonly invades only those sites imme-
diately adjacent to existing clones. Lodgepole
establishes only from seed; in some portions
of its range the seed-bearing cones open
strictly in response to heat, usually that from
fire (serotiny). Lodgepole seedlings do not
root deeper than spruce, but do develop
much more extensive root systems (Noble
1979), which should facilitate .seedling survi-
val on drier, exposed sites. Additionally,
lodgepole seedlings are not damaged or in-
hibited by the high light intensities (such as
in imforested sites) to which spruce and fir
seedlings arc less tolerant (Ronco 1970).
The relative proportions of spruce and fir
in a stand vary geographicallv, altitudinally.
March 1980
SCHIMPF ET AL.: SpRl'CE-FiR SUCCESSION
and locally (Daubcnmire 1943. Peet 1978).
As a species, fir tolerates a wider ranye ot
site conditions than spruce, and seems ca-
pable of becoming established on a greater
variety of substrates (Fowells 1965). Fir seed-
lings are more shade tolerant than spruce
(Fowells 1965). The composition of a voung
spruce-fir stand depends heavily upon which
species seeded abundantly just prior to a pe-
riod favorable for establishment. In the
northern Rocky Mountains good fir seed
crops are more frequent than good spruce
crops (LeBarron and Jemison 1953). In the
Wasatch Mountains the converse is true,
though the cohort of young fir is often larger
because of better fir survival (T. W. Daniel,
pers. comm.).
Both spruce and fir grow slowly by com-
pari.son to lodgepole or aspen. Both have nar-
row pyramidal crowns, fir the more so, and
may have few live branches in the lowest 5 m
of mature closed stands. Individuals of either
species seldom exceed 40 to 45 m in height.
Fir suffers heavy mortality at 125 to 175
years because of its greater susceptibility to
root rot, but spruce often lives 300 to 500
years. Both species are highly susceptible to
fire, due to their low crowns when young and
their thin bark.
The combination of irregular seed produc-
tion and irregular occurrence of suitable con-
ditions for tree establishment leads to an in-
frequent incidence in spruce-fir stands of the
inverse J-shaped population age structure
classically associated with climax commu-
nities (Alexander 1974, LeBarron and Jemi-
son 1953, Whipple and Dix 1979). Stands
may be even-aged or multi-aged, or may in-
clude suppressed individuals of considerable
age that show released growth only when a
gap opens in the canopy.
Injurious organisms may strongly affect the
dynamics of spruce-fir stands. Damage from
the bark beetle (Dryocoetes confii.siis Swaine)
and associated fungi often results in a reduc-
tion in the proportion of fir at a stand age of
125 to 175 years (Schmid and Hinds 1974).
Spruce becomes more susceptible to spruce
bark beetle {Dendroctonus rufijH'imi.s Kirbv)
(Schmid and Frye 1977) damage as it be-
comes older and larger in diameter, at about
250 years. Infestations erupt above endemic
levels when blowdowns of live spruce are fol-
lowed In temperatures and moisture levels
conducive to survival of the larvae; repeated
episodes lead to a stand of spruce so few and
large that they are extremely susceptible to
attack and blowdown (Dixon 1935, Miller
1970, Schmid and Hinds 1974). Stand deva.s-
tation results and the pace of succession rises
again.
Both spruce and fir seeds suffer higher
pregermination mortality in litter of their
own species than in litter from the other spe-
cies (Daniel and Schmidt 1972). Originally
described as Ceniculodendwn pyrifonnc Salt,
the infectious fungu.'^ now appears to be the
imperfect state of Caloscypha fiil^em (Pers.)
Boudier (Paden et al. 1978, Wicklow-Howard
and Skujins 1979). Such self-inhil)ition may
account in part for the relative rarity of ma-
ture monospecific stands of either spruce or
fir in the Rocky Mountains.
A Subalpine Site in the
Northern ^Vasatch Moimtains
The Wasatch xMountains of northern Utah
and southeastern Idaho are a major com-
ponent of the central Rocky Mountains. A
site for intensive study of ecosystem proper-
ties in relation to succession was selected in
the Utah State University School Forest, in
Cache and Rich counties, about 15 km south
of the Utah-Idaho border (Fig. 1). Stands in-
vestigated ranged between about 2550 m and
2650 m in elevation. The site is atop an un-
dissected plateaulike ridge of gentle topogra-
phy. Adjacent areas are lower in elevation
and are not a source of cold air drainage onto
the study site. Soils throughout the site are
derived from the Knight formation of the
Wasatch group, a Tertiary red conglomerate
of quartzite, sandstone, and shale (Veatch
1907). Tliis parent material occurs exten-
sively south of the study site (Stokes 1963).
The site contains no lakes or permanent
streams. Photoperiods range from 15 h 2 min
on 1 June to 13 h 9 min on 1 September,
with a 15 h 15 min maximum at the solstice.
Daily solar radiation totals (horizontal sur-
face) as high as 768 cal cm - have been mea-
sured (Eaton 1971).
The vegetation consists predominantly of
spmce-fir forests of moderate age, containing
Great Basin Naturalist
Vol. 40, No. 1
lll*>W
4a2N
4j<^JV
ii 1 1 1 1 1 1 1 1 1 1 1 1 1 rti
Fig. L Map of northern Utah, locating th. .School Forest (SF«). Garden Citv Su„n„il .CC ^ ' and the Inuus ot-
unshacled lexeept Hear l.akei and honndaries at I.5()(), 2 KM), and 2700 m.
March 1980
SCHIMPF ET AL.: SpRl CE-FiR SUCCESSION
onlv a few widely spaced spruce older than
275 vears. The oldest known trees are 367
vears (spruce) and 278 years (fir) (T. W. Dan-
iel, pers. conmi.). Scattered individuals or
small groups of lodgepole or aspen occur in
the spmce-fir stands. Within this broad for-
ested area are small meadows fringed with
aspen clones on various fractions of their per-
imeters (Fig. 2). Young spruce and fir are fre-
quently observed in these aspen stands but
seldom in the meadows, except on the north
margins of conifer stands, including the occa-
sional small clumps of mature spruce or fir
within the meadows. No significant logging
has taken place on the site. Cattle and sheep
have grazed the site since around 1900. The
zonal climax vegetation of the studv area be-
longs to the Abies lasiocdipa / Pcdicularis
mcemosa climax comniunit\ tvpe and habitat
type (Henderson et al. 1976).
The life-form composition of the vegeta-
tion is rather simple, consisting mainlv of
herbs and, except in the meadows, trees. Indi-
viduals of shrubbv species are uncommon;
shrub biomass is less than 1 percent that of
herbaceous biomass in meadows and under
aspen, and less than 5 percent under spruce-
fir. Small tree species and lianas are absent.
These features distinguish the sere under
study from those on most other forest succes-
sion research sites, especially those in the de-
ciduous forest biome. Cryptogams occur, but
are not a conspicuous element of anv stratimi
of the vegetation, except for periodic emer-
gence of basidiocarps in the forests. The gen-
eral aspect of the several categories of vege-
tation is illustrated in Figure 3. For a
discussion of the compositional and environ-
mental relationships of this forest to other lo-
cal forests, see Henderson et al. (1976) and
Lawton (1979).
A preliminary stud) of the structure of
over 100 stands on environmentally equiva-
lent sites in the School Forest (Sperger and
Henderson, unpubl.) indicated that there are
probably four major pathways of succession
leading to spruce-fir forests (Fig. 4). Pathway
"1" represents succession following the de-
struction by fire of a forest containing signifi-
cant aspen root biomass. The aspens sucker
within a short period of time to produce an
aspen-dominated stand. Spruce and fir sub-
secjuently invade, and eventuallv outlive and
replace the aspen.
Pathway "2" occurs in the lower elevation-
al range of the habitat type around the edge
ot the School Forest area. If aspen is not lo-
cally abundant, then lodgepole is the postfire
pioneer, provided that a local .seed source ex-
ists. Lodgepole cones are not serotinous in
this area, so seeding is from adjacent stands.
Spruce and fir establish and grow more slow-
ly than lodgepole; thus, a pine-dominated
ecosystem exists for some time prior to
spmce-fir stand recovery. Spruce and fir may
establish soon after fire, without site amelio-
ration by aspen or lodgepole, if a significant
quantity of unburned woody material re-
mains as protection. This is pathwav "3,"
where the climax species establish without
preclimax tree species. The climax stand
structure typically takes less time to develop
than through pathways "1" and "2."
Pathway "4," the most common, and that
which we studied intensively, begins with
long-persistent meadows, probably not of fire
origin. These are eventually invaded by a.s-
pen suckers followed by spruce and fir. Typi-
cally the fringe of aspen clones is dis-
continuous around the meadow, yielding
iiregular patterns of meadow invasion (Fig.
2). Once aspen is present, the time frame of
events resembles that of the other three path-
ways. However, the sere as a whole mav take
hundreds of years longer, because of the
long-persistent meadow stage.
Pathway "4" was studied by sinmltaneous
investigation of plots characterized as mead-
ow, aspen, fir, and spruce-fir. The fir-domi-
nated plots represent a stage, containing
some spruce, which sometimes occurs be-
tween occupancy by aspen and the t\pical
spruce-fir mixture. The inference of these
plots as a chronosequence seems reasonable,
based on the minimal relief of the studv site
and its uniform soil parent material. Because
the time since the meadows were last forest-
ed could not be determined, the stands can-
not be positioned on an absolute time axis.
Common herbaceous species in each stage
are listed in Table I. These represent samples
from one series of stands at Big Meadow and
should not be interpreted as ranks for the
study site in general. Eriogonum is semi-
shnibbv rather than wholh' herbaceous. A
Great Basin Naturalist
Vol. 40, No. 1
Fii;. 2. Aoiial pliolo iiicliicliii<4 Ihc Ikul'^i-r station in the Stliool Forest. Lower dianrani inclieates type ol forest
various sections of tlie photo. .Xrrow jxjints to tlie weather station.
March 1980
SCHIMPF ET AL.: SpRUCE-FiR SUCCESSION
substantial change in species composition oc-
curs along the siiccessional secjuence, with no
species being abundant in more than two
stages. The niunber of species is lowest in the
meadow and highest in the preclimax aspen
and fir stages, based on equal sampling effort.
The proportion of short-lived species in the
species list declines onlv slightly along the
sere. The semishrub Vacciniuin scopdiiuin
(Leiberg), perhaps the most abundant under-
storv species in the Rocky Mountain sub-
alpine /.one (Daubenmire 1978), was not
found on the site.
Mannnal species of the four stages are list-
ed in Table 2. The Northern Pocket Gopher
{Tliomoniy.s falpoidcs) is abundant in the
meadows, producing considerable distur-
bance of the upper portions of the soil hori-
Fig. 3. Photographs showing general physiognomy of four stages of succession: meadow, aspen, fir, spruce-fir. Pho-
tos courtesy of D. .\ndersen.
8
Great Basin Naturalist
Vol. 40, No. 1
zons. The amount of bare ground between
meadow plants is substantial and largely due
to gopher activity. Gophers are less abundant
under aspen and rare imder conifers. Their
effect on conifer invasion rate is not well un-
derstood, for though they can directly de-
stroy young spruce or fir, their activities may
also reduce herbaceous species competition
with tree seedlings. Gophers may also create
more favorable .seedbeds for conifers by ex-
posing mineral soil.
The dominant mammals in the coniferous
stages are the Snowshoe Hare {Lepus ameri-
canus) and the Red Squirrel (Tamiasciiirus
hudsonicus). Fall and winter browsing by
hares may retard conifer stand development
(Baker et al. 1921); an individual may con-
sume some 300 g (fresh weight) of woody
stems 4 mm or less in diameter daily in win-
ter (Pea.se et al. 1979). Hare population den-
sities in this area fluctuate little by com-
pari.son with their boreal counterparts
(Dolbeer and Clark 1975).
The Red Scjuirrel harvests spruce and fir
DISTURBANCE (fire)
GRASS -FORB MEADOW
ASPEN
LONG
PERSISTING
MEADOW
LODGE POLE
PINE
ASPEN
FIRS SPRUCE
SPRUCES FIR
SPRUCE / FIR
SPRUCE DOMINATED
CLIMAX WITH FIR
AS A SUBORDINATE.
SPARSE UNDERSTORY
OF Pedicularis racemosa
Fi^. 4. Summarv ol types of succfssioiial patli\\a\>
tli()in;lil to \w operative on liie Scliool Forest.
cones before the seeds are shed, transporting
them to large caches at shady bases of trees.
Finlev (1969) concluded that Red Squirrels
harvest almost all the cones produced in
vears of poor to average seed output, and
that only in high seed production years are
enough .seeds dispersed that significant tree
recruitment is possible. Further information
on the mammalian component of the ecosys-
tems is in Andersen et al. (1980).
Bird .species of the four stages are listed in
Table 3. Only the spruce-fir data are from
the site; preceding stages were inventoried in
more extensive stands nearby. Meadows fea-
tiue low numbers of both species and individ-
uals. Conspicuously absent from the co-
niferous stands is the Gray (or Canada) Jay
(Perisoreus canadensis), a common per-
manent resident throughout most of the
Rocky Mountain .subalpine zone. Similarity
in composition of the vertebrate species
among stages (Jaccard coefficient of commu-
nitv) and its variation between years in the
case of the avifauna are shown in Figure 5.
Further avian information is available in
Smith and MacMahon (submitted).
The comparisons of species similarities for
all vertebrates (Fig. 5) across the .sere show
that the most mature stage, spiTice, is most
different from the least mature stage, mead-
ow. The most similar stages are the two con-
ifers, spruce and fir. All pair-wise serai stage
comparisons of mammals, though showing
these patterns, are more similar than those
for birds. The implication is that the birds, as
we would expect, respond more dramatically
to the physiognomic changes attendant to de-
veloping from meadow to a deciduous forest
and finally to a coniferous forest. Mammals,
on the other hand, respond to the presence or
absence of trees, but do not vary as much
with tree species or leaf habit. The verte-
brate species count of 13-16 mammal .species
and 20-30 bird species for each of oiu' serai
stages correspond to mammal and bird spe-
cies counts from similar communities through
North America and also northern Europe
(e.g., Erskine 1977. Han.sson 1974, Jiirvinen
and Viiisanen 1976, Sabo and Whittaker
1979).
The most abundant insects in the meadows
are .species of aphidids and thripids. Follow-
ing a winter drought, the aphids declined
March 1980
Sc:himpf et al.: Spruce-Fir Succession
9
precipitously in 1977, when two species of
cicadelhds were the most abundant. A cica-
delHd is the most abundant in the aspen un-
derstorv, followed bv three species of thrip-
ids. In the aspen canopv the most common
species is a serpentine leaf miner (Gracila-
riidae), followed bv an aphidid, a cicadellid,
and a blotch mining gracilariid. In the con-
ifer understory the most common species is
an aphidid, followed by two thripids and two
more aphidids. During 1977 the aphids were
scarce and a cicadellid was the most abun-
dant. In the conifer canopy, the most abun-
dant species are an encyrtid, a mirid, a thrip-
id, and an eriosomatid. The insects on spruce
were observed to be verv similar to those on
fir.
Among soil and litter metazoan in-
vertebrates, numbers increase markedly
along the serai sequence. The relative abun-
dance of Collembola and the plant-feeding
nematodes (Tylenchida, Dorylaimida) does
not change much. The forested stages harbor
increased proportions of bacterivorous nema-
todes (Rhabditida), detritivorous mites (Ori-
batei), and predaceous mites (Mesostigmata).
Oligochaetes are essentially absent, a charac-
teristic of the region (Gates 1967), and gas-
tropods are also rare on the site. Populations
of protozoans were not estimated. Soil micro-
organisms also exhibit large absolute increas-
es in number through successional time and
are generally highest in density in the conifer
litter. Estimates of abundance for several
Table 1. Herbaceous species comprising 5 percent or more of the mean daily herbaceous bioinass in samples at
the Big Meadow succession stages. 1977. Numbers following a taxon correspond to its biomass rank, with percent of
the herbaceous biomass indicated for top-ranked species. Number of various categories of species samples and her-
baceous biomass in relation to tree leaf biomass are also included. Only herbaceous tissue of the semishrub Eriogo-
nititi is included.
Meadow
.^spen
Fir
Spruce-Fir
Achillea miUefoUiim L. ssp. kinulosa (Nutt.) Piper
Agropyron trachijcaiilum (Link) Malte var. lati^himc
(Scribn. & .Smith) A. \. Beetle
Agropijron tracJiijcauhnn (Link) Malte var. ghiiicuiu
(Pease & Moore) Malte
Aster engelmannii (Eat.) Grav
Bwmiis cciriudttts Hook & .\rn.
Dcscurainia nchdidsonii (Sweet) Schidz var. sonnet
(Robins.) C. L. Hitchc.
Erigeron speciosus (Lindl.) D. C. var. niacninthiis (Nutt.)
Croncj.
Eriogonum herarleoidcs Nutt.
Cilia aggregata (Pursh) Spreng.
Ligustiriim filicinitni Wats.
Lnpinits argenteus Pursh var. ruhricanlis (Greene)
Welsh
Pediciilahs raceinosa Dugl. var. alba (Pennell) Cronq.
Poa nervosa (Hook.) Vasey var. wheeleri (Vasev) C. L.
Hitchc.
Potentilld argiita Pursh var. convallaria (Hvdb.) Tli. Wolf
Riidheekia ocridenlidis Nutt. var. ocridentcdis
'^eneeio crasstiliis (Jrav
Seneeio serra Hook.
Stellaria jamesiana Torr.
Trisetum spicatum (L.) Richter
Total number of species sampled
Total number of annual species sampled
Total number biennial/short-lived perennial species
sampled
Herbaceous standing crop
H.S.C. -(- tree leaf standing crop
1 (29%)
25%)
Rank
3
1 (25%)
1 (26%)
7
3
6
2
6
41
62
58
49
8
10
7
8
2
4
3
2
100%
2%
0.1%
0.06%
10
Great Basin Naturalist
Vol. 40, No. 1
phylogenetic and functional groups are sum-
marized in Table 4.
Methods of Site Analysis
Field studies were conducted during 1976,
1977, and 1978. From 1970 through 1976 air
temperatures and precipitation had been re-
corded at a station (hereafter the Badger sta-
tion) near the edge of a small meadow on the
study site at 2650 m elevation (Fig. 2). Mean
monthly values recorded in Lomas (1977)
were used as dependent variables in multiple
linear regression models that employed as in-
dependent variables the values for the same
months at nearby lower elevation stations, for
which long-term means are available. For
both precipitation and temperature models
these stations are Logan, USU (elevation
1458 m, 27 km SW of the site) and Laketown
(1825 m, 16 km E-SE); data from Richmond
(1426 m, 25 km W-NW) were also used in
the precipitation models (see Fig. 1). These
data are published in the corresponding years
of Climatological Data, Utah (U.S. Weather
Bureau, U.S. Department of Commerce,
Washington, D.C.). The published mean val-
ues for these stations for the 1941-1970 peri-
od were entered into the appropriate regres-
sion models to produce an estimated monthly
mean temperature or precipitation total for
MAMMALS
1976-77
BIRDS
1976
BIRDS
1977
Fig. 5. Diagrams showing degree of affinity among
vertebrate assemblages in the four serai stages. Number
between stages is the Jaccard coefficient of community.
M = meadow, A = aspen, F = fir. S = spruce-fir.
T.\BLE 2. .Mammal species observed or trapped in various examples of the serai stages at the School Forest from
1976 through 1978.
Meadow
.\spen
Fir
Spruce-Fir
Cervus canadensis (Elk)
Cletliriotwinys ^apperi (Boreal Redback Vole)
Erctliizon dorsatuin (Porcupine)
Eutamki.s niiniiutt.s (Least Chipmunk)
Eiitaiiiias tind)rinus (Uinta Chipmunk)
Glaucomijs sabrinus (Northern Flying Scjuirrel)
Lagurus curtatus (Sagebrush Vole)
Lepus umericanus (Snowshoe Hare)
Mu.stela crminea (Shorttail Weasel)
Mustela frenata (Longtail Weasel)
Neotoma cinereu (Bushytaii Woodrat)
Odocoileus heiniontts (Mule Deer)
Pcromijsnts manicttlatus (Deer Mouse)
.Sorc.v sp. (Shrew)
Spcnnophilus latemlis (Colden-maiitled Stjuirrei)
Tainiascittrus hudsoninis (Red Sfjuirrel)
Ttiomomys talpoidcs (Northern Pocket Copher)
Zapus princeps (Western Jumping .Mouse)
Total
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
■p
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
X
X
X
X
X
X
X
13
16
14
1.3
March 1980
SCHIMPF ET AL.: SpRUCE-FiR SUCCESSION
11
the Badger station for the same 30-vear peri-
od.
Air temperature, relative humidity, and
precipitation were recorded and pan evapo-
ration measured during the summers of 1976,
1977, and 1978 at two other stations (eleva-
tion 2560 m) 1.5 km \-NE of the Badger sta-
tion. One station was located in a meadow
about 20 m from the edge of an aspen stand;
an identical one was situated about 80 m into
the forest from the first, under a well-devel-
oped spnice-fir canopy. These are referred to
as Big Meadow (meadow) and Big Meadow
(conifer), respectively. Instruments used were
tipping bucket recording precipitation
gauges (unshielded), recording hvgrothermo-
graphs in standard shelters, and U.S. Depart-
ment of Agriculture Class A evaporation
pans. Pan evaporation was measured at ir-
regular intervals which averaged about 7
days. The instruments were mounted on a
platform about 2.5 m above ground, higher
than normal because of the deep snows
which cover the area. Snowpack dynamics
were inferred from periodic observations at
the site, in conjunction with measurements
made by the U.S. Department of Agriculture,
Soil Con.servation Service, at Garden City
Summit, 8 km north of the site (elevation
2400 m) (Fig. 1).
Stem xylem water potential (predawn) of
conifers along a 10 X 50 m transect from
meadow through aspen to conifer was mea-
sured with a FMS pressure bomb (Waring
and Cleary 1967) on 3 August 1977. At least
three different branches of each spruce or fir
were measured; these trees were all about 1
m tall. Tree species population structure
along this same transect was assessed by
aging stems and estimating their heights.
Ages were determined by counting terminal
bud scale scars or annual growth rings in xy-
lem cores. In the latter case, the age at the
coring height (1.37 m) was corrected to total
age by adding 5 years for aspen and 20 years
for fir or spruce, values based on average
number of bud scale scars at that height.
Soil pits were dug to 1 m depth in two ex-
amples of each stage of the sequence in the
same stands being sampled for other ecosys-
tem attributes. Each soil horizon was identi-
fied, measured, and sampled prior to phvsical
and chemical analvsis in the laboratory. Per-
cent coarse fragments was estimated bv eye
in the field. Composition of the fine particle
fraction was determined by the hydrometer
method and with sieves. The following chem-
ical parameters were measured: pH (saturat-
ed paste), organic carbon (Walkley-Black
procedure), cation exchange capacity (sodium
acetate method), total cations (ammonium
acetate), potassium and phosphorous (sodium
bicarbonate), iron (DTPA extraction), and to-
tal nitrogen (Kjeldahl). Each of the eight pe-
dons studied was classified to the Great
Group level using the system of USDA Soil
Conservation Service (Anonymous 1975).
Soil moisture was monitored in 1977 and
1978 at the Badger station as a continuation
of previous work (Lomas 1977). Volume per-
cent moisture was measured with a Troxler
neutron probe in five tubes in a meadow and
ten tubes in a spruce-fir stand. Values were
recorded for each 30.5 cm increment of a 122
cm deep profile.
Results
Estimated 1941-1970 monthly mean tem-
peratures and precipitation totals for the
Badger station are reported in Table 5, along
with the multiple coefficient of determina-
tion for the regression model by which each
estimate was derived. These estimates are
plotted against the monthly means
(1941-1970) for three weather stations in the
Rocky Mountain subalpine zone with long-
term records (Figs. 6, 7). Most of the precipi-
tation at Badger is received as snow, with
nearly 30 percent of the annual precipitation
falling in December and January. Rainfall de-
creases to very low amounts in Julv, rising
somewhat in August. Long-term snowpack
records for Garden City Summit are present-
ed in Figure 8. Meadow peak snowpack
depths at the study site averaged about 40-50
cm greater than those at Garden City Sum-
mit during the corresponding winters.
Selected temperature, humidity, and pre-
cipitation data from the Big Meadow station
for the smnmers of intensive studv are pre-
sented in Table 6. Pan evaporation for these
summers is plotted in Figure 9 against days
after snowmelt in the meadows; these can be
converted to calendar dates from the snow-
melt dates in Table 6. The evaporation
12
Great Basin Naturalist
Vol. 40, No. 1
curves were integrated planimetrically over
various time intervals (Table 7). Over the in-
terval from 43 to 73 days after snowmelt,
evaporation in the conifer forest averaged 38
percent (1976), 46 percent (1977), and 46
percent (1978) as great as that in the mead-
ow. Soil moisture trends at the Badger station
during the three summers of intensive study
are represented in Figure 10.
From these data we can generally charac-
terize the three summers during which eco-
system properties were analyzed. During
1976 perhaps the most favorable conditions
for plant growth and development occurred,
with above average July rain, low evapo-
ration, and a moderately long frost-free peri-
od. The summer of 1977 followed an extraor-
dinarily dry winter, resulting in low soil
moisture content. The growing season began
very early and was much longer than for the
other two summers. June and August were
warmer than in 1976 and 1978, but not July.
Pan evaporation was very high, perhaps in
part because it was measured during the
longer days of early summer, since the satura-
tion deficits were not especially great. Ex-
tremely high August rainfall resulted from a
rare deep continental intrusion of a tropical
storm. The 1978 frost-free season began late
and ended early. Temperatures and evapo-
ration rates were intermediate, and satura-
tion deficits were relatively low in June and
July but high in August. Less rain fell than
during the other two summers.
Tree water potential and forest height
structure along the meadow-aspen-conifer
Table 3. Status of avian species observed on study areas in 1976 and 1977. P = permanent resident; B = sum-
mer breeder; F = feeder in serai stage, but not breeder; V = mitirating or wandering visitor; W = winter resident.
Species
Meadow
Aspen
Fir
Spruce-Fir
Turkey Vulture
Goshawk
Cooper's Hawk
Sharp-shinned Hawk
Marsh Hawk
Red-tailed Hawk
Golden Eagle
Sparrow Hawk
Blue Grouse
Ruffed Grouse
Mourning Dove
CJreat Horned Owl
Pygmy Owl
Common Nighthawk
Poor-will
Broad-tailed Hununingl)ird
Rufous Hiuiimingbird
Red-shafted Flicker
Yellow-bellied Sapsucker
WilHamson's Sapsucker
Hairy Woodpecker
Downv Woodpecker
Northern Three-toed Woodpecker
Lewis" Woodpecker
Dusky Flycatcher
Western Wood Pewee
Olive-sided Flycatcher
Violet-green Swallow
Tree Swallow
Steller's Jay
Black-billed Magpie
Clark's Nutcracker
Common Raven
Black-capped C^hickadee
Monntairi (Chickadee
rare permanent
resident
rare permanent
resident
rare permanent
resident
V
V
V
V
B
V
rare permanent
resident
V
V
W
W
W
P
W
W
F
V
V
V
p
p
p
V
V
V
V
F
B
V
V
F
F
V
V
F
B
B
B
V
B
B
P
P
P
B
V
P
V
V
B
B
B?
B
B
B
B
V
F
B
P
P
V
V
V
V
V
V
V
V
V
V
B
P
P
March 1980
SCHIMPF ET AL.: SpRUCE-FiR SUCCESSION
13
transect is plotted in Figure 11, and age
structure in Figure 12. For tabulated age
data, see Daniel et al. (1979:286). It can be
seen that the water stress of conifers one to
two meters tall increases as the aspen is re-
placed bv niatiue spruce and fir.
The eight pedons sampled are character-
ized in Table 8. Their taxonomic assignments
may be interpreted as follows: Cryochrepts
are soils with cold mean annual temperatures
showing little development. Crvoborolls have
dark surfaces, cold mean annual temper-
atures, and little development of the profile.
Paleboralfs are similarlv cold, but have an ar-
gillic horizon (accumulation of clav) deep in
the profile. This extensive development is in-
ferred to be the product of a different cli-
mate in the past.
Cone crops on the School Forest have been
estimated annually since 1947 (T. W. Daniel,
Table 3 coutiuuccl.
pers. comm.). In 1976 fir and spruce had very
low cone abundance, but in 1977 both bore
heavy crops. In 1978 the spruce crop was
very low, and fir bore a moderate crop.
Discussion
The greater similarity of summer air tem-
peratures than of precipitation patterns
among the geographically disparate spruce-
fir sites in Figures 6 and 7 lends support to
the assertion by Daubenmire (1956) that
lower growing season temperatures dis-
tinguish this zone from other Rocky Moun-
tain forests. A variety of precipitation re-
gimes permits the existence of spruce-fir
forests, from the summer-dominated precipi-
tation south and east in the Cordillera to the
winter-dominated patterns north and west.
Few other data on humidity or pan evapo-
Species
Meadow
.\spen
Fir
Spruce-Fir
White-breasted Nuthatch
Red-breasted Nuthatch
Brown Creeper
House Wren
.\merican Robin
Townsend's Sohtaire
Hermit Thrush
Mountain Bluebird
Colden-crowned Kinglet
Ruby-crowned Kinglet
Northern Shrike
Warbling \'ireo
Orauge-Crowued Warbler
Audubon's Warbler
MacGillivray's Warbler
Wilson's Warbler
Townsend's Warbler
Western Meadowlark
Brewer's Blackbird
Western Tanager
Black-headed Grosbeak
lazuli Bunting
Cassin's Finch
.\merican Goldfinch
Pine Grosbeak
Gray-crowned Rosv Finch
Pine Siskin
Red Crossbill
White-winged Crossbill
Green-tailed Towhee
\ esper Sparrow
IDark-eyed Junco (3 races)
Chipping Sparrow
Brewer's Sparrow
White-crowned Sparrow
Lincoln's Sparrow
B
B
P
P
B
P
B
B
B
B
V
B
B
B
W
P
V
B
V
B
B
B
B
V
V
B
B
V
B
V
V
V
B
V
B
B
B
V
B
B
V
w
P
P
V
B
B
B
B
B
V
B
V
V
B
F
B
B
B
F
B
B
B
B
V
B
B
V
V
14
Great Basin Naturalist
Vol. 40, No. 1
ration in this zone have been pubHshed. The
three-year monthly averages for saturation
deficit at Big Meadow (Table 6) are 9 per-
cent greater (June), 53 percent greater (July),
and 28 percent greater (August) than those
we computed from one year of temperature
and humidity records for a spnice-fir site in
the Front Range in Colorado (Marr 1967).
Pan evaporation rates in 1918 and 1919 in
the subalpine of Arizona (Pearson 1931) were
far less than ours (Table 7) in both forested
and exposed sites. The Arizona site receives
large amounts of summer rain, and may expe-
rience lower saturation deficits. Summer davs
o
o
UJ
a:
UJ
Q.
u
— BRIGHTON
--B-- BADGER
LEADVILLE
— -SUMMIT
Fig.
zones:
mates;
JFMAMJJASOND
6. Mean niontlilv temperatures during 1941-1970 period for four sites in the Rocky .Mountain spruce-fir
Badger station, Utah; Brighton, Utah; Leadville, Colorado; Summit, Montana. Means for Badger are esti-
those for the other sites are measured parameters.
Table 4. Relative abundance of phylogenetic and fimctional groups of soil microorganisms. Total number of or-
ganisms in the phylogenetic and functional group coimts do not agree because different methods were used; func-
tional groups are also not nuitually exclusive. Soil samples are from topmost 5 cm.
l(P organisms
g"^ dry substrate
Percent of total
Aerobic bacteria
Streptonivcetes
Fungi
Anaerobic bacteria
Sample type
Summer/Fall
Summer/Fall
Summer/Fall
Summer/Fall
Summer/Fall
Meadow soil
2.9/5.6
66/64
32/34
2/1
1/1
-\spen soil
4.9/9..3
68/68
29/29
3/2
0.2/0.2
Fir soil
7.7/14.7
77/78
20/20
3/2
0.2/ < 0.1
Spruce-fir soil
7.5/10.7
77/70
20/26
3/4
<0.1/<0.1
Fir litter
.39.6/28.0
86/84
13/14
1/2
<0.1/<0.1
Spnice-fir litter
15.0/22.0
77/84
21/13
2/2
<0.1/<0.1
March 1980
SCHIMPF ET AL.: SpRUCE-FiR SUCCESSION
15
15
1
BRIGHTON
o lOH
I-
<
^ 5-
a:
a.
0-
LEADVILLE
F
M
A
M
A S 0 N D
Fig
zone
^.7. Mean monthly precipitation totals during 1941-1970 period for four site.s in the Rockv Mountain spruce-fi
Badger station, Utah; Brighton. Utah; Leadville, Colorado; Summit, Montana. Means for Badger are estimates
those tor the other sites are measured parameters.
Table 4 continued.
Proteolytic
Summer/Fall
l(fi organisms g-^ dry substrate
Hemicellulolytic
Summer/Fall
Chitinolytic
Summer/Fall
Lipolytic
Summer/Fall
Cellulolytic
Summer/Fall
0.8/1.3
0.8/.3.0
1.5/1.8
1.2/1.4
3.6/3.7
1.8/2.0
0.1/0.02
0.3/0.08
0.9/0.4
1.0/0.4
2.6/1.3
1.2/1.4
0.3/0.6
0..3/0.7
0.4/0.3
0.4/0.3
0.6/0.5
0.6/0.4
0.08-0.04
0.2/0.07
0.3/0.1
0.3/0.1
0.4/0.8
0.4/0.2
0.1/0.01
().()6/().()3
0.07/0.09
0.09/0.06
0.2/0.2
0.3/0.2
16
Great Basin Naturalist
Vol. 40, No. 1
80
^^
E
e
60
5 40-
o
Q.
<
u 20
<
a.
40
—J—
50
60
70
80
90
00
DAYS AFTER SNOWMELT
Fig. 8. Pail evaporation at the Big Meadow conifer (C) and meadow (M) stations during summers 1976, 1977,
1978. Time is expressed as days after snowmelt in the meadow. Evaporation is plotted as mm wlr* at the midpoint
of the measuring period, with curves fitted bv eve. Ciuves not labeled bv vear are three-vear averages.
a.
LU
Q
O
<:
Q_
O
C/)
200 1
1 50 -
100 -
50
DEC
JAN
FEB
MAR
APR
MAY
Fig. 9. Time course of snowpack depths at ('.arden City Summit. Broken lines indicate records from the 196()s,
and solid lines indicate records from the 197()s. Years labeled are those in which the particular snowpack melted.
March 1980
SCHIMPF ET AL.: SpRUCE-FiR SUCCESSION
17
would also be shorter there. These com-
parisons, coupled with the paucity of summer
rain at our site (Fig. 7), lead us to venture
that ours is a comparatively dry example of
spruce-fir forest.
Our pan evaporation statistics indicate that
animals and plant shoots in the lower synusia
of the forest should have lower rates of water
loss than those in the meadows (Table 7, Fig.
8). Because saturation deficits were nearly
the same at both Big Meadow stations, the
differences nnist be largely due to differences
in net radiation or mean wind velocity. The
evaporation rates measiued in the meadow
compare favorably with those estimated for
tlie Badger station in 1970 bv use of Pen-
mans combination method (Eaton 1971).
Conifers of comparable small size exliibit
greater water stress in conifer-dominated
stands than in aspen stands (Fig. 11). We in-
terpret this to mean that competition for soil
water from large conifers is more severe than
from aspen or small conifers. From the pan
evaporation data (Fig. 8) we would expect
greater evaporative stress under aspen rather
than conifers, so the low xylem potentials un-
der conifers must be due to inability to ob-
tain sufficient soil water. This is probably an
important factor limiting recruitment in ma-
tiue spruce-fir stands.
The tree population age structures along
the transect (Fig. 12) indicate that aspen in-
vades the meadows at an average of about 19
cm yr', and is followed in about 20 years by
successful conifer establishment. The results
o
o
>
50
25
0-305 cm
0
50-1
- 25-
50
25
A. ^^M77
^C __M.C76
30.5-61.0 cm
^^ M77
^^ C76
"I r
:)U-
^ M78 61.0-91.5 cm
'^?-^^^^^=^'^^^-i'»S3E_,^^^J^J^
M76
25-
^077
C76
0^
T r 1 1
1
JUN JUL AUG SEP OCT
Fig. 10. Soil moisture in the meadow and conifer
stages at Badger station during the summers of 1976,
1977, and 1978. Volume percent water content is ex-
pressed for four .successive 30.5 cm horizons of the soil
profile.
T.\BLE 5 Estimated mean monthly temperatures and monthly precipitation totals for the Badger station during
the period 1941-1970. R^ is the coefficient of determination for the multiple regression model bv which each value
was estimated.
Month
Precipitation, cm
R2
13..5
0.74
8.9
0.03
12.7
0.73
10.4
0.39
7.1
0.81
5.2
0.72
1.2
0.61
3.0
0.68
3.4
0.95
6.2
0.68
9.2
0.90
14.3
0.73
Temperature, °C
R2
[anuarv
Fehniarv
March
April
May
June
Julv
.\ugust
September
October
.November
December
Annual total or mean
95.1
-10.2
-9.0
-5.3
-0.2
2.9
8.7
14.5
13.4
8.6
3.9
-5.0
-9.7
1.1
0.64
0.55
0..39
0..38
0.69
0.79
0.45
0.96
0.61
0.53
0.62
0.79
18
Great Basin Naturalist
Vol. 40, No. 1
presented are in general agreement with oth-
er transects on the study site. Altered slopes
in the aspen age vs. transect distance rela-
tionship suggest that advance of the clones
was slowed during the 1930s and accelerated
in the past decade. The 1930s were generally
warmer and drier than the succeeding period,
and the precipitation since 1970 (Lomas
1977) generally exceeded the 1941-1970 esti-
mated means.
Aspen dominates a plot on the School For-
est for 100 to 150 years before spruce or fir
assumes dominance (Fig. 12) and hastens the
demise of aspen through shading. Persistence
of occasional aspen stems in more open por-
tions of the spruce-fir canopy is important
from the standpoint of regeneration poten-
tial. The transition from aspen to spruce-fir
occurs over a relatively short time and dis-
tance (Fig. 12), explaining why we were
i2 -20n
< -15-
LU
O
_J
X
-10-
LU
^ -5i
0
spruce
Spruce y
oFlrV
r30
h20 E
X
LlI
hlO ^
10
To
30^
— I —
40
— I —
50
DISTANCE FROM MEADOW (m)
Fit;. 11. Average tree height of aspen, fir, and spruce populations, and water potential of short conifers along a
meadow-aspen-conifer transect. Water potential (^) of fir and spruce of about one meter height was measured .3
August 1977. Values are plotted against distance from the edge of a meadow.
Table 6. Climatological data from Big Meadow weather stations. Precipitation is average of meadow and conifer
station totals. Snowmelt is first date when meadow snow cover v.as estimated to be less than 10 percent. Saturation
deficit is computed from weekly mean temperature and mean relative humidity for the meadow station.
.\ir tempe
ature
, °c
Snowmelt
Mean
High
Stage/year
June
July
August
Meadow
1976
7.8
13.7
9.9
24.7
June 4
1977
10.9
12.7
11.7
23.3
Mav 15
1978
-
12.6
10.9
25.6
June 16
Conifer
1976
8.7
14.6
10.6
27.8
1977
12.0
13.9
12.4
26.7
1978
8.8
13.9
12.0
26.1
March 1980
SCHIMPF ET AL.: SpRUCE-FiR SUCCESSION
19
unable to find mixed aspen-conifer stands
sizeable enough to study. Tabulation of the
ages of trees by species indicates that the in-
vasion of spruce and fir occurs with less tem-
poral regularity than aspen invasion of mead-
ows (Daniel et'al. 1979:286).
The great similarity in soil physical and
chemical properties among the four stand
types (Table 8) strengthens our view that the
stands differ mainly as a consequence of suc-
cession and not because of differences in site
potential. Most of the edaphic differences ex-
isting among stages are manifested near the
soil .surface, reflecting the influences of the
current resident biota. The absence of char-
coal in the upper few cm leads us to con-
clude that fire has not been a major force in
the meadow ecosystems during the past few
centuries.
10 20 30 40
DISTANCE FROM MEADOW (m)
Fig. 12. .\ge stnicture of aspen, fir, and spruce populations along a nieadow-aspen-tonifer transect. Mean ages are
plotted against distance from the edge of a meadow.
Table 6
continued.
Last
First
freeze
Precipitation, cm
Saturation deficit,
g nr3
freeze
July August
June
July
August
June 26
May .30
June 26
June 27
May 31
Jime 26
Aug. 26
Aug. 28
Aug. 16
Aug. 26
Aug. 27
Aug. 15
4.0 2.3
2.2 12.7
0.5 3.5
5.10
4.42
3.54
7.49
5.65
5.21
4.35
5.13
5.49
20
Great Basin Naturalist
Vol. 40, No. 1
We must, then, invoke reasons other than
fire for the existence of the meadows we
studied. We .suggest that we are .seeing the
disappearance of subalpine meadows that
were chmatic chmax during a colder period.
If this is true, then the sequence we describe
is botii allogenic (warmer temperatures al-
lowing forest expansion) and autogenic (cli-
max forest establishment facilitated by aspen
invasion) in character. It is important to de-
termine whether these meadows are old or
yoimg. Even though phvsiognomy and ener-
getics may be similar in pioneer meadows
and mature meadows, the life history charac-
teristics of the organisms and the structure of
the communitv mav well differ due to the
length of time a stand has been in existence
(MacArthur and Wilson 1967).
There is evidence from around the globe
for cooling during this earlier period (Brav
1971), though we know of no published evi-
dence for colder conditions in the immediate
vicinity of our site. A dendrochronological
study of a site 375 km SW of ours indicates
that temperatures were cooler than at pres-
ent from 1541 to 1780 (LaMarche and Stock-
ton 1974). This corresponds to the dearth of
trees older than 275 years on our site, though
their absence could be interpreted as the re-
.sult of insect devastation (Miller 1970).
Whipple and Dix (1979) also found fewer
.spmce between ages of 300 and 400 years in
Colorado than they expected.
Whether this cooling was sufficient to
change the general aspect of our studv site
we can only gue.ss. There is general agree-
ment that the cooling was greatest at high
latitudes and altitudes (Bray 1971). The limit-
ing summer air temperatures for normal
growth of spruce or fir are not well under-
stood; Wardle (1968) presented evidence that
the limit lies between 11 and 12 C (July) for
spruce in Colorado. A July decline of 2.5 C
(Table 5) might have been sufficient, in com-
bination with the level topography of our
site, to produce an open subalpine parkland
of herbaceous meadows containing scattered
trees (Billings 1969). The wide dispersion of
old conifer individuals on the School Poorest is
reminiscent of the tree patterns in such park-
lands. Forest expansion since this colder peri-
od may have been anomalously rapid; if our
site reflects climatic trends in llie Rocky
Mountains as a whole, then the years 1870 to
1945 may have been the most warm-moist
75-year period since 1130 (Bradley 1976).
In conclusion, we would like to reempha-
size the integral role played by preclimax
trees in the structuring of the climax forest.
The successional change we describe is not
merely a consequence of the passage of time
and the differential growth rates of preclimax
and climax species (Dmry and Nisbet 1973).
Species characteristic of the climax are not
"present but inconspicuous" early in succes-
sion; they establish extremely slowly and only
on the north margins of their stands in the
absence of aspen or lodgepole.
This is illustrated in two discrete subalpine
basins we have observed. Birch Creek and
Summit Creek South Fork, 17 km W of our
study site. Lodgepole is absent throughout
this western portion of the mountain range at
this latitude. These two basins are unusual in
that they are also devoid of aspen in the sub-
alpine zone. Aspen may have been locally ex-
tirpated during the Pleistocene, when both
basins held small alpine glaciers (De Graff
1976). Now aspen is unable to invade from
adjacent topographic units because its in-
ability to reproduce by seed. Aspen is present
at lower elevations in these drainages, but
perhaps only as the late-leafing form which is
not known to occur at higher elevations (Cot-
tam 1954). The spruce-fir stands of these two
topographic units, at the same elevations as
our study site, are small discrete groves in a
Table 7. Estimated pan evaporation rates at the Big
Meadow stations, expressed as mm wk"'. Means for vari-
ous intervals were obtained by planimetric integration
ot the turves in Fisrure 8.
Stage
Period
Rate
Meadow
1976, 4.3-73 days after melt
40
1977, 4.3-73 davs after melt
.56
1978. 4.3-73 days after melt
43
1976, entire curve
39
1977, entire ciuve
50
Three-year mean, entire cur\e
42
(>oniter
1976. 43-73 da>s after melt
15
1977, 43-73 davs after melt
26
1978, 43-73 days after melt
20
1976, entire curve
16
1977, entire curve
23
Three-year mean, entire curve
17
March 1980
SCHIMPF ET AL.: SpRUCE-FlR SliCC.ESSIOX
21
matrix oi herbaceous veij;etation. We h\ poth-
esize that this pattern represents exactly the
result we would expect for the allo<Tenic ex-
pansion of spruce-fir forest from subalpine
parkland in the absence of aspen or lodge-
pole.
Acknowledgments
The following contributed data or other in-
formation used in this paper: D. Andersen, L.
Bennett, T. Daniel. D. Factor, J. Habeck, G.
Hart, P. Lawton, R. Lawton, G. Reese, M.
Schwartz, J. Skujins, K. Smith, R. Sperger, N.
West, M. Wolfe, D. Zeeman. Many of them
provided constructive comments on an ear-
lier draft of this paper as did R. Bayn and G.
Waagen. We particulary want to thank T.
W. "Doc" Daniel for sharing his wealth of
field experience in the Utah State University
School Forest. R. Bavn, L. Finchum, and B.
Peitersen were instrimiental in the prepara-
tion of the manuscript. This research was
supported by Grant DEB 78-05328 from the
National Science Foundation to James A.
MacMahon.
Literature Cited
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in the central and southern Rockv Mountains:
The status of our knowledge. L'.S. Dept. Atiric.
For. Serv. Res. Pap. R.\l-121.
.\\i)ERSEN, D. C, J. A. M.\cM.\Ho.\, .WD M. L. Wolfe.
1980. Herbivorous mammals along a montane
sere: CommunitS' structure and energetics. ].
Mammal 61: In press.
Anonymous. 197.5. Soil ta.\ononi\. L'.S. Dept. .Vgric. Soil
Cons. Serv. Agric. Hndbk. 4.36.
B.\KER, F. S., C. F. KoRSTi.w. .\.\i) \. J. Fetheroi.k.
1921. Snowshoe rabbits and conifers in tiie
Wasatch Mountains of Utah. Ecology 2: .3()4-:310.
BujjNcs. W. D. 1969. Vegetational pattern near alpine
timberlines as affected bv fire-snowdrift inter-
actions. Vegetatio 19: 192-207.
Bh.xdlev, ]\. S. 1976. i'recipitation histors ol the Kockx
.Mountain states. \\'est\ic\\ Press. Bouldci. (Colo-
rado.
Brav. J. B. 1971. Vegetatioiud distribution, tree growth
and crop success in relation to recent climatic
change. \d\. Kc-ol. Bes. 7: I77-2'>.'5.
Co.N.NEi.i.. J. H.. AM) B. (). Si.ATYKH. 1977. .Mechanisms
ot succession in natural conniumities and their
role in cominunit\ stabilitx and organization.
Amer. \atur. Ill: 1119-1144.
(.'oTiAM. W. P. 19.54. Prevernal leafing of aspen in I'tah
mountains. |. Arnold Arbor. .35: 239-2.50.
Danu:!.. T. W.. J. A. lli;i,\is. am) F. S. Baker. 1979.
Principles of sihicullure. 2d ed. McCirau-Hill,
\e\v '^'ork.
I)\\u 1.. T. \\'.. a.m) J. Sc:iiNnnr. 1972. Lethal and non-
lethal effects of the organic horizons of forested
soils on the germination of seeds from several as-
sociated conifer species of the Rockv Mountains.
Can. J. For. Res. 2: 179-184.
l)\i luwuHK. R. I". 1943. V'egetational zonation in the
Bock\ Mountains. Bol. Rev. 9: .32.5-.393.
19.56. Climate as a determinant of vegetation dis-
tribution in eastern Washington and northern
Idaho. Ecol. Monogr. 26; 131-1.54.
1978. Plant geography, with special reference to
North Xnierica. Academic Press. New York.
l>i: ('.RAFF, ). V. 1976. Quaternarv geomorphic features
of the Bear River Range, north-central Utah. Un-
published thesis. Utah State L'niv.. Logan.
Dixon. 11. 19.35. Ecological studies on the high plateaus
of Utah. Bot. Caz. 97: 272-320.
DoLREER, R. A., A.M) \\ . R. Ci.ARK. 1975. Population
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Mountains. J. Wildl. Manage. .39: .5.3.5-.549.
l>Ri R^. \\ . H.. AM) 1. (]. T. \isBET. 1973. Succession. J.
.\rnold Arlior. .54: .331-368.
Di NWHJDiE. P. W. 1977. Recent tree invasion of sub-
alpine meadows in the Wind River Mountains,
Wyoming. Arc. Alp. Res. 9: 393-399.
Eaton. F. D. 1971. Soil inoisture depletion, actual and
potential e\apotranspiration in an Engelmann
spruce-subalpine fir forest. Unpublished thesis,
Utah State Uni\ .. Logan.
Erskine. .\. J. 1977. lihds in boreal Canada: (Commu-
nities, densities and adaptations. Can. Wildl.
Serv. Rep. Series No. 41.
FiM.E'i . R. B.. Jr. 1969. Cone caches and middens of Ta-
niia.'icitints in the Rock\ .Mountain region. Pages
2.3.3-273 iti J. K. Jones, Jr., ed. Contributions in
mammalogv. Univ. Kansas Mus. Nat. Hist. .Misc.
Pnbl. 51, Lawrence, Kansas.
FowEi.i.s. H. A. 1965. Silvics of forest trees of the United
States. U.S. Dept. Agric. Agric. Hndbk. No. 271.
(^ATFs. C;. E. 1967. On the earthworm fauna of the
(ircat American Desert and adjacent areas. Crcal
Basin Natur. .37: 142-176.
Hansso.n, L. 197 L Small mannnal productivity and con-
sumption in spruce forest and reforestation in
south Sweden. Oikos 25: 1.53-1.56.
Hei.i.mfrs, H., M. K. Centiie, a.nd F. Ronco. 1970.
Temperature affects growth and development of
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\\ . YoiNt.. AM) .\. YoiNCiBEOon. 1976. Prelimi-
narv forest habitat types of northwestern Utah
and adjacent Idaho. Dept. Forestry and Outdoor
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I1()H\. H. S. 1971. The adaptive geometr\' of trees.
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[vHM.NEN, O., AM) R. \. V.\is.\NE.\'. 1976. Between-vear
component of diversit\ in communities of breed-
ing land birds. Oikos 27:.34-.39.
22
Great Basin Naturalist
Vol. 40, No. 1
Table 8. Physical and
chemical characteristics, and
classifications
of two
pedons in each of four
stages of succes-
sion in the School Forest.
Roots: f = few
, c = common, m = many.
vf =
very fine, f = fine, m
= mediiun, c =
coarse.
Cation
Percent
Percent
Percent
exchange
sand,
coarse
organic
capacity
Location, depth, cm
silt, clay
fragments
Texture
pH
carbon
meq/lOOg
Big Meadow (Meadow \)
0-9
.30,53,17
30 gravel
10 stones
gravelly
.silt
loam
63
1.9
14.2
9-38
30,53,17
30 gravel
10 stones
gravelly
silt
loam
6.0
1.5
15.7
38-ft3
33,47,20
30 gravel
10 stones
gravelly
loam
5.8
1.1
13.8
63-98
45,.37,18
40 gravel
5 cobbles
gravelly
loam
5.5
0.3
8.5
98-100
4633,21
40 grave!
5 cobbles
gravelly
loam
5.4
0.2
9.5
Doc's Meadow (Meadow B)
0-9
35,45,20
less than 10
loam
6.3
1.7
14.8
9-26
.35,43,22
less than 10
loam
.5.8
1.4
14.2
26-40
.35,42,23
10 gravel
loam
5.5
0.7
11.5
40-67
45,.33,22
30 gravel
gravelly
loam
5.2
0.3
10.1
67-100
58,26,16
25 gravel
gravelly
fine
sandy loam
5.3
0.2
62
Big Meadow Fir (Fir A)
0-20
.33,56,11
20 gravel
gravelly
.silt
loam
5.8
1.7
18.7
20-62
47,.38,15
30 gravel
10 cobbles
gravelly
loam
5.7
0.7
9.1
62-88
.3.3,.5611
30 gravel
5 cobbles
gravellv
silt
loam
5.2
0.5
12.2
88-100
52,31,17
40 gravel
gravelly
very fine
sandy loam
5.3
0.3
7.9
Harts Fir (Fir B)
0-10
43,4.3,14
25 gravel
gravelly
loam
5.7
2.5
9.9
10-.35
40,44,16
30 gravel
gravellv
loam
.5.8
1.0
8.8
.35-62
40,4.3,17
.30 gravel
gravelly
loam
.5.6
0.8
8.4
62-100
.37,.32,31
30 gravel
gravellv
clay
.5.4
0.3
19.4
Big Meadow Aspen (.\spen A)
4-0 litter
0-6
6-27
organic
.34,44,22
organic
,5.6
30 gravel gravelly 5.6
20 stones loam
25.3
1.7
29.4
19.4
March 1980
SCHIMPF ET AL.: SPRUCE-FiR SUCCESSION
23
Table 8 continued.
64
Percent
base
saturation K, ppm
210
P, ppm
16.3
Fe, ppm
Roots
Charcoal
24
nnf
nif
July percent
total
nitrogen
& depth, cm Great group
0.11 top 3
Crvohoroll
54
146
52 69
72 33
72 44
6.5
3.9
3.5
3.1
31
40
26
24
cvf
cf
cvf
cf
ff
ff
0. 1 1 25
0.05 50
0.03 100
64
128
16.0
.34
cf
0.(W top 3
54
69
7.7
45
ff
+
0.06 25
65
51
3.9
55
ff
4-
-
42
3.3
47
fvf
+
0.05 50
71
22
3.2
37
0.03 100
Cr\oc]irept
Paleboralf
68
73
82
87
122
50
42
31
5.4
5.0
2.6
34
44
20
nivf
inf.fin
fc
cvf
cf.fm
fc
cvf
cf
ff
1..35 litter
0.19 top 3
0.11 25
0.07 50
0.05 100
Paleboralf
-
182
42.4
-
cfjni
fc
+
1.. 39 litter
0.19 top 3
64
200
.37.7
56
cf.fm
fc
+
0.06 25
63
148
32.8
51
ff.fm
fc
+
0.04 .50
87
161
21.0
32
fm
+
0.04 1(K)
Crvoboroll
62
400 +
192
.30.9
4.4
34
29
invt
mf
mill
nnf
lilt. mill
mc
0. 1 1 top 3
0.17 25
24
Great Basin Naturalist
Vol. 40, No. 1
Kaufmann, M. R., and a. N. Eckard. 1977. Water po-
tential and temperature effects on germination of
Engelmann spruce and lodgepole pine seeds. For.
Sci. 2.3: 27-33.
La.Marche, V. C, Jr., and C. W. Stockton. 1974.
Chronologies from temperature-sensitive bristle-
cone pines at upper treeline in western United
States. Tree-ring Bull. 34: 21-45.
Langenheim, J. H. 1962. Vegetation and environmental
patterns in the Crested Butte area, Gunnison
County, Colorado. Ecol. Monogr. 32: 249-285.
Table 8 continued.
Lawton, p. M. 1979. An investigation of the environ-
mental relationships of selected forest habitat
types in northern Utah. Unpublished thesis, Utah
State Univ., Logan.
LeBarron, R. K., and G. M. Jemison. 1953. Ecology and
silviculture of the Engelmann spnice-alpine fir
type. J. For. 51: 349-355.
Lo.viAS, D. A. 1977. Soil water depletion following clear-
cutting small plots in a spnice-fir forest in north-
ern Utah. Unpublished thesis. Utah State Univ.,
Logan.
Cation
Percent
Percent
Percent
exchange
sand,
coarse
organic
capacity
Location, depth, cm
silt, clay
fragments
Texture
pH
carbon
meq/lOOg
24-44
46,39,15
40 gravel
10 cobbles
gravelly
loam
5.6
0.5
10.1
44-100
54,29,17
40 gravel
5 stones
5 cobbles
gravelly
fine
sandy loam
5.5
0.4
10.1
Hart's Aspen (.\spen B)
0-9
43,43,14
20 gravel
gravelly
loam
.5.7
1.5
11.2
9-27
44,42,14
20 gravel
gravelly
loam
5.8
1.3
9.5
27-49
43,43,14
.30 gravel
gravelly
loam
.5.9
0.7
7.6
49-75
44,42,14
30 gravel
10 cobbles
gravelly
loam
,5.6
0.7
7.3
7,5-100
42,4,5,13
10 gravel
loam
5.3
0.6
12.8
Big Meadow Spruce-fir (Spruce-fir A)
3-0 litter
0-19 ,39,48,13
■30 gravel
gravelly
loam
,5.8
1.7
12.9
19-47
47-72
43,43,14
62,18,20
72-100 70,8,22
Sinks Road Spruce-fir (Spruce-fir B)
5-0 litter
0-11 29,57,14
30 gravel
10 cobbles
20 stones
less than 10
gravelly
loam
coarse
sandv
clay loam
less than 10 coarse
sandy
clay loam
15 gravel
silt
loam
.5.5
6.5
5.0
1.0
0.4
0.5
1.3
10.4
7.1
6.2
II -,35
,35-60
60-85
85-l(K)
29,.57.14
40,46,14
43,44,13
.54.,32,14
20 gravel
,30 gravel
•5 cobbles
,30 gravel
5 cobbles
40 gravel
5 cobbles
gravel I V
silt loam
gravelly
loam
gravelly
loam
gravelly
very fine
sandv loam
,5.4
5.4
.5.4
0.7
0.6
0.5
0.3
8.3
7.7
6.4
5.1
March 1980
SCHIMPF ET AL.: SpRUCE-FiR SUCCESSION
25
LfivE, D. 1970. Subarctic and subaljMiie: W'Iumc and
what? Arc. Alp. Res. 2; 63-73.
M.\c..\rthur, R. H., .a..\d E. O. Wilson. 1967. The theo-
ry' of island biogeography. Princeton Univ. Press,
Princeton, New Jersey.
Marr. ]. \\'. 1967. Ecosystems of the east slope of the
Front Range in Colorado. Univ. Colorado .Stu.,
Ser. in Biol. \o. 8, Bonlder, Colorado.
Miller, P. C. 1970. .\ge distributions of .spruce and fir
in beetle-killed forests on the White River
Table 8 continued.
Plateau, Colorado, .^iner. .Midi. Natur. 83-
206-212.
Noble, D. L. 1979. Roots of lodgepole pine seedlings
reach depth of only 3 to 4 inches their first sea-
son. U..S. Dept. .\gric. For. Serv. Res. Note RM-
.363.
Noble, D. L., anp R. R. Alexa.nder. 1977. Environmen-
tal factors affecting natural regeneration of En-
gelniann spruce in the central Rockv Mountains.
For. Sci. 23; 420-429.
Percent
l)ase
saturation K, ppm
P, ppm
Fe, ppm
Roots
Charcoal
July percent
total
nitrogen
& depth, cm Great group
64
54
51
4.8
3.2
36
33
cvf
cf.fm
fc
ff
0.09 50
0.06 100
81
86
75
80
.56
51
68
61
286
220
179
172
132
142
90
48
31
35.0
29.0
25.6
24.3
5.6
10.7
9.0
5.5
4.2
27
34
34
42
19
54
51
28
mf.nini
-f-
cf,cm
+
cf,cm
+
cf.cm
+
ff,fm
+
mvf
mf,cm
cc
mvf
mf,cm
cc
cvf
cf.ftn
fc
ff
Crvoboroll
0.17 top 3
0.06 25
0.03 50
0.03 1(K)
1.29 litter Cryochrept
0.19 top 3
0.07 25
0.07 .50
0.03 1(X)
Cryochrept
1.25 litter
38
93
8.8
69
mvf
mf,cm
fc
0.15 top 3
.36
64
2.1
40
cvf
cf.fc
+
0.08 25
39
71
1.4
34
cf.cm
+
0.04 50
47
33
1.1
29
cf.cm
86
.30
1.2
23
cf,cm
0,03 U)()
26
Great Basin Naturalist
Vol. 40, No. 1
Noble, D. L., and F. Ronco, Jb. 1978. Seedfall and es-
tablishment of Engelmann spruce and subalpine
fir in clearcut openings in Colorado. U.S. Dept.
Agric. For. Serv. Res. Pap. RM-200.
Odum, E. p. 1969. The strategy of ecosystem devel-
opment. Science 164: 262-270.
OosTiNG, H. J., AND J. F. Reed. 1952. Virgin spruce-fir
forests in the Medicine Bow Mountains, Wyom-
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Paden, J. W., J. R. Sutherland, and T. A. D. Woods.
1978. Caloscypha fulgens (Ascomyce-
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seed pathogen Geniculodendron pijrifomie (Deu-
teromycotina: Hyphomycetes). Can. J. Hot. 56:
2.375-2.379.
Patten, D. T. 1969. Succession from sagebrush to mixed
conifer forest in the northern Rocky Mountains.
Amer. Midi. Natur. 82: 229-240.
Pearson, G. A. 1931. Forest types in the southwest as
determined bv climate and soil. U.S. Dept. Agric.
Tech. Bull. 247.
Pease, J. L., R. H. Vowles, and L. B. Keith. 1979. In-
teraction of snowshoe hares and woody vegeta-
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Peet, R. K. 1978. Latitudinal variation in southern
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.331-339.
Sabo, S. R., and R. H. Whittaker. 1979. Bird niches in
a subalpine forest: .\n indirect ordination. Proc.
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ScHMiD, J. M., AND R. H. Frye. 1977. Spruce beetle in
the Rockies. U.S. Dept. .\gric. For. Serv. Gen.
Tech. Rep. RM-49.
ScHMiD, J. M., AND T. E. Hinds. 1974. Development of
spruce-fir stands following spruce beetle out-
break. U.S. Dept. Agric. For. Serv. Res. Pap. RM-
131.
Smith, K. G., and J. A. MacMahon. Bird communities
along a montane sere; Community structure and
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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.
Stokes, W. L. 1963. Geological map of northwestern
Utah. Washington, D.C.
Veatch, A. C. 1907. Geography and geology of a por-
tion of southwestern Wyoming. U.S. Geol. Survey
Prof. Paper 56.
Wardle, P. 1968. Engelmann spruce {Picea engehnannii
Engel.) at its upper limits on the Front Range,
Colorado. Ecology 49: 48.3-495.
Waring, R. H., and B. D. Cleary. 1967. Plant moisture
stress: Evaluation by pressure bomb. Science 155:
1248-1254.
Whipple, S. A., and R. Dix. 1979. Age structure and
successional dynamics of a Colorado subalpine
forest. Amer. Midi. Natur. 101: 142-158.
Wicklow-Howard, M. C, and J. Skujins. 1979. In-
fection of Engelmann spruce seeds by Gen-
iculodendron pijriforme in western North Ameri-
ca. Mycologia (in press).
UTAH FLORA: MALVACEAE
Stanley L. Welsh'
.■\bstract.— This paper is the third in a series dealing with a revision of the flora of Utah. Treated herein are 9
genera and 23 species, including both coniinonly cultivated, escaped, and indigenous representatives. Proposed new
taxa include Sphuemkcdnwssiilariifolia (H. and \.) Rydh. var. inoorei Welsh, Sphaeralcea leptophijUa (Gray) Rydb.
var. jancac Welsh, and Sphacnilcca f)sorak>idcs Welsh.
This third paper in the .series leading to a of specimens examined by me is indicated
revision of the flora of Utah deals with the following the discussion of each species. The
small but significant and taxonomically diffi- number in parenthesis is the number collect-
cult Mallow family. Especially complex are ed by me.
members of the genus Sphaeralcea, ably mon-
ographed by T. H. Kearney (1935), and re- Malvaceae Juss.
viewed for Utah by J. A. M. Jefferies (1972). Mallow Familv
As with previous treatments, the work con-
siders not only indigenous species and weeds Herbs or, less commonly, shrubs, usually
or established escaped species, but those in- pubescent with branched or stellate hairs, an-
troduced species which are commonly grown nual, biennial, or perennial, with mucila-
as ornamentals or for other uses. Casually ginous juice; leaves alternate, simple, mostly
grown species, such as the okra. Hibiscus es- palmately veined, stipulate; flowers perfect
culentus L., have been excluded. Althaea, Hi- (or imperfect), regular, solitary or in thyrsoid
hiscus, and Malta are included entirely on cymes, or more or less racemose or pan-
the basis of cultivated ornamentals and weeds iculate, sometimes with an involucel of sepal-
which have become established in the state. like bractlets; sepals 5, more or less per-
Malva neglecta is a pest of cultivated areas. sistent; petals 5, separate, adnate to the
Iliamna, Malvastrum, Sida, Sidalcea, and staminal sheath; .stamens numerous, united by
Spliaeralcea are represented entirely by in- the filaments (monadelphous); ovary superior,
digenous species. Abutilon has one species in- 3- to many-loculed; fruit a capsule or a schiz-
troduced and the other native. The number ocarp.
1. Involucel lacking 2
— Involucel of 1 or more bractlets, or if lacking (as in some Spliaeralcea
specimens), then the flowers orange (grenadine) 3
2(1). Petals white, pink, or lavender; plants of moist sites, usually at middle and
higher elevations Sidalcea
— Petals yellow or pink to red; plants of cultivated lands or of arid sites, u.sually
at lower elevations Abutilon
3(1). Petals orange or rarely purplish pink; indigenous perennial herbs of arid
habitats at middle and lower elevations Spliaeralcea
— Petals variously colored, but not orange; indigenous or adventive perennial,
biennial, or annual plants or various distribution 4
4(3). Flowers ro.se pink or rarely white; plants indigenous, 7-15 dm tall, perennial,
of middle and higher elevations Iliamna
'Life Science Museum and Department of Botany and Range Science, Brigham Young University. Provo. Utah 84602.
27
28 Great Basin Naturalist Vol. 40, No. 1
_ Flowers white, pink, rose, yellow, or other hues; plants differing in one or
more ways from above '^
5(4) Flowers mostly 6-10 cm broad, opening flat; plants tall adventive or cultivated
biennials Althaea
— Flowers less than 6 cm broad or, if broader then the plants shrubby 6
6(5). Style branches 5, elongate; fruit a capsule; plants low annuals or shrubs
Hibiscus
— Style branches more than 5, short; fruit a schizocarp; plants annual or biennial
7(6). Style branches filiform, with elongate stigmatic lines; plants annual or biennial
Malta
— Style branches with capitate or truncate stigmas 8
8(7). Petals yellow, or orange to pink or red; plants annual with awned carpels or
subshrubs with unawned carpels Abutilon
— Petals yellowish white to lavender or whitish; carpels few to many, not awned;
plants spreading annuals or herbaceous perennials 9
9(8). Petals yellow white; leaves reniform-orbicular, merely crenate-serrate Sida
— Petals lavender or whitish; leaves palmately cleft, with rounded lobes
Malvastrum
.\butilon Mill.
leafy panicles; involucel lacking; calyx 5-
Plants herbaceous, annual or perennial, cleft; corolla yellow to orange pink or red;
with stellate or simple hairs; leaves alternate, fruit truncate-cylindric or subglobose, the
petioled, cordate at base, not or only obscu- carpels smooth sided, dehiscent nearly to the
rely lobed; flowers solitary and axillary or in base; ovules 2 or more per carpel.
Plants perennial, with slender spreading or trailing branchlets; carpels 5, lacking
awn-beaks; plants rare, known only from Washington County A. parvidum
Plants annual, with robust erect stems; carpels usually more than 10, each with
a long divergent awn; plants uncommon, in agricultural regions A. theophrasti
Abutilon parvulum Gray. Perennial, the
stems slender and spreading or trailing, the
caudex woody, grayish tomentose with min-
ute stellate hairs, the branchlets pilose; leaves
0.5-5 cm long, ovate, cordate basally, den-
tate and sometimes obscurely 3-lobed; pe-
duncles slender, axillary, 1 -flowered, longer
than the leaves; calyx lobes ovate-acuminate,
reflexed in fruit; petals orange pink to red or
sometimes yellowish, 4-6 mm long; carpels 5,
somewhat tomentose, to 8 mm long. Known
in Utah only from Veyo, Washington County
(Meyer 4111), Colorado to California, and
south to Texas and Mexico, 1(0).
Abutilon theophrasti Medic. Velvet leaf.
Annual, the stems robast, erect, velvety and
cinereous with short, soft hairs; leaves 3-10
cm long (from sinus to apex) and as broad or
broader, orbicular-ovate, cordate at the base,
abruptly acuminate at the apex, velvety pub-
escent; peduncles shorter than the leaves, one
to few flowered; calyx lobes broadly ovate-
acuminate; petals yellow, to about 6 mm
long; carpels 10 or more, each with a long di-
vergent awn. Adventive weedy species of dis-
turbed or cultivated areas, occasional in Utah
and Washington counties (to be expected
elsewhere); widespread in North America;
native to Europe; 3(0).
Althaea L.
Plants herbaceous, biennial, with coarse
stellate hairs; leaves alternate, petiolate.
March 1980
Welsh: Utah Flora, Malvaceae
29
cordate at the base, lobed; flowers sohtarv or
in racemes; invohicel of 6-9 bractlets, con-
nate at the base; calyx 5-cleft; corolla of vari-
ous colors; fruit flattened wheellike, invested
by the calyx, the numerous carpels separating
at maturity.
Althaea rosea Cav. Hollyhock. Coarse
biennials to 20 dm tall or more, the stems
erect, stellate-hairy; leaves (3-) 5- to 7-lobed,
mostly 3-15 cm long (from sinus to apex) and
often much broader; flowers shortly pedicel-
late, 6-12 cm wide or more, variously col-
ored, often rose to pink or lavender, or some-
times white, usuallv with a dark center; calyx
lobes triangular, investing the fruit at matu-
rity, the involucel calyxlike; carpels numer-
ous, stellate along the margins, and reticulate
on the sides, 5-7 mm long. Cultivated orna-
mental, persisting and escaping, to be ex-
pected in all counties in Utah; widespread in
North America; introduced from China;
15(0).
Hibiscus L.
Plants herbaceous or woody, annual or
perennial, with stellate or simple hairs; leaves
alternate, petiolate, obtuse to truncate or
cordate basally, lobed to incised; flowers ax-
illary, solitary; involucel of 5-10 distinct
bractlets; calyx 5-cleft, more or less accres-
cent in fruit; fruit a loculicidal capsule, the
carpels 5; seeds several in each locule.
1. Plants annual; calyx strongly veined; petals cream colored, with a purple center
H. trionum
Plants shrubs; calyx herbaceous, not distinctly veined; petals variously colored,
but usually rose pink to lavender H. si/riacus
Hibiscus syriacus L. Althaea; Rose-of-Sha-
ron. Shrubs, 20-40 dm tall or more, glabrous
or softly stellate-hairv; leaves 2.5-8 cm long,
1.5-6 cm wide, triangular-ovate to rhombic,
strongly 3-ribbed, commonly 3-lobed; flowers
axillary, 4-7.5 cm wide; bractlets usually 5,
linear, about as long as the calyx, glabrous to
obscurely hairy; corolla variously colored and
often double; fruit oblong-ovoid, to 25 mm
long. Cultivated ornamental, rarely per-
sisting; widely cultivated in North America;
introduced from eastern Asia; 3(i).
Hibiscus trionum L. Flower-of-an-Hour.
Annual, commonly 1.5-5 dm tall, the lower
branches often prostrate, coarsely hispid-stel-
late to glabrate; leaves 3-lobed or more com-
monly 3- to 5-parted, the main lobes cuneate
basally, the middle lobe the largest; flowers
solitary, axillary, mostly 3-6 cm wide; bract-
lets usually 10, linear, often coarsely hispid,
much shorter than the fruiting calyx; corolla
cream colored to yellowish, with a purple
Ltnter, closing in shade. Weedy species of
Lultivated land at lower elevations; wide-
spread in North America; adventive from
:entral Africa; 8(i).
Iliamna Greene
Plants herbaceous, perennial, sparingly and
minutely stellate-hairy; leaves alternate, pe-
tiolate, cordate to truncate basally, the mar-
gin lobed; flowers in thyrsoid panicles; in-
volucel of 3 narrow, persistent bractlets;
calyx 5-cleft; fruit a loculicidal capsule, the
carpels many; seeds usually 3 in each locule.
Wiggins, I. L. 1936. A resurrection and revi-
sion of the genus lUamna Greene. Contr.
Dudley Herb. 1: 213-229.
Iliamna rivularis (Dougl.) Greene. Wild
Hollyhock. {Malta rivularis Dougl. ex.
Hook.; Sphaeralcea rivularis (Dougl.) Torr.
ex. Gray; Phymosia rivularis (Dougl.) Rydb.).
Perennial, the stems few to many from a
woody caudex, mostly 7-15 dm tall, minutely
stellate-puberulent, green; leaves 3- to 7-
lobed, cordate to truncate basally, 2.5-15 cm
long (from petiole apex to tip), 2-16 cm
broad, the lobes triangular, crenate-serrate,
finely stellate; pedicels mostly less than 1 cm
long; bractlets linear-lanceolate, shorter than
the calyx; calyx lobes 3-5 mm long (to 8 mm
30
Great Basin Naturalist
Vol. 40, No. 1
long in fruit); petals rose pink (rarely white),
20-37 mm long; carpels 6-10 mm long in
fniit, hispid and stellate. Along streams, on
foothills, in mountain bnish, ponderosa pine,
aspen, and spruce-fir communities,
1440-2900 m elevation, in Daggett, Davis,
Duchesne, Iron, Juab, Piute, Salt Lake, San-
pete, Sevier, Summit, Tooele, Utah, Wasatch,
and Weber counties; Colorado, Idaho, Ne-
vada, and Washington; 40(vi).
Malva L.
Plants herbaceous, annual, biennial or per-
ennial, from taproots, the pubescence simple
to branched or stellate; leaves alternate, pe-
tiolate, usually more or less cordate basally,
commonly lobed; flowers in axillary clusters
(sometimes solitary) or in subterminal pan-
icles; involucel of 3 narrow to broad per-
sistent bractlets; calyx 5-cleft; fruit a schiz-
ocarp, the carpels mostly 10-15.
1. Petals commonly 1.5-2 cm long; bractlets of involucel ovate to oblong . M. sylvestris
Petals asually less than 1 cm long; bractlets of involucel linear to narrowly
lanceolate 2
2(1). Stems prostrate spreading from the caudex; leaves obscurely lobed; plant a
common weedy species M. neglecta
Stems erect; leaves definitely lobed; plant cultivated, rarely escaping
M. verticillata
Malva neglecta Wallr. Cheeses; Mallow.
Annual or biennial, the stems prostrate-
spreading, commonly 1-6 dm long, stellate-
hairy; leaf blades reniform-orbicular, 0.6-3
cm long (from sinus to apex) or more, and
much broader, crenate and not at all to only
shallowly 5- to 7-lobed, the petioles to 20 cm
long or more; flowers clustered (or solitary)
in the axils; bractlets linear; calyx (3) 4-6 mm
long at anthesis, the lobes acuminate; petals
white to pink or lilac, about twice as long as
the sepals; carpels hairy, roimded on the
back. Weeds of disturbed sites and cultivated
land, in much of Utah (specimens known
from Cache, Iron, Kane, Salt Lake, San Juan,
Summit, Utah, and Washington counties);
widespread in North America; adventive
from Eurasia; 22(ii). Note: Two other species,
M. parviflora L. and M. rotundifolia L.,
might be present in Utah. They are similar to
M. neglecta but have petals subequal to the
sepals. Malva parviflora has glabrous petal
claws, whereas in M. rotundifolia the claws
are bearded.
Malva sylvestris L. High Mallow. Biennial,
the stems ascending, mostly 3-10 dm tall,
rough hairy to glabrate; leaf blades 3-8 cm
long or more and often broader, orbicular to
cordate or reniform, crenate and with 5-7
lobes, the petioles to 10 cm long or more;
flowers clustered in the leaf axils; bractlets
ovate to elliptic; calyx 5-7 mm long at an-
thesis, the lobes short and broad; petals
15-20 mm long, rose purple; carpels glabrous
or nearly so, sharp edged. Cultivated orna-
mental, rarely escaping (Utah Co., Larsen
7152 BRY); widespread in North America;
adventive from Europe; 1(0).
Malva verticillata L. Curled Mallow. An-
nual, the stems erect, mostly to 10 dm tall or
more, sparingly stellate-hairy; leaf blades
mostly 1.5-7 cm long and as broad or broad-
er, orbicular to reniform, undulate-crisped
and distinctly 5- to 7-lobed, long-petioled;
flowers solitary or clustered, subsessile or
some pediceled; bractlets linear to narrowly
lanceolate; calyx 3.5-5 mm long, the lobes
acuminate; petals white, only somewhat sur-
passing the sepals; carpels glabrous, the edges
rounded. Cultivated ornamental, rarely es-
caping (Washington Co., Galway in 1934
BRY); widely scattered in the United States;
adventive from the Old World; 1(0). Our ma-
terial belongs to var. crispa L.
Malvastrum Gray
Plants herbaceous, annual, stellate-hairy;
leaves alternate, petiolate, the blades sub-
cordate to tmncate basally, palmately lobed;
flowers solitary in the axils or in terminal
bracted clusters; involucel of usually 3 slen-
March 1980
Welsh: Utah Flora, Malvaceae
31
der bractlets; calyx 5-cleft, the lobes long-
acuminate; carpels 10-15; fruit a schizocarp.
Malvastrum exile Gray. {Malveopsis exile
(Gray Kuntze; Eremalche exile (Gray)
Greene; Sphaeralcea exile (Gray) Jepson). An-
nual, the stems spreading-decumbent to pro-
strate, branching from near the base, 0.3-4
dni long, rather sparingly stellate-hairy; leaf
blades suborbicular, 0.8-3.2 cm wide, palma-
tely 3- to 5-cleft, with rounded or cuspidate
teeth; petioles 1-5 cm long; bractlets narrow-
ly lanceolate to sublinear; calyx 3-5 mm
long; petals whitish to pinkish or lavender,
only somewhat surpassing the sepals; carpels
transversely wrinkled. Open sites in black-
brush and creosote brush communities,
850-1200 m elevation, in Garfield (report
probably erroneous) and Washington coun-
ties; Arizona and southern California; 6(0).
SiDA L.
Plants herbaceous, perennial, from spread-
ing rhizomes, densely stellate-canescent;
leaves alternate, petiolate, crenate-serrate,
not or obscurely linear, deciduous bractlets;
calyx 5-lobed; carpels 5-10, 1 -seeded; fruit a
schizocarp.
Sida hederacea (Dougl.) Torr. Alkali-Mal-
low. {Malva hederacea Dougl.; A4. californica
Presl.; Disella hederacea (Dougl.) Greene).
Perennial, the stems from elongate rhizomes,
decumbent to prostrate, the surface obscured
by overlapping stellate hairs, 1-4 din long;
leaf blades reniform to orbicular, often
oblique, dentate, obscurely if at all lobed, the
petioles 0.3-2.5 (3) cm long; bractlets sub-
linear; calyx 5-7 mm long; petals yellowish
(fading orange), 10-12 mm long; carpels reti-
culate on the sides. Saline meadows and
seeps, at lower elevations in Emery, Salt
Lake, Tooele, Uintah, and Utah counties (and
probably elsewhere); Washington south to
California, Texas, and Mexico; 6(i).
SiDALCEA Gray
Plants herbaceous, perennial, from tap-
roots or short rhizomes, usually stellate and
somewhat hirsute; leaves alternate, petiolate
often dimorphic, the lowermost merely pal-
mately lobed, the upper ones commonly cleft
and with linear lobes; flowers borne in semi-
spicate racemes, of two types, those of plants
with perfect flowers the largest; involucel
lacking; calyx 5-cleft; carpels 5-10, 1-seeded,
tardily separating.
Hitchcock, C. L. 1957. A study of the per-
ennial species of Sidalcea. Univ. Wash.
Publ. Biol. 18: 1-79.
Roush, E. M. F. 1931. A monograph of the
genus Sidalcea. Ann. Mo. Bot. Card. 18:
117-244.
1. Petals white or merely pinkish-tinged, often drying yellow; anthers bluish pink;
plants rhizomatous; stems hirsute below S. Candida
Petals pink to lavender; anthers usually yellow to white; plants rhizomatous or
not; stems hirsute to glabrous or tomentose below 2
2(1). Plants from rather fleshy taproots; stems commonly hirsute below; calyx
hirsute with pustulose hairs (at least in part) S. neomexicana
- Plants often rhizomatous; stems stellate to glabrous below; calyx seldom with
pustulose hairs S. oregana
Sidalcea Candida Gray. Plants from slen-
der rhizomes, the stems 4-10 dm tall,
glabrous to hirsute with simple hairs below,
more or less stellate above; leaf blades 6-20
cm wide, the basal ones shallowly 5- to 7-
lobed and coarsely crenate, the upper ones
divided into 3-5 entire segments; calyx 7-10
mm long, variously stellate-hairy and glandu-
lar puberulent; petals white to pinkish, often
drying yellow, 12-20 mm long; carpels about
3 mm long. Stream banks, lake shores, and
seeps, 1410-2750 m, in Beaver, Garfield,
Grand, Iron, Millard, Piute, Salt Lake, San
Juan, Sevier, Summit, Uintah, Utah, and
Wasatch counties; Wyoming and Colorado
west to Nevada and south to New Mexico.
Our materials have been treated as belonging
to two more or less and at least partially sym-
patric varieties; 25(vi).
32
1.
Great Basin Naturalist
Vol. 40, No. 1
Calyx rather uniformly hairy from base to apex of the lobes; plants of wide
distribution S. Candida var. Candida
Calyx more hairy at the base than on the lobes, the lobes often subglabrous;
plants mostly from mountainous portions of middle Utah
S. Candida var. glahrata
Var. Candida. (S. Candida var. tincta Cock-
erell). Known from Beaver, Grand, Iron, Salt
Lake, San Juan, Summit, and Wasatch coun-
ties; Colorado, New Mexico.
Var. glahrata C. L. Hitchc. Known from
Iron, Millard, Piute, Salt Lake, Sevier, Sum-
mit, and Uintah counties; Wyoming, Colo-
rado, and Nevada.
Sidalcea neomexicana Gray. Plants from
enlarged taproots or fascicled roots, the stems
2-9 (10) dm tall, hirsute below (or rarely
glabrous) with simple or bifurcate hairs; leaf
blades L5-11 cm wide, the basal ones cre-
nate to shallowly 5- to 7-lobed, the cauline
ones divided usually into 5 laciniate to entire
segments; calyx 5-10 mm long, usually with
some simple pustulose hairs interspersed with
stellate ones; petals rose pink (fading blue-
purple), 11-19 mm long; carpels 2-3 mm
long. Wet Meadows, stream banks, and seeps,
at 1370 to 2150 m in Box Elder, Garfield,
Juab, Piute, Salt Lake, Sanpete, Sevier, Sum-
mit, Utah, and Wasatch counties; Oregon,
Idaho, and Wyoming south to California,
Arizona, and Mexico.
Hairs of lower stem nearly all simple; calyx coarsely and rather densely hirsute
to coarsely hairy, lacking appressed stellate hairs; upper stems usually
glabrous S. neomexicana var. neomexicana
Hairs of lower stem often forked; calyx often with fine appressed stellate hairs
in addition to the coarse ones; upper stems often stellate hairy
S. neomexicana var. crenulata
Var. crenulata (A. Nels.) C. L. Hitchc. (S.
crenulata A. Nels., type from Juab, Utah; S.
neomexicana ssp. crenulata (A. Nels.) C. L.
Hitchc). Known from Box Elder, Juab, Salt
Lake, Sanpete, Sevier, Summit, Utah, and
Wasatch counties; Oregon, Idaho, and Ne-
vada; lO(ii).
Var. neomexicana. Known from Box Elder,
Garfield, Piute, San Juan, Sevier, Utah, and
Wasatch counties; Wyoming, Colorado, Ari-
zona, and Mew Mexico; Mexico; 12(ii).
Sidalcea oregana (Nutt.) Gray. {Sida ore-
gana Nutt. ex T. & G.; S. nervata A. Nels.).
Plants from a taproot, lacking or rarely with
rhizomes, the stems 3-11 dm tall or more,
glabrous or usually appressed-stellate hairy
below, appressed-stellate above; leaf blades
2.5-17 cm wide, the basal ones shallowly 5-to
7-lobed and coarsely crenate, the cauline
ones deeply lobed, with 3-7 coarsely toothed
to entire lobes; calyx 3.5-9 mm long, various-
ly stellate-hairy and sometimes bristly; petals
7-23 mm long, pale pink to rose pink (fading
blue purple); carpels 2.5-3 mm long. Mead-
ows, stream banks, and open woods, at 1680
to 2750 m in Cache, Juab, Salt Lake, San-
pete, Summit, Utah, Wasatch, and Weber
counties; Washington and Idaho south to
California, Nevada, and Utah. Our materials
belong to var. oregana; 32 (ii).
Sphaeralcea St. Hil.
Plants herbaceous, perennial, from tap-
roots or rhizomes, glabrescent to canescent
with stellate hairs; leaves alternate, petiolate,
sometimes dimorphic, the lowermost merely
toothed or palmately lobed (rarely entire),
the upper ones cleft to entire; flowers borne
in racemose to thyrsoid cymes; involucel of 3
or fewer filiform bractlets; calyx 5-cleft; car-
pels 8-20, the seeds 1 or 2 per carpel; fruit a
schizocarp, the mature fruit segments divided
into a basal indehiscent, reticulate portion
and an apical dehiscent portion.
March 1980 Welsh: Utah Flora, Malvaceae 33
Jefferies, J. A. M. 1972. A revision of the Kearney, T. H. 1935. North American species
genus Sphaeralcea (Malvaceae) for the of Sphaemlcea, Subgenus Eusphaeralcea.
state of Utah. UnpubHshed thesis. Brigham Univ. Calif. Fubl. Bot. 19(1): 1-102.
Young University. 92 pp.
1. Inflorescence racemose, rarely with more than one flower per node or, if more,
as in S. caespitosa, then the plants restricted to Millard County 2
— Inflorescence thyrsoid to thyrsoid-glomerate, with usually more than one
flower per node; distribution various 5
2(1). Leaf blades only slightly, if at all, 3- to 5-lobed, the margins irregularly cre-
nate-dentate; hairs with rays radiating in more than a single plane; plants sel-
dom more than 1.5 dm tall, known only from western Beaver and Millard
counties S. caespitosa
— Leaf blades distinctly 3- to 5-lobed, -parted, or -divided; hairs of rays radiating
in a single plane (except in S. coccinea); plants often 1.5 dm tall or more, of
different distribution 3
3(2). Leaves trifoliolate, the leaflets linear to narrowly oblanceolate and entire, or
the upper ones simple and entire; plants of southeastern Utah S. leptophylla
— Leaves various, but if trifoliolate then the leaflets oblanceolate and entire to
toothed, if the uppermost simple then toothed or lobed; distribution various 4
4(3). Lowermost leaves simple and entire or trifoliolate, or some broadlv toothed or
lobed; involucel present; rays of hairs radiating in one plane; plants of eastern
Wayne County S. psoraloides
— Lowermost leaves usually 3- to 5-lobed, the lobes usually toothed or again
lobed; involucel present or lacking (caducous); rays of hairs radiating in several
planes; plants of broad distribution S. coccinea
5(1). Plants only sparingly pubescent, the herbage bright green 6
— Plants moderately to densely pubescent, the herbage yellowish, whitish, or
grayish 8
6(4). Leaves 3- to 5-parted or -divided, the lobes with narrow, regularly pinnatifid
margins, the teeth at nearly right angles to the vein; carpels often with
transparent lacunae, 4-6 mm high; plants rare, of southern Utah only S. rusbiji
— Leaves variously lobed, divided, or parted, the lobes with broader margins ir-
regularly toothed or lobed, but not as above; carpels with opaque lacunae,
3-4.5 mm high 7
7(6). Leaves slightly lobed, the margins unevenly toothed or, in .some, deeply parted
to divided with the margin coarsely and irregularly lobed, the base subcordate
to cuneate; plants of northern Utah S. munroana
— Leaves 3- to 5-parted or -divided, the margins regularly cleft, lobed, or
toothed, the base subcordate to deeply cordate; plants mostly of southern Utah
S. grossulariifolia
8(5). Inflorescence loosely thyrsoid (appearing paniculate), leafy; flowers not numer-
ous at each node; peduncles generally elongate; calyx surpassing the fruit; car-
pels with reticulae extending onto back of carpel; plants of southwestern Utah .
S. ambigua
— Inflorescence contracted thyrsoid-glomerate; flowers often numerous at each
node, not especially leafy; calyx often shorter than the fruit; carpels with
reticulae confined to lateral face of carpel; plants of various distribution 9
34
Great Basin Naturalist
Vol. 40, No. 1
9(8). Leaves 3- to 5-cleft, -parted, or -divided; carpels with well-defined reticiilae on
less than half of carpel face; plants of all but the northeastern one-fourth of
\JlJ^y^ S. grossiilariifolia
Leaves shallowly 3- to 5-lobed; carpels with well-defined to nearly obscure re-
ticulae on the lower one-third of the carpel; plants mainly of eastern and
southern Utah, scattered elsewhere S. parvifolia
Sphaeralcea ambigua Gray. Stems arising
from a woodv caudex, several to numerous,
.3-10 dm tall, whitish to yellowish canescent;
leaf blades 1-6 cm long (from sinus to apex),
0.8-5 cm wide, thickish, usually rugose, with
veins prominent beneath, ovoid, deltoid, or
nearlv orbicular, the base cordate to deeply
cordate, obscurely to definitely 3- to 5-lobed,
the lobes crenate; inflorescence an open pan-
icle, sometimes narrowly thyrsoid; pedicels
usually shorter than the calyx; calyx uni-
formly pubescent to glabrate, 6-20 mm long
at anthesis, the lobes lanceolate to acumi-
nate; petals 15-22 mm long, orange to or-
ange pink (fading pinkish); carpels 12-16 mm
high, the indehiscent portion comprising
about one-third of the carpel, prominently
reticulate. Creosote bush-blackbrush and
mixed warm desert shrub communities,
670-1070 m, in Washington Co.; Nevada,
Arizona, and California; and Mexico. Our
material belongs to var. ambigua; 10(i).
Sphaeralcea caespitosa M. E. Jones. Jones
Globemallow. Stems solitary or more com-
monly few to several from the summit of a
branching woody caudex, 0.2-2.5 dm tall,
whitish to grayish canescent; leaf blades
1.2-5.5 cm long, 1.2-6 cm wide, thickish, not
nigose, veins apparent but not especially
prominent, ovate to deltoid or orbicular, the
base tnmcate to obtuse, obscurely if at all
lobed, crenate to crenate-dentate; in-
florescence thyrsoid, the flowers tightly clus-
tered or solitary; pedicels shorter than the ca-
lyx; calyx uniformly stellate, the rays of hairs
not radiating in a single plane, the lobes
lance-acuminate; petals 15-21 mm long, or-
ange; carpels 12-14, 4-6 mm high, the in-
dehiscent portion forming slightly more than
one-third of the carpel, reticulate on the
sides. Mixed desert shrub communities (shad-
scale, rabbitbmsh, winterfat), mainly on Sevy
Dolomite formation, at 1370-1750 m, in Mil-
lard and Beaver counties; endemic; 20(iii).
Sphaeralcea coccinea (Nutt.) Rydb. Com-
mon Globemallow. {Malva coccinea Nutt.;
Cristaria coccinea (Nutt.) Pursh; Sida coc-
cinea (Nutt.) DC; Malvastrum coccineum
(Nutt.) Gray.; Sida dissecta Nutt.; M. c. var.
dissectum (Nutt.) Gray; M. dissectum (Nutt.)
Cockerell; S. dissecta (Nutt.) Rydb.; S. coc-
cinea ssp. dissecta (Nutt.) Kearney; S. coc-
cinea var. dissecta (Nutt.) Kearney; M. c. var.
elatum Baker; M. elatiim (Baker) A. Nels.; S.
elata (Baker) Rydb.; S. c. ssp. elata (Baker)
Kearney; S. c. var. elata (Baker) Kearney; M.
cockerellii A. Nels.; M. micranthiim W. & S.).
Stems solitary or few to many from the apex
to a woody caudex, or less commonly from
creeping rhizomes, 0.6-4.2 dm tall, white to
yellowish canescent; leaf blades 1.1-3.7 cm
long, 1.2-5.2 cm wide, usually wider than
long, ovate to cordate-ovate in outline, the
base often cordate, usually 3-to 5-lobed, with
main divisions cleft almost or quite to the
base, the lobes usually again toothed or
lobed; inflorescence racemose, sometimes
paniculate, rarely thyrsoid; pedicels shorter
than the calyx; calyx uniformly stellate, the
rays or hairs not radiating in a single plane,
the lobes lance-acuminate; petals 8-15 mm
long, orange; carpels 8-14, 2-3 mm high, the
indehiscent part forming two-thirds or more
of the carpel, reticulate on the sides and on
the back. Blackbrush, shadscale-greasewood,
sagebRish, juniper-pinyon, mountain brush,
and ponderosa pine communities, 920-2750
m, in all counties (except Morgan and
Wasatch?); Saskatchewan and Alberta south
to Arizona, New Mexico, and Texas. Our ma-
terials have been recognized as belonging to
vars. dissecta and elata, but the segregation
of these entities appears to have been wholly
arbitrary, with intermediates more numerous
than the supposed taxa; 152(xviii).
Sphaeralcea grossulariifolia (H. & A.)
Rydb. Gooseberry-Leaved Globemallow.
{Sicki grossulariifolia H. & A.; Malvastrum
March 1980
Welsh: Utah Flora, Malvaceae
35
grossulariifoUwn (H. & A.) Gray; S. pedata
Torr., in Gray; S. g. ssp. pedata (Torr.)
Kearney; S. g. var. pedata (Torr.) Kearney).
Stems few to many from a woody caiidex,
1-10 dm tall or more, whitish to yellowish
canescent to subglabrous and green; leaf
blades 1.3-5 cm long, 1.3-5 cm wide, usually
longer than wide, ovate to cordate-ovate in
outline, the base cordate to truncate or ob-
tuse, usually 3- to 5-lobed, the main division
asually cleft or parted to irregularly toothed;
inflorescence thyrsoid, with usually more
tlian one flower per node; pedicels shorter
than to much longer than the calyx; calyx
luiiformly stellate, the rays of hairs not radi-
ating in a single plane, the lobes ovate to
lance- acuminate; petals 8-20 mm long, or-
ange or rarely rose pink; carpels 10-14,
2.5-4.5 mm high, the indehiscent portion
forming from two-fifths to three-fifths of the
carpel, reticulate on the sides. Blackbrush,
shadscale, rabbitbrush, sagebrush, juniper-
pinyon, and less commonly mountain brush
communities, 920-2450 m, in Beaver, Box El-
der, Cache, Emery, Garfield, Grand, Iron,
Kane, Juab, Millard, Morgan, Piute, Salt
Lake, San Juan, Sanpete, Sevier, Tooele,
Utah, Wasatch, Washington, and Wayne
counties; Washington, Oregon, Nevada, Cali-
fornia, and Arizona. Two infraspecific taxa
have been segregated, largely on the basis of
form of the leaf blades. Intergradation of the
phases seems to be complete. Further, S.
grossulariifolia appears to form intermediates
with S. coccinea, S. parvifolia, and the more
northern S. munroana. A phase with green
herbage and thin leaves occurs along Glen
Canyon. It seems to represent a taxonomic
imit worthy of recognition.
1. Herbage bright green; leaves thin-textured; plants of eastern Kane and
Garfield, and western San Juan counties S. grossulariifolia var. moorei
Herbage gray green to whitish canescent; leaves thick-textured; plants
widespread S. grossulariifolia var. grossulariifolia
Var. grossulariifolia. This is the common
and widely distributed phase of the species in
Utah. The report by Kearney (I.e., p. 90) of S.
digitata (Greene) Rydb. apparently belongs
here; 115(xii).
Var. moorei Welsh var. nov. Plantae siniilis
var. grossulariifoliae sed differt in folii et
caules virides et folii tenues. Holotype: Kane
County, Utah, east side of Last Chance Bay,
Lake Powell, Entrada Sandstone, S. L. Welsh
and N. D. Atwood 11597, 2 May 1972 (BRY).
Additional specimens: Kane County, mouth
of Escalante River, Lake Powell, S. L. Welsh
and G. Moore 11810, 5 June 1972; do. Wil-
low Tank, D. A. White 111, 4 May 1962; do,
Escalante Canyon, S. L. Welsh and G. Moore
11827, 5 June 1972; do, N. D. Atwood and R.
Allen 3211, 24 August 1971; do, Hole-in-the-
Rock, B. F. Harrison 12112, 14 May 1953;
San Juan Co., 1 mi. E of Hole-in-the-Rock, S.
L. Welsh and C. A. Toft 11869, 16 June
1972; do. Three Garden, Lake Powell, ca 1
mi. N of confluence with San Juan Arm, S. L.
Welsh 12420, 5 May 1974; do. Comb Wash,
S. L. Welsh and N.'D. Atwood 9972, 6 June
1970 (all at BRY). This variety is named to
honor Glen Moore, botanist, teacher, collabo-
rator, and collector.
Sphaeralcea leptophylla (Gray) Rydb,
{Malvastrum leptophyllum Gray). Stems few
to many from a woody caudes, 2.0-5.5 dm
tall, grayish canescent to yellow green
throughout; leaf blades 1.0-3.2 cm long, di-
gitately 3-lobed, the lobes entire, linear to
oblanceolate, 1-4 mm wide, or the upper
leaves simple and linear; inflorescence race-
mose, elongate, usually with one flower per
node; pedicels from much shorter to longer
than the calyx; calyx uniformly stellate, the
rays of hairs radiating in a single plane, the
lobes lance-attenuate; petals 8-12 mm long,
orange; carpels 7-9, 3-3.5 mm high, the in-
dehiscent portion forming two-thirds-three-
fourths of the carpel, coarsely reticulate,
ridged, or tuberculate on the back. Black-
brush and mixed semidesert shrub commu-
nities, 1200-1520 m, in Garfield, Grand, and
San Juan counties; New Mexico, Arizona,
Texas, and Mexico. Two distinctive phases
are recognizable among our materials; they
can be distinguished as follows:
36
1.
Great Basin Naturalist
Vol. 40, No. 1
Plants grayish canescent, the hairs obscuring the surface of stems, leaves, and
calvx lobes; leaf lobes narrowly oblong to linear; distribution as for the species
S. leptophylla var. leptophylla
Plants green to yellow green, the hairs widely separated, not obscuring stems,
leaves or calyx lobes; leaf lobes, at least of lowermost leaves, oblanceolate to
spatulate- known only from San Juan County, Utah S. leptophyUa var. janeae
Var. leptophylla. This is the common form
of the species. It is known from Garfield,
Grand, and San Juan counties, Utah, and
from New Mexico and Arizona; 7(iii).
Var. janeae Welsh var. nov. Plantae similis
var. leptophylhi sed differt in folii caules et
calvces virides et lobos foliorum inferiorum
oblanceolatos vel spathulatos. Holotype: San
Juan County, Utah, along White Rim road,
north of Turks Head, on sandy slopes in
blackbnish communitv, Canyonlands Nation-
al Park, S. L. Welsh 7064, 17 May 1968
(BRY). This variety is named to honor Jane
.\rdis Murray Jefferies, student of Sphae-
ralcea in Utah.
Sphaeralcea miinroana (Dougl.) Spach in
Gray. Munroe Cdobemallow. {Malta miin-
roana Dougl. in Lindl.; Nuttallia miinroana
(Dougl.) Nutt.; Malvastnirn munrooniim
(Dougl.) Gray; S. suhrhomhoidea Rydb.; S. m.
ssp. suhrhomhoidea (Rydb.) Kearney; S. m.
var. s. (Rydb.) Kearney). Stems several to
many from a branching woody caudex, 1.8-7
dm tall or more, yellowish green to some-
what grayish canescent, the foliage usually
bright green; leaf blades 1-6 cm long, 0.8-6
cm wide, ovate to orbicular or rhombic in
outline, the base truncate to obtuse or sub-
cuneate, usually 3- to 5-lobed, the sinuses
shallow to very deep, the main divisions
merely toothed or the lateral ones incised; in-
florescence narrowly thyrsoid, usually with
more than one flower per node; pedicels usu-
ally much shorter than the calyx; calyx uni-
formly stellate, the rays of hairs not radiating
in a .single plane, the lobes deltoid-ovate to
ovate; petals 8-15 mm long, orange; carpels
10-1.3, 2.5-3 mm high, the indehiscent por-
tion forming about half the carpel, reticulate
on the sides. Mixed desert shrub, or more
commonly, in sagebrush and mountain brush
conununities, 1370-2450 m, in Box Elder,
Cache, Duchesne, Emery, Summit, Tooele,
Uintah, Utah, and Wasatch counties; Mon-
tana, Idaho, Washington, Wyoming, Nevada,
and California. This entity is much like both
S. parvifoUa and S. grossulariifolia. The green
color of herbage is diagnostic of S. miinroana
from both, except for the var. moorei which is
not sympatric with S. miinroana; 21(ii).
Sphaeralcea parvifoUa A. Nels. Nelson
Globemallow. (S. marginata York, ex Rydb.;
S. arizonica Heller, ex Rydb.). Stems few to
many from a branching woody caudex,
1.5-10 (11) dm tall, grayish canescent, the fo-
liage gray green or only somewhat yellow
green; leaf blades 1.0-5.5 cm long, 1.2-5.2
cm wide, ovate to orbicular, reniform, or
cordate-ovate, the base cordate to truncate or
obtuse, usually shallowly 3- to 5-lobed, the si-
nuses usually shallow, the lobes crenate-den-
tate; inflorescence commonly narrowly thyr-
soid, usually with more than one flower per
node; pedicels usually shorter than the calyx;
calyx uniformly stellate, the rays of hairs not
radiating in a single plane, the lobes lance-
ovate to deltoid; petals 7-15 mm long, or-
ange; carpels 10-12, 3-4 mm high, the in-
dehiscent part forming from one-fourth to
one-third of the carpel, faintly reticulate on
the sides. Blackbrush, salt desert shrub, sage-
liRish, pinyon-jiuiiper, and mountain brush
communities, at 850 to 2700 m, in Box Elder,
Cache and Tooele counties, where probably
of recent introduction, and in Duchesne,
Emery, Garfield, Grand, Iron, Kane, Piute,
San Juan, Sevier, Tooele, Washington, and
Wayne counties, where likely indigenous;
Nevada, Arizona, New Mexico, and Califor-
nia. Sphaeralcea parvifoUa has been com-
pared by Kearney (I.e.) with S. amhigua,
which it resembles. The relationship of S.
parvifoUa in Utah seems to lie with the
largely sympatric S. gro.s.sularii folia; 144(xxii).
Sphaeralcea psoraloides Welsh sp. nov.
Stems few to many from a branching caudex,
1.4-2.4 dm tall or more, sparsely yellowish
canescent, the foliage yellow green; leaf
blades 1.3-3.5 cm long, 0.4-3.8 cm wide, ob-
lanceolate to cuneate-ovate in outline,, cu-
March 1980
Welsh: Utah Flora, Malvaceae
37
neate to obtuse or rounded basally, trifoliol-
ate or simple to 3-lobed below, deeply 3- to
5-cleft above, the lobes entire to few toothed
or lobed, usually more than 5 mm wide; in-
florescence racemose, the flowers solitary in
tlie upper axils; calvx uniformlv stellate, the
rays of hairs radiating in a single plane, the
lobes lance-acuminate; petals 10 (8-12) mm
long, orange; carpels 10 (fruit unknown).
Ephedra-Gmijia commimity on Entrada silts-
tone, 1500 m, in Wavne Countv; endemic.
Plantae similis S. leptophylla sed differt in
foliolos oblanceolata vel laminas super iores
confluentes et lobatos; e S. coccinea laminis
inferioribus simplicibus vel trifoliolati digi-
tatis distinguenda.
Caules pauci vel multi e caudicibus raniifi-
cantibus 1.4-2.4 dm alti vel plures flavidi-ca-
nescentes parce folia et caules luteo-virides;
laminae foliorum 1.3-3.5 cm longae 0.4-3.8 .
cm latae oblanceolata ad cuneati-ovatas cu-
neatae ad obtusas vel rotundatas basaliter tri-
foliolatae vel simplicia ad trilobata infra 3-5
fissa profimde supra lobis intergris ad pauci-
dentatis vel pauci-lobatis plerumque plus
quam 5 mm latis; inflorescentiae racemosae,
flores solitari in axilas supras; calyces stellati
uniformiter, radius pilos radiantibus in plan-
item singularem, lobus calycis lanci-acumi-
natis; petala 10 (8-12) mm longa, aurantiaca;
carpeli 10 (fructus ignotus). Holotvpe:
Wavne County, Utah, Salt Wash, ca 17 mi.
due WMW of Hanksville, T27S, R8E, Sec.
24, at 1500 m, on Entrada siltstone, Gmi/ia-
Ephedra connnunitv, S. L. Welsh 13348, 1
June 1976 (BRY). Paratvpe: do, S. L. Welsh
13345, 1 June 1976 (BRY).
Sphaeralcea riisbyi Gray. Stems few to
many from a caudex, or rarely sub-
rhizomatous, mostly 2-6.5 (8.5) dm tall, yel-
lowish green to somewhat grayish canescent;
leaf blades 1.3-3 cm long, 1.2-4 cm wide,
ovate to orbicular in outline, the base trun-
cate-obtuse to prominently cordate, parted to
divided or merely cleft, the lobes again
toothed (the teeth spreading at nearly right
angles); inflorescence thrysoid to paniculate,
with more than one flower per node; pedicels
usually shorter (to much longer) than the ca-
lyx; bractlets often dark red; calyx imiformly
stellate (more densely so than on the her-
bage), the rays of hairs not radiating in a
single plane, the lobes ovate to lance-ovate;
petals 9-18 mm long, orange; carpels 10-12,
4-6 mm high, the indehiscent part forming
from one-fourth to two-fifths of the carpel,
finely reticulate on the sides. Blackbrush,
creosote brush, and mixed warm desert shrub
communities, 820-1070 m, in Washington
County; Arizona. S. rusbiji forms apparent
intermediates with phases of S. grossularii-
folia and S. pa wi folia; 4(0).
UTAH FLORA: MISCELLANEOUS FAMILIES /
Stanley L. Welsh'
Abstract.— Considered in this treatment are the families Aquifoliaceae, Canabinaceae, Ericaceae, Krameriaceae,
Magnoliaceae, Moraceae, Oleaceae, Pyrolaceae, Resedaceae, Tamaricaceae, and Tiliaceae. These 11 families include
61 cultivated, escaped, and indigenous species.
The flora of Utah is both large and diverse.
A portion of the diversity is due to the pres-
ence of a large number of cultivated species
in many plant families. Floras of regions have
traditionally avoided inclusion of strictly cul-
tivated species. Only those taxa which escape
and become acclimated have been treated.
Included are the cultivated plants and those
species which escape. The present treatment
covers all taxa in common cultivation, and es-
pecially those which are represented in re-
gional herbaria. In Table 1 a list is presented
of the families treated herein, the numbers of
genera and species, and whether cultivated
or indigenous.
The list heavily favors the cultivated
and/or escaped species, and, because of the
status of cultivated species collections, the
treatment is likely to be incomplete. It is
presented herein for use by students of the
flora who want to know the names of culti-
vated and of native plant species.
Aquifoliaceae
Holly Family
Evergreen Shrubs or small trees: leaves al-
ternate, simple, coriaceous, armed with spiny
teeth; stipules minute, caducous; flowers usu-
ally imperfect, regular, small and inconspic-
uous, solitary or few in axillary cymes; sepals
usually 4, more or less connate basally; petals
usually 4, distinct or slightly connate basally;
stamens or staminodes usually 4 (-9), alter-
nate with the petals; pistil 1, the ovary supe-
rior, 3- to many-loculed, the carpels as many
as the locules; fruit a globose, berrylike drupe
with 2-8 bony 1 -seeded divisions.
Ilex L.
Evergreen; leaves thick and shining; flow-
ers small, mostly in few-flowered axillary
cymes; staminodia usually present in pistil-
late flowers, a rudimentary pistil present in
most staminate flowers; fruit usually brightly
colored. (Note: Members of this family are
known in Utah in cultivation only).
Flowers in axillary clusters on branches of the previous year /. oquifolium
Flowers in solitary cymes on branches of the current year /. opaca
Ilex aquifolium L. English Holly. Tall
shnibs to small trees of ornamental plantings,
rare in Utah; introduced from the Old
World; 1(0).
Ilex opaca Art. American Holly. Low to
moderate shrubs of ornamental plantings, oc-
casional in Utah; introduced from the eastern
United States; 1(0).
Cannabinaceae
Hemp Family
Plants herbaceous, with watery juice;
leaves alternate or opposite, palmately
veined and lobed or divided to essentially
compound; stipules persistent; flowers imper-
'Life Science Museum and Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602.
38
March 1980
Welsh: Utah Flora, Miscellaneous Families
39
feet, the plants dioecious, regular, the stami-
nate in open racemes or panicles, the pistil-
late in dense clusters; sepals 5, connate in
pistillate flowers and enclosing the ovary;
stamens 5; pistil 1, 2-carpelled, the styles 2;
fruit an achene.
Plants strong-smelling, stout, erect herbs; leaves palmately 5- to 9-parted .. Cannabis
Plants rough-stemmed clambering vines; leaves coarsely 3- to 7-lobed Humulus
CaNxNabis L.
Plants dioecious or rarely some mon-
oecous; leaves palmately lobed to parted and
apparently compound, alternate or the lower
opposite; flowers small, inconspicuous, the
staminate in leafy panicles in upper axils;
sepals 5, oblong; stamens 5; pistillate flowers
in small clusters on leafy branches from up-
per axils, each flower subtended and enclosed
by an acuminate bract, the calyx barelv
lobed, surrounding only the base of the
ovary; stigmas 2, elongate; fruit a lenticular
achene, enclosed within the accrescent bract.
Small, E. and A. Cronquist. 1976. A practical
and natural taxonomv for Cannabis. Taxon
5: 405-435.
Cannabis sativa L. Hemp, Marijuana,
Hashish, Pot, Grass. Plants 6-20 dm tall or
more, the stems simple or much branched;
leaves long petioled, the blades 3- to 7-part-
ed, the segments oblanceolate to elliptic, at-
tenuate to accuminate apically, sharply ser-
rate, mostly 4-12 cm long and 0.4-1.8 cm
wide, scabrous and more or less glandular
and pubescent; achenes mostly 3.5-4.5 mm
long. Cultivated historically in Utah for fiber
Table 1. Families, genera, and species treated.
No. Genera
No.
Species
Family
Cultivated Indigenous
.Aquifoliaceae
1
2
Canabinaceae
2
1
1
Ericaceae
6
0
11
Krameriaceae
1
0
2
Magnoliaceae
2
4
Moraceae
3
4
Oleaceae
6
14
4
Pyrolaceae
3
0
7
Resedaceae
1
1
Tamaricaceae
1
3
Tiliaceae
1
7
TOTAL
27
36
25
produced from the stems, the commercial
source of hemp; currently sporadic, or else
grown illegally for its intoxicant properties.
Utah materials are sufficiently rare as to give
only hints as to the classification below the
species level. It seems likely, however, that,
at least historically, two phases have been
grown in the state (for a complete review see
Small and Cronquist 1. c). Nineteenth-cen-
tury plantings for hemp likely belonged to
ssp. sativa, demonstrated to have only limited
intoxicant ability. At least some of the recent
introductions clearly belong to ssp. indica
(Lam.) Small & Cronq. (C. indica Lam.),
which has demonstrated high intoxicant lev-
els.
Humulus L.
Plants herbaceous, twining, perennial
vines; stems scabrous; leaves opposite, broad-
ly 3- to 5-lobed; flowers small, inconspicuous,
the staminate in axillary panicles; sepals 5,
distinct; stamens 5; pistillate flowers in short
spikes, in pairs, with each pair subtended by
a foliaceous bract; calyx membraneous, un-
lobed, closely covering the ovary; stigmas 2,
elongate; fruit an achene enclosed by the
persistent calyx and accrescent bracts.
Humulus americanus Nutt. American
Hop. Plants twining, the stems to 20 dm long
or longer; leaves ovate to orbicular in out-
line, deeply cordate ba.sally, mostly 3-15 cm
long and 2.8-16 cm wide, the lobes serrate to
doubly so, attenuate to acimiinate apically,
rough-hairy above, glandular-dotted beneath;
fruiting spikes usually 2-3.5 (4) cm long at
maturity. Twining over shrubs and other veg-
etation at lower and middle elevations in
Duchesne, (iarfield. Grand, Millard, Piute,
Salt Lake, Summit, Uintah, Utah, Wasatch,
Washington, and Weber counties, and prob-
ablv throughout Utah; widespread in North
America. The hop of commerce, H. hipulus
40
Great Basin Naturalist
Vol. 40, No. 1
L., or European hop, is grown in the United
States, where it has escaped and persists.
Though not definitely known for Utah, the
European hop might occur here. It can be
distinguished by its unlobed leaves, or when
lobed, the terminal lobe is less than twice
longer than broad; 27(v).
Ericaceae
Heath Family
Shrubs or subshnibs; leaves simple, alter-
nate, sometimes leathery or persistent; flow-
ers perfect, regular, axillary, in terminal clus-
ters, or solitary; sepals mostly 4 or 5, distinct
or more or less connate; petals mostly 4 or 5,
connate or distinct, the corolla rotate to fun-
nelform or urn shaped; stamens as many as
the corolla lobes and alternate with them or
twice as many, the anthers dehiscent by ter-
minal pores or by longitudinal slits; pistils 1,
the ovary superior or inferior, usually with
4-10 carpels and locules; styles 1, the stigma
capitate or lobed; fruit a capsule or a berry.
1. Ovary inferior or apparently so 2
- Ovary superior 3
2(1). Plants prostrate shrublets, rooting along the stems; ovary superior but sur-
rounded by the fleshy calyx when ripe and apparently inferior
Gaultheria (hiimifusa)
- Plants erect or ascending, rooting only at the base; ovary inferior Vciccinium
3(1). Flowers borne in terminal corymbs, white, the segments of the corolla much
longer than the short tube; leaves punctate below with yellow-glandular dots ...
Ledum
- Flowers solitary and axillary or in axillary racemes, rarely terminal, pink to
lavender, the segments of the corolla much shorter than the tube; leaves
lacking glandular punctae 4
4(3). Corolla broadly saucer shaped or ratate, not constricted at the apex Kahnia
Corolla campanulate to urn shaped, often more or less constricted at the
throat 5
5(4). Corolla campanulate; anthers lacking appendages; fruit a capsule embedded in
a fleshy calyx Gaultheria {humifusa)
Corolla urn shaped; anthers 2-awned; fruit a berry Arctostaphylos
Arctostaphylos Adams
Evergreen prostrate to ascending or erect
shnibs, often with purplish to orange brown,
smooth bark; leaves alternate, simple, entire,
leathery-thickened; flowers in terminal pan-
icles or racemes, perfect, regular; sepals usu-
ally 5; petals usually 5, united almost to the
tips; corolla urn shaped; stamens usually 10,
included; anthers opening by falsely terminal
pores, each with 2 hornlike appendages;
ovary superior, usually 5-loculed; fruit fleshy,
berry like, 1- to several-seeded.
Adams, J. E. 1940. A systematic study of the
genus Arctostaphylos. J. Elisha Mitchell
Soc. 56: 1-62.
Eastwood, A. 1934. A revision of Arctosta-
phylos with keys and descriptions. Leaft.
West. Bot. 1: 105-127.
Plants with creeping-prostrate stems; leaves obovate-spatulate, commonly less
than 1.5 cm long X. uva-ursi
Plants with stems ascending to erect; leaves mostlv ovate to lanceolate or
elliptic, often more than 2 cm long 2
March 1980
Welsh: Utah Flora, Miscellaneous Families
41
2(1). Calvx and pedicels puherulent with spreading glandular hairs; twigs and leaves
puberulent throughout with spreading hairs; plants of Washington County
A. pringlei
- Calyx glabrous or nearly so; twigs and leaves puberulent or sessile to sparingly
stipitate-glandular or almost or quite glabrous; plants of various distribution 3
3(2). Twigs and axis of inflorescence white-puberulent, not glandular; plants of
Washington and Kane counties A. piingens
Twigs and axis of inflorescence glandular to glandular-puberulent; plants
widely distributed A. patulu
Arctostaphylos patula Greene. Green-leaf
Manzanita. {Uva-iirsi patiila (Greene)
Abranis; A. piingens var. phityphylla Gray;
A. platyphyUa (Gray) Kuntze; A. obtiisifolia
Piper; A. patula var. incarnata Jeps.; A. pi-
nctorum Rollins; A. parryana var. pinetorum
(Rollins) Weislander & Schreiber). Rounded
shrubs with gnarled stems to 15 cm long or
more, the bark smooth, cinnamon to reddish
brown or purplish in color; branchlets
glandular-puberulent and sometimes with
long-spreading hairs as well; leaf blades (0.8)
1.8-4.7 cm long, (0.6) 1.5-4 cm wide, ovate
to elliptic, lanceolate, or orbicular, obtvise to
acute apically, rounded to truncate basally,
glabrous or glandular, yellow green; petioles
pubescent like the twigs; inflorescence pan-
iculate, the axis and bracts glandular-pub-
enilent and sometimes with some long hairs;
pedicels glabrous; sepals glabrous; corolla
pink to white, 5-8 mm long; ovary glabrous;
fruit 8-11 mm thick, depressed-globose,
glabrous, white to brown, with nutlets sepa-
rable or not. Usually associated with ponde-
rosa pine at 1520 to 2830 m in Beaver, Duch-
esne, Garfield, Iron, Juab, Kane, Millard, San
Juan, Sanpete, Sevier, Summit, Tooele, Uin-
tah, Utah, Wasatch, and Washington coun-
ties; Colorado, Nevada, Oregon, Arizona, and
California. Arctostaphylos patula and A.
platyphyUa both date as species from the
same year, 1891; the question of which has
priority is difficult to ascertain; 64(x).
Arctostaphylos pringlei Parry. Pink-
bracted Manzanita. Rounded, erect shrubs to
20 dm tall or more, the bark smooth, dull red
l)rown; branchlets densely glandular-hairy
with long-spreading hairs; leaf blades (1.2)
1.8-4.2 cm long, (0.4) 0.8-2 cm wide, elliptic
to lance-elliptic or lanceolate, obtuse to
acute apically, truncate to rounded or obtuse
basally, glandular-pubescent, gray green; pe-
tioles pubescent like the twigs; inflorescence
paniculate or racemose, the axis and bracts
glandular-hairy; corolla pink, 6.5-8.5 mm
long; ovary glandular-hairy; ovary glandular-
hairy; fruit 6-10 mm thick, ovoid, glandular-
hairy, red, with nutlets inseparable. Oak-juni-
per community, 1840-2750 m, in Washing-
ton County; Arizona, California and Baja
California; 4(o).
Arctostaphylos pungens H.B.K. Mexican
Manzanita. Erect or ascending, rounded
shrubs to 20 dm tall or more, the bark
smooth, red brown; branchlets canescent
with a dense pubescence; leaf blades 1.6-4.7
(6) cm long, 0.5-3.2 cm wide, ovate to ellip-
tic or oblong, rounded to acute apically,
acute to rounded basally, puberulent on one
or both sides, bright green; petioles pub-
escent like the twigs; inflorescence pan-
iculate, the axis and bracts canescent; pedi-
cels glabrous; sepals glabrous; corolla pink to
white, 5.5-8.5 mm long; ovary glabrous; fruit
5-8 mm thick, depressed-globose, glabrous,
brownish red, with nutlets separable or not.
Pinyon, juniper, live oak communities,
920-2750 m, in Washington and Kane (At-
wood .3538 BRY) counties; California, Ari-
zona, New Mexico, Texas; Mexico; 19(iii).
Arctostaphylos uva-ursi (L.) Spreng. Kin-
nikinnick, Bearberry, Sandberry. {Arbtitus
uva-ursi L.; Uva-ursi procumhens Moench;
Mairania uva-ursi (L.) Desv.; U. buxifolia S.
F. Gray; A. officinalis Wimm. & Grab.; A.
procumhens in Mey. & Elkan; U. uva-ursi
(L.) Britt. in Britt. & Br.; A. media Greene; A.
uva-ursi var. coactilis Fern. & Macbr.; A.
uva-ursi var. adenotricha Fern. & Macbr.).
Prostrate shrub with stoloniferous rooting
steins, mat-forming, the branches ascending,
the intemodes usually apparent, puberulent
and sometimes glandular, the bark exfoliating
exposing dull brown imder bark; leaf blades
42
Great Basin Naturalist
Vol. 40, No. 1
(0.6) 1-2.7 (3) cm long, 0.3-1.2 wide, oblan-
ceolate to spatulate, rounded apically, cu-
neate to acute basally, glabrous or pub-
enilent, especially on the margins, green;
inflorescence racemose, the axis and bracts
glandular; pedicels glabrous or sparingly
puberulent; sepals glabrous; corolla pink to
white, 4-5.2 mm long; ovary glabrous; fruit
6-11 mm thick, globose, bright red, with sep-
arable nutlets. Ground layer in coniferous
forests, at 2140-3350 m, in Daggett, Duch-
esne, Garfield, Salt Lake, Sevier, Summit,
Uintah, and Wasatch counties; Alaska and
Yukon east to the Atlantic and south to Cali-
fornia, New Mexico, Illinois, and Georgia;
Eurasia; 15(ii).
Gaultheria L.
Prostrate shrubs, the branches rooting;
leaves alternate, thin, serrulate; flowers ax-
illary, solitary, perfect, regular; calyx 5-
lobed, united, enlarging and becoming fleshy
at maturity; corolla campanulate, the lobes
shorter than the tube; stamens usually 10, in-
cluded, the filaments flattened, tapering to
the apex; anthers opening by terminal pores,
not awned; ovary superior, usually 5-loculed;
fniit a loculicidally dehiscent capsule en-
closed by the fleshy expanded calyx.
Gaultheria humifusa (Grab.) Rydb. Al-
pine Wintergreen. {Vaccinium humifusum
Grab.; G. myrsinites Hook.). Prostrate,
scarcely woody plants with creeping, rooting
stems to 2 dm long, glabrous or puberulent;
leaves 0.6-1.5 cm long, 0.4-1.3 cm wide, oval
to ovate or elliptic, rounded to obtuse api-
cally and basally, serrulate; flowers solitary,
axillary; calyx glabrous; corolla 3-4 mm long,
campanulate, pink; fruit 5-7 mm thick, sub-
globose, red. Ground layer in coniferous for-
ests and margins, 2900-3350 m, in Duchesne
and Summit counties, and possibly elsewhere;
Colorado westward to California and north
to Alberta and British Columbia; 5(i).
Kalmia L.
Low shrubs with puberulent branches;
leaves opposite, evergreen, leathery, decur-
rent, entire, revolute, glaucous beneath; flow-
ers in terminal leafy-bracted corymbs or soli-
tary, perfect, regular; calyx 5-lobed, the
segments almost distinct; corolla bowl
shaped, the lobes shorter than the tube, the
tube with 10 pouches in which the anthers
are enclosed in bud; stamens usually 10, the
filaments flattened, hairy below; anthers
opening throughout, unawned; ovary superi-
or, 5-loculed; fruit a septicidally dehiscent
capsule.
Kalmia microphylla (Hook.) Heller. Bog
Laurel. {K. glauca var. microphylla Hook.; K.
polifolia var. microphylla (Hook.) Rhed.).
Erect slender shrubs, 0.7-1.5 dm tall; leaves
0.6-1.8 (3) cm long, 0.2-0.8 (1.2) cm wide,
lance-oblong to elliptic, revolute, shining and
green above, grayish beneath; corymbs most-
ly 2- to 6-flowered, the pedicels 1-3 cm long;
sepals glabrous, ciliate; corollas 11-14 mm
broad, pink; capsules 4-6 mm broad. Alpine
meadows and lake margins, 2900-3800 m, in
Daggett (?), Duchesne, Summit, and Uintah
counties; Alaska and Yukon south to Califor-
nia and Colorado; ll(i).
Ledum L.
Erect or spreading shrubs with glandular-
pubenilent branchlets; leaves alternate, ever-
green, leathery, entire, revolute, pale below;
flowers in terminal corymbs, perfect, regular;
calyx small, the segments almost distinct; co-
rolla rotate, the 5 petals distinct or nearly so;
stamens usually 5-10, the filaments almost
filiform, usually hairy below; anthers opening
by terminal pores, unawned; ovary superior,
5-loculed; fruit a septicidally 5-valved cap-
sule, opening at the base. Note: At least some
species of this genus are poisonous to live-
stock.
Ledum glandulosum Nutt. Trapper's Tea.
Plants mostly 5-15 dm tall, the branchlets
puberulent and glandular dotted; leaves
1.1-3.4 (4) cm long, 0.4-1.4 (1.8) cm wide, el-
liptic to oblong, rounded to acute apically
and basally, green above, pale to grayish be-
neath, glandular, the margin more or less
revolute; flowers white, the segments to 5
mm long or more; pedicels commonly 1-2.5
cm long, puberulent near the base; capsules
3-6 mm long, puberulent and glandular.
Meadows, stream banks, and bogs in open
forest, 2600-3050 m, in Duchesne^ Salt Lake,
March 1980
Welsh: Utah Flora, Miscellaneous Families
43
Summit, and Uintah counties; British Cohim-
bia east to Montana and south to Cahfornia,
Nevada, and Wyoming; 9(o).
Vaccinium L.
Decumbent-ascending to erect shrubs;
leaves alternate, deciduous, or more or less
evergreen, entire or serrulate, flat, green or
pale beneath; flowers solitary, axillary, or in
terminal clusters, perfect, regular; calyx 4- to
6-lobed, united at the base; corolla urn
shaped or campanulate, the 4-6 lobes shorter
than the tube; stamens 8-12, the fllaments
usually glabrous; anthers opening by pores at
the ends of tubular beaks, usually 2-awned;
ovary inferior, usually 4-locular; fruit a sever-
al-seeded berry.
Camp, W. H. 1942. A survey of the Ameri-
can species of Vaccinum, subgenus Eu-
vaccinium. Brittonia 4: 205-247.
1. Branches bright green and angled; plants often less than 3 dm tall 2
- Branches neither bright green nor angled, or sometimes irregularly angled
when dry; plants often more than 3 dm tall 3
2(1). Fruit red; grooves of branches usually glabrous; leaves often less than 12 mm
long V. scoparium
- Fniit blue black or black; grooves of branches usually puberulent; leaves often
over 12 mm long V. myrtillus
3(1). Flowers in clusters of 2-4, or solitary; leaves entire; calyx deeply lobed, the
lobes persistent in fruit V. occidentale
Flowers solitary in leaf axils; leaves more or less serrate; calyx shallowly lobed,
the lobes deciduous in fruit 4
4(3). Plants mostly 1-3 dm tall; leaves serrate above the middle and uncon-
spicuously below the middle, mainly 1-3 (4) cm long, oblanceolate to obovate .
V. caespitosiwi
Plants mostly 4-7 dm tall or more; leaves serrate to the base or nearly so,
commonly 2-6 cm long, elliptic to ovate V. membranaceum
Vaccinium caespitosum Miehx. Dwarf
Huckleberry. Plants mostly 1-3 dm tall;
twigs brownish, somewhat angled, pub-
erulent or glabrous; leaves 0.7-4 cm long,
0.3-2 cm wide, oblanceolate to obovate, ob-
tuse or less commonly acute to rounded api-
cally, usually cuneate basally, serRilate from
tip to below the middle; flowers solitary, ax-
illary, whitish to pink, the corollas 5-6 mm
long, twice as long as thick; calyx obscurely
lobed, the lobes deciduous in fruit; berries
blue glaucous, subglobose, 5-8 mm broad,
edible and good. Streamsides, meadows, and
rock outcrops, 2,227-3,416 m elevation, in
the Uinta Mountains and Boulder Mountains,
in Daggett, Duchesne, Garfield, Summit, and
Uintah counties; Alaska and Yukon east to
Newfoundland and New Hampshire, and
south to California and Colorado. Materials
from Utah have previously passed under the
names V. membranaceum Dougl. (see below)
and V. globulare Rydb. The latter is not
known for the state; 10(i).
Vaccinium membranaceum Dougl. Moun-
tain Huckleberry. Shrubs mostly 3-7 dm tall
or more; twigs brownish, glabrous or pub-
CRilent; leaves 1.8-7 cm long, 1-3.4 cm wide,
elliptic or less commonly ovate or obovate,
acute to obtuse apically, acute to rounded
basally, serrate almost throughout; flowers
solitary, axillary, yellowish pink, the corollas
about 6 mm long, about one-third longer
than broad; calyx obscurely lobed, the lobes
deciduous; berries purple, not glaucous, 7-9
mm broad, edible and good. Slopes in aspen-
conifer and spruce-fir woods, 2,500 to 2,775
in, in Cache, Carbon, Duchesne, Salt Lake
(?), and Summit counties; British Columbia
southward to California, Idaho, and Mon-
tana; 6(0).
Vaccinium myrtillus L. Dwarf Billberry.
(V. oreophilum Rydb., in part, the type from
44
Great Basin Naturalist
Vol. 40, No. 1
the Uinta Mountains.) Plants mostly 0.5-3 dm
tall; twigs seldom numerous and broomlike,
green, sharply angled, puberulent; leaves
1.1-3.9 cm long, 0.6-1.6 cm wide, ovate to
lanceolate or elliptic, acute to obtuse api-
cally, obtuse to roimded basally, serrulate al-
inos't or quite from base to apex; flowers soli-
tary, axillary, pink, the corollas 4-5 mm long;
calyx shallowly lobed; berry usually bluish,
5-8 mm broad. Ground layer in coniferous
forests, 2750-3200 m, in the Uinta, Wasatch,
and LaSal moimtains (Daggett, Duchesne,
San Juan, Summit, and Uintah counties),
where evidently not common; British Colum-
bia and .\lberta south to Arizona and New
Mexico; Eurasia. Vaccinium myrtillus is a
near congener of the very common V. scopa-
riwn and can be distinguished by the larger
size of its leaves and flowers and by the pub-
erulent stems; 6(i).
Vaccinium occidentale Gray. Western
Huckleberry. Plants mostly 2-6 dm tall, the
twigs round, usually glabrous; leaves 0.6-2.1
cm long, 0.4-1.2 cm wide, oblanceolate,
rounded to obtuse apically, acute basally, en-
tire; flowers 2-4, or less commonly solitary in
the axils, pinkish, the corollas 3.5-6 mm long;
calvx definitely lobed, the lobes persistent in
fniit; berries blue, glaucous, 4-6 mm thick.
Meadows, streamsides, and forest margins,
2750-3100 m, in the Uinta Mountains in
Daggett (?), Duchesne, Summit, Uintah, and
Wasatch counties; British Columbia south to
California and Idaho; 12(iv).
Vaccinium scoparium Leiburg. Grouse-
berry. (V. myrtillus var. microphijUum Hook.;
V. microphijllwn (Hook.) Rydb., not Rein.; V.
enjthrococcum Rydb.). Plants mostly 1-2.5
dm tall, the twigs numerous, broomlike,
sharply angled, usually glabrous; leaves
0.6-1.3 cm long, 0.3-0.7 mm wide, ovate, ob-
tuse to acute apically, roiuided to obtuse ba-
sally, serrulate throughout; flowers solitary,
axillary, pinkish, the corollas 2.5-3.5 mm
long; calyx very shallowly lobed; berry bright
red, drying red purple, 4-6 mm thick. Com-
mon component of ground layer in con-
iferous forests and forest margins, 2450-3200
m, in the Uinta Mountains in Daggett, Duch-
esne, Summit, Uintah, and Wasatch counties;
British Columbia and Alberta south to Cali-
fornia and Colorado; 20(i).
Krameriaceae
Ratany Family
Shrubs, with divaricate branches; herbage
grayish pubescent; leaves alternate, simple,
entire extipulate; flowers perfect, irregular,
solitary, axillary; pedicels usually with 2 op-
posite foliacious bracts; sepals 4 or 5, un-
equal; petals 5, the upper 3 long clawed, dis-
tinct or partially connate and often purplish
in color, the 2 others broad, thick, sessile,
usually greenish and glandlike; stamens 4,
free or adnate to claw of upper petal, the an-
thers dehiscent by pores; ovary superior, 1-
loculed; ovules 2; fruit an indehiscent pod,
armed with prickles.
A family of the Western Hemisphere of a
single genus with about 25 species from
South America to southern United States.
Krameria L.
A single genus with characteristics of the
family.
Branchlets modified as thickened thorns 0.8-1.2 mm in diameter at base; spines
of fruit barbed at apex only K. graiji
Branchlets not modified as thorns or if so then less than 0.6 mm in diameter;
spines of fruit with barbs scattered or, rarely, barbless K. pawiflora
Krameria grayi Rose & Painter. White Ra-
tany. Shrubs branched, 2.5-6 dm tall and as
wide; leaves 5-7 (25) mm long, 1-3 mm
wide, lance-ovate to lanceolate, elliptic or
oblong, more or less spinulose-tipped, tomen-
tose on both surfaces; pedicels not glandular-
pubescent; upper petals 2.5-3.5 mm long,
0.3-0.5 mm wide, yellowish with a purplish
tip; sepals 4.5-6.5 mm long, villous-pilose
dorsally, pilose to glabrate within, purplish;
prickles of the fruit 2-6 mm long at maturity,
each with a whorl of barbs at the apex; pods
March 1980
Welsh: Utah Flora, Miscellaneous Families
45
subglobose, 6-10 mm in diameter, hirsute
over the surface and on bases of prickles.
Blackhmsh and creosote bush communities at
670-1170 m in western Washington Co.; Cal-
ifornia, Nevada, Arizona, New Mexico,
Texas, and Mexico; 2(i).
Krameria parvifolia Benth. Range Ratany.
(K. glandulosa Rose and Painter; K. parvi-
folia var. glandulosa (Rose and Painter)
Macbr.; K. iniparata (Britton) Macbr.)
Shrubs, intricately branched, 2-6 dm tall and
as wide; leaves 3-15 mm long, 0.3-1 mm
wide, linear to oblong, callous- to spinulose-
tipped, tomentose on both surfaces; pedicels
glandular or not; upper petals 2.5-2.8 mm
long, 0.7-1.2 mm wide, yellowish; sepals 4-6
mm long, strigulose dorsally, glabrous within,
pinkish to purplish; prickles of fruit 2-5 mm
long, retrorsely barbed along the rachis; pods
subglobose, 5-9 mm in diameter, pilose-hir-
sute on the surface. Joshua tree, blackbrush,
creosote bush, and bursage communities,
750-1600 m, in Washington Co.; California,
Nevada, Arizona, New Mexico, Texas, and
Mexico. The materials demonstrate variation
in glandular condition of pedicels, sepals, and
bracts. The variation seems to be haphazard,
with little or no correlation with other fea-
tures or with ecology. Hence, included herein
as synonyms are those names involved with
recognition of glandular and nonglandular
phases; 16(i).
Magnoliaceae
Magnolia Family
Deciduous or evergreen trees or shrubs;
leaves alternate, simple, entire or lobed, stip-
ulate, the stipules enclosing the buds, de-
ciduous or caducous, and leaving a circular
scar; flowers regular, perfect, solitary, termi-
nal and axillary, large and showy, the floral
parts spirally arranged; sepals often 3, the
petals 6 to many; stamens numerous, sepa-
rate, hypogynous, the anthers 2-loculed; pis-
tils several to many, each 1-loculed and 1-
carpelled; style 1, the stigma 1; fruit a follicle
or samara.
1. Leaves lobed, truncate or broadly retuse at the apex; flowers borne after the
leaves Liriodendron
Leaves entire, acute, or acuminate; flowers borne before or after the leaves
Magnolia
Liriodendron L.
Trees, the leaves large and 4-lobed; flowers
large, inconspicuously colored; sepals 3, soon
reflexed; petals 6, ascending to erect, forming
a tuliplike corolla; anthers extrorse; pistils
many, en masse becoming conelike, the indi-
vidual samaras eventually deciduous.
Liriodendron tulipfera L. Tulip Tree; Yel-
low Poplar. Deciduous, cultivated trees to 40
m tall or more, the tmnks to 10 dm in diam-
eter or more; leaves long-petioled, the blades
6-15 cm long and almost as wide; flowers
solitary, terminal; sepals green; petals 3.7-6
cm long, yellow green, with a basal orange
spot within; samaras narrow, 3-4 cm long.
Occasional shade tree in more moderate low
elevation portions of Utah; introduced from
the eastern United States; 8(o).
Magnolia L.
Trees or shrubs; leaves large, entire; flow-
ers large, conspicuous or inconspicuous; se-
pals 3, colored like the petals; petals 6-12,
erect or spreading; anthers introrse; pistils
many, en masse becoming conelike, the indi-
vidual follicles finally dehiscent.
Plants shrubs or small trees, deciduous; flowers showy, cream to pink or
suffused with rose or lavender, borne before the leaves appear M. soulangeana
Plants moderate to large trees, deciduous or evergreen; flowers greenish and
inconspicuous or, if showy, then white in color and the trees evergreen 2
46
Great Basin Naturalist
Vol. 40, No. 1
Plants evergreen, the leaves dark green, leathery; flowers white M. grandiflora
Plants deciduous, the leaves not both dark green and leathery; flowers greenish
ygllovv ^- acuminata
Magnolia acuminata L. Cucumber-tree.
Deciduous trees to 30 m tall or more; leaves
deciduous, short-petioled, the blades 8-25 (3)
cm long and 4-15 cm wide; flowers solitary,
terminal; perianth greenish yellow, 5-8 cm
long. Cultivated shade tree, uncommon,
hardy in the major cities of the state; in-
troduced from the eastern United States; 2(o).
Magnolia grandiflora L. Bull Bay. Ever-
green trees to 30 m tall; leaves evergreen,
short-petioled, the blades mostly 8-20 cm
long and 3-8 cm wide; flowers solitary, ter-
minal; perianth white, mostly 8-12 cm long.
Cultivated ornamental, uncommon, not
hardv except in favorable sites in moderate to
warm portions in Utah; introduced from
southeastern United States; l(o).
Magnolia soulangeana Soul. Showy Mag-
nolia. Shrubs or small trees to about 4 m tall;
leaves deciduous, short-petioled, the blades
mostly 8-14 cm long and 3.5-10 cm wide;
flowers solitary, terminal; perianth cream to
pink or suffused with rose or lavender, 6-12
cm long or more. Cultivated ornamental, oc-
casional in more moderate climatic areas of
Utah; a hybrid of M. denadata Descr. and M.
lili flora Descr., both native of China; 3(o).
MORACEAE
Mulberry Family
Deciduous trees or shrubs with milky juice;
leaves alternate, simple, pinnately or pal-
mately veined, entire, serrate, or lobed, stipu-
late, the stipules small and distinct or each
pair forming a cap over the bud and leaving
a scar around the stem; flowers imperfect,
minute, regular, borne in cymes or much
modified inflorescences; perianth of usually 4
sepals; staminate flowers with usually 4 (2 in
Ficus) stamens, the filaments distinct; pistil-
late flowers with or without a 4-lobed per-
ianth; pistil 1, the ovary superior to inferior,
1-loculed, the styles and stigmas 2 (1 in Ma-
dura)., fruit a multiple (Morus, Madura) or a
syconium (Ficus).
1. Fniit a fleshy hollow receptacle with flowers borne inside (syconium); leaves
palmately veined and lobed; cultivated plants of Washington County, and of
greenhouses elsewhere Ficus
- Fruit a multiple (formed of several flowers and a common axis); leaves various .
2
2(1). Leaves crenate-serrate, palmately veined and often palmate lobed as well;
flowers, both sterile and fertile, borne in catkinlike spikes; fruit seldom more
than 1 cm thick Morus
Leaves entire, pinnately veined, not lobed; flowers borne in dissimilar in-
florescences, the sterile in racemes, the fertile in globular heads; fruit globular,
more than 5 cm thick Madura
Ficus L.
Trees or large shrubs; leaves alternate,
simple, palmately veined and lobed, the stip-
ules forming a circular scar around the stem;
flowers minute, numerous, borne inside a hol-
low receptacle which ripens to form a syco-
nium; staminate perianth 2- to 6-parted, with
1 or 2 stamens; pistillate perianth reduced or
lacking; receptacles perfect or imperfect;
fruits of individual flowers of achenes.
Ficus carica L. Common Fig. Deciduous
trees to 5 m tall, rarely more, often sprawling
in age; leaves prominently veined, thick, to
25 cm long or more and to 20 cm broad, 3- to
5-lobed, the lobes undulate-serrulate; fruits
obovoid, mostly 2.4-4.5 cm long and 2-3 cm
thick. Cultivated fruit plant in Washington
March 1980
Welsh: Utah Flora, Miscellaneous Families
47
(and formerly Garfield, at Hite) County, frost
sensitive elsewhere except under glass; in-
troduced from the Mediterranean region of
the Old World. This is the fig of commerce;
5(iii).
Maclura Nutt.
Dioecious trees with hard yellow wood;
leaves entire, the stipules minute, the scar not
encircling the stem; staminate flowers nu-
merous in loose, peduncled, axillary heads or
umbels, the calyx 4-parted and with 4 sta-
mens; pistillate flowers coherent in dense,
globose, axillary heads, the calyx 4-lobed, the
single filiform style very long; fruit a globose
multiple.
Maclura pomifera (Raf.) Schneid. Osage
Orange. {Toxylon pomiferum Raf.) Trees to
10 m tall, rarely more; stems usually armed
with stout thorns 1-2 cm long; leaves petio-
late, the blades 5-10 cm long and 1.8-6.5 cm
wide, ovate, entire, rounded to obtuse ba-
sally, attenuate to acuminate apically; clus-
ters of staminate flowers 2.5-3.5 cm across;
heads of pistillate flowers 2-2.5 cm across;
multiple fruit mostly 8-14 cm thick. Culti-
vated ornamental and botanical curiosity of
low elevation regions in Utah, long per-
sisting; introduced from the eastern states.
The wood of this tree is very strong, and has
served as a source of bows for American In-
dians and others; 5(i).
Morus L.
Dioecious trees; leaves palmately veined,
serrate to dentate, sometimes lobed; stipules
lanceolate, the scar not encircling the stem;
flowers monoecious or dioecious, those of
both sexes borne in stalked, axillary, catkin-
like clusters; calyx 4-parted; stamens 4; styles
2, deeply parted; fruit a multiple.
Leaves glabrous above and beneath or pubescent beneath only along main
veins and/or in vein axils; our common mulberry M. alba
Leaves pubescent over much of the lower surface, scabrous above; rarely
cultivated M. nisra
Morus alba L. White Mulberry. Cultivated
ornamental and shade tree to 10 m tall or
more; leaves obliquely ovate and crenate-ser-
rate or irregularly lobed, mostly 3.5-14 cm
long and 2.5-10 cm wide, truncate to sub-
cordate basally, acute to acuminate apically,
glabrous above and below except along veins
and in vein axils; fruit 1-2 cm long and 0.6-1
cm thick, white, pink, red purple, or nearly
black. Persisting and occasionally escaping in
most of Utah at lower elevations; introduced
from China; widespread in North America.
This plant was introduced to southern Utah
to provide food for silkworms in an attempt
to develop a silk industry. The fruit is edible,
but is consinned mainly by birds. Reports
from Utah of red mulberry, M. rubra L., be-
long here. Red mulberry is easily recognized
by the densely hairy lower and scabrous up-
per leaf surfaces. So-called fruitless phases
are known; 24(v).
Morus nigra L. Black Mulberry. Small
trees to about 10 m; leaves cordate-ovate,
crenate-serrate, seldom lobed, 5-20 cm long,
3-15 cm wide, cordate basally, obtuse to
acuminate apically, scabrous above and hairy
over veins and at least some intervein areas
below; fniit 1-2.5 cm long and to 1 cm thick,
purple to black. Sparingly cultivated orna-
mental, mainly in warm regions of Washing-
ton County; widely cultivated in temperate
regions of the earth for its fruit; introduced
from Asia; 5(o).
Oleaceae
Olive Family
Trees or shrubs; leaves opposite (or rarely
alternate), simple or pinnately compound,
stipulate; flowers perfect or imperfect, borne
in axillary or terminal racemose, paniculate,
or thyrsoid inflorescences; calyx commonly 4-
48
Great Basin Naturalist
Vol. 40, No. 1
lobed or absent; corolla usually of 4 united or
distinct petals, or lacking; stamens 2, distinct;
pistil 1, the ovary superior, 2-carpelled and
2-loxuled; style 1, or lacking, the stigmas 1 or
2; fruit a berry (Ligustrinn), drupe
(Forestiera), loculicidal capsule {Syringa, For-
sythia), circumscissile capsule (Menodora), or
samara {Fraxiniis).
1. Leaves pinnately compound; fruit a samara Froxinus
Leaves simple, or rarely compound; fruit various 2
2(1). Leaves ovate to orbicular, crenate-serrate; fruit a samara; plants indigenous
Fraxiniis
Leaves various, but seldom ovate to orbicular and crenate-serrate; fruit a
drupe, capsule, or berry; plants cultivated or indigenous 3
3(2). Shrubs with yellow flowers appearing before the leaves; plants cultivated
Forsythia
Shmbs, subshrubs, or trees with flowers variously colored, but if yellow then
not as above, and appearing with or after the leaves (before in Forestiera) 4
4(3). Corolla none or rudimentary, the flowers often unisexual; fruit a drupe; shrubs
of stream banks in southeastern Utah Forestiera
- Corolla well developed, the flowers perfect; fruit a berry or a capsule 5
5(4). Corolla yellow; fruit a membranous, circumscissile capsule; plants indigenous
subshrubs of southern Utah Menodora
- Corolla commonly lavender to red, purple, white, or cream; fruit a loculicidal
capsule or a berry; plants cultivated shrubs or trees 6
6(5). Flower clusters usually less than 6 cm long; flowers white to cream; fruit a
berry Ligustrwn
Flower clusters usually 6-30 cm long or more; flowers lavender to red, purple,
lilac, white or cream Syringa
Forestiera Poir.
Sprawling indigenous shrubs; leaves oppo-
site, simple, serrate to entire; flowers incon-
spicuous, polygamo-dioecious, borne sessile
or in cymes, appearing before the leaves; ca-
lyx minute, unequally 5- to 6-cleft, or lack-
ing; corolla lacking, or rarely with 2 or 3 pet-
als; stamens 2 or 4; ovary 2-loculed, with 2
ovules per locule; style slender; stigma 1;
fniit a dnipe.
Forestiera pubescens Nutt. Desert Olive.
(F. neo77iexicana Gray; Adelia neomexicana
(Gray) Kuntze; A. parvifolia Gov.). Shrubs to
2 m tall or more; leaves (0.8) L5-5.5 cm
long, (0.3) 0.5-2 cm wide, oblanceolate to el-
liptic, entire to serrulate; staminate flowers
sessile; pistillate flowers pedicellate; drupe
5-7 (8) mm long, ellipsoid, blue black. Sandy
terraces along the Colorado and San Juan riv-
ers and tributaries, 1280-1750 m in Grand
and San Juan counties; California eastward to
Oklahoma and Texas, and south to
Chihuahua. The fruit is eaten by fox and by
coyotes, and tlie purple-stained, stone-laden
fecal pellets are to be found far from the riv-
ers. Long known as F. neomexicana, our ma-
terials form a portion of a complex whose
definition includes those portions known as F.
pubescens, and that name has priority; 8(iii).
Forsythia Vahl
Cultivated shrubs; leaves opposite, simple
or some compound, entire to serrate; flowers
perfect, showy, borne in axillary cluster of
3-5, or solitary, appearing before the leaves;
calyx 4-lobed; corolla 4-lobed, campanulate;
stamens 2, inserted at corolla base; ovary 2-
loculed, with several ovules per locule; fruit
a loculicidal capsule, with many winged
seeds (ours seldom fruiting).
March 1980 Welsh: Utah Flora, Miscellaneous Families 49
Forsythia suspensa (Thunb.) Vahl. C.old- Fraxinus L.
en-bell. {St/rin^a suspciisa Thunb.). Shrub to
2 m tall or more; branchlets somewhat 4- Deciduous, cultivated and/or indigenous
angled; leaves 6-10 cm long, ovate to lanceo- trees or .shrubs; winter buds often prominent,
late, acute apically, cuneaie to rounded ba- g^y to brown or black; leaves opposite, pin-
.sally, usually serrate; flowers to 25 mm long, nately compound (simple in F. anomala);
golden yellow; fruit lance-ovoid, to 15 mm flowers perfect or uni.sexual, inconspicuous,
long, .seldom developing. Cultivated orna- borne in panicles; calyx 4-lobed or lacking;
mental, common, persisting but not spread- toroUa lacking or of 2 or more, usually dis-
ing at lower elevations throughout Utah; tinct petals; stamens commonly 2; ovary 2-lo-
widespread; introduced from China. Numer- <-uled; styles 1; stigmas 1 or 2; fruit a samara,
ous horticultural varieties are present; 2(o).
1. Leaves normally simple, sometimes with 1 or 2 leaflets below the terminal one;
indigenous shrubs or .small trees of eastern and southern Utah F. anomala
— Leaves normally pinnately compound with 5-9 or more leaflets; trees, either
indigenous or cultivated 2
2(1). Branchlets, petioles, and axis of panicle commonly spreading hairy, seldom
glabrous; leaflets usually 5 or fewer; trees, indigenous in southwestern Utah,
cultivated elsewhere F. velutina
2. Branchlets, petioles, and axis of panicle variously hairy or glabrous, but seldom
spreading hairy; leaflets usually 7 or more; trees, cultivated and sometimes
escaping 3
3(2). Flowers appearing after leaves formed; corolla present F. ornus
— Flowers appearing before leaves formed; corolla lacking 4
4(3). Fiiiit with calyx persisting as a campanulate cap; anthers oblong; leaflets
usually 5-7 5
— Fruit with calyx early deciduous or lacking (except in F. qiiadrangulata);
anthers often cordate; leaflets u.sually 9-11 or more 6
5(4). Petiolules of middle and lower mature leaflets wingless nearly their entire
length; winter buds black; leaf scars horseshoe shaped; wing of fruit terminal,
not or only slightly decurrent F. americana
— Petiolules of middle and lower mature leaflets winged nearly to the base; win-
ter buds brown; leaf scars semicircular or shield shaped; wing of fruit decur-
rent to below the middle F. pennsylvanica
6(4). Branchlets 4-sided, 4-angled; bark broken into plates; flowers with a minute,
deciduous calyx F. qtiadrangulata
— Branchlets terete, not or only slightly 4-angled; bark smooth or irregularly
roughened; flowers with calyx lacking 7
7(6). Leaflets glabrous or somewhat hairy along veins beneath; commonly cultivated
tree F. excelsior
— Leaflets definitely pubescent beneath, especially along the veins, the long red-
dish hairs extended onto and along the leaf rachis; unconunon to rarely
cultivated tree F. nigra
Fraxjnus americana L. White Ash. Moder- usually 7 (5-9) 6-15 cm long, petiolulate,
ate to large trees; branchlets terete, green to ovate to lanceolate, acuminate apically, cu-
brown, glabrous; winter buds black; leaflets neate to rounded ba.sally, entire to serrate,
50
Great Basin Naturalist
Vol. 40, No. 1
glaucous beneath and usually glabrous; an-
thers oblong, apiculate; calyx persistent; co-
rolla lacking; samaras (20) 25-35 (50) mm
long, 4-7 mm wide, the wing not decurrent
along the terete base. Shade tree of lower
elevations in Utah; introduced from eastern
North America; 5(o)
Fraxinus anomala Torr. ex Wats. Single-
leaf Ash. Shrub or small tree, commonly
2.5-4 m tall, usually with many stems;
branchlets 4-angled; leaves glabrous, ovate,
crenate-serrate to subentire, 1.5-6.5 cm long,
1-6 cm wide, acute to obtuse or subcordate
basally, acute to rounded or emarginate api-
cally, sometimes 2- or 3-foliolate or transi-
tional to simple; flowers usually perfect; an-
thers oblong: calyx campanulate, persistent;
petals lacking; samaras winged almos^ to the
base, 12-27 mm long, 5-11 mm wide. Mixed
desert shrub, mainly on rimrock or along
drainages, and in pinyon-juniper woodland,
900-2150 m, in Emery, Garfield, Grand,
Iron. Kane, San Juan, Uintah, Washington,
and Wayne counties; Colorado, New Mexico,
Arizona, and California; 80(xv).
Fraxinus excelsior L. European Ash. Mod-
erate to large trees; branchlets terete,
glabrous; winter buds black; leaflets 7-11,
5-12 cm long, .sessile, ovate to oblong or lan-
ceolate, acuminate apically, cuneate basally,
serrate, green Ijeneath, glabrous except along
midrib, the hairs sometimes extending to the
rachis; flowers polygamous; anthers ovoid;
calyx lacking; corolla lacking; samaras 25-35
(40) mm long, 5-11 mm wide, the blade de-
current almost or quite to the base of the
flattened body. Shade tree of habitations and
streets at lower elevations throughout Utah;
introduced from Europe; 11(0).
Fraxinus nigra Marsh. Black Ash. Moder-
ate trees; branchlets terete, glabrous; winter
buds black; leaflets 7-11, mostly 6-12 cm
long, sessile, lanceolate to oblong, obtuse to
rounded basally, long-acuminate apicallv,
serrate, green and glabrous except reddish-
hairy along veins, the pubescence extending
along the leaf rachis; flowers dioecious; an-
thers oblong; calyx lacking; corolla lacking:
siunaras mostly 25-35 nun long and 6-10 mm
broad, the blade decurrent to the ba.se of the
flattened body. Sparingly cultivated shade
tree at lower elevations in at least the major
population centers; introduced from eastern
North America; 4 (o).
Fraxinus ornus L. Flowering Ash. Small to
moderate trees; branchlets terete; winter
buds gray to brownish; leaflets usually 7
(7-11), mostly 2.5-7 cm long, petiolulate,
lance-ovate to obovate (terminal one),
rounded to obtuse basally, acuminate api-
cally, crenate-serrate, glabrous except along
midrib; flowers perfect; calyx present, per-
sistent, with 4 triangular-acuminate, spread-
ing lobes; petals present, linear; samaras
20-25 mm long, 3-6 mm wide, the blade ter-
minal on the terete base. Rarely cultivated
shade and ornamental tree of lower eleva-
tions in Utah; introduced from Europe; 2(o).
Fraxinus pennsylvanica Marsh. Red Ash.
Moderate trees; branchlets terete, pubescent
to glabrous, sometimes glandular; winter
buds olive to brown; leaflets usually 7 (5-9),
6-15 cm long, petiolulate, lanceolate to
lance-oblong, acuminate apically, acute to
obtuse or rounded basally, serrate to entire,
green and glabrous or hairy (especially along
the veins) beneath; anthers oblong, apiculate;
calyx campamulate, persistent; corolla lack-
ing; samaras 27-40 (50) mm long, the blade
decurrent to the middle of the terete body or
below. Common shade tree of lower eleva-
tions throughout Utah, persisting and escap-
ing, in Box Elder, Cache, Davis, Iron, Juab,
Millard, Salt Lake, Utah, and Washington
counties; introduced from eastern North
America. The escaped plants have become
established along streams and on lake mar-
gins at lower elevations. Much of our mate-
rial has glabrous branchlets and petioles, and
has been designated as F. pennsylvanica var.
lanceolata (Borkh.) Sarg. (F. lanceolata
Brokh.). This phase is known as green ash; 28
(o).
Fraxinus quadrangulata Michx. Blue Ash.
Small to moderate trees; branchlets sharply
4-angled, glabrous; winter buds black; leaflets
7-11, mostly 5-12 cm long, petiolulate, lan-
ceolate to ovate-lanceolate, acute to rounded
basally, acute to acuminate apicallv, serrate,
glabrous except along the midrib or rarely
hairy over the lower surface; flowers perfect;
calyx minute, caducous; corolla lacking; an-
thers cordate-oblong, blunt; samaras 20-40
(50) nmi long, the blade decurrent to the base
of the flattened body. Sparingly cultivated
March 1980
Welsh: Utah Flora, Miscellaneous Families
51
shade tree, at lower elevations in Utah; in-
troduced from eastern North America; l(o).
Fraxinus vehitina Torr. Velvet Ash, Ari-
zona Ash. [F. pcnnsylvcmica Marsh, ssp. veln-
tina (Torr.) G. N. Miller] Moderate trees;
hranchlets terete, denselv spreading hairv to
merelv sparingly so, or glabrous; winter buds
brown; leaflets 3-5 (or leaves simple), lan-
ceolate to ovate, elliptic, or orbicular, pe-
tiolulate, cuneate to acute basally, acuminate
to rounded apicallv, serrate, glabrous or hairv
over the lower surface; flowers imperfect; ca-
lyx campanulate, persistent; corolla lacking;
anthers oblong, apiculate; samaras 16-34 mm
long, 4-6 mm wide, the blade decurrent
about half way along the terete body. In-
digenous tree of stream courses and flood
plains in Washington and Iron counties, and
cultivated there and elsewhere in Utah; Ari-
zona and New Mexico. The phase with co-
riaceous leaflets has been treated as var. co-
riacea (Wats.) Rehd. (F. coriacea Wats.), but
seems not to be worthy of taxonomic recogni-
tion, at least in Utah; 25(ii).
Note: The shrubby Fraxinus dipetala
Hook. & Am. is reported for Utah in
Kearney & Peebles, 1961. Flora of Arizona,
Supplement p. 1063. The related F. cuspi-
data Torr. is known from adjacent Mohave
and Coconino counties, Arizona, and might
occur in Utah. Both species have corollas
present; the former has two petals and the
latter has four.
LiGUSTRUM L.
Shrubs; leaves opposite, simple, entire;
flowers perfect, white, showy through small,
borne in terminal penicles, appearing after
the leaves; calyx 4-toothed; corolla 4-lobed,
funnelform; stamens 2, inserted on the co-
rolla tube; ovary 2-loculed, 1- or 2-seeded;
fruit a berry.
Ligustrum vulgare L. Common Privet. De-
ciduous or semievergreen shrub to 3 m tall or
more, with puberulent to glabrate branchlets;
leaves 2-6 cm long, 0.8-2 cm wide, oblong to
elliptic or ovate-lanceolate, glabrous; panicle
dense, 3-6 cm long; corolla tube shorter than
the lobes, white; anthers exserted; fruit 6-8
mm long, black, ovoid to subglobose. Culti-
vated hedge plant throughout Utah at lower
elevations, persisting and escaping; in-
troduced from Europe; 3(o).
Menodora Humb. & Bonpl.
Subshrubs; leaves alternate or the lower-
most opposite, simple, sessile or nearly so;
flowers perfect, arranged in cymes; calyx 5-to
15-lobed; corolla yellow, subrotate, 5- to 6-
lobed; stamens 2, inserted on the corolla
tube; ovary 2-loculed, with 2-4 ovules per
locule; style slender, the stigma capitate;
fiiiit a circumscissile capsule.
Steyermark, J. A. 1932. Revision of the genus
Menodora. Ann. Missouri Bot. Card. 19:
87-176.
Menodora scabra Gray. Plants erect or as-
cending, commonly 2-3.5 dm tall, woody at
the base only; leaves 0.5-2.9 cm long, 0.2-0.5
cm wide, narrowly elliptic to oblong or lan-
ceolate, glabrous or scaberulous; calyx mi-
nutelv puberulent, the lobes linear; corolla
bright yellow, subrotate, the lobes 5-9 mm
long; capsule 8-12 mm thick, membranous;
seeds 4-5 mm long. Pinyon-juniper commu-
nity, known in Utah only from Garfield, San
Juan, and Washington counties; California,
Arizona, New Mexico, Texas, and Mexico;
3(1).
Syringa L.
Shrubs or small trees; leaves opposite,
simple, petiolate; flowers perfect, in terminal
or lateral panicles; calyx campanulate, 4-
toothed to nearly truncate, persistent; corolla
tubular, the limb 4-lobed and rotate or nearly
so; stamens 2, inserted on the corolla tube;
ovary 2-loculed, each locule with usually 2
ovules; style with a 2-lobed stigma; fruit a
loculicidal capsule.
Flowers cream to whitish, borne in large panicles; corolla tube 1-2.2 mm long,
only half as long as the calyx; fragrance musky, not that of lilac; plants
flowering in summer, often treelike 2
Flowers lilac, violet, purpli.sh, or white; corolla tube mostly 6-12 mm long or
more, several times longer than the calyx; fragrance usually of lilac; plants
commonly shrubs, flowering in spring or .summer 3
52 Great Basin Natuhalist Vol. 40, No. 1
2(1). Leaves ovate, rounded or subcordate basally, the veins prominent on the lower
surface S. amiirensis
— Leaves lanceolate to elliptic or ovate-lanceolate, obtuse to cuneate basally, the
veins not prominent S. pekinensis
3(1). Panicles from terminal buds; leaves of current season borne on branch with
panicle; plants flowering in summer S. villosa
— Panicles from lateral (or terminal) buds, the terminal buds often lacking; leaves
of current season not borne on the branch with panicle; plants flowering in
springtime 4
4(3). Leaves ovate to cordate, the base subcordate to obtuse; our most common
species S. vulgaris
— Leaves lanceolate to elliptic or ovate, obtuse to cuneate basally; common to
uncommon 5
5(4). Leaves mostly less than 4 cm long, some often irregularly lobed; individual
panicles short, mostly 7 cm long or less S. persica
— Leaves often over 4 cm long, entire; individual panicles usually 8-12 mm long
S. X chinensis
Sijringa amiirensis (Rupr.) Rupr. Amur Li-
lac. Shrubs or small trees to 5 nun tall or
more; leaf blades 3.5-13 cm long, 1.3-8 cm
wide, ovate, rounded to obtuse of short
acuminate basally, acuminate apically, the
lower surface hairy to glabrous, the veins
prominent; petioles mostly 1-2 cm long; pan-
icles 10-15 cm long, the clusters of panicles
usually much longer; flowers cream to white;
stamens exserted. Sparingly cultivated orna-
mental of lower elevations in Utah; in-
troduced from Japan; flowering in summer;
4(0).
Sijringa \ chinensis Willd. Chinese Lilac.
Shrub to 4 m tall or more, with spreading
and often arching branches; leaves 2.5-8 cm
long, 1.5-4 (5) cm wide, ovate-lanceolate, ob-
tu.se to cuneate basally, acuminate apically,
glabrous, the veins not prominent; petioles
0.5-1.5 cm long; panicles mostly 8-12 cm
long, the clusters of panicles much longer;
flowers purple lilac, or otherwise; stamens in-
cluded. Commonly cultivated ornamental al-
most throughout Utah; introduced from the
Old World. This plant is evidently of hybrid
origin, having resulted from a cross between
S. persica and S. vulgaris, (j.v.; flowering in
springtime; 3(o).
Syringa pekinensis Rupr. Peking Lilac.
Shrub or small tree to 5 m tall or more, with
spreading branches; leaves 5-12 cm long, 2-4
(6) cm wide, lanceolate to ovate, cuneate ba-
sally, acuminate apically, glabrous, the veins
not prominent; petioles 1.5-3 cm long; pan-
icles mostly 8-15 cm long, the clusters of
panicles to 30 cm long or more; flowers
cream to yellow white; stamens exserted. Un-
common, cultivated ornamental in northern
Utah, but to be expected elsewhere; in-
troduced from China; flowering in early sum-
mer; l(o).
Syringa persica L. Persian Lilac. Shrub to
2 m tall, with upright to arching branches;
leaves 1.5-6 cm long, 0.6-3 cm wide, lan-
ceolate to elliptic, sometimes lobed, cuneate
to obtuse basally, acute to acuminate api-
cally, glabrous, the veins not prominent; pet-
ioles 0.5-1 cm long; panicles mostly 3-7 cm
long; flowers usually lilac but purple phases
are known; stamens included. Uncommonly
cultivated ornamental, especially in northern
Utah; introduced from Asia Minor; flowering
in springtime; 5(o).
Syringa villosa Vahl. Shrub to 3 m tall,
rarely more, with erect branches; leaves 4-15
cm long (or more), 2.5-9 cm wide, ovate to
elliptic, acute basally, abruptly acuminate
apically, spreading hairy below, especially
along the prominent veins; petioles 0.8-2 cm
long; panicles mostly 10-18 cm long; flowers
March 1980
Welsh: Utah Flora, Miscellaneous Families
53
pink lilac to white; stamens included. Spar-
ingly but widely planted ornamental, mainly
in northern Utah; introduced from China;
tlowering in summer; 4(o).
Syringa vulgaris L. Common Lilac. Shrubs
to 4 m tall or more, the branches usually
erect; leaves 3-12 cm long. 1.5-8 cm wide,
ovate to cordate, cordate to rounded, trun-
cate or obtuse basally, acute to acuminate
apically, glabrous; petioles 0.8-3 cm long;
panicles mostly 10-20 cm long; flowers lilac
or white, seldom purple; stamens included.
Abundantly cultivated ornamental, long per-
sisting, in most of Utah; introduced from Eu-
rope; flowering in springtime. Many horticul-
tiual forms are known; 7(o).
Pyrolaceae
VVintergreen Family
Suffrutescent or herbaceous perennials;
leaves simple, alternate, opposite, or appear-
ing whorled, evergreen or much reduced and
lacking chlorophyll; flowers usually perfect,
regular, or irregular; calyx with 4 or 5 more
or less distinct sepals; corolla with 4 or 5
more or less distinct petals (united in Ptew-
spora): stamen twice as many as the petals,
the anthers pendulous, opening by appa-
rently terminal pores or by slits, or the an-
thers erect, awnless or 2-awned; pistil 1;
ovary superior, 4- or 5-loculed; style 1; fruit a
capsule.
1. Plants lacking chlorophyll; leaves reduced and scalelike, reddish, brownish,
purple, or yellowish when fresh, often drying dark Ptewspom
Plants with chlorophyll (rarely without); leaves not reduced to scales, except
rarely, commonly evergreen 2
2(1). Flowers solitary, the petals rotate or nearly so Moneses
Flowers few to several, the petals concave 3
3(2). Stems leafy, though short, the leaves apparently whorled; flowers corymbose;
staminal filaments dilated near the base; styles very short or lacking
ChimaphUa
Stems leafy at base only; flowers in elongate racemes; filaments not especially
dilated at the base; styles in most species over 2 mm long Pyrola
Chimaphila Pursh
Low shrubs from creeping rhizomes, the
stems erect or ascending; leaves evergreen,
leathery, apparently whorled or some alter-
nate; flowers (1) 2-several, borne in peduncu-
late, umbellate corymbs; sepals usually 5, dis-
tinct nearly to the base, persistent; petals
usually 5, distinct, rotate-cam panulate; sta-
mens usually 10, the filaments dilated and cil-
iate near the base; anthers awnless, opening
by falsely terminal pores on short tubes;
ovary superior, 5-lobed and 5-loculed; fruit a
loculicidallv dehiscent capsule.
Cimaphila umbellata (L.) Bart. Pipsis-
sewa. Prince's Pine. {Pyrola innbellata L.; C.
occklentalis Rydb.; C. umbellata ssp. occiden-
talis (Rydb.) Hulten). Plants (1) 1.5-2.5 (3)
dm tall, the stems glabrous, only somewhat
woody; leaves 1.5-4.5 (6) cm long, 0.5-1.5 (2)
cm wide, elliptic to oblanceolate, cuneate ba-
sally, sharply serrate, shining above, pale be-
neath, glabrous; peduncles 4-7 (10) cm long,
glabrous or minutely glandular-puberulent,
often suffused with red purple; pedicels
glandular-puberulent or merely puberulent;
flowers 1-6 or more, umbellate-corymbose;
sepals erose-ciliate; petals 5-7 mm long,
pink; stamens with expanded bases ciliate;
capsules 5-7 mm broad. Coniferous forests,
2300-2750 m, in Duchesne, Summit, Uintah,
and Washington counties; .\laska, southward
to California and Mexico, east to New
Mexico and Colorado, and in the eastern
United States; Eurasia. Our materials are
referable to var. occidentalis (Rvdb.) Blake;
4(0).
54
Great Basin Naturalist
Vol. 40, No. 1
MoNESES Salisb.
Rhizomatous herbs; leaves with chloro-
phyll, leathery, persistent, mainly basal, but
sometimes opposite or in whorls; flowers soli-
tary, nodding, borne on a long peduncle; se-
pals usually 5, persistent; petals usually 5, dis-
tinct, spreading; stamens usually 10, the
filaments tapering to the apex, the anthers
awnless, nodding, opening by means of ap-
parently terminal pores; ovary superior, 5-
loculed, the stigma borne on an elongate,
glabrous style; fruit a loculicidal capsule.
Moneses uniflora L. Single Delight, Wax-
flower. (M. reticulata Nutt.; M. uniflora var.
reticulata (Nutt.) Blake). Plants 0.4-1.7 dm
tall; leaves (including petioles) 0.8-4 cm long,
0.6-2 cm broad, serrate to crenate-serrate;
peduncles 3-15 cm long, usually with 1 or 2
bracts along its length; flowers 1.3-2.5 cm
broad, white to cream; sepals 1.5-2.5 m long,
ciliate; petals 7-11 mm long, spreading; style
2-4 mm long; capsule 5-8 mm broad. Moist
sites in coniferous forest, 2450-3050 m, in
Beaver, Carbon, Duchesne, Emery, Juab, Salt
Lake, and Utah counties; widely distributed
in North America; Eurasia; 9(ii).
Pterospora Nutt.
Plants herbaceous saprophytes, devoid of
chlorophyll, tall, reddish or purplish brown,
the stems arising from a bulbous cluster of
coralloid roots; leaves alternate, simple,
scalelike, colored like the stems; flowers nu-
merous, borne in an elongate raceme, nod-
ding; calyx 5-lobed; corolla urn-shaped, the
tube much longer than the lobes; stamens 10,
the filaments flattened, tapering to the apex,
glabrous, the anthers with 2 recurved awns,
dehiscent almost throughout; ovary superior,
5-loculed, the stigma borne on a short thick
style; fruit a loculicidal capsule.
Pterospora andromeda Nutt. Pinedrops.
Plants erect, the stems simple, 2-8.5 (10) dm
tall, reddish brown, succulent, arising from a
cluster of roots to 5 cm in diameter, glandu-
lar-hairy, leafy only near the base; racemes
3-35 cm long or more; flowers 5-8 mm long,
nodding, axillary; pedicels 5-15 mm long, re-
curved; sepals oblong, glandular; corolla pale
yellow, depressed urn-shaped; capsule 8-12
(14) mm broad, 5-lobed, depressed globose.
Coniferous forest, 2300-2900 m, in Daggett,
Duchesne, Garfield, Grand, San Juan, Sum-
mit, Uintah, and Washington counties, and to
be expected at higher elevations elsewhere;
widely distributed in North America; 15(iii).
Pyrola L.
Rhizomatous herbs; leaves with chloro-
phyll, leathery, persistent, all basal or appar-
ently so, or rarely lacking and the plants then
partially or completely saprophytic; flowers
regular to irregular, borne in terminal ra-
cemes; sepals 5, united at the base; petals 5,
distinct, usually concave, deciduous; stamens
10, the filaments tapering to the apex, the an-
tliers unawned, pendulous, opening by means
of apparently terminal pores; ovary superior,
4-loculed, the stigma borne on a straight or
curved style; fruit a loculicidal capsule.
Copeland, H. F. 1947. Observations on struc-
ture and classification of the Pyroleae.
Madrono 9: 65-102.
1. Styles straight or nearly so; pores of anthers sessile; stigma usuallv much
broader than the style 2
Styles bent or curved; pores of anthers usually borne on short tubes; stigmas
only slightly broader than the styles 3
2(1). Styles 2 mm long or less, not (or seldom) exserted from the flower; flowers not
secund; petals pinkish to cream P. minor
Styles over 2 mm long, exserted from the flower; flowers secund; petals
greenish white P. secunda
3(1). Flowers pink to purplish; sepals longer than broad P. asarifolia
Flowers pale, greenish yellow; sepals broader than long P. virens
March 1980
Welsh: Utah Flora, Miscellaneous Families
55
Pyrola asarifoUa Michx. Liver-leaf Win-
tergreen. {P. rotimdifolia var. bracteata
(Hook.) Gray; P. asarifoUa var. hractcata
(Hook.) Jeps.; P. rotimdifolia var. purpurea
Bunge; P. asarifoUa var. purpurea (Bunge)
Fern.; P. iiiearnata Fisch. in DC; P. asari-
foUa var. inearnata (Fisch.) Fern.; P. asari-
foUa var. ovata Farw.; P. uliginosa T. & G. ex
Torr.; P. rotimdifolia var. uliginosa (T. & G.)
Gray; P. asarifoUa var. uUginosa (T. & G.)
Farw.; P. ekita Nutt.; P. bracteata var. hiUi J.
K. Henry). Plants 1.3-4 dm tall; leaves basal
or essentially so, the blades 1.3-7.5 cm long,
1.1-7.3 cm wide, oval, rotund, elliptic, or
obovate, subcordate to rounded, obtuse, or
acute basally, rounded to obtuse or emargi-
nate apically, entire to serrulate; petioles 1-9
cm long; racemes mostly 2- to 12-flowered;
pedicels 3-8 mm long; sepals longer than
broad, 1.5-4 mm long; petals pink to pur-
plish, 5-7 mm long; anthers pink, the pores
on short tubes; style curved, with a flaring
collar below the stigma. Coniferous and de-
ciduous woods, often along streams, or less
commonly in meadows, 1750-2750 m, in
Daggett, Duchesne, Emery, Garfield, Grand,
Iron, Juab, Piute, Rich, Salt Lake, Summit,
Uintah and Washington counties (and likely
elsewhere); Alaska east to Newfoundland and
south to California, New Mexico, South Da-
kota, and New England; Asia. Varietal status
of Utah materials is not clear; .30(iv).
Pyrola minor L. Lesser Wintergreen.
{Amelia tninor (L.) Alef.; Erxlebenia minor
(L.) Rydb.; P. minor var. conferta C. & S.; P.
conferta (C. & S.) Fisch. ex Ledeb.). Plants
0.8-2.4 dm tall; leaves basal, the blades (0.4)
1.1-3.3 cm long, (0.6) 0.9-2.5 cm broad, oval,
elliptic, or ovate, obtuse to rounded or sub-
cordate basally, obtuse to rounded apically,
crenate to subentire; petioles 0.2-3 cm long;
racemes mostly 5- to 13-flowered; pedicels
2-3 mm long; sepals 1-1.5 mm long, erose to
subentire; petals pale pink to cream, 3.5-4.5
mm long; anthers with pores sessile; style
straight, very short, not exserted from the co-
rolla, with a more or less distinctive collar
below the stigma. Wet stream sides and other
moist sites, usually in coniferous forests,
2150-2750 m, in Beaver, Daggett, Duchesne,
Garfield, Juab, Salt Lake, Sevier, Siunmit,
Uintah, and Washington counties; Alaska and
Yukon east to Greenland and south to Cali-
fornia and Colorado; circumboreal; ll(i).
Pyrola secimdti L. One-sided Wintergreen.
{Ramisehia secimda (L.) Garke; Actinocijclus
secundus (L.) Klotzsch; P. secimda var. obtii-
sata Turcz.; Orthilia secimda var. obtusata
(Turcz.) House; P. secunda var. pumila
Paine; P. secimda f. eiicycla Fern.). Plants
0.6-1.8 (2.1) dm tall; leaves basal or rarely
some cauline, or sometimes with a naked
stem below the leaves, the blades 1.3-4 (5)
cm long, 1-3 cm wide, ovate, oval, elliptic,
or orbicular, obtuse ro rounded basally, acute
to obtuse or rounded apically, crenate-ser-
rate; petioles 0.6-2 cm long; racemes mostly
4- to 15-flowered, the flowers secund; pedi-
cels 2-5 mm long; sepals 0.5-1.5 mm long;
petals greenish white, 4-6 mm long; anthers
with pores sessile; style straight, exserted
from the corolla, lacking a collar. Ground
layer in usually coniferous forests, 2000-3350
m, in Box Elder, Carbon, Daggett, Duchesne,
Garfield, Juab, Kane, Piute, Salt Lake, San
Juan, Sanpete, Summit, Uintah, Utah, and
Washington counties, broadly distributed in
North America; Eurasia. Segregation of our
materials into the various proposed in-
fraspecific categories seems unwarranted;
33(vi).
Pyrola virens Schweigg. in Schweigg. &
Koerte. Greenish Wintergreen. {P. chlorantha
Sw.; P. chlorantha var. saximontana Fern.; P.
virens var. saximontana (Fern.) Fern.; P. chlo-
rantha var. paucifolia Fern.; P. virens f.
paucifolia (Fern.) Fern.; P. chlorantha f.
paucifolia (Fern.) Camp). Plants 0.9-2.5 dm
tall; leaves basal, the blades 0.6-3.5 cm long,
0.5-3 cm broad, elliptic, oval, or obovate, ob-
tuse to rounded basally, rounded to obtuse
apically, crenate-serrate to subentire; petioles
0.8-6 cm long; racemes mostly 2- to 9-flow-
ered; pedicels 3-8 mm long; sepals 0.5-1.5
mm long; petals greenish yellow, 5-7 mm
long; anthers yellowish, the pores on elongate
tubes; style curved, with a flaring collar be-
low the stigma. Coniferous or deciduous
woods, often in moist sites, 2150-2750 m, in
Daggett, Duchesne, Piute, Salt Lake, Sum-
mit, and Uintah counties; widely distributed
in North America; Eurasia; 8(i).
56
Great Basin Naturalist
Vol. 40, No. 1
Resedaceae
Mignonette Family
Annual or perennial herbs with watery
sap; leaves alternate, simple, or pinnately to
siibpalmately divided; flowers perfect, ir-
regular, borne in terminal racemes; sepals (4)
5-6 (8), distinct; petals (4) 5-6 (8), unequal in
size, the upper one the largest, appendaged;
stamens 8 or more, borne on the upper side
of a rounded disk, the anthers 2-loculed; pis-
til 1, the ovary superior, 1-loculed, with usu-
ally 3 (2-6) carpels; style lacking; fruit a cap-
sule, usually open at the tip before maturity.
Reseda L.
Erect or ascending annual or perennial
herbs from a taproot; leaves alternate; flow-
ers greenish yellow; sepals subequal; petals
imequal; pistils 1, the carpels usually 3, open
toward the apex.
Reseda lutea L. Yellow Mignonette. Plants
simple or much branched, glabrous; leaves
pinnatifid or subpalmately divided; flowers
greenish yellow, numerous, borne in elongate
racemes; petals usually 6, each commonly
with 3 connate or distinct appendages; ovary
and capsule usually with 3 apical lobes. Cul-
tivated ornamental; rarely escaping in Utah;
1(0).
Tamaricaceae
Tamarisk Family
Shrubs or small to moderate trees; leaves
alternate, scalelike, exstipulate, entire; flow-
ers mostly perfect, regular, borne in spikelike
racemes arranged in panicles; sepals 4 or 5,
overlapping; petals 4 or 5, separate, more or
less overlapping, arising from the base of a
nectiferous disk; stamens usually as many or
twice as many as the petals, the anthers 2-lo-
culed; pistil 1, the ovary superior, unilocular,
usually 3 or 5 carpelled, the placentation bas-
al; stigmas 2-5, separate; ovules 2 per pla-
centa; fruit a capsule, the seeds comose.
Baum, B. R. 1967. Introduced and natural-
ized tamarisks in the United States and
Canada (Tamaricaceae). Baileya 15:
19-25.
Tamarix L.
Deciduous or evergreen shrubs or trees,
the branchlets deciduous; leaves clasping or
sheathing; flowers small, shortly pedicelled;
petals white to pink or lavender, inserted be-
low the disk; capsules dehiscent by 3-5
valves.
1. Leaves sheathing; evergreen trees of moderate size, restricted to Washington
County T. ophylla
Leaves not .sheathing, at most merely clasping; deciduous trees of small size or
merely shrubs of broad distribution 2
2(1). Flowers 4-merous, or the stamens sometimes more than 4; stamens emerging
gradually from the di.sk-lobes; plants uncommon both in cultivation and as
e.scapes T. pairiflora
Flowers 5-merous, or the stamens sometimes more than 5; stamens inserted un-
der disk near the margin between the emarginate lobes; plants abundant,
cultivated and otherwise T. ramosissima
Tamarix aphylla (L.) Karst. Athel Tamar-
isk. {Thuja aphylla L.) Trees to 10 m tall and
6 dm in diameter or more, the bark reddish
brown to gray; branchlets jointed; leaves
sheathing, minute, evergreen; bracts longer
than the pedicels; flowers 5-merous; sepals
entire, the inner ones slightly larger; petals
elliptic-oblong to ovate, 2-2.2 mm long,
early deciduous or with 1 or 2 persisting; sta-
minal filaments inserted between the disk
lobes. Cultivated sparingly in Washington
County, where it seldom flowers; native to
•Africa and the Middle East; introduced in
California, Nevada, Arizona, and Texas; 2(i).
Tamarix parviflora DC. Small-flowered
Tamarisk. Shrubs or small trees to 5 m tall;
bark brown to deep purple; branchlets not
jointed; leaves merely sessile, not .sheathing.
March 1980
Welsh: Utah Flora, Miscellaneous Families
57
deciduous with the branchlets; bracts longer
than the pedicels, more or less translucent;
flowers 4-merous; sepals erose-denticulate,
the outer two keeled and acute, the inner flat
or slightly keeled and obtuse; petals oblong
to ovate, 1.9-2.3 mm long, persistent; stainin-
al filaments arising gradually from disk-lobes.
Cultivated and naturalized along streams and
seeps, in Emery, Kane, Utah, and Washing-
ton counties, and to be expected elsewhere;
introduced from southern Europe and now
widespread in Canada and the United States;
7(i).
Tamarix ramosissima Ledeb. Branched
Tamarisk; Salt Cedar. (T. gaUica authors, not
L.; T. pentandra authors, not Pall.). Shrubs or
small trees to 6 m tall, or rarely more; bark
reddish brown; branchlets not jointed; leaves
merely sessile, not sheathing, deciduous with
the branchlets; bracts longer than the pedi-
cels, scarious but scarcely translucent; flow-
ers 5-merous; sepals erose-denticulate, the
outer 2 narrower than the inner, all more or
less acute; petals obovate, 1-1.8 mm long,
persistent; filaments inserted under the disk
near the margin between the emarginate
lobes. Cultivated and naturalized along seeps,
streams, and reservoirs, almost throughout
Utah (Carbon, Davis, Duchesne, Emery, Gar-
field, Grand, Juab, Kane, Millard, San Juan,
Sevier, Tooele, Uintah, Utah, Wasatch,
Washington, Wayne, and Weber counties);
introduced from Eurasia, now widespread in
the southern United States; 99(xix).
Tiliaceae
Linden or Bas.swood Family
Trees; leaves alternate, simple, serrate to
obscurely lobed, usually oblique, stipulate;
flowers regular, perfect, borne in cymes; sep-
als 5, distinct or more or less connate; petals
5, alternate with the sepals; stamens numer-
ous, the filaments free or connate in bundles
of 5-10; ovary superior, 5-loculed; fruit dru-
paceous.
TiLIA L.
Cultivated trees; leaves long-petioled, the
blades obliquely cordate, serrate or doubly
so, sometimes obscurely lobed; flowers in
long-peduncled cymes, the peduncle adnate
at its base to a ligulate bract; sepals 5; petals
5; stamens numerous, distinct or in 5 clusters,
sometimes bearing petaloid staminodia oppo-
site the petals; ovary 5-loculed, the stigma 5-
lobed; fruit subglobose, 1- to 3-seeded.
1. Branchlets and petioles densely white-hairy; leaf blades white stellate hairy
beneath T. tomentosa
Branchlets and petioles glabrous or nearly so; leaf blades variously pubescent
or glabrous 2
2(1). Leaf blades hairy (sometimes thinly so) over the lower surface and usually
along the veins beneath '^
Leaf blades glabrous beneath, except in vein axils 5
3(2). Leaf blades densely white or brown stellate hairy beneath T. heterophylla
Leaf blades variously hairy but the surface not obscured by hairs 4
4(3). Hairs of lower leaf surface stellate, at least some; flowers with staminodes
T. neglecta
Hairs of lower leaf surface all simple; flowers without staminodes
T. platijpliyllos
5(2). Leaf blades deflnitely glaucous beneath, usually less than 8 cm long; flowers
lacking staminodes T. cordata
- Leaf blades green or merely pale green beneath, the largest usually more than
8 cm long; flowers with or without staminodes 6
58
Great Basin Naturalist
Vol. 40, No. 1
6(5). Flowers with staminodes; leaves serrate to doubly serrate with long-acuminate
teeth, the largest blades on flowering stems to 10 cm long or more
T. atnericana
Flowers without staminodes; leaves serrate with short acute teeth, the largest
blades on flowering stems usually less than 10 cm long T. europaea
Tilia atnericana L. American Linden.
Moderate to large trees of streets and other
ornamental plantings, common in Salt Lake,
Utah, and Weber counties, and probably
grown elsewhere; indigenous to the eastern
states and Canada; 9(o).
Tilia cordata L, Small-leaved European
Linden. Small to large trees of ornamental
plantings; common in Box Elder, Cache,
Juab, Salt Lake, Utah, and Weber counties;
widely cultivated in North America; in-
troduced from Europe; 12(o).
Tilia X europaea L. Common or European
Linden. Moderate to large trees of ornamen-
tal plantings, uncommon in Utah; indigenous
to Europe. This tree is reputed to be a hybrid
derivative of T. cordata x T. platijphyllos;
2(0).
Tilia heterophylla Vent. White Basswood.
Large ornamental trees, uncommon in Utah;
indigenous to the eastern United States; 2(o).
Tilia neglecta Spach. Moderate to large or-
namental trees, uncommon in Utah; in-
digenous to the eastern United States and
Canada. This taxon resembles, and apparent-
ly intergrades with, T. atnericana, with which
it is very closely allied; 2(o).
Tilia platyphyllos Scop. Large-leaved
Linden. Moderate to large ornamental trees,
common in Salt Lake, Utah, and Weber
counties, and probably elsewhere; indigenous
to Europe; 8(o).
Tilia tomentosa Moench. Silver Linden.
Moderate to large ornamental trees, moder-
ately common in Cache, Juab, Salt Lake,
Utah, and Weber counties; indigenous to
eastern Europe and Asia Minor; 5(o).
THE TAXONOMIC STATUS OF THE ROSY BOA
LICHANURA ROSEOFUSCA (SERPENTES: BOIDAE)
John R. Ottley', Robert W. Murphy-, and Geoffrey V. Smith'
Abstract. — Evidence is presented indicating that lAchanura roseofusca and Lkhanum triiirgata are conspecific.
Data include the report of an intermediate specimen from El Arco, Baja California Norte, a site midway between
the previously known peninsular ranges of the two species; captive hybridization provides additional support for the
conclusion.
The close relationship of the boas Lich-
anurci triiirgata Cope and Lichanura roseo-
fusca Cope has long been recognized, due
principally to the overlap of most scale char-
acters and because the desert boa L. roseo-
fusca gracia Klauber appears to be an inter-
mediate between the two species (Klauber
1931). The problem in establishing their rela-
tionship stems from the rather broad gaps be-
tween their known ranges in central Baja
California and southwestern Arizona.
Subsequent to the description of L. r.
gracia, Klauber (1933) reported a single spec-
imen from Guaymas, Sonora. This specimen
agrees exactly with L. trivirgata in coloration
but has scale counts resembling those of L.r.
gracia. He stated that the specimen might be
considered an intergrade of L. trivirgata and
L.r. gracia. This is somewhat surprising since
he restricted L. trivirgata to the cape region
of Baja California, thus necessitating a trans-
gulfian dispersal of trivirgata to facilitate
hybridization. Gorman (1965) reemphasized
the wide variation in meristic characters
within the genus, as first demonstrated by
Stejneger (1891), and referred to Klauber s
(1933) scale counts and color descriptions as
evidence indicating that the populations
from southern Arizona, Sonora, and southern
Baja California are all one form, L. trivirgata
(all have three primary stripes of chocolate
brown on a light drab background).
The variation seen in the genus led Klau-
ber (1931, 1933) to speculate that we might
be dealing with a single, polytypic species, L.
trivirgata. He suggested, however, that before
such a designation be considered we should
await the collection of more material from
regions of potential hybridization.
Gorman (1965) and Bostic (1971) com-
mented on new material from the range gaps
and stated that the basis was yet lacking for
uniting the two species because of the great
uniformity of L. trivirgata throughout its
range and the absence of obvious intergrades.
The range gaps were shown to be separations
of approximately 160 km (100 miles) in both
central Baja California and southwestern Ari-
zona. In spite of these appraisals, several au-
thors (Miller and Stebbins 1964, Lowe 1964,
Soule and Sloan 1966) have proposed, in ad-
vance of adequate evidence, to imite the two
species. The needed evidence is reported in
this paper.
During the summer of 1979, an unusual
specimen of L. trivirgata (Fig. 1) was collect-
ed at the town known as El Arco, Baja Cali-
fornia Norte (28°02'N, 113°27'W). The spec-
imen, taken as it was crossing the road in
front of the military base on 17 July at 2225
hours by Kenneth A. Stockton, is unique for
two reasons. First, its coloration" and scale
counts are intermediate between the two spe-
cies. Second, the geographic location of El
Arco is midway between the previously re-
ported limits for the two species (Bostic
'Life Science Museum, Brighani Young University, Provo, Utah 84602.
^Department of Biology. UCLA, Los Angeles, California 90024.
'Alia Mira .\nimal Clinic, VLsta, California 9208.3.
Color characters with numbers refer to the color-name charts by Kelly (1958).
59
60
Great Basin Naturalist
Vol. 40, No. 1
1971). Scale counts are as follows: 224 ven-
trals, 48 subcaudals, 41 dorsal scale rows,
15-14 supralabials, 15-16 infralabials, and
10-11 oculars. The specimen is an adult male
measuring 577 mm total including the 85 mm
tail. The coloration and color pattern consists
of three primary stripes of deep brown (No.
56) on a ground of light gray olive (No. 109).
When one considers all these characters, the
El .\rco specimen appears to be the obvious
intergrade spoken of by Gorman (1965). Al-
though this report essentially closes the range
gap on the Baja California peninsula, a gap
vet remains between the Kofa Mountains and
Organ Pipe Cactus National Park in south-
western Arizona. Fowlie (1965) has indicated
in a range map that trivirgata and grcicia
overlap in the region of the Growler Moun-
tains southwest of Ajo. If two subspecies are
in fact found together in the area, we would
expect to see the effects of intergradation. No
such evidence has ever been reported or are
we aware of any specimens that substantiate
such a claim. We must therefore (juestion the
validity of Fowlie's range for gracia in the
Ajo region.
Notes on Captive Breeding
Recent captive breeding experiments have
produced enlightening results. In April 1975
a male L.t. roseofusca from San Diego, Cali-
fornia, was bred to a female L.t. trivirgata
from Cabo San Lucas, Baja California Sur.
On 7 August 1975 three young were born,
two of which died within a few days; how-
ever, the third specimen, a male, is alive at
the time of this writing and in our possession
(Fig. 2). Coloration and color pattern consist
of three primary stripes of medium brown
(No. 56) on a light olive gray (No. 112) back-
ground. The stripes are moderately serrated,
yet fairly uniform. Scale counts are as fol-
lows: ventrals 2.31, subcaudals 47, dorsal scale
rows 41, supralabials 14-14, infralabials
17-15, and oculars 10-10. Another cross, in-
volving a male L.t. trivirgata from near San
Bartolo, Baja California Sur, and a female of
the same subspecies from the vicinity of Esta-
cion Ortiz, Sonora, occurred in March 1976.
Four young were born on 29 July 1976. A fe-
male from that litter (Fig. 3) yet remains in
our possession. Coloration and pattern are of
Fig. 1. Dorsal view of a Lkhunum triim^ala x wscofusai iiitcnnfcliato tioin El Arco. Baja California \ortt
March 1980
Ottley et al.: Rosy Boa
61
Fig. 2. Dorsal view of a Lichaniini tritirgata x roscufiisca hybrid; male parent is a L. t. wscofttsca from San Diego,
California, and the female parent is a L. t. triiirgata from Cabo San Lucas, Baja California Sur.
Fig. .3. Dorsal view of a Lichanura t. triiin^dta transgnlfian cross; male parent is a L. t. trivirgata from near San
Bartolo, Baja California Sur, and the female parent is a L. t. trivirgata from Estacion Ortiz, Sonora.
62
Great Basin Naturalist
Vol. 40, No. 1
three primary stripes of chocolate brown on
a cream ground. The stripes are uniform with
shghtly serrated edges. Scale counts are as
follows: ventrals 220, subcaudals 44, dorsal
scale rows 38, supralabials 13-13, infralabials
15-14, and oculars 10-11. The female L.t. tri-
virgata transgulfian cross and a male desig-
nated as L.t. gracia from near Punta Prieta,
Baja California Norte, were observed cop-
ulating on 16 May 1979. On 24 October 1979
three young were bom, all males, each bear-
ing well-delineated medium brown stripes
and a ground of color intermediate between
the parents.
Acknowledgments
We thank Wilmer W. Tanner and Kent M.
Van De Graaff for their constructive criti-
cisms and comments in reviewing this paper,
Vickie R. Ottley for typing the manuscript,
and Lawrence E. Hunt, Kenneth A. Stockton,
and Dale M. Stockton for their help and
companionship in the field. Scientific collec-
ting permit 30/832/79 was issued by Ignacio
Ibarrola Bejar, director general of the De-
partamento de la Conservacion de la Fauna
Silvestre.
Summary
In coloration the El Arco specimen and the
captive bred trivergata x roseofusca hybrid
are very similar, differing only in the latter
having moderately serrated stripes. These
data support our consideration of the El Arco
specimen as an intermediate. Since El Arco is
situated in a geographical region midway be-
tween "pure" trivergata and roseofusca, in-
dicating a continuous range, we find no alter-
native to considering the two taxa as being
conspecific. The binomial Lichanura trivir-
gata Cope has priority over L. roseofusca
Cope by publication date. Accordingly, we
recognize L. trivirgata as a single, polytypic
species with four subspecies (L. t. trivirgata
Cope, L. t. roseofusca Cope, L. t. gracia
Klauber, and L. t. bostici Ottley).
Literature Cited
BosTic, D. L. 1971. Herpetofauna of the Pacific Coast of
north central Baja CaUfomia, Mexico, with a de-
scription of a new subspecies of PhyUodactylus
xanti. Trans. San Diego Soc. Nat. Hist.
16(10):237-263.
FowLiE, J. A. 1965. The snakes of Arizona. Azul Quinta
Press, Fallbrook, California, 164 pp.
Gorman, G. C. 1965. The distribution of Lichanura tri-
virgata and the status of the species. Herpetolo-
gica 21(4):2a3-287.
Kelly, K. L. 1958. ISCC-NBS color-name charts illus-
trated with centroid colors. National Bureau of
Standards, NBS Circular 533.
Klauber, L. M. 1931. A new subspecies of the Califor-
nia Boa, with notes on the genus Lichanura.
Trans. San Diego Soc. Nat. Hist. 6(20):305-318.
19.33. Notes on Lichanura. Copeia (4):214-215.
Lowe, C. H. 1964. The vertebrates of Arizona: Anno-
tated check lists. Tucson, University of Arizona
Press.
Miller, A. H., and R. C. Stebbins. 1964. The lives of
desert animals in Joshua Tree National Mon-
ument. Berkeley and Los Angeles, University of
California Press.
Ottley, J. R. 1978. A new subspecies of the snake Lich-
anura trivirgata from Cedros Island, Mexico.
Great Basin Nat. .38:411-416.
SouLE, M., AND A. J. Sloan. 1966. Biogeography and dis-
tribution of the reptiles and amphibians on is-
lands in the Gulf of California, Mexico. Trans.
San Diego Soc. Nat. History 14(11):137-156.
Stejneger, L. 1891. On the snakes of the California
genus Lichanura. Proc. U.S. Nat. Mus.
14:511-515.
HESPEROPERLA HOGUEI, A NEW SPECIES OF STONEFLY
FROM CALIFORNIA (PLECOPTERA: PERLIDAE)
Richard \V. Baiiinaiin' and Bill P. Stark-
.\bstr.\ct.— a new species of Hcspcropcila is named from northern California. The adult male and female,
nymph, and egg are described and figured. Hespcroperla lioouci adds a second species to this previously monotypic
genus.
The genus Hcsperoperla Banks (1938) was
not accepted by other workers until the re-
cent world catalog (lilies 1966). Studies by
Needham and Claassen (1922), Claassen
(1940), and Frison (1942) placed six species in
synonymy luider Acroneiiria pacifica Banks,
including Hesperoperia obscura (Banks), the
designated type species.
Stark and Gaufin (1976), in their revision
of the Perlidae, confirmed that Hesperoperia
was indeed a separate genus with one valid
species, Hesperoperia pacifica (Banks).
In the fall of 1976, Charles L. Hogue of
the Natural History Museum, Los Angeles
County, sent a distinctive female perlid to
the senior author for identification. When it
proved to belong to Hesperoperia, the help of
numerous colleagues, including Bill P. Stark,
was enlisted.
The types of several species in the A. pa-
cifica synonymy were examined, including:
A. pacifica Banks, A. nigrita Banks, A. pu-
mila Banks, and A. obscura Banks. The spe-
cific name nigrita suggested that its type
might be dark and distinctive, but this was
not the case. The type locality of A. pumila.
Three Rivers, California, which is near Vis-
alia, is a short distance south of the known
range of this new species, but the specimen
was positively H. pacifica.
Several additional specimens were sub-
sequently found that confirmed that this was
an undescribed species of Hesperoperia with
a known range that included most of north-
em California.
Both species of Hesperoperia occur in the
same streams with no intergradation; thus
they are sympatric species.
Hesperoperia hoguei, n. sp.
Figs. 1-9
Male.— Macropterous. Length of fore-
wings (18-20 mm; length of body 16-18 mm.
Dorsum of head mostly yellow, with brown
U-shaped pattern connecting ocelli, posterior
lateral margins brown behind compound
eyes, sometimes with faint brown area near
frontal margin. Pronotum with broad yellow
median stripe, lateral margins dark brown,
rugosities distinctive and slightly lighter than
lateral margins (Fig. 4). Legs brown. Wings
light brown, veins brown. Abdomen yellow
with narrow brown lateral stripes. Tergum
10 with median tergite, segment covered
with medium-length hairs. Sternum 9 with
large quadrangular hammer. Paraprocts
sclerotized, broad basally, apex narrow and
pointed (Fig. 2). Aedeagus with two bands of
large spinules, apical band broad, consisting
of 12 or more closely set spinule rows, basal
band narrow, interrupted on both dorsomesal
and ventromesal surfaces; small spinules pres-
ent in patches near base and at dorsal apex
(Figs. 5-6).
Female.— Macropterous. Length of fore-
wings 26-28 mm; length of body 24-26 mm.
Color pattern similar to male. Subgenital
plate strongly produced, with more darkly
sclerotized area along mesoposterior margin
'Monte L. Bean Life Science Museum and Department of Zoology, Brigham Young University, Provo, Utah 84602.
'Department of Biological Sciences, Mississippi College, Clinton, Mississippi 39056.
63
64
Great Basin Naturalist
Vol. 40, No. 1
Fig. 1. Hesperoperht /logi/ei, mature nymph, habitus.
March 1980
Baumann, Stark: California Stonefly
65
2
3
11
5
Figs. 2-6. Hesperoperla hogtici: (2) male terminalia, dorsal; (3) female terminalia, ventral; (4) adult, head and pro-
notum; (5) aedeagiis, dorsal (.50X, inset lOOX); (6) aedeagiis, lateral (SOX, inset lOOX).
66
Great Basin Naturalist
Vol. 40, No. 1
(Fig. 3). Vagina, spermathecum, and acces-
sory glands membranous.
Egg.- Outline oval; cross-section circular.
Collar stalked, margin flanged and irregu-
larly incised; collar end of egg with regular
indentations which end at terminal margin.
Chorion smooth. Micropyles arranged cir-
cumlinearly in apical end opposite of collar
(Figs. 7-9).
Nymph.— General color dark brown, pat-
terned with yellow markings. Occiput with
an irregularly spaced row of spinules. Post-
ocular fringe present. Head mostly dark, with
distinctive inverted yellow W-shaped pattern
located anterior to compound eyes, frontal
margin entirely yellow. Pronotum with later-
al setae sparse or absent. Abdominal terga
with numerous small intercalary spinules.
Cerci with fringe of spines at segmental
joints, never longer than segments; few tiny
intersegmental spinules present. Proventricu-
lus with teeth in 12 longitudinal bands; acces-
sory bands and structures absent. Thoracic
and anal gills present (Fig. 1).
Diagnosis.— Hespemperla fwguei can be
easily separated from H. pacifica in the nym-
phal stage by the difference in head pattern
and the abdominal spinulation. Hesperoperla
pacifica exhibits a large, inverted, mushroom-
shaped pattern anterior to the compound
eyes that terminates in an enlarged base on
the mesoanterior margin, and H. hoguei bears
an inverted W-shaped pattern and a separate
broad yellow band running the full width of
the anterior margin. Intercalary spinules are
entirely absent from the abdominal terga of
mature H. pacifica nymphs but are numerous
in H. hoguei specimens. Claassenia sabulosa
nymphs also have an inverted W-shaped pat-
tern on the head, but they possess a complete
occipital ridge.
Adults of H. pacifica have a plain yellow
brown pattern on their head and pronotum.
Those of H. hoguei have a broad yellow me-
dian area set off by distinctive dark lateral
margins. The external genitalia are quite sim-
ilar, but the basal spinule band on the ae-
deagus of the males is different. In H. pacifi-
ca the band is only broken ventrally, but in
H. hoguei the band is broken both dorsally
and ventrally.
Types.- Holotype: $ , Gibson Creek, 800
ft, 1 mile west of Ukiah, Mendocino Co., Cal-
ifornia, 6-IX-1976, C. L. Hogue. Allotype:
Toadtown, 3000 ft, 4 miles SW Stirling City,
Butte Co., California, 9-IX-1976, C. L.
Hogue.
Paratypes: Butte Co.: Paradise, 25-V-1966,
Lowe, 2 females (CSUC); Tehama Co.: Big
Chico Creek, Hwy. 32, 14-IX-1979, G. L.
Boles, 13 males, 7 females (GLB) (BYU). Ad-
ditional specimens: Butte Co.: Big Chico
Creek, 580 ft, V4 mile below Salt Springs,
Bidwell Park, P-VIII-1972, M. W. Kainu,
nymph (UCD); 2 miles SW Stirling City, 20-
VI- 1979, J. A. Stanger, nymphs (BYU).
Plumas Co.: Sulphur Creek, Hwy. 89, 5-VII-
1979, B. P. Stark and K. W. Stewart, nymphs
Figs. 7-9. Hesperoperla hoguei: (7) egg, lateral (200X); (8) egg, collar end {400X); (9) egg, micropyles (700X).
March 1980
Baumann, Stark: California Stonefly
67
(BPS). SJiasta Co.: unnamed spring tributary
to Lake Britton, 20-IX-1978, G. L. Boles,
nvniphs (GLB); South Fork Bear Creek, 12-
v'lI-1979, G. L. Boles, nymphs (GLB). Te-
hama Co.: Big Chico Creek, Hwv. 32, 14- V-
1978, G. L. Boles, nymph (BYU),' same data,
2-IX-1978, nymphs (2 females emerged 28-
IX- 1978) (GLB) (BYU).
Holotype and allotype deposited at the
Natural History Museum, Los Angeles Coun-
ty-
Etymology.— This species is named in
honor of Dr. Charles L. Hogue, Senior Cura-
tor of Entomology, Natural History Museum,
Los Angeles County. He has collected many
interesting stoneflies during his studies on the
torrenticolus insects of the New World.
Acknowledgments
We thank the following individuals for the
help that we received during this study: Ger-
ald L. Boles, California Department of Water
Resources, Red Bluff, California (GLB); Dr.
Charles L. Hogue, Natural History Museum,
Los Angeles County, California (LACM); Dr.
David H. Kistner, California State University,
Chico, California (CSUC); Dr. Robert O.
Schuster, University of California, Davis,
California (UCD); Jean A. Stanger, Brigham
Young University, Provo, Utah (BYU); Dr.
Kenneth W. Stewart, North Texas State Uni-
versity, Denton, Texas.
The drawings were made by Connie A. Be-
van Bhagat.
Literature Cited
Banks, N. 1938. A new genus of Perlidae. Psyche
45:136-1.37.
Claassen, p. W. 1940. A catalogue of the Plecoptera of
the world. Mem. Cornell Agr. Exp. Station
232:1-235.
Prison, T. H. 1942. Descriptions, records and systematic
notes concerning western North .American stone-
flies (Plecoptera). Pan-Pac. Entomoi. 18:61-73.
Illies, J. 1966. Kataiog der rezenten Plecoptera. Das
Tierreich, Berlin, 82, 632 pp.
Needham, J. G., AND P. W. Claasse.n. 1922. The North
American species of the genus Acronetiriti (Order
Plecoptera). Canad. Entomoi. 54:249-255.
Stark, B. P., and A. R. Gaufin. 1976. The nearctic gen-
era of Perlidae (Plecoptera). Misc. Pubi. Entomoi.
Soc. Amer. 10:1-77.
REPRODUCTION IN THREE SYMPATRIC LIZARD SPECIES
FROM WEST-CENTRAL UTAH
John B. Andre' and James A. MacMahon'
.\bstr.-vct.- Data on reproduction by the lizards Uta stanshiiriana. Ciotaphytus colkiris, and Cnciindophorus tigris
are presented from a communitv where they are sympatric in west-central Utah. Data are compared to a previous
studv of these species at the same site and to data from other sites in the United States.
Lizard reproductive data from geographi-
callv separated populations are important to
ecologists attempting to explain a highly var-
iable species characteristic. Reproduction by
Uta stansburiana has been well documented
in the literature (Fautin 1946, Medica and
Turner 1976, Nussbaum and Diller 1977,
Tanner 1965, Tinkle 1961, 1967, Turner et
al. 1970, 1973, 1976, Worthington and Ar-
vizo 1973, Parker 1974, Parker and Pianka
1975, Tinkle and Hadley 1975, and Goldberg
1977). Reproductive studies of Ciotaphytus
coUaris and Cnetnidophorotis tigris are scarce
(Fautin 1946, Turner et al. 1969, Pianka
1970, Burkholder and Walker 1973, Parker
1973, and Vitt and Ohmart 1977). This paper
presents data on the reproduction by three
lizard species {Uta stansburiana stansbu-
riana, Crotaphytus collaris bicinctoris, and
Cnemidophorus tigris tigris), coexisting in
west-central Utah.
We are cognizant of the limitations im-
posed by the short period covered by our col-
lections. Despite this, there is clearly a need
for carefully collected data on reproduction
by lizards, or other taxa, so that one might
gather such data into a body of information
used to address general evolutionary theorv
(Tinkle 1969a, b. Tinkle et al. 1970).
Methods
Lizards were collected in Tule Valley, Mil-
lard County, Utah (lat. 39°13'N,' long.
113°27'W). Tule Valley, bordered on the east
and west by mountain ranges, is typical of
the basin and range topography of the Great
Basin Desert (MacMahon 1979). Bajadas (coa-
lesced alluvial fans) slope from the bases of
both mountain ranges to the playa that cov-
ers most of the valley floor.
The study site was located in the Tetra-
dy?nia glabrata and Atriplex confertifolia
communities described by Fautin (1946). The
common plant species were T. glabrata, A.
confertifolia, Chrysothamnus viscidiflorus,
Artemisia spinescens. Ephedra nevadensis,
Ceratoides (Eurotia) lanata, and Hilaria
jamesii. The substrate was mostly small rocks
embedded in packed soil, with localized
areas containing large boulders, which were
used as basking /perching sites by C. coUaris.
Lizards were collected (shot) throughout
the day, at three-week intervals between 1
April and 29 August 1976. Specimens were
preserved in 10 percent formalin within two
hours of collection. Analysis of reproductive
state (for females) and measurement of snout-
vent length (SVL) were made in the labora-
tory.
The sex of each specimen was determined
by dissection. The reproductive tracts of the
females were removed and the number of
corpora lutea, yolked follicles and/ or oviduc-
al eggs were recorded. Estimates of clutch
size were based on the number of yolked fol-
licles > 2.5 mm diameter and/ or oviducal
eggs and corpora lutea for U. stansburiana
'This paper is a contribution from the Department of Biology and the Ecology Center, Utah State University, Logan, Utah 84322. Reprint requests sh
be sent to James A. MacMahon. Present address for John B. Andre is Cape Romain National Wildlife Refuge, R.R. 1, Box 191, Awendaw, South Car,
should
rolina
68
March 1980
Andre, MacMahon: Utah Lizards
69
and volked follicles > 5.0 mm diameter
and/or oviducal eggs and corpora liitea for
C. coUam and C. tigris.
Results
Mean clutch size and mean SVL of sex-
ually mature females of each species are list-
ed in Table 1. The relationship between
clutch size and SVL is illustrated for U.
stanshuriona in Figure I. Analyses of similar
data for C. coUaris and C. tigris showed no
significant correlation (F-tests). The line in
this figure was determined by linear regres-
sion, the correlation coefficient is given for
the data set.
Uta stanshuriana females reach sexual
maturity in their second growing season (10
mo. old) at about 40 mm SVL (the smallest
female having yolked follicles was 37.0 mm
SVL, see Table 2). Most U. stansburiana
emerged from hibernation bv the first week
of April and bred shortly after this time.
Yolked follicles and oviducal eggs were pres-
ent from 1 April to 16 May; only oviducal
eggs were found from 6 Jiuie to 29 June.
From 17 July through 29 August no yolked
follicles or oviducal eggs were found in the
females collected. Though yolked follicles
and oviducal eggs were present in the fe-
males from April to the end of June, females
contained the most oviducal eggs between 24
April and 15 May. While we believe that fe-
males laid one or two clutches of eggs in
1976, our data are not exten.sive enough on
this point. Turner et al. (1970) have warned
of the problem of determining clutch fre-
quency with too few observations.
Crotaplujtus coUaris females are sexually
mature at about 85 mm SVL. Yolked follicles
were present in the single specimen collected
15 May. All females collected in the first
week of June contained oviducal eggs. At the
end of June no females contained yolked fol-
licles or oviducal eggs.
Cnemidophorus tigris females attain sexual
maturity abovit 73 mm SVL. Specimens col-
lected during the first and last weeks of June
CLUTCH
SIZE
5
.Ufa
•
4
,
t •
•
•
4 4
3
•
10 9
5
f% \ • I V •
2
,
t-rr
^4 4
V #^#^ *• V •
• •
1
y = -6.448+0.216 X
r = 0.605
36
40
42
44
46
48
50
SNOUT-VENT LENGTH (mm)
Fit;. 1. Relationship hctufi-ii tlutcli si/c iiiuiiil)t'n and S\'I, mini) tor I'. sldiisbitrUm
Table 1. Clutch size and SVL of adult female Vta stansburiana, Crotaphytus collaris and Cnemidophorus tigris.
N
SVL (mm)
Clutch size
Species
X
Range
X Range
Uta stanshuriana
Crotaphytus coUaris
Cnemidopliorus tigris
96
13
15
44.61
90.69
a3.13
.39.5-49.0
85.5-99.0
73.0-96.0
2.99 2-5
5.38 3-7
3.07 2-5
70
Great Basin Naturalist
Vol. 40, No. 1
contained yolkfed follicles and/or oviducal
eggs. After mid-July, no females contained
yolked follicles or oviducal eggs.
The first hatchlings observed were: U.
stansburiana, 17 July; C. collaris, 9 August;
C. Tigris, 7 August.
Discussion
Mean clutch size for U. stansburiana was
2.99 (range 2-5). Fautin (1946) reported a
mean clutch size of 4.1 (range 3-5) from the
vicinity of our study site. For northern popu-
lations (Tooele Coimty, Utah) Parker and
Pianka (1975) found a mean clutch size for
Vta of 4.6; Nussbaum and Diller (1977) in
Oregon found 3.3. Both Parker and Pianka
(1975) and Nussbaum and Diller (1977) re-
ported that Vta produces one or two clutches
per season. At our site, one, and for some fe-
males perhaps two, clutches of eggs were laid
by Uta in 1976.
Parker and Pianka (1975) also reported
that oviducal eggs were present during a
three- to four-month period; our data agree.
The relationship between SVL and clutch
size indicates that larger females produce
larger clutches. An F test shows a significant
relationship between SVL and clutch size
(0.05 level). Other workers report the same
relationship between SVL and clutch size for
Uta stansburiana from other parts of its
range (Tinkle 1961, Turner et al. 1973, Park-
er and Pianka 1975, and Goldberg 1977).
By mid-July Uta is in postreproductive
condition: the reproductive tracts of both
males and females have decreased in size.
This size decrease is accompanied by an in-
crease in the size of the fat bodies. Fat bodies
continue to enlarge as the growing season
progresses.
Little information exists on the reproduc-
tion of C collaris. In southern New Mexico,
Parker (1973) reported a mean clutch size of
5.3 (range 3-7). Robison and Tanner (1962)
reported a mean clutch size of 6.7, but the
lizard was collected from many different
parts of its range. Our data show a mean
clutch size of 5.38 (range 3-7).
The relationship between SVL and clutch
size is illustrated in Figure 2. A loose correla-
CLUTCH ^
SIZE 5
88 90 92 94 96
SNOUT-VENT LENGTH (mm)
Fig. 2. Relationship between clutch size (number) and SVL (mm) for C. collaris.
98
lOO
Table 2. Monthly clutch size and SVL of adult female Uta stansburiana.
Month
April
May
June
July
August
SVL (mm)
Clutch size
N
X
Range
X
Range
21
43.17
37.0-47.5
2.39
2-4
13
44.38
41.5-48.0
3.46
2-5
20
42.73
38.0-47.5
2.45
2-4
20
42.80
38.0-49.0
2.47
2-4
22
43.50
40.0-48.0
2.76
2-4
March 1980
Andre, MacMahon: Utah Lizards
71
tion exists, with larger females producing
larger clutches; an F test (0.05 level) is not
significant.
Yolked follicles and/ or oviducal eggs were
present from mid-May to mid-June in C. col-
laris females. By the end of June this species
is postreproductive; the reproductive tracts
are decreasing in size and the fat bodies are
enlarging.
Cneinidophoms tigris becomes active near
the end of April. Mating occurs shortly after.
Yolked follicles and/or oviducal eggs were
present from June to the first week of July.
Bv mid-July this species is postreproductive;
they exhibit small reproductive tracts and en-
larging fat bodies.
Mean clutch size was 3.07 (range 2-5),
whereas Fautin (1946) reported a mean
clutch size of 6.7 (range 5-9). The relation-
ship between SVL and clutch size is similar
to that of C. coJIaris, with larger females ten-
ding to produce more eggs (Fig. 3), although
an F test (0.05 level) is not significant. Pianka
(1970) and Vitt and Ohmart (1977) report a
relationship between SVL and clutch size
that is "loosely correlated" for C. tigris.
Acknowledgments
This work was made possible by the
US /IBP Desert Biome fimded by the Nation-
al Science Foundation (Grant GB32139).
Linda Finchum typed the manuscript. Robert
Bayn executed the figures.
Literature Cited
BuRKHOLDER, G. L., AND J. M. Walker. 1973. Habitat
and reproduction of the desert whiptail lizard,
Cncmidophonis tigris Baird and Girard in south-
western Idaho at the northern part of its range.
Herpetolot^ica 29:76-8.3.
Fautin. R. W. 1946. Biotic communities of the northern
desert shrub biome in western Utah. Ecol. Mon-
ogr. 16;251-31().
Goldberg, S. R. 1977. Reproduction in a mountain pop-
ulation of the side-blotched lizard, Uta stansbttr-
iana (Reptilia, Lacertilia, Igiianidae). J. Herpetol.
11:31-35.
MacMahon, J. A. 1979. North American deserts: Tlieir
floral and faiuial components. Pages 21-82 in R.
Perry and D. Goodall, eds. Arid land ecosystems:
Their structure, fvmctioning and management.
Vol. 1. Cambridge University Press, Cambridge.
Medica, p. a., and F. B. Turner. 1976. Reproduction
by Uta stansburiana (Reptilia, Lacertilia, Igua-
nidae) in southern Nevada. J. Herpetol.
10:123-128.
Nussbaum, R. a., and L. V. Diller. 1977. The life his-
tory of the side-blotched lizard, Uta stansburiana
Baird and Girard, in north-central Oregon.
Northwest Sci. 50:24.3-260.
Parker, W. S. 1973. Notes on the reproduction of some
lizards from Arizona, New Mexico, Texas and
Utah. Herpetologica 29:258-264.
1974. Home range, growth and population den-
sity of Uta stansburiana in Arizona. J. Herpetol.
8:135-139.
P.ARKER, W. S., AND E. R. PiANKA. 1975. Comparative
ecology of populations of the lizard Uta stansbur-
iana. Copeia 1975:61.5-6.32.
PiA.NiCA, E. R. 1970. Comparative autecology of the liz-
ard Cneinidophorus tigris in different parts of its
geographic range. Ecology 51:703-719.
RoBisoN, W. G., Jr., a.nd W. VV. Tanner. 1962. A com-
parative study of the species of the genus Crota-
phi/tus Holbrook (Iguanidae). Brigham Young
Univ. Sci. Bull., Biol. Ser. 2(1):1-21.
CLUTCH
SIZE
Cnemidophorus
72
76
80
84
88
y = -1.423 +0.051 X
r= 0.333
92
96
SNOUT- VENT LENGTH (mm)
100
Fig. 3. Relationship between clutch size (number) and SVL (mm) for C. tigris.
72
Great Basin Naturalist
Vol. 40, No. 1
Tanner, W. W. 1965. A comparative population study
of small vertebrates in the uranium areas of the
Upper Colorado River Basin of Utah. Brigham
Young Univ. Sci. Bull., Biol. Ser. 7(1): 1-31.
Tinkle, D. W. 1961. Population structure and reproduc-
tion in the lizard Uta stansburiana stejnegeri.
Amer. Midi. Natur. 66:206-234.
1967. The life and demography of the side-
blotched lizard Vta stansburiana. Misc. Publ.
Mus. Zool. Univ. Michigan 132:1-182.
1969a. The concept of reproductive effort and its
relation to the evolution of life histories of liz-
ards. Amer. Natur. 103:501-516.
1969b. Evolutionary implications of comparative
population studies in the lizard Uta stansburiana.
Pages 133-154 in Systematic biology, Proc. of In-
ternational Conf., National Acad. Sci. Publ. No.
1692, Washington, D.C.
TiNKJLE, D. W., .\ND N. F. H.\uLEY. 1975. Lizard repro-
duction effort: caloric estimates and comments
on its evolution. Ecology 56:427-4.34.
Tinkle, D. W., H. M. Wilbur, and S. G. Tilley. 1970.
Evolutionary strategies in lizard reproduction.
Evolution 24:5.5-74.
Turner, F. B., G. A. Hoddenbach, P. A. Medica, and J.
R. La.n.nom. 1970. The demography of the lizard,
Uta stansburiana Baird and Girard, in southern
Nevada. J. Anim. Ecol. .39:50,5-519.
Turner, F. B., P. A. Medica, and B. W. Kowalewsky.
1976. Energy utilization by a desert lizard (Ufa
stansburiana). US/IBP Desert Biome Monogr.
No. 1. Utah State Univ. Press, Logan.
Turner, F. B., P. A. Medica, J. R. Lannom, and G. .\.
Hoddenbach. 1969. A demographic analysis of
fenced populations of the whiptail lizard, Cne-
midophorus tigris, in southern Nevada. South-
west. Nat. 14:189-202.
Turner, F. B., P. A. Medica, and D. D. Smith. 1973.
Reproduction and survivorshp of the lizard, Uta
stan.iburiana, and the effects of winter rainfall,
density and predation on these processes. US/IBP
Desert Biome Res. Memo. RM 74-26, Utah State
Univ., Logan.
ViTT, L. J., and R. D. Ohmart. 1977. Ecology and repro-
duction of lower Colorado River lizards: II. Cne-
midophorus tigris (Teiidae), with comparisons.
Herpetologica 33:223-234.
WoRTHiNGTON, R. D., AND E. R. Arvizo. 1973. Density,
growth and home range of the lizard Uta
stansburiana stejnegeri in southern Dona Ana
County, New Mexico. Great Basin Nat.
.33:124-128.
HAPLOPAPPUS ALPINUS (ASTERACEAE): A NEW SPECIES FROM NEVADA
Loran C'. Anderson'
Abstr.\ct.— The new species, llaplopapptis alpinus of section Toncstus, is forniallv described and illustrated. It is
endemic to the high mountains of central Nevada. Vegetative and floral morphology of related species is detailed.
The new species appears to he most closely related to H. exirnius but also demonstrates close affinity to H. aberrans
(all three are diploids with n = 9).
In North America, Haplopappus (Aste-
raceae) contain.s about 95 .species represented
in 17 sections. Additional species— including
the type species, H. ghttinosus—Sire in South
.\merica. Chronio.somally, two major group-
ings can be identified in the genus (Anderson
at al. 1974). One group is generally her-
baceous and chromosomallv based on x = 4,
5, or 6. The other, more woody group, is
based on x = 9. These groupings are poorly
distinguished by growth form. Woodiness in
the "herbaceous group" is seen in sections
Isoconia and Hazardia (correlated with their
xeromorphy?), and reduced woodiness is seen
in the "shrubby group" in sections Tonestus
and Stcnotus where their growth forms are
apparently related to their montane or alpine
habitats.
The generic integrity of this assemblage
has been challenged. Many feel that Hap-
lopappus is unnatural and should be broken
up (Shinners 1950, Anderson 1966, Turner
and Sanderson 1971, Clark 1977, Urbatsch
1978); some would raise each section to ge-
neric standing. Others feel many of the sec-
tions are interrelated; i.e., the South Ameri-
can taxa and Hazardia (Grau 1976). Jackson
(1966) has demonstrated genetic relationship
among many of the sections of the "her-
baceous group" through intersectional hybr-
idizations; he has recently (pers. comm.)
hybridized South American taxa with those
of .section Hazardia.
Until more is known about the biology of
Haplopappiis (especially the South American
taxa), Hall's con.servative generic treatment
(1928) seems better at the moment than the
alternative of elevating each section to ge-
neric status. Possibly a half dozen genera are
represented in the North American material,
but I can not envision a precise treatment
now.
Description of a new species in section To-
nestus presents a problem. That section is in
the x = 9 group, whereas the type species,
H. ghitinosus, is n = 5 (personal count in
1971 from Kew Garden material; Grau 1976).
So, in retaining the wider generic inter-
pretation at this time, a species of Hap-
lopappus will be named that most likely will
be transferred to another genus when the
complex is better known. Deferring descrip-
tion of this species until a comprehensive ge-
neric revision is available might possibly
withhold additional data that would be sup-
portive of the ultimate revision.
Methods and Materials
Fresh and dried materials were processed
as in Anderson, 1964. Five heads were mea-
.sured for involucral and floral data. Cytolo-
gical methods are those of Anderson, 1966.
Plant materials were collected personallv
in the field or .supplied by Sherel Goodrich.
Vouchers for various measurements and
chromosome counts are at FSU.
Taxonomy
Haplopapfnis alpinus L. C. Anderson & S.
Goodrich, sp. nov.
'Department of Biological Science, Florida State University, Tallahassee, Florida .33206.
73
74
Great Basin Naturalist
Vol. 40, No. 1
Herba perennis et lignosa, 0.5-2.0 dm alta;
stirpes glandulosae; folia in basi obovata vel
oblanceolata, serrata vel dentata, 3-7 cm
longa, 10-36 mm lata, folia caulina aliquan-
temis angustiora et serrata, 3-5.5 cm longa,
8-18 mm lata; inflorescentia vel mon-
ocephala vel cyma paucis cum capitibus; in-
volucra 10-12 mm longa, circa 7 mm lata,
phyllariis 21-28, exterioribus ovatis et folio
similibus et glandulosis, interioribus bracteis
angustioribus; disci florum 29-55, flavi, co-
rollis 5.8-7.1 mm longis, lobis circa 1.3 mm
longis, lineis stigmaticis saepissime longitu-
dine paribus styli appendicibus; achaenia 4-5
mm longa et pubescentia.
Type: Nevada, Nye Co., granitic rocks at
10,600 ft on 11,077-ft peak on Toiyabe Crest
between Washington Creek and Aiken
Creek, 24 air mi SSW of Austin, 21 Jun 1979,
L. C. Anderson 4885 (BRY-holotype!, FSU!,
m^!, UC!).
Perennial herb, woody only at base, short
rhizomatous, (0.5)0.7-1.0(2.0) dm tall; stems
branched only in the inflorescence or mon-
ocephalous, densely glandular-pubescent; fo-
liage dark green, glandular-pubescent, basal
leaves obovate to oblanceolate, petiolate, ser-
rate to deeply toothed above the middle, 3-7
cm long, 10-36 mm wide, moderately vis-
cous, cauline leaves oblanceolate to spatu-
late, cuneate or clasping the stem, saliently
dentate, 3-5.5 cm long, 8-18 mm wide; in-
florescence usually monocephalous (open
sites) or with up to 5 heads in an elongate or
flat-topped cyme (deep crevices or protected
sites); heads campanulate to hemispheric,
10-12 mm long, 7-10 mm wide (pressed),
phyllaries 21-28, outer ones nearly as long as
involucre, leaflike, broadly ovate, 3-nerved,
glandular, slightly spreading, obtuse with
small mucro, inner bracts narrower, lanceol-
ate-spatulate, with finely ciliate margins,
acuminate-cuspidate; ray flowers absent; disk
flowers (29)35-50(55), golden-yellow, corollas
(5.8)6.4-7.1(7.6) mm long, lobes (1.0)1.3(1.6)
mm long, lanceolate, slightly spreading to re-
curved; anthers about 2.6 mm long, appen-
dages 0.6 mm long, style branches slender,
stigmatic lines nearly as long as style appen-
dages; achenes cylindric to hisiform, 4-5 mm
long, pubescent, pappus dull white, 6-7 mm
long; n = 9 (Fig. 1). Infrequent on boulders,
talus, or rocky summits near treeline (primar-
ily on light-colored granites but occasionally
on basalt, andesite, metamorphics, or lime-
stone), 9,000-11,000 ft, Toiyabe and To-
quima mountains of southern Lander and
Nye counties, Nevada. Mid-July-September.
Additional specimens examined: Nevada,
Lander Co., peak between Aiken and Carsely
Creek, S. Goodrich 12137 (FSU, UTC); Nye
Co., type locality, S. Goodrich 12126 (FSU,
NY, UTC), head of left fork San Juan Creek,
S. Goodrich 11997 (BRY, UTC), S. Goodrich
12006 (FSU, UTC), McLeod Creek, S. Good-
rich 13437 (BRY, FSU), crest between Tim-
Fig. 1. Representative, but somewhat stout, specimen
of//, alpinus; drawn largely from Goodrich 12233 (FSU).
March 1980
Anderson: A Nevada Haplopappus
75
blin Creek and Marysville Canyon, S. Good-
rich 12226 (BRY), toiyabe crest at French
VABM, S. Goodrich 12233 (FSU), right fork
Stewart Creek, S. Goodrich 13502 (BRY), top
of Shoshone Mtn., Toquima Range, S. Good-
rich 6 F. Smith 13267 (FSU). All collections
but the last came from the Toiyabe Range.
Goodrich (pers. comm.) also reports seeing a
population on Mt. Jefferson at 11,000 ft at
the head of left fork of Barker (Shipley)
Creek in the Toquima Range. The species is
found for about 23 miles along the crest of
the Toiyabe Range from the Lander-Nye
County line south to the head of Stewart
Creek and is reported for two sites in the To-
quimas.
At the type locality, H. alpinus occurs on
windswept slopes above the treeline with H.
macronema, ChrysotJiamnus viscidiflorus,
Erigeron compositus, and Eriogonian iim-
bellatU7n. Some sites are at or just below
treeline, where the species occurs on rocks in
scattered Pinus flexilis or Cercocarpus ledi-
foUiis. Other alpine endemics from central
Nevada that have been found in the vicinity
of H. alpinus are Draba arida, Eriogonum
ovalifolium var. caelestinum, Geranium
toquimense, Hackelia sp. nov., Senecio sp.
nov., and Smelowskia holmgrenii.
Relationships and Phytogeography
This species belongs to section Tonestus
and is related to H. aberrons and H. eximius
(Figs. 2-4). Comparative floral features for
the three species plus the more distantly re-
lated H. peirsonii are given in Table 1; all are
distinctive. The Nevadan endemic is like H.
abcrrans in its eradiate heads of similar size;
however, the latter has differently shaped
cauline leaves that overtop the racemosely
disposed, turbinate to narrowly campanulate
heads. Also, the bracts are more numerous,
narrower, and somewhat squarrose in H.
abcrrans compared to H. alpinus. The new
species is more similar to H. eximius in leaf
size and shape, and, although the phyllaries
are fairly similar, the campanulate heads
differ with those of H. eximius, being radiate
with shorter involucres. Haploppapus alpinus
also differs from H. eximius in the following
minor floral features: H. alpinus has stouter
corolla tubes (as in Chrijsothamnus spathu-
latus versus C. viscidiflorus; illustrated in An-
derson 1964); its pappus is shorter than the
corolla length, whereas pappus equals corolla
length in H. eximius; and it has longer corolla
lobes.
Original meiotic chromosome counts for
the taxa include: H. alpinus, n = 9 {Ander-
son 4885, the type collection); H. abcrrans, n
= 9 {Anderson 3660 from Blaine Co., Idaho);
and H. peirsonii, n = 45 (Anderson 4326
from Inyo Co., California). The count for H.
peirsonii agrees with the earlier count by
Stebbins, who also reported H. eximius as a
diploid (Howell 1950). The other counts rep-
resent first reports for those species. Meiosis
appeared normal with pairing as bivalents in
all instances. Pollen stainability for Anderson
'■JU
Figs. 2-4. Flowering heads and individual outer bracts; heads scaled to 5 mm bracket, individual bracts slightly
enlarged. Fig. 2. H. aherrans (Anderson 3660, FSU). Fig. 3. //. alpinus (C^oodricli 12137, FSU). Fig. 4. //. eximius
[Anderson 4320, FSU).
76
Great Basin Naturalist
Vol. 40, No. 1
4885 was 97.7 percent, and there were no in-
dications of apomixis in H. alpinus.
Haplopappus aberrans was originally de-
scribed as an aberrant member of section
Macronema and was later tentatively as-
signed to Tonestus (Hall 1928), but until now
it has been considered poorly placed in To-
nestus (Cronquist 1955). With the addition of
H. alpinus, the section houses H. aberrans
more comfortably.
Haplopappus alpinus is .somewhat inter-
mediate between H. aberrans and //. eximius
morphologically and geographically. Harper
et al. (1978) consider the Great Basin moun-
tain ranges flori.stically as islands in the sur-
rounding desert possibly populated by migra-
tions from the "mainland" mountain systems
of the Rockies or Sierras. The introduction of
H. alpinus from eximiuslike precursors from
the Sierra Madre seems very plausible. The
possible origin of H. aberrans in the Saw-
tooth Mountains of Idaho from alpinuslike
stock poses an interesting situation. Hap-
lopappus aberrans occurs in the Rocky
Mountain "mainland" system (in which H.
hjallii and H. pijgmaeus of Section Tonestus
occur). Still, its affinities lie with H. alpinus
and H. eximius. The species could represent
the culmination of a migration from the
western Sierra mainland across the Great Ba-
sin deserts to the Sawtooths and the eastern
mainland. These mountain groups were less
isolated in the relatively recent past (Harper
et al. 1978), and such a migration is plausible.
Billings (1978) suggests the alpine flora in
the Great Basin may have resulted from "up-
ward evolution" of preadapted desert species
of lower elevations. This doesn't .seem to ap-
ply to H. alpinus or other Tonestus taxa. Bil-
lings further suggests that due to reduced
habitat diversity in Great Basin mountains
there is a trend toward edaphic endemism.
Again, H. alpinus does not follow the trend;
it has been collected on granite, basalt, meta-
morphics, andesite, and limestone in the geo-
logically diverse Toiyabe Mountains. Clearly,
these alpine areas of central Nevada do need
more vegetational work, as Billings (1978)
observed. Our ideas of that region may be
greatly changed with further study; endem-
ism apparently is not as low as Harper et al.
(1978) record. Goodrich, for example, has
found several imdescribed endemics in his
current survey of the region.
T.\BLE 1.— .\veraged floral data (and ranges entered parenthetically) for .selected taxa of Haplopappus, section To-
nestus.
Taxon anc
i collection
H. aberrans
Anderson
3660
H. alpinus
H. eximius
H. peirsonii
Feature
Anderson
4885
Goodrich
12137
Anderson
4320
Anderson
4899
Anderson
4326
Involucre
Bract number
Length, mm
42.0 (37-49)
11.0(10-12)
24.2 (21-25)
10.6(10-11)
23.8 (21-28)
11.0(10-12)
27.2 (25-30)
8.5 (8-9.5)
29.0 (24-35)
9.0 (8.2-10)
27.0 (23-29)
15.2 (13-16)
Rav flowers
Flower number
Flower length, mm
Flower width, mm
-
-
-
13.6(11-19)
10.5
2.2
15.0 (12-20)
10.2
2.1
21.8(21-23)
10.4
4.4
Disk flowers
Flower number
Flower length, mm
Lobe length, mm
Stigmatic area— total
style branch, %
38.0 (29-48)
7.6 (7.0-8.2)
1.0(0.9-1.1)
51.7 (50-54)
41.6 (.35-49)
6.3(5.8-7.1)
1.3(1.0-1.6)
50.0 (43-55)
44.4 (29-55)
7.1 (6.4-7.6)
1.3(1.2-1.4)
48.1 (45-50)
59.2 (54-69)
7.1 (6.6-7.4)
1.1(1.0-1.2)
56.3 (44-62)
51.6(44-60)
6.4 (6.0-7.0)
1.0(0.7-1.2)
53.7 (50-56)
55.8 (45-75)
8.4 (7.0-9.5)
1.1(1.1-1.2)
.38.1 (33-43)
March 1980
Anderson: A Nevada Haplopappus
77
Acknowledgments
Appreciation is expressed to Sherel Good-
rich for guiding me to the alpine collection
sites and for supplying habitat data and sev-
eral collections. Melanie Darst prepared the
line drawings; Dr. Walter Forehand is
thanked for assistance with the Latin diag-
nosis. This study was supported by National
Science Foundation grant DEB 76-10768.
Literature Cited
Anderson, L. C. 1964. Taxonomic notes on the Chrijso-
thamnus viscidiflorus complex (Astereae. Com-
positae). Mandrono 17:222-227.
1966. Cytotaxonomic studies in Chitj.sothdmiuis
(Astereae, Conipositae). .\nier. ]. Bot.
53:204-211.
.\nderson, L. C, D. W. Kyhos, T. Mosquin, A. M.
Powell, .\nd P. H. Raven. 1974. Chromosome
numbers in Conipositae. IX. Haplopappus and
other .\stereae. .\mer. J. Bot. 61:66.5-671.
BiLLLNGS, D. W. 1978. .\lpine phytogeography across the
Great Basin. Great Basin Nat. Mem. 2:105-117.
Clark, W. D. 1977. Chemosystematics of the genus
HaMrdia (Conipositae). J. Ariz. .\cad. Sci. 12:16.
Cro.nquist, .v. 1955. Conipositae. V. In: C. L. Hitch-
cock, A. Cronquist, M. Ownbey, J. W. Thompson
(eds.). Vascular plants of the Pacific Northwe.st,
Univ. of Washington Press, Seattle.
Grau, J. 1976. Chromosomenzahlen von Siidanierikanis-
chen Haplopappus ,\rten. Mitt. Bot. .Miinchen
12:40.3-410.
IL\LL, H. M. 1928. The genus Haplopappus, a phyloge-
netic studv in the Conipositae. Carnegie Inst.
Publ. .389:i-.391.
Harper, K. T., D. C. Freeman, W. K. Ostler, and L.
G. Klikoff. 1978. The flora of Great Basin moun-
tain ranges: diversity, sources, and dispersal ecol-
ogy. Great Basin Nat. Mem. 2:81-103.
Howell, J. T. 1950. Studies in California Aplopappus.
Leafl. West. Bot. 4:84-88.
Jackson, R. C. 1966. Some intersectional hybrids and
relationships in Haplopappus. Univ. Kans. Sci.
Bull. 46:47.5-488.
Shlnners, L. H. 1950. Notes on Texas Conipositae, IV,
V. Field & Lab. 18:25-42.
Turner, B. L., and S. Sanderso.n. 1971. Natural hybridi-
zation between the Composite "genera" Machae-
ranthera and Haplopappus (sec. Blepharodon).
Amer. J. Bot. .58:467.
Urbatsch, L. E. 1978. The Chihuahuan Desert species
of Ericaincria (Compositae: .\stereae). Sida
7:298-303.
MISCELLANEOUS PLANT NOVELTIES FROM ALASKA, NEVADA, AND UTAH
Stanley L. Welsh' and Sherel Goodrich'
\BSTRACT - Described as new to science are Ahwnia ar^ilhsa Welsh & Goodrich, from Utah and Colorado; An-
drosace alaskana Gov. & Standi, var recdae Welsh & Goodrich, from Kohlsaat Peak, Alaska; Lepidium osten Welsh
& Goodrich, from Beaver Gounty, Utah; Lygode.mia entrada Welsh & Goodrich, from Grand Connty, Utah; Pedto-
cactm des-painii Welsh and Goodrich, from Emery County, Utah; and Senecio toiyabensis Welsh and Goodrich, from
the Toiyabe Range in Nye Coimty, Nevada.
Several undescribed and unnamed taxa
have accumulated at the herbarium of Brig-
ham Young University in recent years.
Mainly they represent materials which have
been treated within other taxa, or they are
oddities of exceedingly limited areal extent.
Some have been known for long periods of
time. Others are only recently discovered. All
are unique in one or more ways and of suffi-
ciently limited apparent distribution as to be
candidates for inclusion on lists of sensitive
species.
Abronia argillosa Welsh & Goodrich, sp.
nov.
Plantis similis Abronia fragranti Nutt. et
Abronia elliptica in habitas sed differt in an-
thocarpus alarum nuUis rostro nuUo gracilior-
ibus, in receptaculo conico et fructibus in-
structis superioris foliis floribus et
anthocarpis glabris, et floribus paucioribus.
Plantae perennes e caudicibus ramifican-
tibus gracilibus (6) 15-30 cm altae; caules
glabri ad basim nibellos saepe frondosi om-
nino; folia (5) 15-35 mm longa, 3-35 mm lata
elliptica cvato obovata vel suborbiculares
glabra; pedunculi 1-8 cm longi glabri vel
puberulentes raro; bractae 7-15 mm longae,
6-15 mm latae ovales vel orbiculares sca-
riosae glabrae vel ciliolatae; flores 15-22 in
quoque inflorescentiam; perianthi tubus
10-15 mm longus viridis glaber vel raro pub-
enilentus limbus ca. 6 mm latus albidus; re-
ceptaculiun breve conicum ferens fnictus in
dimidio superiore anthocarpus sine alis sine
rostris plicatus leviter vel nullus 7-9 mm
longus 3-4 mm latus scariosus glabri rugosus
alborostratus; semina 2.5-3 mm longa 1 mm
lata.
Holotype: Utah, Grand Co., T22S, R24E,
Sec 18, ca. 6 miles due south of Cisco at ca.
4300 feet elev., on Mancos Shale Formation,
in an Atriplex community, S., E., and M.
Welsh 16689, 30 May 1979 (BRY, 8 isotypes
to be distributed).
Paratypes: Utah, Grand Co., Fifteen miles
east of Thompson, B. F. Harrison et al 10403,
16 June 1941 (BRY); first escarpment north of
Thompson, west of Sego Canyon, S. L. Welsh
6943, 1 May 1968 (BRY); T22S, R24E, Sec 7,
ca. 4 mi. south of Jet. 50-6 and U-128, S. L.
Welsh and K. Taylor 14637, 28 April 1977
(BRY); T18S, R25E, Sec 27, ca. 20 mi. NE of
Cisco, S. L. Welsh 14916, 8 June 1977 (BRY).
Uintah Co., Ca. 2 mi. S of Dragon, S. L.
Welsh 5379, 13 May 1966 (BRY). Colorado,
Mesa Co., ca. 5 mi. W of Mack along US
Hwy 6-50, L. C. Higgins and S. L. Welsh
1034, 14 June 1967 (BRY); ca 13 Km due NW
of Mack, T9S, R104W ca. Sec 11, A.
Cronquist 11427, 25 May 1976 (BRY;NY).
The clay verbena, Abronia argillacea
Welsh and Goodrich, is restricted to the
Grand River Valley, and less commonly in
the drainage of the White River in east-cen-
tral Utah and west-central Colorado, where it
occurs on heavy soils derived from Mancos
Shale and Green River formations. It seems
probable that the taxon was taken much pri-
or to the specimens cited above, but was
'Life Science Museum and Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602.
78
March 1980
Welsh, Goodrich: Plant Novelties
79
KHT
Fig. L Abronia argillosa: A, habit of plant.
80
Great Basin Naturalist
Vol. 40, No. 1
overlooked due to the superficial resem-
blance to phases of A. elliptica A. Nels. and
to A. fragrans Nutt. ex Hook.
The plants are glabrous or essentially so
and possess orbicular leaves which are
glaucous. The genus has been revised by Gal-
loway (1975), who distinguishes A. elliptica
(common and widespread in Utah) from A.
fragrans (cited from San Juan County only)
by technical characters of the anthocarps.
The following key will distinguish A. argil-
losa from those taxa.
1. Anthocarps with 2 wings, these folded together to form a deep groove; plants
of western Colorado and westward A. elliptica A. Nels.
— Anthocarps wingless, or, if winged, then the wings not folded together; plants
of eastern Utah and eastward 2
2(1). Anthocarps beaked, deeply grooved or narrowly winged; leaf blades variable
in shape, glandular-puberulent to glandular-pubescent; plants usually of sandy
soils in San Juan Co., Utah, and eastward in Colorado
A. fragrans Nutt. ex Hook
— Anthocarps beakless, slightly or not at all folded; leaf blades mainly orbicular,
glabrous; plants usually of clay soils in Uintah and Grand counties, Utah and
Mesa Co., Colorado A. argillosa Welsh & Goodrich
Androsace alaskana Gov. & Standi, ex Hul-
ten
var. reedae Welsh & Goodrich, var. nov.
Similis Androsace alaskana Gov. & Standi,
in scapis numerosis floribus sessilibus solitariis
vel binatum, sed differt scapis numerosioribus
brevioribus gracilioribus pubescentioribus,
foliis integris parvioribus dense villosis, et flo-
ribus parvioribus.
Herbae annuae vel biennis; scapi 25-40 in
quoque rosellam 10-45 mm longi filiformes
0.3-0.5 mm in diametrum pubentes parce vel
dense pilis furcatis apprime infra flores juxta;
folia 5-10 mm longa 1-3 mm lata linearia vel
spathulata integra vel denticulata cum 1-2
dentibus glabra infra vel pilis ad costam juxta
apicem supra dense villosa pilis multi-
cellulosis furcatis vel simplicibus apprime ul-
tra medium; flores solitari vel imparibus ali-
quando bractaea subtenda tubus calycis
subaequali; tubus calycis 2-3 mm longus,
dentibus circa 1.5 mm longis; corolla alba
tubo calyce subaequali lobus circa 2 mm
longis; capsulae maturae ignotae.
Holotype: Alaska, Lat. 62°12'N., Long.
152°47'W, ca. 2 mi. SSW of Kohlsaat Peak,
near VABM 5048, at 4900 feet (1495 m)
elev., on rocky ridge top, K. Reed 5857, 29
June 1977 (BRY, isotype at Leningrad).
Par.\type: Yukon Territory, Canada,
Mount St. Elias Quad.: Outpost Mt. at south
end of Kluane Lake, 60°56'N, 138°22'W, at
ca. 2140 m., D. F. Murray 3014, 22 Julv 1969
(BRY; ALA).
The materials herein segregated as var.
reedae represent the slender peduncled, sub-
entire- to entire-leaved, smaller, flowered
high elevation phase of A. alaskana in interi-
or Alaska and southwestern Yukon. Tliat the
morphological differences noted in the pro-
tologue might be the result of ecological re-
sponse has been considered. The tendency to
entire leaves and less pronounced ciliate mar-
gins, along with slender peduncles and flow-
ers that seem to average smaller, indicate a
syndrome of characteristics which should re-
ceive at least some taxonomic recognition.
The plant is named to honor the collector
of the holotype, Katherine Reed of Ancho-
rage, Alaska.
Lepidium ostleri Welsh & Goodrich, sp. nov.
Habitu Lepidio nana S. Wats sed differt in
inflorescentia longiore floribus numer-
osioribus foliis longioribusque pubentioribus
et sepalis pubentioribus.
Herbae perennes pulvinatae caespitosae
caudice ramoso folium basim marcescentibus;
caules fnictifri 10-35 mm alti hirsuti; folia
4-15 mm longa hirsuta linearia et integra vel
basalia 3-5 lobatis vulgo caulina nulla vel
pauca; racemi circa 1 cm longi in florem et
March 1980
Welsh, Goodrich: Plant Novelties
81
mm
Fig. 2. Androsace alaskana: A, adaxial side of leaf; B, abaxial side of leaf; C, elose-iip of stem and inflorescence; D,
habit of plant.
82
Great Basin Naturalist
Vol. 40, No. 1
:pi.^ I mm
mm '•'iiLv-M /•;
Fig. 3. Lepidiiim ostleri: A, habit of plant; B. close-up of leaves showing variation; C, close-up of flower; D, close-
up of fniit; E, close-up of inflorescence.
March 1980
Welsh, Goodrich: Plant Novelties
83
1-2 cm longi in fructem 5-25 floribus; pedi-
celli in fructem 2-3 mm longi; sepala 1.3-2
mm longa obtusa hyalina aspre pilosa saepe
purpurascentia; petala 2-3 mm longa alba
purpiirascentia; fnictus 2.5-3 mm longa,
2.3-2.5 mm lata ovata late usque ad 1 mm
crassa sinus 0.2 mm profundus; stylus 0.4-0.7
mm longus.
Holotype: Utah, Beaver Co., T27S,
R13W, Sec 23 (SW'/4), San Francisco Moun-
tains, at Frisco, on rocky ridge, in pinyon-
jimiper community, K. Ostler and D. Ander-
son 1258, 6 June 1978 (BRY).
Paratypes: Utah, Beaver Co., T27S,
R13W, Sec 23 (NW1/4), near Frisco, 6900
feet, rocky slopes, K. Ostler and D. Anderson
1210, 1 June 1978 (BRY); do, T27S, R13W,
Sec 16 (SE^/i), San Francisco Mountains, near
Utah Hwy 21, at 5800 ft, on west slope, dry
limestone, Atriplex community, K. Ostler
1415, 19 June 1978 (BRY); do, west slope of
San Francisco Mountains, bristlecone pine,
ponderosa pine, Douglas fir, white fir com-
numity, K. Ostler 1588, 5 July 1978 (BRY).
The obvious relationship of this perennial
dwarf species lies with the Lepidium nantim,
an endemic of Nevada. The longer racemes,
greater flower number, and dense pub-
escence constitute the most important diag-
nostic features. The petals of Lepidium ostleri
are white, while those of L. nanum are yel-
low to cream yellow. While probably of less
importance than other features, the color of
the petals becomes important when taken
with the differences in raceme, flower num-
ber, and pubescence features.
The species is named in honor of its collec-
tor. Dr. Kent Ostler, an enthusiastic collector
and botanist.
Lygodesmia entrada Welsh & Goodrich, sp.
nov.
Ab Lygodesmia grandiflora Nutt. in caul-
ibus rigidioribus ramosissimis foliis brevior-
ibus et paucioribus et radiis albis differt.
Herbae perennes caudice subterraneo
ramosissimo omnino usque ad 45 cm altae;
folia integra linearia vel acicularia 5-30 mm
longa; pedunculi potius numerosi bracteati
elongati 12-20 cm longi in capitulum termi-
nans; bracteae involucrorum hyalinae exte-
riores 5-10 mm longae fimbrillatae interiores
circa sex 16-18 mm longae apex puberulus;
radii albi circa 3 cm longi; pappus barbel-
latus sordidus setae 10-15 mm longae; ach-
enia costata glabra.
Holotype: Utah, Grand Co., T24S, R19E,
Sec 25, Tusher Canyon, ca. 15 mi. due WNW
of Moab, 4800 feet elev., Entrada Sandstone
Formation, juniper community, S. L. and S.
L. Welsh 16725, 3 June 1978 (BRY, four iso-
types to be distributed).
This white-flowered material has been
identified by A. S. Tomb (pers. comm.) as a
probable triploid assignable to Lygodesmia
arizonica Tomb. The triploid hypothesis can-
not herein be questioned because of lack of
knowledge concerning the cytological nature
of the plants in question. However, despite
the ultimate disposition of these plants fol-
lowing future determinations of chromosome
numbers, the strikingly different morpholo-
gical features dictate taxonomic recognition,
if for no reason other than the fact that the
plants are so different from other plants of
Lygodesmia in Utah.
A second collection at BRY, here assigned
to L. entrada, is J. S. Allen 132, from north of
Courthouse Wash Ridge in Arches National
Park. Dried flower remnants appear to be
pink, but the tall nidularius habit and defi-
nitely ligneous stems and branches are appar-
ent.
Lygodesmia arizonica Tomb is a low her-
baceous plant usually of more southern distri-
bution in Utah. Even in late anthesis the
stems are herbaceous and lack the character-
istic bird's nest appearance of L. entrada.
Lygodesmia entrada differs from L. grandi-
flora in ways similar to those discussed for L.
arizonica.
Pediocactus despainii Welsh & Goodrich, sp.
nov.
Ab Pediocacto hradyi L. Benson differt in
spinis paucioribus brevioribus gracilioribus et
floribus colorum.
Plantae carnosae hemisphericae depressae
3-6 cm in diametrum 4-8 cm longae; tuber-
culi ovata numerosa ordinata in serialia cir-
cularia vel spiralia; areolae spinis 8-14; spin-
ae 2-5 mm longae serialia stellatim; pilis
coactis instructis interdum; flores 2.5-4 cm
diametrum fragrantes ad apicem gerenti; se-
pala numerosa; petala numerosa albida suf-
fusa rosea et flava; stamina numerosa lutea
84
Great Basin Naturalist
Vol. 40, No. 1
Fig. 4. Lygodesmia entrada: A, habit of plant; B, close-up of achene.
March 1980
Welsh, Goodrich: Plant Novelties
85
cm
B
<c:^
Fig. 5. Pcdiocactus despainii: A, habit of plant; B, habitat of plant.
86
Great Basin Naturalist
Vol. 40, No. 1
omnino; stigmata numerosa lutea omnino;
fmctus 8-9 mm diametrum 10-12 mm longus
operculo circumscissili umbone 8-9 mm dia-
metnim 5.5-6 mm alto; fructus corpi 8-9
mm diametrum 5.5-6 mm longus findens
longirostronim coloris viridis ad porphyreus;
seminum 3-3.5 mm longum 2-2.5 mm latum
varicosis tuberculatibus minutibus.
Holotype: Utah, Emery Co., San Rafael
Swell, Despain 266a, 5 May 1978 (BRY).
Paratypes: Utah, Emery Co., San Rafael
Swell, Despain 445, 15 May 1979 (BRY); do,
E. Neese & K. Thome 504, 7 May 1979.
The Despain pediocactus is a diminutive
cactus of very local distribution in the San
Rafael Swell of Emery County, Utah. Exact
locality is not given so as to provide a mea-
sure of protection of this species from ama-
teur and commercial fanciers. The species is
compared in the protologue to P. bradyi,
from which it differs as indicated. It is similar
to the newly described P. winkleri (Heil
1979) from Wayne County, but differs inter
alia in the monocephalous nature, stems
which average larger (3-6 cm in diameter vs.
2-2.6 cm), larger flowers (2.5-4 cm broad vs.
1.7-3 cm), and shorter capsules (5.5-6 mm
long vs. 7-9 mm).
The species is named in honor of its discov-
erer, Mr. Kim Despain, student of the flora of
the San Rafael Swell.
Senecio toiyabensis Welsh & Goodrich, sp.
nov.
Ab Senecio fremontii T. & G. foliis integris
angustis et statura elata differt.
Herbae perennes caudicibus ramificantibus
ligneis; caules 15-60 cm altae glabri vel pub-
escentes parce; folia accrescentia sursum
magniora linearia integra vel denticulata 2-8
cm longa 2-7 mm lata; bracteae diminutae;
inflorescentia corymbosa; bracteae in-
volucromm ca 13, 6-8 mm longae margines
hyalini apices acuti pilosi breves vel glabri
raro rubentes interdum; radii 8 vel pauciores
circa 1 cm longi lutei; achenia scabra.
Holotype: Nevada, Nye Co., Toiyabe Na-
tional Forest; Toiyabe Range, just under the
crest of the range on the leeward side, above
Timblin Cr., 35 air miles SW of Austin,
100-500 yards N of French VABM, T13N,
R42E, near center of Sec. 4, 10,500 feet, in
cracks of metamorphic rocks and talus, with
Artemisia michauxiana, Penstemon watsonii,
Eriogonwn microthecwn, Philadelphiis micro-
plujllus, and Sphaeromeria cana, Goodrich
12235, 30 Aug. 1978 (BRY; numerous iso-
types to be distributed).
Paratypes: Nevada, Nye Co., Toiyabe Na-
tional Forest, Toiyabe Range, along or near
the crest of the range between San Juan-
Tierney Creeks and McLeod Cr., T14N,
R43E, in or near W1/2 of Sec. 11, 9800-10,000
feet, on metamorphic precambrian outcrops,
talus and rocky ground, Goodrich & Schlatter-
er 12156, 10 Aug. 1978 (BRY); do, north rim
of Aiken Creek, very near the Lander-Nye
Co. marker; T15N, R43E, Sec. 17, NE1/4 of
SEV4, 10,800 feet, on rocky slope; growing
with Cymopterus petraeus, Haplopappus
macronema, Penstemon speciosus, Oryzopsis
hymenoides, and scattered Pinus flexilis,
Goodrich 12138, 5 Aug. 1978 (BRY); Nevada,
Lander Co., east side of Bunker Hill, 16.5 mi.
187° from Austin, N39n5'25-35" W117°7'
10-20", 11,000 ft., steep, rocky limestone
slopes, Goodrich 13338, 10 July 1979 (BRY).
The Toiyabe groundsel is a near congener
of S. fremontii from which it differs in having
linear entire leaves (not ovate or obovate to
oblanceolate and dentate). The leaves are 2-7
mm wide, compared to 1-4 cm wide in S.
fremontii. Stems are mostly erect and are
1.5-6.0 cm tall when in an thesis.
In the key to group X. Triangulares by
Barkley (1978), S. toiyabensis would require
modification as follows:
2. Plants taprooted, or with a sub-
rhizomatous caudex surmounting a tap-
root.
3. Leaves linear, entire; plants 1.5-6
dm tall, restricted to the Toiyabe
Range, Nevada S. toiyabensis
3. Leaves ovate or obovate to ob-
lanceolate, dentate; plants 1-3(4)
dm tall, distribution not as above
S. fremontii
2. Plants variously fibrous rooted or with a
persistent caudex but not taprooted.
Literature Cited
Barkley, T. M. 1978. Senecio. Fl. N. Amer. Series 11,
Part 10: 50-139. New York Bot. Card., New York.
Galloway, L. A. 1975. Systematics of the North Ameri-
can desert species of Ahronui and Thptcrocahjx
(Nyctaginaceae). Brittonia 27:328-347.
March 1980
Welsh, Goodrich: Plant Novelties
87
Fig. 6. Senccio toiyabensis: A, close-up of flower; B, habit of plant.
Great Basin Naturalist
Vol. 40, No. 1
Heil, K. D. 1979. Three new species of Cactaccae from
southeastern Utah. Cact. & Succulent J. (U.S.) 51:
25-30.
Hitchcock, C. L. 1936. The genus Lepidium in the
United States. Madroiio 3: 265-320.
To.MB, .\. S. 1979. Novelties in Lygodesmia and Stephan-
omeria (Compositae-Cichorieae). Sida 3(7):
530-5.32.
Welsh, S. L. 1974. Anderson's flora of Alaska and adja-
cent parts of Canada. Brigham Young University,
Provo, Utah.
Welsh, S. L., and J. L. Reveal. 1977. Utah flora: Brassi-
caceae (Cruciferae). Great Basin Nat. 37:
279-365.
NEW GENERA AND NEW GENERIC SYNONYMY IN SCOLYTIDAE (COLEOPTERA)
Steplieii L. Wood'
Abstract.— New generic synonymy in the world fauna of Scolytidae includes: Acanthotomictis Blandford
{ = Isophthorus Schcdl), Acranttt.s Broun { = Chaetophorotis Fuchs, Chaetopteliiis Fuchs), Cosmudercs Eichhoff
{ = Erioschiilia.s Schedl), Ernoporicus Berger ( = Ernopoceius Balachowsky), Ernoporus Thomson { = Euptiliits Schcdl),
Hyliirdri'ctonii,s Schedl ( = .\'(//og(>/JHiw,y Schedl), Ozopanon Hagedorn {Dnjocoetiops Schedl), Scolijtogenes Eichhoff
( = Cnjpliahmiorplius Schauhiss). Stepluinopodius Schedl {^Crijphdloniinictes Browne), and Xijlechintis Chapuis
(^Sijuaina.'iinulus Nimberg). Genera new to science and their tvpe-species include: Anaxylchont.'i {Tomicits tnin-
ccittis Erichson), Apoxi/lchorus (Xt/lcborus mancus Blandford), Crifphulogenes {Cri/pluilo^enes euphorbiae Wood), Er-
nochidius {Cn/phahis corpulcntu.s Sampson), lladrodcniius (Xijlebonis globus Blandford), Leptoxt/lebonis (Phloeo-
trootis sordicauda Motschulsky), Micropenis (Xijlebonis theae Eggers), Taphrodasus [Xijlebonis penorthi/liis Schedl),
and Taiirodemiis (Xijlebonis sharpi Blandford). The new name Hijliirdn'ctoniis corticiiiiis is presented to replace H.
araiicariae (Schedl 1972). Dnjocoetes coffeae Eggers is transferred to Eiilepiops. The following genera are treated in a
revised context: Cn/togeniiis. Dnjocoetes, Eiilepiops, Enioporiciis, Enwponis, Xijlebonis, and Xylechinus. Cnjphalo-
genes euphorbiae and C. exiguus (Sri Lanka) are named as new to science.
In a review of the genera of Scolytidae in
the world fauna, several problems that relate
to synonymy were encountered. The new
svnonvmv listed in the above abstract is re-
ported here in order that names might be
used in their new context before the generic
revision is completed. In addition, several
genera are treated in a sense somewhat dif-
ferent from the traditional. The basis for
these departures is established. The genera
are treated alphabetically for convenience of
reference. They include representatives from
the subfamily Hylesininae, tribe Tomicini
{Acmntiis, Hyhirdrectonus, Xylechinus) and
from the subfamily Scolytinae, the tribes
Ipini {Acanthotornicus), Dryocoetini {Cijrto-
genitis, Eulepiops, Ozopemon), Xyleborini
(Xyleborus), and Cryphalini {Cosmoderes, Er-
noporictis, Ernoporus, Scolytogenes, Steph-
anopodius). Nine new genera represent the
tribes Xyleborini {Anoxyleborus, Apoxyle-
borus, Hadrodcmius, Leptoxyleborus, Micro-
perns, TapJirodasus, Taurodemus) and
Cryphalini (Cryphalogenes, Ernocladius).
Xylechinosomus Schedl is removed from syn-
onymy with Pteleobius. The new name Hy-
hirdrectonus corticinus is presented to re-
place the junior homonym H. araucariae
(Schedl 1972). The species Cryphalogenes eu-
phorbiae and C. exiguus (Sri Lanka) are
named as new to science.
Acanthotornicus Blandford
Acanthotornicus Blandford, 1894, Trans. Ent. Soc. Lon-
don 1894:89 (Type-species: Acanthotornicus spin-
osus Blandford, monobasic)
Isophthorus Schedl, 1938, Archiv Naturgesch. 7(2): 173
(Type-species: Isophthorus quadrituberculatus
Schedl, present designation). .Vpir synonymy
In the original description of Isophthorus
Schedl, two .species were definitely included
and a third species was doubtful, but a type-
species was never designated. Since then,
Schedl has transferred all three species else-
where. To anchor the generic name, Isoph-
thorus quadritidierculatus Schedl is here des-
ignated as the type-species of Isophthorus.
Because this species and Myeloborus bico-
nicus Schedl have been transferred to Acan-
thotornicus and the unrelated, doubtfiil spe-
cies, Pityophthorus heteae Hagedorn, has
been transferred to Cryptocarenus, the fix-
ation of a type-species requires that Isoph-
fliorus be placed in .synonymy under Acan-
thotornicus.
'Life Science Museum and Department of Zoology. Brigham Young University. Provo. Utah 84602. Scolytidae contribution number 69.
89
90
Great Basin Naturalist
Vol. 40, No. 1
Acrantus Broun
Hfliminis Broun, 1881, Manual of New Zealand Coleop-
tera 2:720 (Tvpe-species: Homanis inunduliis
Broim, monobasic). Preoccupied
Acrantus Broun, 1882, Ann. Mag. Nat. Hist. (5)9:409.
Replacement name
Chaetophorus Fuchs, 1912, Morphologische studien iiber
Borkenkafer, II. die europiiischen Hyiesinen, p.
46 (Tvpe-species: Hylesiniis vestitus Mulsant 6c
Re\', monobasic). Preoccupied
Clwetoptelius Fuchs, 191.3, in Reitter, Wiener Eut. Zeit.
.32(Beiheft):4.3. {Replacement name). Xeic synon-
!/'"!/
The names Acrantus Broun and Chaetop-
telitis Fuchs have been treated as synonyms
of Pteleobius Bedel (Schedl 1963:262) and
Pseudohijlesinus Swaine (Schedl 1966:75), re-
spectively. However, in a review of the char-
acters of the type-species of these genera, it
was demonstrated (Wood 1978) that Pte-
leobius must be placed in the tribe Hylesinini
and that Chaetopteliiis and Pseudohijlesinus
belong in the tribe Tomicini. For that study,
Schedl's (1963:262) placement of Acrantus
was not challenged.
In a subsequent review of the genera of
Tomicini, di.ssection demonstrated that Ho-
marus mundulus Broun, type-species of Acr-
antus, clearly belongs to the Tomicini and is
quite unrelated to Pteleobius. Furthermore,
Pseudohijlesinus totally lacks pronotal aspe-
rities, it has three distinct sutures on the an-
tennal club, and the male frons is not noticea-
bly impres.sed. Acrantus, Chaetopteliiis, and
Xijlechinosomus all have numerous pronotal
asperities, two or four poorly marked sutures
on the antennal club, and the male frons
strongly impressed and, thus, form a group
quite distinct from Pseudohijlesinus. Xijle-
chinosomus, which Schedl (1966:75) also
placed in synonymy with Pteleobius, has the
antennal club less elongate, less strongly
compressed, and (apparently) with four ob-
•scure sutures and the frontal rectangle much
more elongate. Acrantus and Chaetopteliiis
have the antennal club more elongate,
strongly flattened, and marked by two su-
tures and the frontal rectangle comparatively
broad. Biological differences also support the
continued recognition of Xijlechino.wmus.
However, I can find no characters that sepa-
rate Acrantus and Chaetoptelius. For this rea-
son, Chaetoptelius is placed in synonvmv un-
der Acrantus, as indicated above.
Acrantus includes mundulus and vestitus,
cited above, and most if not all of the species
from New Zealand, Australia, New Guinea,
and neighboring areas placed by Schedl in
Leperisinus and Xijlechinus.
Anaxyleborus, n. gen.
This genus is distinguished from Eii-
wallacea Hopkins and allied genera by the
truncate, concave elytral declivity which has
a complete, .sharply elevated, circum-
declivital costa from base to apex. The discal
interstrial punctures are uniseriate; in the su-
perficially similar Apoxijleborus they are con-
fused.
Description.— Antennal club with one su-
ture visible on posterior face, anterior face
with segment 1 corneous, 2 conspicuous,
sometimes rather large. Procoxae contiguous.
Protibia armed by more than 11 socketed
teeth. Declivity and discal punctures on in-
terstriae as described in above diagnosis.
Type-species: Tomicus truncatus Erichson.
Species assigned previously to the Xijle-
borus truncatus group belong here.
Apoxyleborus, n. gen.
This genus is distinguished from Tauro-
demus by the presence of only four to seven
socketed teeth on the protibia, by the
obliquely truncate elytral declivity, with an
abrupt (not acute) circumdeclivital costa, and
the face flat to weakly concave. It is distin-
gui-shed from the superficially similar Ana-
xyleborus by the rather widely separated pro-
coxae, by the strongly confused interstrial
punctures on the disc, and by the less dis-
tinctly concave elytral declivity.
Description.— Body stouter than 1.9 times
as long as wide. Antennal club with .segment
1 corneous, with no sutures evident on poste-
rior face, apical margin of segment 1 on ante-
rior face acutely elevated into a continuous
co-sta forming a complete circle. Procoxae
moderately separated. Protibia armed by four
to seven socketed teeth. Elytral disc with in-
terstrial punctures strongly confused, declivi-
ty as described in above diagnosis.
Tvpe-species: Xijleborus mancus Bland-
ford.
March 1980
Wood: American Bark Beetles
91
Species assigned previously to the Xyle-
bonis mancus group belong here.
Cosmoderes Eichhoff
Cosmoderes Eichhoff, 1878, preprint of Mem. Soc. Roy.
Sci. Liege (2)8:49.5 (Type-species: Cosmoderes
moiiilUcollis Eichhoff, monobasic)
Erioschidias Schedl, 1938, Trans. Roy. Soc. S. Austraha
62:42 (Type-species: Crijphalus sctistriattis L^a,
subsequent designation by Wood, 1960, Insects of
Micronesia 18(1):21). New sijnoiuit)nj
The Beeson Collection at the Forest Re-
search Institute, Dehra Dim, India, contains
series of three species that were placed by
Beeson in Cosmoderes. One, from Samsingh,
Kalimpong, Bengal, is labeled monillicoUis
Eichhoff; the other two bear manuscript
names not yet validated. Beeson 's private
notes, of which two volumes treating Scoly-
tidae are in my possession, contain no in-
dication under this name that he saw the
type of moniUicoUis. However, elsewhere in
his notes there are several indications that he
saw the Eichhoff Collection at Hamburg be-
fore it was destroyed during World War II.
Blandford also saw the Eichhoff Collection,
but there is some doubt (Blandford 1894:86)
that he actually examined the type of mon-
illicoUis.
Both the Beeson and Blandford specimens
are congeneric with Erioschidias Schedl. Bee-
son's .specimens of monillicoUis match the dis-
tinctive characters of Eichhoff's description
in every detail. It is, therefore, propo.sed that
Erioschidias be placed in synonymy under
Eichhoff's name, as indicated above.
Cryplialogenes, n. gen.
This genus is distinguished from the closely
allied Scolytogenes Eichhoff by the 3-seg-
mented antennal funicle, by the antennal
club with sutures 1 and 2 weakly procurved,
marked by .setae, and 1 grooved and partly
.septate, and by tlie horizontal venter of the
abdomen.
Description.— Frons convex, not sexually
dimorphic. Eye elongate-oval, entire. Anten-
nal scape elongate, simple; fimicle 3-seg-
mented; club oval, a slight constriction and
groove at suture 1, sutures 1 and 2 moder-
ately procurved, 1 partly septate at lea.st on
lateral half. Pronotum with basal and lateral
margins marked by a fine, raised line; ante-
rior slope asperate, anterior margin armed by
low, poorly formed serrations. Elytral punc-
tures largely replaced by rows of rounded
strial and interstrial granules; vestiture of
rows of -Strial hair and interstrial scales. Pro-
tibia armed by four socketed teeth. Venter of
abdomen horizontal. Sexes subequal in size.
Type-species: Cryphalogenes euphorbiae
Wood.
Cryphalogenes euphorbiae, n. sp.
This species is distinguished from exiguus
Wood by the larger size, by the absence of
reticulation of the pronotum (except in ex-
treme lateral areas of some specimens), and
by the comparatively .smaller pronotal and
elytral granules.
Male.— Length 1.2-1.4 mm, 2..3 times as
long as wide; color dark brown.
Frons broadly convex, very feebly so on
longitudinal axis; surface largely reticulate,
minute punctures moderately, uniformly
abundant, most of them feebly granulate. An-
tennal club slightly longer than scape.
Pronotum 1.0 times as long as wide; sides
almost straight and parallel on basal third,
rather broadly rounded in front; anterior
margin armed by about four to six irregular,
poorly formed serrations; summit near
middle; anterior .slope rather coarsely aspe-
rate, punctured between asperities; posterior
areas smooth, shining (except some reti-
culation present in extreme lateral areas of
some specimens), with close, moderately
large, rounded granules, anterior slope of
each granule bearing a puncture (punctures
usually visible only when light source ceph-
alad). Vestiture of fine, .short, semirecumbent
hair.
Elytra 1.3 times as long as wide, 1.4 times
as long as pronotum; .sides almost straight and
parallel on basal two-thirds, rather broadly
rounded behind; striae not impressed, each
puncture largely replaced by a large roimded
granule as wide as striae, puncture confined
to posterior slope of each granule; interstriae
as wide as striae, .smooth, shining, punctures
largely replaced by rounded granules of same
size and shape as those of striae. Declivity
steep, convex; sculpture as on disc. Vestiture
of rows of fine, short, strial hair and rows of
92
Great Basin Naturalist
Vol. 40, No. 1
erect interstrial scales, each scale slightly
shorter than distance between rows, spaced
within a row by length of scale, each four to
six times as long as wide on disc, two to four
times on declivity.
Female — Similar to male in all respects.
Type locality- Thirty km southeast of
Puttalam, Sri Lanka (Ceylon).
Type material— The male holotype, fe-
male allotype, and 34 paratypes were taken
at the type locality on 18-VI-1975, No. 214,
from Euphorbia antiquonim, by me; 28 para-
types bear the same data except they were
taken 24 km SE Puttalam. Additional para-
tvpes were taken in Sri Lanka as follows: 21
at 5 km SE Naula, 14-VM975; 14 at 48 km
N Naula, 14-VI-1975; 2 at 32 km N Haba-
rana, 12-VI-1975; 1 at 8 km SW Kurunegala,
13-VI-1975; and 1 at 11 km W Kikirawa, 19-
VL1975; all from the same host and collec-
tor.
The holotype, allotype, and half the para-
types are in the U.S. National Museum. The
remaining paratypes are in my collection.
Cryphalogenes exiguus, n. sp.
This species is distinguished from eu-
phorhiae Wood by the smaller size, by the
strongly reticulate pronotum, and by the
comparatively larger pronotal and elytral
granules.
Male.— Length 0.8-1.0 mm, 2.2 times as
long as wide; color dark brown.
Frons as in euphorbiae except more strong-
ly convex, granules smaller, less conspicuous.
Antennal club with septum in suture 1 less
apparent.
Pronotum as in euphorbiae except reti-
culate, shining, granules in posterior areas
proportionately slightly larger.
Elytra as in euphorbiae except interstrial
scales averaging more slender, those on de-
clivity not less than four times as long as
wide.
Female.— Similar to male in all respects.
Type locality.— Thirty km southeast of
Puttalam, Sri Lanka (Ceylon).
Type material- The male holotype, fe-
male allotype, and 43 paratypes were taken
at the type locality on 18-VI-1975, No. 214,
from Euphorbia antiquonim, by me. Addi-
tional paratypes were taken in Sri Lanka dur-
ing 1975 from the same host, by me, as fol-
lows: 13 at 24 km SE Puttalam, 17-VI; 5 at 5
km SE Naula, 14-VI; 4 at 48 km N Naula, 14-
VI. The specimens were taken in indepen-
dent galleries in the same stems with eu-
pliorbiae.
The holotype, allotype, and half the para-
types are in the U.S. National Museum. The
remaining paratypes are in my collection.
Cyrtogenius Strohmeyer
Ki/rtogcniits Strohmeyer, 1910, Ent. Blatt. 6:127 (Type-
species: Kijrtogeniiis bicolor Strohmeyer, mon-
obasic)
Ciirtooenius Strohmeyer, 1911, Ent. Bliitt. 7:116. Valid
emendation
Carposinus Hopkins, 1915, U.S. Dept. Agric. Rept. 99:9,
47 (Tvpe-species: Carposiniis pini Hopkins
= Lepicenis nitidus Hagedorn, original designa-
tion)
Orosiotes Niisima, 1917, Coll. Essays Y. Nawa, p. 1
(Type-species: Orosiotes kumatoensis Niisima,
monobasic)
Metahylastes Eggers, 1922, Ent. Bliitt. 18:165 (Type-spe-
cies: Metahijlastes africanus Eggers, monobasic)
Pelicenis Eggers, 1923, Zool. Meded. Roy. Mus. Nat.
Hist. Leyden 7:216 (Type-species: Lepicerus ni-
tidus Hagedorn, original designation)
Taphrohorus Nunberg, 1961, Ann. Mag. Nat. Hist.
(13)3:617 (Type-species: Taphrohorus vaticae
Nunberg, original designation)
Much confusion exists in the literature rel-
ative to the identity of this tropical genus. It
is characterized by five socketed teeth on the
lateral margin of the protibia, by the posteri-
or face of the antennal club with only one su-
ture, sutures on anterior face procurved, with
the pubescence extending to the base, by the
narrowly separated procoxae, and by the
slightly elevated or armed posterolateral
margin of the elytral declivity. Dryocoetes
differs from it by the recurved suture 1 on
the antennal club, the pubescence never ex-
tending to the base, by the contiguous pro-
coxae, and by the rounded, unarmed, post-
erolateral margins of the elytral declivity.
Both genera are phloeophagous and have
heterosanguineous polygenous breeding
habits in which the male is subequal in size to
the female and assists in the formation of new
parental galleries. Both genera have been
confused with Eulepiops (see below).
March 1980
Wood: American Bark Beetles
93
Ernocladiiis, n. gen.
This genus is distinguished from the closely
allied Ernoporus Thomson by the 3-seg-
mented antennal funicle, by the uniseriate in-
terstrial setae (interstrial ground vestiture al-
ways absent on disc, a few setae sometimes
present on declivity), and by the weakly pro-
curved (often obscure) sutures of the antennal
club.
Description.— Frons dimorphic, moder-
atelv impressed in male, convex in female.
Eye elongate-oval, entire. x\ntennal scape
elongate; funicle 3-segmented; club rather
large, sutures weakly to moderately pro-
curved, aseptate, marked by rows of setae,
grooves present or not. Pronotum with basal
margin marked by a fine, raised line, lateral
margin rounded, without a raised line; aspe-
rities in concentric rows, their bases often
contiguous or even reduced to a continuous
costa. Elytra with basal margins rounded,
strial punctures in rows, sculpture conserva-
tive; vestiture of rows of minute strial hair
and rows of erect interstrial scales, interstrial
ground vestiture absent on disc, a few short
setae in ground cover sometimes present on
declivity.
Type-species: Cn/pJialus corptilentus
Sampson.
Several additional species will be trans-
ferred to this genus as soon as their types can
be examined. Schedl (1940:590) assigned Cry-
phuhis corpulentus to Margadillius, apparent-
ly without appreciating the significance of
tlie emarginate eye or the fine, raised line on
the lateral margin of the pronotum of Marga-
dillius species.
Ernoporicus Berger
Ernoporicus Berger, 1917, Rev. Russc d'Ent. 16:242
(Tvpe-species: Ernoporicus spessivtzevi Berger,
monobasic)
Eocnjphalm Kurentzov, 1941, Acad. Sci. USSR, Koma-
rov Sta. Sci., Orient, p. 230 (Type-species: Eocnj-
pluilus seinenovi Kurentzov, monobasic)
Ernopocerus Balachowsky, 1949, Fauna de France
5():211 (Type-species: Ernoporus caitcasicus Lind-
emann, subsequent designation by Wood, 19.54,
Univ. Kansas Sci. Bull. 36:986). Xinc sijnonyiny
The complex of genera allied to Ernoporus
Thom.son have been poorly known and erro-
neously classified, largely due to the paucity
of material for study. Following an exam-
ination of the type-species of Ernoporicus,
Eocryphalus, and Ernopocerus, it was con-
cluded that these three congeneric species
have the ba.sal and lateral margins of the pro-
nottun rounded (without a fine, raised line),
the procoxae narrowly separated, the eye
short and entire, the antennal funicle 4-seg-
mented, and the antennal club with the su-
tures procurved and marked only by setae or
obsolete (never septate). Ernoporus kanawhae
Hopkins of North American and E. fagi (Fab-
ricius) and a few species from A.sia also be-
long here. The genus Ernoporus is quite dif-
ferent, as indicated below.
Ernoporus Thomson
Ernoporus Thomson, 1859, Skandinaviens Coleoptera
Svnoptiskt Bearbitade, p. 147 (Type-species: Bos-
trichus tilkic Panzer, original designation)
Cryphalops Reitter, 1889, Wiener Ent. Zeit. 8:94 (Type-
species; Cn/phalus lederi Reitter =Bostrichus
tiliae Panzer, monobasic)
Sfcphanorhopalus Hopkins, 1915, U.S. Dept. .\gric.
Rept. 99:35 (Type-species: Stephanorhopalus
nulodori Hopkins, amended to melodori by
Schedl, 1966, Ent. Abh. Mus. Dresden .35:19,
original designation)
Euptilius Schedl, 1940, Mitt. .Miincher Ent. Ges. .30:.590
(Type-species: Ernoporus concentralis Eggers,
original designation). New synonymy
Ernoporus Thomson has the basal and lat-
eral margins of the pronotum marked by a
fine, rai.sed line, the procoxae contiguous,
most pronotal asperities arranged in con-
centric rows, the antennal funicle 4-seg-
mented, the antennal club .sutures strongly
procurved to obsolete, and the elytral vesti-
ture abundant and conhused. Most of the .spe-
cies occur in tropical .\sia except for tiliae,
the type-species. In a review of the genera
belonging to this complex, it was found that
Ernoporus concentralis Eggers falls well with-
in the range of variability for Ernoporus. Be-
cause concentralis is the type-species of Eu-
ptUius Schedl, it is, therefore, necessary to
place Schedl's genus in synonymy as in-
dicated above. The structure of the pronotum
indicates that this genus is quite di.stinct from
Ernoporicus, as noted above.
94
Great Basin Naturalist
Vol. 40, No. 1
Eulepiops Schedl
Eulepiops Schedl, 19.39, J. Fed. Malay St. Mus. 18{,3):344
(Tvpe-species: EiiUpiops glaber Schedl, mon-
oba.sic)
This genus has been confused with Cyrto-
genius Strohnieyer and Dnjocoetes Eichhoff.
It differs by the protibia bearing only three
socketed teeth on the lateral margin, by the
posterior face of the antennal club with two
sutures indicated, the anterior face with su-
ture 1 straight to recurved and always on the
basal fourth. The male is either unknown or
dwarfed, deformed, flightless, and does not
participate in the formation of new parental
galleries. Reproduction is either by con-
sanguineous polygyny or possibly by some
form of parthenogenesis. The habit is myelo-
phagy for the only species observed. Dnjo-
coetes coffeae Eggers and its allies belong to
this genus.
Hadrodemius, n. gen.
This genus is distinguished from Eccoptop-
tems Eichhoff by the tibiae being of normal
size and all bearing socketed teeth, by the
normal metatarsi (not compressed), by the
declivity being restricted to the posterior half
of the elytra, and by the convex to moder-
ately impressed, unarmed elytral declivity.
Description.— Body very stout, less than
1.8 times as long as wide, usually black. An-
tennal club with posterior face unmarked by
sutures, on anterior face costa marking apical
margin of corneous area usually forming a
complete ring. Scutellum visible only on an-
terior declivous slope of elytral margins. De-
clivity and tibiae as described in above diag-
nosis.
Type-species: Xylebarus globus Blandford.
Members of the Xyleborus globus species
group should be referred here.
Hylurdrectonus Schedl
Hylurdrectomis Schedl, 1938, Trans. Roy. Soc. S. .'Vustra-
lia 62;4() (Type-species: Hylurdrectonus piniarius
Schedl, monobasic)
Xylogopinus Schedl, 1972, Papua New Guinea .\5;ric. J.
23:64 (Type-species: Xylogopinus araucariae
Schedl = Hylurdrectonus corticinus Wood, mon-
obasic). New synonymy
A review of long series of Hylurdrectonus
piniarius Schedl, H. araucariae Schedl
(1964a:213), and Xylogopinus araucariae
Schedl indicates the absence of characters
that will separate these two genera. Con-
sequently, it is necessary to place Xylogo-
pinus in synonymy under the older name as
indicated above. This act creates homonymy
as indicated below.
Hylurdrectonus corticinus, new name
Xillogopinus araucariae Schedl, 1972, Papua New Guin-
ea Agric. J. 23:64 (Bulolo, Morobe Distr., New
Guinea)
A long series of this species was collected
near Bulolo and compared to the holotype
and paratypes in the Forest Research Labora-
tory collection at Bulolo. As indicated above,
this species must be transferred to Hylurdrec-
tonus. The transfer makes this species a jun-
ior homonym of H. araucariae Schedl, 1964.
Tlie new name Hylurdrectonus corticinus is
proposed to replace H. araucariae (Schedl
1972).
Leptoxyleborus, n. gen.
This genus is distinguished from the allied
Theobortis Hopkins and Coptoborus Hopkins
by the declivity commencing anterior to the
middle of the elytra, its lower half broadly
impressed and either flat or shallowly con-
cave. If the discal interstrial punctures are
uniseriate, then the declivital surface is den-
sely covered by small, confused scales; if the
declivital setae are hairlike, then the discal
interstrial punctures are confused.
Description.— Antennal club with two su-
tures indicated on posterior face, anterior
face with segment 2 comparatively large,
sclerotized, convex, apical portion beyond
segment 2 flat to concave. Protibiae and
metatibiae each armed by six or seven sock-
eted teeth. Anterior coxae contiguous. Scutel-
lum visible. Declivity as described in above
diagnosis.
Type-species: Phloeotrogus sordicauda
Motschulsky.
Other species placed previously in the
Xyleborus sordicauda group also belong here.
Micropenis, n. gen.
This genus is distinguished from Taphro-
dastis Wood by the convex elvtral declivity
tliat lacks a circumdeclivital costa, by the ab-
March 1980
Wood: American Bark Beetles
95
sence of declivital scales, and by the strial
pimctures that are arranged in definite rows.
Description.— Body slender, at least two
times as long as wide, color yellowish or red-
dish brown. Posterior face of antennal club
with at least one suture visible, apical margin
of corneous area never costate. Scutellum not
visible. Strial punctures usually in rows. De-
clivity convex, variously sculptured, without
a costa.
Type-species: Xyleborus theae Eggers.
Members of the Xijleborus theae species
group should be referred here. The name Mi-
cropenis was originally coined by F. G.
Browne for this group for use in an unpub-
lished manuscript a decade ago.
Ozopeinon Hagedorn
Ozopcmon Hagedorn, 1908, Deutsche Ent. Zeitschr.
1908:382 (Type-species: Ozopemon regius Hage-
dorn, monobasic)
Dryocoetiops Schedl, 1957, Ann. Mus. Roy. Congo Beige,
Tervairen, Ser. 8, Sci. Zool. .56:13 (Type-species;
Ozopemon laevis Strohmeyer, monobasic). New
sijnonymy
A series of Ozopemon laevis Strohmeyer
was compared to Eggers's series of this spe-
cies and to representatives of eight species of
Ozopemon. Although the sculpturing of the
pronotum is somewhat unique for the genus,
this species appears to fall well within the
limits of variability for Ozopemon. For this
reason, Dryocoetiops is placed in synonymy
as indicated above.
Scolytogenes Eichhoff
Scolytogenes Eichhoff. 1878, preprint of Mem Soc. Roy.
Sci. Liege (2)8:475, 497 (Type-species: Scolyto-
genes danvinii Eichhoff. monobasic)
Cryphcilomorphus Schaufnss, 1890 (1891), Tijdschr. Ent.
34:12 (preprint 1890 by Martinus N'ijhoff, Hagg)
(Type-species: Cryphahnurpbiis conimiinif: Schau-
fnss, monobasic). \eiv synonymy
Eggers (1929:53) examined the type-speci-
mens of the type-species of Scolytogenes and
Lepicerus and compared them to the type-
specimens of Negritus major Eggers and A'.
minor Eggers. He concluded that N. major
and N. minor were congeneric with Scoly-
todes danvinii Eichhoff. The holotype of S.
darwinii apparently was lost when the Stettin
Museum was damaged during World War II.
In the absence of that type, direct com-
parisons are not now possible; however, if it
is assumed that Eggers was correct in his ob-
servations, then N. major and S. danvinii are
congeneric. My examination of the lectotype
of A', major and .syntypes of N. ater (type-spe-
cies of Negritus) demonstrates that these spe-
cies are congeneric; consequently, Negritus
must be a jimior synonym of Scolytogenes.
Because N. major and A\ ater are also consid-
ered congeneric with Cryplmlomorphus com-
munis Schaufuss (type-species of Cryphalo-
morplius) (Schedl 1957:152), it must also be
concluded that Cryphalomorphus is a junior
synonym of Scolytogenes.
(Note added in press: The list of types in
the Schedl Collection at the Vienna Museum,
just received, includes the type of S. darwinii.
It will be examined as soon as arrangements
can be completed.)
Stephanopodius Schedl
Stephanopodius Schedl, 1941, Rev. Zool. Bot. Afr. .34:.396
(Type-species: Stephanoderes dispar Eggers, sub-
sequent designation by Schedl, 1961, Rev. Ent.
Mozambique 4:6.33)
Cryplwlomimus Browne, 1962, West .\frican Timber Bo-
rer Research Unit Rept. 5:75 (Type-species: Hy-
pocryphahts ghanaensis Schedl. original designa-
tion)
Cryphcdmomimetes Browne, 1963, Ann. Mag. Nat. Hist.
(1.3)6:242 (Replacement name). Xcii synonymy
When Schedl named Htjpocryphalus gha-
naensis and then later (Schedl 1964b:305)
transferred this species from Cryphalomi-
metes back to Hypocnjphalus, he overlooked
some very important characters. In this spe-
cies and in Stephanopodius, the basal margin
of the pronotum bears a fine, raised line, but
the lateral margin is rounded and lacks the
fine, raised line of Hypocryphalus. In addi-
tion, the antennal club is quite different from
Hypocryphalus. The species ghanaensis is
congeneric with Stephanopodius dispar (Egg-
ers) and, as indicated above, should be trans-
ferred to that genus. Cryphalomimetes is,
therefore, a .synonym of Stephanopodius
Schedl and not of Hypocryphalus Hopkins.
Taphrodasus, n. gen.
This genus is distinguished from Micro-
penis Wood by the confused interstrial pimc-
tures, by the presence of scales on the elytral
96
Great Basin Naturalist
Vol. 40, No. 1
declivity, and by the strongly concave decliv-
ity that commences on the basal half of the
elytra and is marked on its lateral margins in
such a way as to form a blunt, elongate, cir-
cumdeclivital costa.
Description.— Body slender, at least 2.0
times as long as wide, color reddish brown.
Posterior face of antennal club with one su-
ture visible, apical margin of corneous area
never costate. Scutellum not visible. Strial
punctures on disc confused. Declivity as de-
scribed in above diagnosis.
Type-species: Xylehorus percorthylus
Schedl.
Taurodemus, n. gen.
This genus is distinguished from Xijleborus
Eichhoff by the moderately to rather widely
separated procoxae, by the rather stout body,
by the presence of 9 to 12 socketed protibial
teeth, and by the distinctive sculpture of the
sulcate elytral declivity.
Description.— Body stout, less than 1.9
times as long as wide. Antennal club with
segment 1 corneous, without any sutures evi-
dent on posterior face, apical margin of seg-
ment 1 on anterior face acutely elevated into
a continuous costa forming a complete circle.
Procoxae moderately to rather widely sepa-
rated. Protibia armed by 9 to 12 socketed
teeth. Elytral declivity moderately to very
strongly sulcate on at least basal half, lateral
margins armed by at least one major spine
and several smaller tubercles.
Type-species: Xijleborus sharpi Blandford.
The following species are transferred from
Xylehorus to Taurodemus: bicornutus Wood,
ebenus Wood, (Bostrichus) flavipes Fabricius,
godmani Blandford, pandulus Wood, {Am-
phic(iranus) perebeae Ferrari, salvini Bland-
ford, sanguinicollis Blandford, sharpi Bland-
ford, splendidus Schaufuss, (Bostrichus)
varians Fabricius, and varus Wood.
Xyleborus Eichhoff
Xijleborus Eichhoff, 1864, Berliner Ent. Zeitschr. 8:37
(Type-species: Bostrichus monographus Fabricius,
subsequent designation bv Hopkins. 1914, Proc.
U.S. Nat. Mus. 48:131)
The genus Xyleborus Eichhoff, as inter-
preted in recent years by Schedl, contains
more than 1400 nominate species, that is, vir-
tually all the species in the tribe Xyleborini.
However, the diversity of characters and
habits within this group suggests the exist-
ence of several distinct clusters of species and
species groups that could and should be char-
acterized as genera. The difficulty in frag-
menting the group piecemeal, as has been at-
tempted by some workers, is that when one
group is removed and elevated to generic
rank, the remainder becomes unclassifiable
on a logical, phylogenetic basis. In order to
remedy this situation, a classification is being
composed, based on such constant features as
the location of mycetangia, structure of the
antennal club, form and armature of the
tibiae, characters of the scutellum, and many
other features. A deliberate effort is being
made to avoid use of adaptive characters
such as the surface sculpturing of the pro-
notum and elytra.
Tentatively, 27 groups are being given ge-
neric status within the Xyleborini. Those de-
scribed previously include: Ambrosiodmus
{ = Browneia, Phloeotrogus), Arixyleborus
{=Xyleboricus), Cnestus { = Tosaxyleborus),
Coptoborus { = Streptocranus), Cryptoxyle-
borus, Dryocoetoides, Eccoptoptcrus { = Eu-
rydactylus, Platydactylus), Euwallacea, Ka-
lantanius, Mesoscolytus, Notoxyleborus,
Premnobius { = Prernnophilus), Pseudoxyle-
borus, Sampsonius, Schedlia, Theoborus,
Webbia { = Prowebbia, Pseudowebbia, Xely-
borus), Xyleborinus, Xyleborus {=Anaeritus,
Anisandrus, Boroxylon, Bufonus, Coptodryas,
Cyclorhipidion, Heteroborips, Phloeotrogus,
Progenius, Terminalinus, Xyleborips), and
Xylosandrus. The above is mentioned to es-
tablish a context into which the seven genera
in this tribe, described in this article, can fit.
The seven include: Anaxyleborus, Apoxylc-
borus, Hadrodemius, Leptoxyleborus, Micro-
penis, Taphrodasus, and Taurodemus.
Xylechinus Chapuis
Xylechinus Chapuis, 1869, Synopsis des Scolvtides, p. 36
(Type-species: Dcndroctontis pilosiis Knoch)
Squamasinuhis Nunberg, 1964, Ann. Hist. -Nat. Mus.
Nat. Hungarici, Pars Zool. 56:431 (Type-species:
S(iu(im(isinulus chiliensis Nunberg, original desig-
nation). New synonymy
When the holotype of Squamasinuhis chi-
liensis Nunberg and several allied species
March 1980
Wood: American Bark Beetles
97
from South American were examined, no
characters could be found that distinguish
this genus from Xylechinus Chapuis. As
nearly as can be determined at the present
time, the genus Xylechinus consists of 14
Central and South American, 2 North Ameri-
can, 5 Asian, and 1 European species. Schedl
has referred four New Guinean and Austra-
lian species to this genus, all of which appar-
ently should be transferred to Acrantus. One
African species placed in Xijlechinus by
Schedl apparently belongs elsewhere.
Literature Cited
Blandford, W. F. H. 1894. The Rhynochophorous Co-
leoptera of Japan. Part III. Scolytidae. Trans.
Ent. Soc. London, pp. 53-14L
Eggers, H. 1929. Zvir Synonymic der Borkeiikafer
(Ipidae, Col.). Wiener Ent. Zeit. 46:41-55.
Sc;hedl, K. E. 1940. Zur Einteilung und Synonvmie der
Cryphalinae (Col. Scoivt.). Mitt. Miichner Ent.
Ges. .30:583-591.
1957. Bark- and timber-beetles from South .Af-
rica. Ann. Mag. Nat. Hi.st. (12)10:149-159.
1963. Zur Synonymie der Borkenkafer IX. Ent.
Abh. Mus. Tierk. Dresden 28:257-268.
1964a. On some Coleoptera of economic impor-
tance from New Guinea and Australia. Pacific In-
sects 6:211-214.
1964b. Zur Synonymie der Borkenkafer XV. Rei-
chenbachia 3:303-317.
1966. Neotropi.sche Scolytoidea VIII. Ent. Arb.
Mus. Frey 17:74-128.
Wood, S. L. 1978. A reclassification of the subfamihes
and tribes of Scolytidae (Coleoptera). Ann. Soc.
Ent. France (N.S.).' 14:95-122.
THE BACTERIUM THIOPLOCA INGRICA ON WET WALLS
IN ZION NATIONAL PARK, UTAH
Samuel R. Riishforth', Sheril D. Burton", Jeffrey R. Johansen', and Judith A. Grimes'
.\bstract.— Hanging gardens and wet wall habitats have been studied for the past several years in many arid
regions of the Intermountain West. One unusual large wet wall habitat in Zion National Park was found to be cov-
ered with a mucilaginous red-colored growth of the filamentous gliding bacterium Thioploca ingrica Visloukh. This is
the only habitat we have examined where the predominant matrix-forming organism was a bacterium rather than an
alga.
Hanging gardens are unusual habitats
found in several areas around the world, par-
ticularly in western North America. Such
habitats form when water percolates vertical-
ly through permeable rock facies (generally
sandstone) imtil it reaches an impervious lay-
er. The water then moves laterally imtil it
exits the rock formation often on a vertical
wall or cliff. Such exit springs often occur
along fairly long horizontal lines to form lin-
ear seep walls or wet walls. Such habitats
rapidly become colonized by a variety of
mesic plant species, some of which are en-
demic to such gardens. Seep walls become
weakened through time, particularly in the
massive mesozoic sandstone formations typi-
cal of areas of southern Utah and northern
Arizona. When this occurs, large portions of
the wall slough away to form grottos that are
.shaded from the sim and are cooler and more
humid than .surrounding areas.
Such hanging garden habitats have been
imder study for several years (Rushforth et al.
1976, Clark and Rushforth, in press, Welsh
and Moore 1965a, 1965b). We have been
particularly interested in the algal species
that colonize hanging garden walls. The algal
floras of such habitats are variable depending
upon several factors, primarily the amount of
water available. The moister walls are almost
always covered with heavy growth of green
or blue green algae that secrete copious
mucilage. Such algal mats are in turn colo-
nized by dozens of other algal species, in-
cluding blue green algae, green algae, eu-
glenophytes, golden algae, and diatoms. Algal
diversity is really quite high in many of these
gardens, and the species are often imusual in
morphology and/or distribution.
We have found that wet wall faces that are
exposed to the sun are often colonized by al-
gae with dominant nonchlorophylous pig-
ments. Thus, on certain walls, the green alga
Palmella miniata Leiblein is common. This
organism is often a deep red color due to
hematochrome pigments, and in turn the gar-
den walls colonized by these organisms are
often a beautiful brick-red color. Likewise,
Scytonema myochrous (Dillw.) C. A. Agardh
and Scytoneina alatiim (Carm.) Borzi are
prevalent on some walls. These organisms are
large, prominently ensheathed blue green al-
gae. The sheath and the cell wall of these al-
gae become colored as the organisms mature
to form yellow or yellow brown filaments
which also color their walls of colonization.
Recently, we have been studying algae and
lichens of Zion National Park. One promi-
nent wet wall in the park is Weeping Rock
(Fig. 1), a famous tourist attraction. This wall
is very large and is usually very moist. It is
heavily colonized by many algal species. In
several places on the wall, bright orange red
patches and streaks are evident. We collected
specimens from these areas expecting to find
Palmella miniata. Upon returning to the lab-
'Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602.
■Department of Microbiology, Brigham Young University, Provo, Utah 84602.
98
March 1980
RUSHFORTH ET AL.: HANGING GaRDENS BaCTERIUM
99
Fig. 1-4. 1. Habit of Tliioplora and algal communities of Weeping Rock, Zion National Park; 2, "Braid" of Thio-
ploca im^rica filaments in a common sheath; .3, Thioplocd with Chamae.siphon species attached; 4, Low magnification
photograph showing heavy epiphytic growth of Chamaesiphon species.
100
Great Basin Naturalist
Vol. 40, No. 1
oratory, we examined the organisms using a
Zeiss RA Research microscope with No-
marski differential phase contrast accessories.
We were surprised to find that 'the red color-
ation was not due to Palmella but to a fila-
mentous prokaryote. Careful study elimi-
nated known blue green algae as the causal
organism. Further examination showed the
organism to be a bacterium in the family
Beggiatoaceae Migula. These bacteria are
gram-negative filaments that are motile by
gliding motion. Three genera are presently
placed in this family: Beggiatoa Trevisan,
Vitreoscilh Pringsheim, and Thioploca Lau-
terbom. In addition, the three genera Bactos-
cilla Pringsheim, Flexoscilla Pringsheim, and
Thiospirillopsis Uphof are possible members
of the family (Leadbetter 1974).
The organisms we collected from Weeping
Rock in Zion National Park may be placed in
the genus Thioploca, based upon the presence
of more than one filament in the sheaths
(Figs. 2-4). Furthermore, the filaments
ranged between the width of 2 and 7 um,
placing them in Thioploca ingrica Visloukh.
The filaments often served as a substrate for
epiphytic algae, particularly Cluimaesiphon
species (Fig. 3).
This observation of T. ingrica is interesting
for several reasons. First, it represents the
only example in the several gardens we have
studied where the predominant mucilaginous,
matrix-forming organism was a bacterium
rather than an alga. Second, it represents the
only reported occurrence of Thioploca from
intermountain western North America that
we are aware of. Third, Thioploca usually ex-
hibits a greenish blue color. However, the
specimens we have collected produce a
bright orange red color on the wall itself and
a paler orange color when examined beneath
the microscope. The nature of this color is
unknown, although Beggiatoaceae are report-
ed to not form carotenoid pigments. And
fourth, the presence of this organism on the
moist walls is itself unusual because all the
reports to date that we are aware of chron-
icle Thioploca species as inhabiting the upper
layers of bottom muds of freshwater and
brackish habitats (Maier 1974). Specifically,
such organisms have been collected from
both oxidizing and reducing environments in
such muds (Perfil'ev et al. 1965).
Literature Cited
Clark, R. L., and S. R. Rushforth. In review. A study
of the algal flora of selected hanging gardens of
Glen Canyon, Utah. Great Basin Naturalist Mem-
oirs.
Leadbetter, E. R. 1974. Beggiatoaceae Migula. Pages
112-115 in R. E. Buchanan, N. E. Gibbons, eds.
Bergey's manual of determinative bacteriology,
8th ed. Williams and Wilkins, Baltimore.
Maier, S. 1974. Thioploca Lauterborn. Pages 113-116 in
R. E. Buchanan, N. E. Gibbons, eds. Bergey's
manual of determinative bacteriology, 8th ed.
Williams and Wilkins, Baltimore.
Perfil'ev, B. V., D. R. Gabe, A. M. Galperina, V. A.
Rabinovich, a. a. Sapotnitskii, E. E. Sherman,
AND E. P. Troshanov. 1964. Applied capillary
microscopy: The role of microorganisms in the
formation of iron-manganese deposits. Izadatels-
tov, Akad Nauk SSSR, Savarenskii Laboratory for
Hydrogeological Problems. Moscow. In Russian
(Transl. Consultants Bureau Enterprise Inc. 1965
New York).
Rushforth, S. R., L. L. St. Clair, T. A. Leslie, K. H.
Thorne, and D. S. Anderson. 1976. The algal
flora of two hanging gardens in southeastern
Utah. Nova Hedwigia 27:231-323.
Welsh, S. L., and G. Moore. 1965a. Plants of Canvon-
lands National Park. Part I. The Needles Region.
Proc. Utah Acad. Sci. 42(1): 160.
Welsh, S. L., and G. Moore. 1965li. Plants of Canyon-
lands National Park. Part II. The Islands in the
Sky Region. Proc. Utah Acad. Sci. 42(1):160-161.
NOTICE TO CONTRIBUTORS
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TABLE OF CONTENTS
Some aspects of succession in the spruce-fir forest zone of northern Utah. David J.
Schimpf, Jan A. Henderson, and James A. MacMahon 1
Utah Flora: Malvaceae. Stanley L. Welsh 27
Utah Flora: Miscellaneous families. Stanley L. Welsh 38
The taxonomic status of the rosy boa Lichamira roseofusca (Serpentes: Boidae). John
R. Ottley, Robert W. Murphy, and Geoffrey V. Smith 59
Hespewperla lio^uei, a new species of stonefly from California (Plecoptera: Perlidae).
Richard W. Baumann and Bill P. Stark 63
Reproduction in three sympatric lizard species from west-central Utah. John B.
Andre and James A. MacMahon 68
Haplopappus alpinus (Asteraceae): A new species from Nevada. Loran C. Anderson .. 73
Miscellaneous plant novelties from Alaska, Nevada, and Utah. Stanley L. Welsh and
Sherel Goodrich 78
New genera and new generic synonvmv in Scolytidae (Coleoptera). Stephen L.
Wood [ '.....'. '. 89
The bacterium Thioplova ingrica on wet walls in Zion National Park, Utah. Samuel
R. Rushforth, Sheril D. Burton, Jeffrey R. Johansen, and Judith A. Grimes 98
HE GREAT BASIN NATURALIST
ume 40 No. 2
June 30, 1980
Brigham Young University
MUS. COMP. ZOOU,
NOV
GREAT BASIN NATURALIST
Editor. Stephen L. Wood, Department of Zoology, Brigham Young University, Provo, Utah
84602.
Editorial Board. Kimball T. Harper, Botany; Wilmer W. Tanner, Life Science Museum;
Stanley L. Welsh, Botany; Clayton M. White, Zoology.
Ex Officio Editorial Board Members. A. Lester Allen, Dean, College of Biological and Agricul-
tural Sciences; Ernest L. Olson, Director, Brigham Young University Press, University
Editor.
The Great Basin Naturalist was founded in 1939 by Vasco M. Tanner. It has been published
from one to four times a year since then by Brigham Young University, Provo, Utah. In gener-
al, only previously unpublished manuscripts of less than 100 printed pages in length and per-
taining to the biological and natural history of western North America are accepted. The
Great Basin Naturalist Memoirs was established in 1976 for scholarly works in biological natu-
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cover.
5-80 650 45971
ISSN 0017-3614
The Great Basin Naturalist
Published at Provo, Utah, by
Brigham Young University
ISSN 0017-3614
Volume 40
June 30, 1980
No. 2
FEEDING ECOLOGY OF GILA BORAXOBIUS (OSTEICHTHYES: CYPRINIDAE)
ENDEMIC TO A THERMAL LAKE IN SOUTHEASTERN OREGON'
Jack E. Williams' and Cynthia D. W'illianis'
.\bstr.\ct.— (w/a hoiaxchiit.s is a dwarf species of cvprinid endemic to a thermal lake in southeastern Oregon.
Despite a relatively depauperate fauna and flora in the lake, 24 food items were foimd in intestines of G. horaxobins.
Ten of the 24 foods, including six insects, were of terrestrial origin. The relative importance of food items fliictuated
seasonally. Diatoms, chironomid larvae, microcrustaceans, and dipteran adults were the primary foods during spring.
In summer, diatoms decreased and terrestrial insects increased in importance. During autumn important foods were
terrestrial insects, chironomid larvae, and diatoms. Diatoms and microcrustaceans increased in importance during
winter. Chironomid larvae were of importance in winter, when the importance of terrestrial food items decreased
.substantially. Similar food habits were observed between juveniles and adults, except that adults consumed more
gastropods and diatoms and juveniles consumed more copepods and terrestrial insects. Gila howxobius feeds op-
portunistically with individuals commonly containing mostly one food item. Fish typically feed by picking foods
from soft bottom sediments or from rocks. However, fish will feed throughout the water column or on the surface if
food is abundant there. Gila boraxohiiis feeds throughout the day, with a peak in feeding activity just after sunset. .\
daily ration of H.l percent boch' weight was calculated for the species during June. .\ comparison of food habits
among G. boraxobius and populations of G. alvordensis during the summer shows that all are opportimistic in feed-
ing, but that G. boraxobius relies more heavily on terrestrial foods.
The Borax Lake chub, Gilo boraxobius, is a
rare species of cvprinid fish endemic to a
thermal lake in the Alvord Basin of south-
eastern Oregon. The restricted habitat of G.
boraxobius is threatened bv geothermal
energy development in the Alvord. In recog-
nition of this problem, the species is listed as
"threatened" by the Endangered Species
Committee of the American Fi.sheries Society
(Deacon et al. 1979) and is currently on the
protected list of the Oregon Department of
Fish and Wildlife.
Gila boraxobius was recently described and
has been diagnosed as a dwarf relative of G.
alvordensis (Williams and Bond, in press).
Typical adult size of Gila boraxobius is 33-50
mm standard length (SL). Individuals longer
than 55 mm SL are rare.
Becau.se no life history information was
known concerning this species, a study was
conducted to determine its food habits rela-
tive to sea.sons, fish length, diel feeding chro-
nology, and food habits of other Alvord Basin
fishes of the genus Gila.
Methods
Feeding ecology of Gila boraxobius was in-
vestigated by analyzing intestinal contents of
fish collected monthly from March 1978 to
June 1979. Fish were collected from the
southwest one-quarter of Borax Lake using a
3 X 5 mm me.sh seine approximately 3 m in
length. Specimens were preserved in 10 per-
cent formalin and transferred to 45 percent
isopropanol after one week. Standard length
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
'Technical Paper 5335, Oregon .Agricultural Experiment Station.
■Department of Fisheries and Wildlife. Oregon State University. Corvallis, Oregon 97331.
101
102
Great Basin Naturalist
Vol. 40, No. 2
the nearest 0.01 g. Gila boraxohius, like other
cvprinids, has no stomach; therefore, contents
of the intestine, from esophagus to anus, were
removed and examined under dissecting mi-
croscopes. Percent frequency of occurrence,
mean number per intestine, mean percent
volume, and a value of relative importance
are reported for each food item. Percent fre-
quency of occurrence is defined as the num-
ber of intestine samples in which one or more
of a given food item is found expressed as a
percentage of all nonemptv intestines exam-
ined (Windell and Bo wen 1978). The total
number of a given food item observed in the
intestines divided by the number of nonemp-
tv intestines examined is the mean number
per intestine. Mean percent volume is de-
fined as the total volume estimates for a giv-
en food item divided by the number of non-
empty intestines examined. Percent volumes
were derived by separating the intestine into
three subsamples and visually estimating the
percent contribution of a given food item in
each sample. The percent contribution of
each subsample to the contents of the entire
intestine was estimated so that the volume of
a <£iven food item relative to all intestinal
contents could be made. Percent frequency
of occurrence, mean number per intestine,
and mean percent volume each contain a bias
which limits their usefulness when used sepa-
rately (Windell and Bowen 1978). For ex-
ample, percent frequency of occurrence
overemphasizes the importance of small food
items that may be ingested frequently but
have a small impact on the volume of food in
the intestine. On the other hand, mean per-
cent volume overemphasizes the importance
of large food items, such as adult insects, that
may be consumed infrequently but have a
large volume. To offset these biases against
each other, an index of relative importance
(RI) is reported for each food item. The rela-
tive importance index combines the percent
frequency of occurrence and mean percent
volume for food item a into an absolute im-
portance index (AI„) as follows:
AT = P^''<-'(-'"t frequency of occurrence +
mean percent volume
RI„ = 100AI„/SAI„, where n is the number
of different food items. The determination of
Rio and AI^ are by methods modified from
George and Hadley (1979).
Invertebrate identification was based on
Pennak (1978).
Food habits of G. boraxohius are summa-
rized as follows: (1) seasonal variation of
foods consumed, (2) foods consumed by dif-
ferent size classes of fish, (3) diel feeding
chronology, and (4) comparison with food
habits of two populations of Gila alvordcnsis.
Changes in food habits with season and fish
size class were determined from monthly col-
lections taken from March 1978 through Jan-
uary 1979. Monthly collections were grouped
into seasons as follows: spring (March-May),
summer (June-August), autumn (Septem-
ber-November) and winter (Decem-
ber-January). To compare changes in food
habits with fish size, two size classes of fish
were compared, 15.0 mm to 32.9 mm SL
(juveniles), and 33.0 mm SL or longer
(adults). Gila boraxobius matures at approx-
imately 33.0 mm SL (Williams and Bond, in
press). Diel feeding chronology was deter-
mined from collections made at 3-hour inter-
vals during a 24-hour period in June 1979.
Fullness of the intestine was determined ac-
cording to the formula defined by Hureau
(1969 in Berg 1979) as follows:
_ weight of ingested food X 100 percent
weight of fish
(Ir = L'indice de repletion = fullness index).
The daily ration of food for G. boraxobius
was estimated from diel trajectories of the
fullness of the intestine. This estimate was de-
rived by the following formula proposed by
Bajkov (1935) and modified by Darnell and
Meierotto (1962) and Eggers (1977):
Rt = 24Sa
where Rj is the daily ration, S is the average
weight of intestinal contents expressed as
percent of body weight during the 24-hour
period, and a is the intestinal evacuation
rate. An intestinal evacuation rate of 0.2 hr'
is assumed (Eggers 1977).
Intestinal contents of Gila alvordcnsis col-
lected from Thousand Creek, Nevada, on 13
June 1978 and G. alvordensis collected from
Serrano Pond, Oregon, on 6 August 1977
were compared to those of G. boraxobius col-
June 1980
Williams, Williams: Borax Lake Chub
103
lected during the summer of 1978. Methods
of collection and food habits analysis for pop-
ulations of G. alvordensis were the same as
those used for G. boraxobius.
Study Area
Fish were collected from Borax ( = Hot)
Lake, Serrano Pond, and Thousand Creek, all
in the Alvord Basin of southeastern Oregon
and northwestern Nevada. The Alvord is an
endorheic basin of semiarid climate, sur-
rounded by fault-block mountain ranges. Bo-
rax Lake'(T37S, R33E, Sec. 14, Harney
County, Oregon) is a 4.1 ha natural lake that
receives water from several thermal springs
in the lake bottom. Water temperature in
Borax Lake is typically 29-32 C, with ex-
tremes of 35 C or greater near spring sources
and 17 C near the lake margin during winter.
The lake is relatively shallow, with a soft, sil-
tv bottom interspersed with rocks and hard
outcroppings. The water is transparent, and
aquatic vegetation is limited to a few areas
along the lake margin. Sodium is the princi-
pal cation, whereas bicarbonate, sulfate, and
chloride are the major anions in Borax Lake
(Mariner et al. 1974). Specific conductance of
the water is 2410. Serrano Pond (T36S,
R33E, Sec. 1, Harney County, Oregon) is a
0.1 ha reservoir that receives water from a
cool spring approximately 60 m distant. Wa-
ter temperature is usually 16-21 C during the
summer. The bottom of this shallow pond is
primarily silt. The water is turbid, and aquat-
ic vegetation is abundant. Thousand Creek
(collection site at T46N, R28E, Sec. 34,
Humboldt County, Nevada) is a small, shal-
low stream rarely exceeding 2 m in width.
The creek often becomes intermittent during
summer months. Water temperature varies
from 15-27 C during the summer, fluctuating
rather closely with air temperature. The bot-
tom is a silt and gravel mix. The water is tur-
bid, and aquatic vegetation is moderately
abundant.
Results
Seasonal Variation in Foods Consumed
Twenty-four different food items were
found in intestines of Gila boraxobius during
this study (Tables 1-4). Ten of the 24 food
items were encountered in all seasons. Many
food items fluctuated seasonally in occur-
rence; however, some insects, especially chi-
ronomid larvae, diatoms, and micro-
crustaceans, were of importance throughout
the year. During the spring, algae, chirono-
mid larvae, copepods, dipteran adults, and
ostracods were the predominant food items
(Table 1). Algae, which was composed almost
wholly of diatoms, was the most frequently
ingested food during the spring, occurring in
over one-half of the intestines. Some of the
diatoms were secondarily ingested with mi-
crocmstaceans; however, in some individuals
diatoms accounted for 70-80 percent of the
intestinal contents by volume. The high vol-
ume suggests that diatoms are not exclusively
the result of secondary ingestion but are a
preferred food item for most fish. The most
common diatom observed in intestines was a
benthic species, Denticida thermalis. Nov-
iciila sp., Synedra sp. Achnanthes lanceolata,
and A. minutissima were observed in lesser
numbers. Both Achnanthes species are ben-
thic, whereas the Navicula and Synedra spe-
cies could be benthic or planktonic forms.
Adult dipterans accounted for the highest
mean percent volume, nearly 20 percent, of
all food items during spring. Several fish fed
on dipteran adults exclusive of other foods.
During May 1978 dipteran adults were heav-
ily utilized, appearing in 19 of 23 intestines
examined. During the summer, diatoms were
less frequently encountered in intestines and
comprised a smaller mean percent volume
than in spring. Chironomid larvae, copepods,
dipteran adults, and gastropods were the
most important food items in summer (Table
2). Gila boraxobius utilized more terrestrial
insects and spiders and fewer micro-
crustaceans and diatoms in summer than in
spring. Terrestrial insects and spiders ac-
counted for approximately 31 percent mean
volume of foods consumed during summer
compared to approximately 21 percent in
spring. During autumn terrestrial insects, chi-
ronomid larvae, and diatoms were the princi-
pal food items (Table 3). In winter Gila
boraxobius relied more heavily on autoch-
thonous food items, utilizing primarily dia-
toms, ostracods, copepods, chironomid larvae
and cladocerans (Table 4). Terrestrial insects.
104
Great Basin Naturalist
Vol. 40, No. 2
which were of importance in spring, summer,
and autumn, seldom appeared in intestines of
G. boraxobius during winter, when they con-
tributed only 2 percent mean volume. Food
items consumed in autumn and winter were
less diverse than in other seasons. Sixteen
food items were observed in fish collected in
autumn and winter, 20 food items during
spring, and 21 during summer. Aquatic in-
sects were important foods throughout the
year, comprising mean volumes of approx-
imately 19 percent, 23 percent, 16 percent,
and 13 percent in spring, summer, autumn,
and winter, respectively. The primary con-
tributor to these high values were chirono-
mid larvae, which consistently exhibited a
high relative importance. Chironomid pupae
and Odonata nymphs were also consumed
throughout the year but were of much less
importance. Coleoptera larvae and aquatic
Coleoptera adults were utilized to lesser de-
grees seasonally. The increased consumption
of copepods, ostracods, and cladocerans in
the winter was dramatic. These micro-
CRistaceans comprised approximately 35 per-
cent mean volume of intestines during win-
ter, but only 16.5 percent, 12 percent and 4.5
percent in spring, summer, and autumn, re-
spectively. Large amounts of inorganic debris
were found in intestines throughout the year.
Tliis was probably ingested accidently while
the fish were feeding on bottom organisms.
Many important foods in Borax Lake, such as
insect larvae, gastropods, diatoms, and prob-
ably many small invertebrates, are benthic.
Observations in Borax Lake and in aquaria
show that G. boraxobius feeds primarily by
rooting around in bottom material and pick-
Table 1. Contents of 71 intestines oi Cihi boraxobius collected during the spring of 1978. ND = no data.
Item ingested
Percent
frequency
of occurrence
Mean number
per intestine
Mean percent
volume
RI
.\lgae'
C'.astropods-
C.astropod eggs
Haplotaxid oligochaetes
Harpacticoid copepods
Ostracods
Cladocerans
Plant seeds
Higher plants
Fish scales
Araneae
Insect eggs
Unidentified insects'
Tekrestrial i\sec;ts
CoUemhola
Thysanoptera adults
Hemiptera adults
Coleoptera adults
Hymcnoptera adults
Diptera adults
Asiatic: insects
Chironomid larvae
C;hironomid pupae
Odonata nvmphs
Klmid larvae
C'oleoptera adults
In()Iu;amc: dehhis
56.14
ND
12.67
15.70
22.81
1.07
6.69
6.73
0.00
0.00
0.00
().(M)
1.75
0.11
0.32
0.47
49.12
44.95
10.38
13..57
40.35
8.39
5.91
10..55
15.79
0.21
0.17
3.64
5.26
0.11
0.06
1.21
12.28
ND
1.72
3.19
3.51
0.11
0.04
0.81
3.51
0.04
0.34
0.88
10.53
12.26
2.15
2.89
26.32
ND
2.50
6..57
0.00
().(H)
0.(X)
0.00
3.51
0.04
0..39
0.89
0.00
0.00
0.00
0.(K)
1.75
0.02
0.21
0.45
3.51
0.04
0.49
0.91
35.09
1.93
19.49
12.45
47.37
12.30
13.71
13.93
7.02
0.14
2.32
2.13
3.51
0.04
0.91
1.01
7.02
0,39
1.76
2.(X)
0.00
{).()()
().(K)
0.(X)
61.40
ND
17.48
-
Total
99.71
'Mostly diatoms
'Mostly PUinorhulla, rarely Phy.m
'Mostly terrestrial forms
June 1980
Williams, Williams: Borax Lake Chub
105
ing up food items. However, if Ijenthic food
items are scarce, or if other foods are abun-
dant, C. homxohius will readilv feed on ma-
terials drifting through water column or on
the surface. Thus, during the summer some
G. boraxohius readilv switched to ingestion of
terrestrial invertebrates. This resulted in the
lowest mean percent volume of inorganic
debris ingested for any season. The shoreline
of Borax Lake provides habitat for many ter-
restrial invertebrates that can enter Borax
Lake. Terrestrial invertebrates are scarce
during winter, reducing the likelihood of
their being a primary food source at that
time.
Gila boraxohius is often a highly exploitive
omnivore, feeding almost entirelv on one
food source. For example, examination of in-
testines of fish collected during May 1978
disclosed the following (percent volumes of
Tahle 2. C'ontents of 70 intestines of Cila boraxob'uts colh
the food item are given in parentheses): one
individual contained 32 gastropods (84 per-
cent volume), a second fish contained 14
adult dipterans (98 percent volume), a third
contained 775 copepods (79 percent volume),
a fourth contained 340 first instar chironomid
larvae (69 percent volume), and a fifth con-
tained 485 insect eggs (64 percent volume)!
Although the preceding is somewhat unusual,
many fish were found with one food item
dominating their intestinal contents.
Foods Consumed by
Different Size Classes of Fish
To study the effect of fish size and age on
foods consumed, we compared intestinal con-
tents of juvenile and adult Gila boraxohius.
Overall food habits of juveniles and adults
were similar. Both consumed large amounts
:ted di
tile summer of 1978. ND = no data.
Item ingested
Percent
frequency
of occurrence
Mean number
per intestine
Mean percent
volinne
RI
■\lgae'
Gastropods-
Gastropod eggs
Haplotaxid oligochaetes
Harpacticoid copepods
Ostracods
Cladocerans
Plant seeds
Higher plants
Fish scales
Araneae
Insect eggs
Unidentified insects'
Terrestrial insects
Collembola
Thvsanoptera adults
Hemiptera adults
Coleoptera adults
H\ nienoptera adults
Diptera adults
Asiatic: insects
C!hironoinid larvae
(Ihironomid pupae
Odonata nvmphs
Elmid larvae
Coleoptera adults
Inorganic debris
31.15
ND
6.42
7.42
32.79
1.07
9.04
8.26
1.64
0.66
1.56
0.63
1.64
0.08
0..30
0.38
47.54
33.30
8.45
11.06
32.79
2.87
3.03
7.08
8.20
0.38
0.31
1.68
13.11
0.25
0.49
2.69
0.00
0.00
0.00
0.(X)
().(X)
0.00
0.00
0.(X)
14.75
0.25
3.21
3..55
3.28
0.69
0.15
0.68
37.70
ND
4.50
8.34
6.56
0.16
0.23
1.34
9.84
0.13
0.45
2.03
0.00
0.00
0.00
0.00
26.23
0.70
5.94
6.36
18.03
0.21
2.27
4.01
39.34
1.62
14.31
10.60
.57.38
4.10
13.79
14.06
9.84
0.11
1.12
2.17
1.64
0.03
0.49
0.42
11.48
0.23
0.85
2.44
18.03
0.31
6.29
4.80
.54.10
ND
15..30
-
Total
98.50
'Mostly diatoms
'Mostly Planorhulla. rarely Phy.sa
'Mostly terrestrial forms
106
Great Basin Naturalist
Vol. 40, No. 2
of terrestrial insects in spring, summer, and
autumn, and algae (mostly diatoms) and mi-
crocm-staceans in winter (Fig. 1). Aquatic in-
sects, primarily chironomid larvae, were im-
portant to juveniles and adults throughout
the year. Despite the overall similarity of
food habits between juveniles and adults,
some differences were noted. Elmid (Coleop-
tera) larvae were consumed almost exclusi-
vely by adults. Elmid larvae were found in
intestines of 15 adults but only one juvenile.
More terrestrial insects were consumed by
juveniles than adults except during winter,
when terrestrial insects were relatively unim-
portant to both groups. The relatively large
size of many terrestrial insects, such as the
commonly consumed muscoid fly adults, did
not deter their ingestion by juvenile Gila
horaxohius. Adults consumed more gastro-
pods than did juveniles during all seasons. In-
testines of adults averaged 8.5 percent mean
volume of gastropods during the year, where-
as intestines of juveniles averaged 2.3 per-
cent. Adults also consumed more diatoms
than did juveniles. Increased relative con-
sumption of diatoms by adults was primarily
evident in summer and autumn, when adults
consumed 9.3 percent and 6.9 percent mean
volume of diatoms, respectively; however,
juveniles consumed 0.1 percent and 1.3 per-
cent mean volume, respectively. The small
volume of gastropods and diatoms ingested
by juveniles was compensated for by inge-
stion of large numbers of copepods. Intestines
of juveniles averaged 13.4 percent mean vol-
ume of copepods during the year, whereas
adults averaged 4.2 percent mean volume of
copepods.
Table 3. Contents of 57 intestines of C.iUi horaxohius collected during the autumn of 1978. ND = no data.
Item ingested
Percent
frequency
of occurrence
Mean number
per intestine
Mean percent
volume
RI
Algae'
Gastropods-
Gastropod eggs
Haplotaxid oligochaetes
Harpacticoid copepods
Ostracods
Cladocerans
Plant seeds
Higher plants
Fish scales
Araneae
Insect eggs
Unidentified insects'
Terrestrial insects
Collembola
Thysanoptera adults
Hemiptcra adults
Coleoptera adults
Hymenoptera adults
Diptera adults
AgUATIC INSECTS
Chironomid larvae
Chironomid pupae
Odonata nymphs
Elmid larvae
Coleoptera adults
I.NORGANIC DEBRIS
50.00
ND
5.03
12.63
10.87
0.50
4.04
3.42
0.00
0.00
0.00
0.00
0.00
0.00
0.00
0.00
34.78
25.46
4.22
8.95
10.87
0.50
0.15
2.53
0.00
0.00
0.00
0.00
39.13
1,57
0..59
9.12
2.17
ND
0.07
0.51
0.00
0.00
0.00
0.00
0.00
0.00
0.00
0.00
2.17
0.04
0.04
0.51
67.39
ND
19.54
19.95
0.00
0.00
0.00
0.00
21.74
0.46
1.69
5.38
13.04
0.22
2.55
3.58
4.35
0.04
0.22
1.05
8.70
0.11
l.,54
2.35
19.56
0.22
3..52
5.30
71.74
5.43
12.00
19.22
13.04
0.65
3.15
3.72
6.52
0.07
1.29
1.79
0.00
0.(K)
0.(X)
0.00
().(X)
0.(X)
0.(K)
0.00
86.96
ND
40.34
-
Total
99.94
'Mostly diatoms
'Mostly Planorhulla, rarely Physa
'Mostly terrestrial forms
June 1980
Williams, Williams: Borax Lake Chub
10'
Diel Feeding Chronology
Feeding chronologv and daily ration were
determined bv the relative weight of material
ingested bv CUla honixohiiis collected during
a 24-hour period in June 1979 (Fig. 2). The
average weight of fish was 1.21 g. Gila horax-
ohius fed throughout the day with peak feed-
ing activity shortly after sunset. Minimal
feeding activities occurred after sunrise. An
increase in feeding activity after sunset has
been observed in Gila hicolor (Snyder 1917).
The average weight of ingested material in
intestines, as determined from 1800-1500
hours, was 2.32 percent of body weight. This
average weight of ingested material (S) was
used to determine the daily ration (Rj) as fol-
lows:
Rt = 24Sa = 24(2.32)(.2) = 11.14
By the above method we calculated that G.
horaxohius ingested 11.14 percent of their
body weight daily. This estimate is larger
than most reported by researchers for other
species. Brett (1979) summarized in-
vestigations made by various researchers who
calculated daily rations that were typically
2-5 percent of body weight. Several studies
have noted increased relative ration with in-
creased temperature (e.g., Brett et al. 1969,
Kinne 1960, Stauffer 1973) and with smaller
fish size (e.g., Brett 1971, Brett and
Shelbourn 1975, Elliott 1975). Brett (1979)
reported that temperature and fish size were
of greatest importance in determining ration
size. The dwarf size of G. horaxohius and its
Table 4. Contents of 62 intestines oi Gila hoiaxobius collected during the winter of 1978-79. ND = no data.
Item ingested
Percent
frequency
of occurrence
Mean nmuber
per intestine
Mean percent
volume
RI
Algae'
Gastropods-
Gastropod eggs
Haplotaxid oligochaetes
Harpacticoid copepods
Ostracods
Cladocerans
Plant seeds
Higher plants
Fish scales
Araneae
Insect eggs
Unidentified insects'
Terrestrial i\sec:ts
C'ollembola
Thvsanoptera adults
Ileniiptera adults
Goleoptera adults
Hynienoptera adults
Diptera adults
AyiATic: insects
('hironomid larvae
(;hir()nomid pupae
Odouata nymphs
Elmid larvae
Goleoptera adults
I.N()R{;ank: debris
94.44
ND
18.81
22.05
18.52
0.54
3.88
4.36
0.00
0.00
0.00
0.(K)
5.56
0.91
2.21
1,51
75.93
16.44
9.96
16.72
77.78
21.65
20.58
19.15
55.56
6.52
4.07
11.61
3.70
0.04
0.02
0.72
5.56
ND
0.35
1.15
1.85
0.04
0.03
0.37
0.00
0.00
().(X)
(),(K)
0.00
0.00
(),()()
0.00
9.26
ND
1.49
2.09
1.85
0.04
0.33
0.42
0.00
0.00
0.00
0.(X)
0.00
0.00
0.(K)
0.00
0.00
0.00
0.(K)
0.(K)
0.00
0.00
().(K)
0.00
1.85
0.02
0.15
0.39
70.37
5.15
10.94
15.83
1.85
0.02
0.22
0.40
5.56
0.07
0.77
1.23
9.26
0.11
0.84
1.97
0.00
0.00
().()()
().(M)
90.74
ND
25.37
-
Total
100.02
'Mostly diatoms
■Mostly PkmorhuUa. rarely Physa
Mostly terrestrial forms
108
Great Basin Naturalist
Vol. 40, No. 2
gastropod
gastroDod
copepod
algae
est racod
copepod
algae
debris
ostracod
other
debris
other
terrestrial insect
JUVEN I LES
ADULTS
SPRING
gastropod 2.5
0.1 algae
copepod
ostracod
other
gastropod
ebris
aquatic
insect
other
teirest rial i nsec t
JUVENI LES
ADULTS
SUMMER
Fig. 1. Comparison of food habits between 128 juvenile and 1.32 adult Cila Inmixohitis. Mean percent volume of
food items are given in circle.
June 1980
Williams, Williams: Borax Lake Chub
109
copepod
terrestrial insect
debris
copepod gastropod
istracod 0.2
debris
0.9 other
JUVENILES
ADULTS
AUTUMN
0.9 gastropod
1
copepo
alqae
est racod
gast ropod
copepo
ost racod
debris
ther
aquatic insect i
1.0 terrestrial insect
debris
f other
2-^ terrestrial insect
JUVENI LES
ADULTS
Wl NTER
Fig. 1 continued.
110
Great Basin Naturalist
Vol. 40, No. 2
habitation in thermal spring waters contrib-
uted to the large daily ration found in this
species. Many researchers determined daily
ration using prepared foods that were per-
haps more nutritious than material ingested
by Gila boraxobiiis, which included such un-
digestible items as insect exoskeletons, gastro-
pod shells, and inorganic debris. The pres-
ence of large amounts of undigestible
material increases S because the fish must in-
gest a larger volume of material to get
enough calories. This results in a larger daily
ration than would be calculated if fish con-
sumed only digestible foods.
Food Habits of Gila ahordensis
Ten food items were foimd in intestines of
Gila alvordensis collected from Thousand
Creek, Nevada, in June 1978 (Table 5). Of
the ten food items, 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 of intestines. Microcrustaceans com-
prised almost 45 percent mean volume of in-
testines. Diatoms accounted for 5 percent
mean volume of intestines. No terrestrial in-
sects were ingested by G. alvordensis from
Thousand Creek.
Eleven food items were found in intestines
of Gila alvordensis collected from Serrano
Pond, Oregon, in August 1977 (Table 6). Of
the eleven food items, chironomid larvae, di-
atoms, cladocerans, and ostracods were of
greatest importance. Chironomid larvae oc-
curred in over three-quarters of the intestines
examined and accounted for approximately
50 percent mean volume of all intestines. Di-
atoms occurred in one-half of the intestines
and accounted for almost 23 percent mean
volume of all intestines. Microcrustaceans
comprised approximately 17 percent mean
volume of intestines. No intestines contained
terrestrial insects. Gila alvordensis from Ser-
rano Pond were highly opportimistic feeders.
Eighty-nine percent of the fish from Serrano
Pond with food in their intestines contained
one item that accounted for more than 50
percent of their intestinal volume. Thirty-
nine percent of fish contained one food item
that comprised 90 percent or more of in-
testinal volume. This exploitive feeding was
focused on chironomid larvae, cladocerans,
or algae. One intestine was exclusively filled
with 2570 cladocerans. Such exploitive feed-
ing was not noted in Gila alvordensis from
Thousand Creek.
A comparison of foods of Gila alvordensis
collected during June and August with foods
of Gila boraxobiiis collected during the sum-
mer shows several differences. Terrestrial in-
sects were important foods for Gila borax-
obiiis during the summer but were absent
from intestines of Gila alvordensis. Gila
boraxobiiis also consumed larger quantities of
other terrestrial foods, such as spiders and in-
sect eggs, than did G. alvordensis. Intestines
of G. alvordensis from Thousand Creek, and
to a lesser extent those from Serrano Pond,
contained much larger amounts of micro-
ciTistacea than did intestines of G. boraxobiiis
during the summer. Diatoms were a major
food item of fish in Serrano Pond during the
simimer, but not of fish in Thousand Creek or
of Gc boraxobiiis. Gila boraxobiiis consumed
a larger number of food items than either
population of G. alvordensis. This is due to
the greater opportimism, including the use of
terrestrial foods, exhibited by G. boraxobiiis.
A larger sample size may also contribute to
the greater diversity of foods utilized by G.
boraxobiiis.
Discussion
Gila boraxobiiis and G. alvordensis have
tentatively been included with G. bicolor in
the subgenus Siphateles (Hubbs and Miller
1972, Williams and Bond, in press). Although
no life history information has previously
been published for G. boraxobiiis or G. alvor-
densis, several researchers have examined
food habits of G. bicolor and concluded that
they are primarily opportunistic omnivores
(Bird 1975, Cooper 1978, La Rivers 1962).
However, differences in food habits between
the coarse gill raker form, G. b. obesa, and
the form with numerous, fine gill rakers, G.
b. pectinifer, have been noted. In describing
habits of the form with coarse gill rakers
from Lake Tahoe, Miller (1951) foimd them
to be primarily benthic feeders, with a diet
June 1980
Williams, Williams: Borax Lake Chub
111
composed of 89 percent bottom organisms.
Snyder (1917) noted that the form with
coarse gill rakers collected from the littoral
zone of Lake Tahoe fed on algae, other plant
material, and insects. Gila b. pectinifer from
Lake Tahoe, with its numerous gill rakers,
fed almost exclusively on midwater micro-
crustacea (Miller 1951). La Rivers (1962) also
reported that G. b. pectinifer contained many
midwater foods in their intestines, primarilv
consuming diatoms and microcrustaceans.
Cooper (1978) reported that the form with
numerous gill rakers (although he referred it
to G. h. obesa) in Walker Lake, Nevada, fed
mostly on zooplankton and filamentous al-
gae. A population complex of G. bicolor in
Eagle Lake, California, that included forms
with both coarse and fine gill rakers fed on a
variety of foods, including zooplankton, plant
material, insect larvae, and surface insects
(Kimsey 1954). There appears to be a definite
correlation between gill raker morphology
Table 5. Contents of 21 intestines of Gila uhordctisis collected 13 June 1978 from Thousand Creek, Nevada.
ND = no data.
Item ingested
Percent
frequency
of occurrence
Mean number
per intestine
Mean percent
volume
RI
Diatoms
Gastropods
Harpacticoid copepods
Ostracods
Cladocerans
Plant seeds
Araneae
AgiL-VTIC INSECTS
Chironomid larvae
Chironomid pupae
Coleoptera adults
I.NORCANIC: DEBRIS
36.36
ND
5.00
7.91
9.09
0.09
0.36
1.81
81.82
6.82
6.82
16.94
72.73
15.82
13.73
16.53
90.91
27.73
24.23
22.01
27.27
0.27
0.45
5.30
9.09
0.09
0.55
1.84
10().(K)
16.73
25.78
24.04
9.09
0.09
0.09
1.75
9.09
0.09
0.68
1.87
lOO.(K)
ND
20.95
-
Total
98.64
Table 6. Contents of 20 intestines of Gila akordensis collected 6 August 1977 from Serrano Pond, Oregon.
ND = no data.
Item ingested
Percent
frequency
of occurrence
Mean number
per intestine
Mean percent
volume
RI
Diatoms
Harpacticoid copepods
Ostracods
Cladocerans
.Araneae
Insect eggs
Unidentified insects
.\quatic insects
Chironomid larvae
Chironomid pupae
Odonata nymphs
Ephemeroptera larvae
Inorganic debris
50.00
ND
22.94
18.16
.33.33
1.67
0.36
8.39
38.89
5.94
1.70
10.11
55.56
253.28
15.34
17.65
5.56
0.06
0.22
1.44
5..56
0.06
0.05
1.40
5.56
ND
0.05
1.40
77.78
20..33
50.41
31.92
22.22
0.33
2.96
6.27
5.56
0.06
0.67
1.55
5.56
0.06
1.33
1.72
55..56
ND
3.96
-
Total
99.99
112
Great Basin Naturalist
Vol. 40, No. 2
and food habits, tho.se with coarse gill rakers
ingesting more benthic food organisms and
those with fine gill rakers ingesting more
zooplankton. Gila boraxobius and G. alvor-
densis possess approximately 16 and 20 short
gill rakers, respectively, agreeing closely with
the gill raker morphology of G. bicolor obesa
form. Although G. boraxobius typically feeds
on benthic organisms, large amounts of dia-
toms, microcnistaceans, and terrestrial insects
are ingested sea.sonally.
The ingestion of terrestrial insects by Gila
is not common. However, several researchers
have found that terrestrial insects comprised
a small part of the diet of Gila (Cross 1978,
Kimsey 1954, Moyle 1976, Sigler and Miller
1963). Terrestrial insects were the primary
foods of G. robusta and G. elegaas longer
than 200 mm SL collected from the Green
River (Vanicek and Kramer 1969). Smaller G.
robusta and G. elegans contained pre-
dominantly aquatic insect larvae. Juvenile
and adult G. boraxobius consumed large
quantities of terrestrial insects. Several re-
searchers (Kimsey 1954, Miller 1951) have
noted that as Gila grow, they switch to larger
food items; however, at least one study (Gra-
ham 1961) found foods of different sized
groups of CAla to be nearly identical. We
found foods of juvenile and adult G. borax-
obius to be very similar, except that adults
exhibited a greater consumption of gastro-
pods and diatoms, and juveniles consumed
more copepods and terrestrial insects. The
hard shells and, to a lesser extent, the rela-
tively large size of gastropods probably con-
tributed to juveniles avoiding them as a food
source. Larger Gila ingest more algae than
do smaller fish in studies by Moyle (1976) and
Vanicek and Kramer (1969). Age II Gila coe-
rulea feed predominantly on filamentous al-
gae, whereas algae were entirely or prac-
tically absent from age I fish (Moyle 1976).
Large adult Gila from the Green River con-
sumed more algae than did smaller fish (Van-
icek and Kramer 1969). Juvenile G. borax-
Tl ME OF DAY
Fig. 2. Feeding throiiologv^ of Cila boraxobius on 5-6 June 1979. N = 72.
June 1980
Williams, Williams: Borax Lake Chub
113
obitis consumed more terrestrial insects than
did the adults, except in winter, when small
amoiuits of terrestrial insects were ingested
by both groups. The reason for juveniles con-
suming large volumes of terrestrial insects is
unknown; apparently the relatively large size
of this food item is not a deterrent.
Both juvenile and adult G. horaxoJyiiis in-
creased consumption of diatoms and micro-
crustaceans in winter. This probably in-
dicates a scarcity of food items during the
winter, as is reflected by finding 24 percent
fewer food items in intestines during winter
than in summer. A factor contributing to the
winter scarcity of food items is a decrease in
the availability of terrestrial foods, causing
concentrated feeding on remaining food
items. For example, the Devil's Hole pupfish,
Cyprinodon diabolis, which inliabits a small
thermal spring, dramatically increases in-
gestion of diatoms during winter due to a
scarcity of preferred foods (Minckley and
Deacon 1975). Although the amount of nutri-
tion derived from consuming diatoms is un-
known, we suspect that the large con-
sumption of diatoms at certain times of the
year by G. bomxobius would indicate that
some nutritive value is gained. Arnold (1971)
found that species of Ci/prinodon derived oil
droplets from ingested diatoms, thus extract-
ing nutritive value. A similar mechanism
could operate in Gila bomxobius.
Examination of summer foods of both G.
ah'ordensis populations showed differences
from the summer foods of G. boraxobius.
During summer Gilo boraxobius relied heavi-
ly on terrestrial food items, whereas popu-
lations of G. alvordensis consumed prac-
tically no terrestrial foods. Forty-three
percent of food items consumed by G. bora-
xobius during the summer were of terrestrial
origin. Surveys of potential food items in Bo-
rax Lake conducted at various times of the
year found that all potential food items were
utilized by G. boraxobius except some adult
hemipteran and coleopteran insects that were
probably too large to be ingested. Also, many
hemipterans possess scent glands that render
them impalatable to predators.
Acknowledgments
The authors are indebted to Carl E. Bond
for his guidance during the course of this
study and for his review of the manuscript.
Stanley V. Gregory provided identification of
diatoms and information on their habitats,
the Oregon Department of Fish and Wildlife
provided collecting permits, and James J.
Long, Kevin M. Howe, and Glen DeMott as-
sisted with field collections. Kevin M. Howe
reviewed the manuscript and provided field
notes of the Serrano Pond area. This informa-
tion is part of the senior author's doctoral dis-
sertation at Oregon State University and is
prepublished by permission of the Graduate
School and the Department of Fisheries and
Wildlife.
Literature Cited
Arnold, J. T. 1971. Behavioral ecology of two pupfishes
(Cyprinodontidae, Genus Cijprinodon) from
northern Mexico. Unpublished dissertation. .Ari-
zona State Univ. 1.33 pp.
B.\]Kov, \. D. 1935. How to estimate the daily food con-
sumption of fish under natural conditions. Trans.
.\m. Fish. Soc. 65:288-298.
Berg, J. 1979. Discussion of methods of investigating the
food of fishes, with reference to a preliminary
study of the prey of Gohiusculus flavescens (Go-
hiidae). Marine Biology. 50:263-273.
Bird, F. H. 1975. Biology of the Blue and Tui chubs in
East and Paulina Lakes, Oregon. Unpublished
thesis. Oregon State Univ. 165 pp.
Brett, J. R. 1971. Satiation time, appetite, and max-
imum food intake of sockeve salmon, Oncor-
hijnchus nerkd. J. Fish. Res. Bd. Can. 28:409-415.
1979. Environmental factors and growth. Pages
599-675 in W. S. Hoar, D. J. Randall and J. R.
Brett, eds. Fish Physiology vol. VIII. .\cademic
Press, New York.
Brett, J. R., and J. E. Shelbourn. 1975. Growth rate of
voung sockeve salmon, Oncorhynchus uerka. in
relation to fi.sh size and ration level. J. Fish. Res.
Bd. Can. 32:2103-2110.
Brett, J. R., J. E. Shelbourn, and C. T. Shoop. 1969.
Growth rate and body composition of fingerling
sockeve salmon, Onrorhijnchus ncrka. in relation
to temperature and ration size. J. Fish. Res. Bd.
Can. 26:2.363-2394.
Cooper, J. J. 1978. Contributions to the life hi.story of
the Lahontan Tui chub, Gila bicolor ohesa (Gi-
rard), in Walker Lake, Nevada. Unpublished
thesis. Univ. Nevada, Reno. 89 pp.
Cross, J. N. 1978. Status and ecology of the Virgin River
roundtail chub, Gila whiista sertiintida (Os-
teichthves: Cvprinidae). Southwestern Nat.
23:519-528.
Dar.nell, R. M., and R. R. Meierotio. 1962. Determi-
nation of feeding chronology in fishes. Trans. .\in.
Fish. Soc. 91:313-320.
Deacon, J. E., G. Kobetich, J. D. Willlams, S.
Contreras, and other members of the
Endangered Species Committee of the
114
Great Basin Naturalist
Vol. 40, No. 2
American Fisheries Society. 1979. Fishes of
North America endangered, threatened, or of
special concern: 1979. Fisheries. 4:29-44.
EcGERS, D. M. 1977. Factors in interpreting data obtain-
ed bv die] sampling of fish stomachs. J. Fish. Res.
Bd. Can. 34:290-294.
Elliott, J. M. 1975. Number of meals in a day, max-
imum weight of food consumed in a day and
maximum rate of feeding for brown trout, Salmo
trtitta L. Freshwater Biol. 5:287-303.
George, E. L., and W. F. Hadley. 1979. Food and habi-
tat partitioning between rock bass (Ambloplites
rupestris) and smallmouth bass (Micropteriis do-
lomieui) yoimg of the year. Trans. Am. Fish. Soc.
180:253-261.
Graham, R. J. 1961. Biology of the Utah chub in Heb-
gen Lake, Montana. Trans. Am. Fish Soc.
90:269-276.
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
Soc. Nat. Hi.st. 17:101-106.
HuREAU, J. C. 1969. Biologie comparee de quelques
poissons antarctiques (Nothotheniidae). Bull. In.st.
Oceanogr. Monaco. 68:1-44.
KiMSEY, J. B. 1954. The life hi.story of the tui chub, Siph-
ateles bicolor (Girard), from Eagle Lake, Califor-
nia. Calif. Fish Game. 40:.395-410.
KiNNE, O. 1960. Growth, food intake, and food con-
version in a euryplastic fish exposed to different
temperatures and salinities. Physiol. Zool.
33:288-317.
La Rivers, 1. 1962. Fishes and fisheries of Nevada. Ne-
vada State Fish Game Comm. 782 pp.
Mariner, R. H., J. B. Rapp, L. M. Willey, and T. S.
Presser. 1974. The chemical composition and es-
timated minimum thermal reservoir temper-
atures of selected hot springs in Oregon. U.S.
Geol. Surv. Open-File Report. Menlo Park, Calif.
Miller, R. G. 1951. The natural history of Lake Tahoe
fishes. Unpublished dissertation. Stanford Univ.
160 pp.
MiNCKLEY, C. O., AND J. E. Deacon. 1975. Foods of the
Devil's Hole pupfish, Cijprinodon diaboUs (Cy-
prinodontidae). Southwestern Nat. 20:105-111.
MoYLE, P. B. 1976. Inland fishes of California. Univ.
California Press, Berkeley. 405 pp.
Pennak, R. W. 1978. Freshwater invertebrates of the
United States, 2d ed. Wiley-lnterscience, New
York. 803 pp.
SiGLER, W. F., AND R. R. MiLLER. 1963. Fishes of Utah.
Utah State Dept. Fish Game, Salt Lake City. 203
pp.
Snyder, J. O. 1917. The fishes of the Lahontan system of
Nevada and northeastern California. U.S. Bureau
Fisheries Bull. .35 (for 1915-16):3.3-86.
Stauffer, G. D. 1973. A growth model for salmonids
reared in hatchery environments. Unpublished
dissertation. Univ. Washington.
Vanicek, C. D., and R. H. Kramer. 1969. Life history of
the Colorado squawfish, Ptijclwcheilus luciiis,
and the Colorado chub, Gila robusta, in the
Green River in Dinosaur National Monument,
1964-1966. Trans. Am. Fish. Soc. 98:19.3-208.
Williams, J. E., and C. E. Bond. Gila boraxobius, a
new species of cyprinid fish from southeastern
Oregon with a comparison to G. alvordensis
Hubbs and Miller. Proc. Biol. Soc. Wash. Vol. 93.
Windell, J. T., and S. H. Bowen. 1978. Methods for
study of fish diets based on analysis of stomach
contents. Pages 219-226 in T. Bagenal, ed..
Methods for assessment of fish production in
fresh waters, .3d ed. Blackwell Scientific Pub-
lications, Oxford.
FIRST RECORD OF THE PALLID BAT (ANTROZOUS PALLIDUS) FROM MONTANA
Jeff Shryer' and Dennis L. Flath'
.\bstract.— a pallid bat {Antrozous pallidiis) was taken 20 km SE Warren, Carbon County, Montana. This repre-
sents a 410 km range extension and a new record for Montana.
The pallid bat {Antrozous pallidiis) has not
been previously reported from Montana. On
the basis of specimens from Grangeville and
Pocatello, Idaho, Hoffman and Pattie (1968)
suggested that the species may occur in
southwestern Montana. Fichter (1964) report-
ed A. pallidus from Boise, Idaho.
On 25 August 1978 we mist-netted an
adult female A. pallidus (Montana Fish,
Wildlife, and Parks Collection NG 748) at an
isolated spring approximately 20 km SE
Warren, Carbon County, Montana. This rec-
ord extends the known range of A. pallidas
approximately 410 km northeast of Pocatello,
Idaho, and approximately 460 km north of
Dinosaur Quarry, Utah, where Knitzsch and
Heppenstall (1955) obtained a specimen.
The new locality is in an alluvial plain
south of the Pryor Mountains, at 1370 m ele-
vation, and lies within the 25-30 cm precipi-
tation zone (U.S. Soil Conservation Service
1977). The area is characterized by scattered
outcrops of sandstone and .shale, with a vege-
tative community dominated by western
wheatgrass {Agropyron smitliii), sagebrush
{Artemisia spp.), and saltbush (Atriplcx spp.).
External measurements of the specimen
are as follows: total length, 110 mm; length
of tail, 37 mm; length of hind foot, 13 mm;
length of ear, 29 mm; and length of forearm,
59 mm. The dentition displaved substantial
wear, indicating the bat was old. This speci-
men is similar to specimens in the University
of Montana Zoological Museum from New
Mexico (MSUZ 10419) and Arizona (MSUZ
12996) in that it is of a similar color— a very
pale fawn. Allen (1864) described A. p. pal-
lidas as having two varieties of color, fawn
and yellowish brown. Bailey (1936) described
A. p. cantwelli as darker than pallidus, with
dark brown ears and membranes. The mem-
branes of our specimen are medium brown.
Other species captured concurrently in-
clude: Myotis lucifugus and M. leihii.
We thank John Ciralli for field a.ssistance
and P. L. Wright for assistance in the identi-
fication of the specimen. This paper is a con-
tribution of the U.S. Bureau of Land Manage-
ment, Montana Nongame Program, and the
Montana Department of Fish, Wildlife, and
Parks, Nongame Program, State Project
5853.
Literature Cited
.\LLEN, H. 1864. Monograph of the bats of North Ameri-
ca. Smithsonian Misc. Coll. 165:68-69.
.\.\o.NYMOUS. 1977. Average annual precipitation for
Montana based on 1941-1970 base. USDA, Soil
Cons. Serv., Portland, Ore. 13pp.
B.\iLEY, V. 1936. The mammals and life zones of Oregon.
North American Fauna 55:390-.393.
Fichter, E. 1964. The pallid bat in Idaho. Tebiwa
7:23-27.
Hoffman, R. S., a.nd D. L. Pattie. 1968. .-^ guide to
Montana mammals: Identification, habitat, distri-
bution and abundance. Univ. Montana Print.
Serv. 1.33 pp.
Krutzsch, p. H., and C. A. Heppenstall. 1955. Addi-
tional distributional records of bats in Utah. J.
Manmial. .36:126-127.
'U.S. Bureau of Land Management, Lewistown District Office. Lewistown, Montana 594.57.
'Montana Department of Fish, Wildlife, and Parks, Montana State University, Box 5, Bozeman, Montana 59717
115
A CHIRACANTHIUM SPIDER BITE
Dorald M. Allied'
.\bstr.\ct.— a bite bv Cliimcanthium mildci L. Koch is described.
In May 1979, Mr. Lee Carson of Provo,
Utah, brought a spider to me which had bit-
ten him on the right index finger. It was sub-
sequently identified by Dr. Wilhs J. Gertsch
as a female Chiracanthium niildei L. Koch.
Mr. Carson was in the process of placing a
pair of Ribbers over his shoes to work in his
garden. The mbbers were kept in his closed
garage, and had been worn a few days pre-
viously. As he inserted his fingers into one of
the rubbers, he felt a sudden pain at the tip
of his finger. Examination disclosed the spi-
der with a web and cocoon in a rubber.
Within 3 to 4 seconds after the bite, the
finger began to ache severely. The pain soon
extended to his upper arm where it remained
for 2 to 3 hours, although it was most con-
centrated in the finger. He described the sen-
sation as a painful "tingling" similar to what
one experiences when his elbow "crazy-
bone" is bumped. No nausea, headache.
swelling, or numbness was experienced. No
evidence of inflammation or necrosis oc-
curred at the site of the bite, and the cheli-
ceral punctures healed rapidly. Pain in the
finger and arm disappeared after about 4
hours.
Dr. Gertsch kindly supplied information on
the spider and its ecology. The species appar-
ently was introduced into the United States,
became a typical house spider in the New
York and Boston area, and subsequently
spread to other areas of the U.S. As early as
1930 Dr. Gertsch collected it at Salina, Utah.
It is known from southern California, and
Kaston (1972, "How to Know the Spiders, p.
221) indicated its distribution as New Eng-
land, New York, New Jersey, Alabama, Mis-
souri, and Utah. Gertsch further stated that
the genus is reputed to have a venom of in-
termediate potence, and that of the Eu-
ropean species is said to be next to their Lat-
rodectus in severity.
'Department of Zoology, Brigham Young University, Provo, Utah 84602.
116
IDENTITY OF NARROW-LEAVED
CHRYSOTHAMNUS VISCIDIFLOR US (ASTERACEAE)'
Loraii ('. Aiidersoir
.\bstr.\ct.— Two ijroups of glabrous, narrow-leaved Chiysotliaiiinits visciclifloius (Asteraceae) are perceived, and
appropriate taxononiie combinations are made, i.e., C. v. ssp. vmidiflorus var. stcnofjlttjlhis and C. v. ssp. axillaris.
The two are fairl\' distinct ideograph icallv, and thev can be separated l)v floral niorpliologv. .\ key to all species of
section Chrtjsothanntus (to which C. liscidiflorus belongs) is given.
Classification of intra.specific variants of
Chrysothamnus liscidiflorus has been prob-
lematic, in part, becau.se floral features seem-
ingly lacked sufficient distinctions. Hence,
vegetative aspects such as stature, vesture,
and leaf dimensions have been used. Experi-
mental -Studies (Anderson 1964) have demon-
strated that in many in.stances plant height,
leaf twisting, and leaf width are expressions
of differing edaphic conditions, droughtiness,
or other environmental parameters and
thereby complicate taxonomic resolution.
Study of interpopulational variation and
the distribution of the narrow-leaved, yellow
rabbitbmsh (C. viscidiflorus ssp. stenophyUus)
reveals that there are two different taxa rep-
resented. One is found sporadically through
the northern latitudes of the western United
States and into southern California (open cir-
cles in Fig. 1). The other taxon (stars. Fig. 1)
is found further .south and is more generally
distributed, i.e., has a more pronounced
"range integrity." In south-central Nevada,
Beatley (1976) reports it is the common
Chrysothamnus of basin floors and foothills,
especially in volcanic areas and on disturbed
sites, usually below 5500 ft. Examination of
populations at possible type locality sites
(type specimens labeled either West Hum-
boldt Mountains or Huntington Valley) for
ssp. StenophyUus shows that the narrow-
leaved plants represent extremes of the
broader-leaved ssp. viscidiflorus.
I have concluded from field observations,
garden culture, and herbarium studies that
the northern elements, which include the
tvpe collection of C. i". ssp. stenophyUus, are
actually environmentally induced variants of
ssp. liscidiflorus. Although quadrinomials are
cumbersome, the following nomenclatural
combination more appropriately identifies
the relationship of these plants:
Chrysothamnus viscidiflorus (Hook.) Nutt.
ssp. viscidiflorus var. stenophyUus (Gray) L.
C. Anderson, comb. nov.
Basionym: Bigelovia douglasii Gray var.
stenophylla Gray. Proc. Am. Acad. Sci.
8:646, 1873. W. ' Humboldt Mtns, Nevada,
Watson 566 (GH, holotype; NY, US, iso-
types).
Synonymy: Chrysothamnus pumilus Nutt.
var. varus A. Nels. Bot. Gaz. 28: 375, 1899.
Centennial Valley, Wyoming, Nelson 1847
(RM, holotype; GH, NY, i.sotypes).
Tlie .southern elements that had previously
been referred to ssp. stenophyUus warrant
subspecific recognition. Their narrow-leaved
characteristic is independent of environmen-
tal conditions. These plants are diploids;
broad-leaved forms of ssp. viscidiflorus that
grow in the same region are tetraploids or
hexaploids (Anderson, 1966, 1971). The only
available name for these narrow-leaved
plants is C. axillaris. In 1964, I noted that C.
axillaris was not specifically distinct from C.
viscidiflorus, and the name was synonomized
luider .ssp. stenophyUus. Munz (1968), in re-
'This shidy was supported by .National Science Foundation Grant DEB 76-10768. Ken VVonilile helped with ijraphics.
•Department of Biological Science, Florida Stale University, Tallahassee, Florida 32306.
117
118
Great Basin Naturalist
Vol. 40, No. 2
ferring to my studies, stated I had made C.
uxillaris a suljspecies of C. viscidiflonis. The
statement was inaccurate at that time— but
prophetic. The combination is now made:
Chrysothamnus viscidiflonis (Hook.) Nutt.
ssp. axillaris (Keck) L. C. Anderson, comb. &
Stat. nov.
Basionvm: Chri/sothamntis axiUaris Keck.
Ahso 4:103, 1958.' Deep Springs Valley, Cali-
fornia, Ferris 6924 (NY, holotype; DS, LL,
POM, isotypes).
Keck (1958), in describing C. axiUaris, re-
lated it to C. albidus and more distantly to C.
greenei. Actually, ssp. axillaris is not close to
C. albidus in relationship, but it is to C.
greenei. In fact, the type collection of ssp. ax-
illaris with very acute phyllaries suggests
some intergradation with C. greenei. Keck
stated that the latter was known only from
eastern Nevada and eastward, but it does ex-
tend through southern Nevada into Inyo
County, California. At its western limit in
California and also in northeastern Arizona,
C. greenei intergrades somewhat with C. vis-
cidiflorus. The feature of vertically aligned
phyllaries noted by Keck (1958) is not con-
sistent for ssp. axillaris (Anderson 1964).
Although ssp. axillaris and var. steno-
plnjllus are fairly distinctive habitally, they
could not be "keyed out" easily vinless refer-
ence was made to geographic distribution
(Fig. 1). Floral morphology was studied in
search of additional distinguishing features.
Methods are those used earlier (Anderson
1964). Detailed floral data and a list of speci-
mens examined are on file at FSU. Significant
comparisons are graphically illustrated in
Figure 2.
Involucral width in Chrysothamnus is gen-
erally strongly correlated with flower num-
ber because more flowers per head require a
broader receptacle. The pattern is evident in
general range for C. v.
sep. viscidiflorus
C. V. ssp. viscidiflorus
var. stenophyllus
C. V. ssp. axillaris
Fig. 1. Range of Clirysolhamnu.^ viscidiflonis ssp. viscidiflonis and ssp. axillahs. Range for ssp. viscidiflonis nearly
equals that of the species; ssp. tanceolatus extends into southern British Columbia and north-central New Mexico.
Distribution of ssp. (ixillaiis (stars) is fairly general through southern parts of Utah and Nevada and adjacent regions,
whereas that of .ssp. viscidiflonis var. stenophijllus (open circles) is sporadic, but mostly north of ssp. axillaris.
June 1980
Anderson: Chrysothamnus
119
C. viscidiflorus ssp. viscidiflorus, wherein
plants averaging 10.8 (with up to 12) flowers
per head have involucral widths over 50 per-
cent of involucral length (Fig. 2); these plants
represent an altitudinal record for the genus
of 12,800 ft in the White Mountains of Cali-
fornia. Previous descriptions of C. t/,s-
cidiflonis listed flower number as about 5
(Hall and Clements 1923). Nearly all plants
of ssp. viscidiflorus with high flower number
come from altitudes over 10,000 ft. Most
populations of the subspecies, including var.
stenophyUus, have 4-6(7) flowers per head
with proportionately narrower involucres.
Heads of ssp. axillaris depart from the bas-
ic correlation of flower number— involucral
width /length ratio. Although they average
fewer than 5 flowers per head, the
width /length ratio for the involucre is high
(Fig. 2). Tlius, ssp. axilluris and var. steno-
phijllus can be distinguished by floral fea-
tures as well as geographically.
Taxonomic interpretation of Chryso-
thamnus section Chrysothamnus (to which C.
viscidiflorus belongs) has been altered consid-
erably since Hall's monograph (Hall and Cle-
ments 1923). Chrysothamnus vaseyi and C.
molestus (C. viscidiflorus var. tnolestus) have
been transferred to section Fulchelli (Ander-
son 1970) and C. gramineus to Petradoria
(Anderson 1963). Additional species have
been recognized in the section. A key to sec-
tion Chrysothamnus as currently understood
is presented here.
-/At-
Fig. 2. Correlation of involucral shape and flower
number in heads of Cltrysothamntis viscidiflorus ssp. vis-
cidiflorus (closed circles, var. viscidiflorus; open circles,
var. stcnoplnjilus) and ssp. axillaris (stars). Vertical axis
represents the involucral width /length relationship ex-
pressed as percent; the horizontal axis is average flower
number per head. Note that ssp. axillaris departs from
the general correlation in ssp. viscidiflorus and other
members of the genus.
1. Flowers white; leaves terete, strongly punctate C. albidus (Jones) Greene
— Flowers yellow; leaves planate or involute, not punctate 2
2(1). Flowers 2-3(4); style branches included in erect corolla lobes; plants mostly
less than 1.5 dm tall C. hurnilis Greene
— Flowers (3)4-6; styles exerted beyond spreading corolla lobes; plants often over
2 dm tall 3
3(2). Style appendages long (40-70 percent of style branch); leaves never twisted or
involute; tall shRibs 4
— Style appendages short (30-45 percent of style branch); leaves frequently
twisted or involute 5
4(3). Leaves lanceolate; achenes densely pubescent C. linifolius Greene
— Leaves spatulate to oblanceolate; achenes sparsely pubescent
C. spathulatus L.C.Anders.
5(3). Phyllaries acuminate-cuspidate; leaves 1-2 mm wide C. greenei (Gray) Greene
— Phyllaries obtuse to acute; leaves 1-10 mm wide (C. viscidiflorus) 6
120
6(5).
7(6).
8(7).
9(7).
10(9).
Great Basin Naturalist
Vol. 40, No. 2
Leaves planate, glabrous; flowers 3.5-4(4.5) mm long
C. V. ssp. planifolius L.C.Anders.
Leaves twisted or pubescent or flowers longer 7
Upper stems, frequently leaves, hairy 8
Stems glabrous; leaves only ciliate 9
Stems and leaves pubenilent; leaves 1-2(4) mm wide
C. V. ssp. puberiilus (D.C.Eat.) H.&C.
Stems hispid near inflorescence; leaves over 2 mm wide, hirsute or glabrous
C. t;. ssp. lanceolatiis (Nutt.) H.&C.
Leaves ± 1 mm wide; flowers 3-4(5); involucres somewhat turbinate
C. r. ssp. axillaris (Keck) L.C.Anders.
Leaves 1-10 mm wide; if 1 mm, flowers 4 or more and involucres narrowly
cylindric 10
Leaves more than 1.5 mm wide; plants up to 1 m tall
C. V. ssp. viscidiflorus var. viscidiflorus
Leaves 1-1.5 mm wide; plants mostly less than 3 dm tall
C. V. ssp. viscidiflorus var. stenophyllus (Gray) L.C.Anders.
Names often applied in C. viscidiflorus,
but not representative as distinct subspecies,
include: (1) elegans, usually misapplied to
certain forms of ssp. puberulus with bracts
with enlarged green tips, but the type speci-
men does not have such bracts and is clearly
part of ssp. lanceolatus; (2) pumilus, low
form that is part of ssp. viscidiflorus; (3) torti-
folius, environmental variant with strongly
twisted leaves, part of ssp. viscidiflorus; and
(4) latifolius, wide-leaved plants from north-
ern Nevada that could be considered a varie-
ty of ssp. viscidiflorus; however, not all wide-
leaved plants of the subspecies would belong
to that variety.
A conceptual distinction between sub-
species and variety exists in Chnjsothamnus
for me. Subspecies is applied to groups of
populations with pronounced geographical
and fairly distinct morphological limits. The
variety can be applied in two ways. It may
represent a sporadic but rather distinctive
morphotype within a given subspecies, such
as ssp. viscidiflorus var. stenophijllus, and it
could possibly be applied to such variation in
species where subspecies are not recognized,
such as C. grcenci var. filifolius for the nar-
row-leaved variant. The second application
(my preferred usage) of variety would be for
elements of a subspecies that have some fair-
ly consistent morphological distinction and
also have relatively sharp geographic limits.
but are clearly subordinate to the subspecies.
An example would be ssp. viscidiflorus var.
latifolius— a the combination were made.
Publication of additional quadrinomials as
needed to clarify relationships in the genus
will be part of my upcoming monograph.
Literature Cited
Anderson, L. C. 1963. Studies on Pctradoria (Conipos-
itae): Anatoniv, cvtologv, taxonoiiiv. Trans. Kans.
Acad. Sci. 66:632-684."'
1964. Taxononiic notes on the CIinjsotlKininus
viscicliflonis complex (Astereae, Conipositae).
Madrono 17:222-227.
1966. Cytotaxonomic studies in Clinisothiimnus
(Astereae, Conipositae). Amer. J. Bot.
53:204-212.
1970. Floral anatomy of Chrysotli(iiuniis (As-
tereae, Conipositae). Sida 3:466-503.
1971. Additional chromosome counts in Chnjso-
tlunnntts (Asteraceae). Bull. Torrev Bot. Club
98:222-225.
Be.\tley, J. C. 1976. Vascular plants of the Nevada Test
Site and central-southern Nevada. Energy Res. &
Dev. ,\dmin., Nat'l Tech. Inf. Service,
Springfield, Va.
Hall, H. M., .\nu F. E. Clements. 1923. The phyloge-
netic method in taxonomy: The North American
species of Artemisia, Chnjsoihamnus, and Atri-
plcx. Carnegie Inst. Publ. 326:l-.355.
K.ec:k, D. D. 1958. Taxononiic notes on the California
flora. Aliso 4:101-114.
Munz, p. a. 1968. Supplement to A California Flora.
I'niv. Calif. Press, Berkeley.
RIBULOSE DIPHOSPHATE CARBOXYLASE ACTIVITIES IN COLD-RESISTANT
COMMON MALLOW, MALVA NEGLECTA WALLR. AND A COLD-SENSITIVE
TOMATO, LYCOPERSICON ESCULENTUM L., ACE 55 VAR.
W illiam H. AiidfiSL-ii' and Jack D. Brotlicrson'
Abstfl^ct.— Coiiinioii mallow (Malta nc^hcta W'allr. ) and tomato (Li/copcisicon csciiUnttim L. var. Ace 55) were
compared as to certain characteristics: C02 fixation properties, ribulose diphosphate carboxyl activities, (RiiDPCase)
photosynthesis, respiration, and compensation points. Significant differences in these factors were observed in all
cases except dark respiration. Mallow enzyme (RuDPCase) activities were higher per unit of enzyme than those of
tomato. The Mallow RuDPCase exhibited slightly higher activity at 5 to 25 C. Mallow leaves retained their capacity
for photosynthesis and respiration after long periods of exposure to subfreezing temperature. The cold adapted mal-
low had a higher C(% compensation point, suggesting a lower efficiency for CO2 fixation. The results suggest that
cold acclimation in common mallow affects photosynthesis but has little effect on respiration.
Several physiological factors are associated
with the development of resistance to winter
injury in plants. Qualitative and quantitative
changes in protein, carbohydrate, and lipid
contents have been observed during cold ac-
climation (Roberts 1969, Cerloff et" al. 1967,
Hochachka and Somaro 1968, Zeller 1951).
However, the in vivo features of observed
biochemical and physiological alterations as-
sociated with cold acclimation in specific in-
stances are not clear. In particular, very little
is known about the intracellular mechanisms
of freezing resistance in broad-leaved plants
that remain conspicuously green and meta-
bolically active throughout the winter
months of cool temperature regions.
Common mallow, Malva neglecta Wallr.,
is an example of a broad-leaved plant that of-
ten remains green and succulent throughout
the winter in north-temperature regions. Its
green leaves can tolerate subfreezing temper-
atures without visible evidence of injury. It
appears that common mallow is capable of
surviving winter cold by some mechanism
other than dormancy, because the plant re-
tains the capacity for photosynthesis and rel-
atively high respiration rates when favorable
conditions are present.
These observations have prompted an in-
vestigation of certain photosynthetic charac-
teristics and CO2 fixation properties in win-
ter-hardened mallow. This paper reports the
activity of purified ribulose diphosphate car-
boxylase (RuDPCa.se) and the capacity of
whole leaves to fix COo from cold-accli-
mated, field-grown mallow and from green-
house-grown mallow and tomato.
Materials and Methods
Plant nuitcriah: Common mallow is a per-
ennial weed characteristic of cultivated
ground, gardens, yards, and waste places
throughout the United States. Introduced
from Eiuope, the weed belongs to the same
plant family as cotton, hollyhocks, rose of
sharon, and the weeds known as velvet-leaf
and flower-of-the-hour. This family (Mal-
vaceae) has flowers which contain a tube of
stamens siurounding the pistil and a ring of
seeds centered in persistent floral parts remi-
niscent of a small flat cheese (thus one of the
plant's common names, "cheese weed"). The
plant's long tap root and its wide distribution
in relation to habitat occupation indicates a
wide ecological amplitude in regard to envi-
ronmental stress factors.
The garden tomato, Lycopersicon esculen-
tiim L., variety Ace 55, cannot tolerate sub-
freezing temperatiues. Tomatoes are warm
'Department of Botany and Range Science, Bni;liam Young University. Provo. Utah 84602.
121
122
Great Basin Naturalist
Vol. 40, No. 2
season plants; the Ace 55 variety yields very
well under high day and cool night temper-
ature regimes.
Method of sampling plants for measure-
ment of photosynthesis and respiration rates:
Plants of mallow and tomato were grown in
the greenhouse at 25 C (76 F) day and 20 C
(68 F) night temperatures. Mallow plants
were also grown in the field near Provo,
where they were exposed to subfreezing tem-
peratures. Plant samples were taken from
greenhouse and field areas during January.
When harvested, plants were collected
whole, petiole ends were cut under water,
and then they were placed in controlled envi-
ronment chambers with the cut ends remain-
ing immersed in water. Only deep green suc-
culent growth was harvested. Photosynthetic
and respiration measurements were then
made repeatedly as described below.
COo fixation methods: Rates of net photo-
synthesis (APS), dark respiration (DR), and
CP2 compensation points (CP) were deter-
mined in mid-January on excised shoots cut
under water. Analysis was made utilizing a
Beckman IR-215 infrared gas analyzer and a
plexiglas controlled-environment chamber.
Apparent photosynthesis (APS) was deter-
mined by the time required for the closed
system's CO2 content to decrease from 315 to
275 jul per 1 of air. Dark respiration (DR) was
determined by the time for the closed sys-
tem's CO2 content to return from 275 to 315
jtxl per 1. The CO2 compensation point (CP)
was determined in a closed system by allow-
ing the plants to fix CO2 until no further
change in CO2 concentration occurred in the
atmosphere of the lighted plexiglas chamber
surrounding the plant. The assimilation
chamber was housed inside a large growth
chamber with lighting provided by 8 cool-
white inforescent tubes, 8 grolux (Sylvania)
inforescent tubes, and 10 25-watt in-
candescent globes. The light was filtered
through 4 cm of water and provided an in-
tensity of 6.05 X 104 ergs per cm per minute
at leaf height. This light intensity has been
indicated to be saturating for tomato at 315
jul CO2 per 1 of air. Chamber parameters
were: temperature -23 ±.6 C; relative hu-
midity—65 ±10 percent; wind speed— 4.0 dm
per minute (3 chamber volumes per minute)
(Brewster 1971).
Preparation of extracts from leaf homoge-
nates and enzyme purification: Fully ex-
panded leaves were washed and their midribs
removed and blotted dry. From this step on,
all procedures were carried out at 5 C. Ap-
proximately 3.0 gm samples of leaf tissue
were groimd manually with cold mortar and
pestle for 5 minutes, using 5 ml of 0.1 M
HEPES (N-2-hvdroxvethvlpiperzaine-N-2-
ethanesulfonic acid) buffer pH 8.00, 0.001 M
EDTA, 0.0001 M DTT, 0.01 M MgClai, 0.025
mM NaHC03, per gram fresh weight leaf tis-
sue. The homgenates were centrifuged for 10
minutes at 20,000 rpm in a Sorvall model
RC-2B centrifuge with the S-34 rotor. The
supernatant was decanted and used as the
crude enzyme extract. The crude enzyme ex-
tract from the low speed centrifugation was
further clarified by centrifugation at 40,000
rpm in a Spinco model L3-50 for 5 minutes.
The supernatant fraction was collected. The
RuDPCase enzyme was purified further by
sedimentation of the extract into a sucrose
step gradient consisting of 2 ml of 5 percent,
2 ml 30 percent and 3 ml 50 percent sucrose
solution in HEPES buffer layered in a centri-
fuge tube. The rapidly sedimenting RuD-
PCase accumvilated in the 50 percent sucrose
layer after sedimentation for 12 hours at
25,000 rpm in a Spinco SW-25 rotor. The su-
crose-enzyme solution was then passed
through a 10 X 1.0 cm Sephadex G-25 col-
umn for further purification.
Measurement of enzyme activities: Deter-
mination of RuDPCase activity was based
upon fixation of ^^COa. into acid stable prod-
ucts. The assay mixture contained 0.01 M
HEPES-S04 buffer (pH 8.00) 0.01 M MgC12,
0.001 M DTT, 0.02 M NaHi^COa, and ribu-
lose-1, 5-bisphosphate in 200 [i\. The enzyme
(30jul) was added to initiate the reaction and
was allowed to proceed for 10 minutes. The
reaction rates were linear over this time peri-
od. The enzyme reaction was stopped by the
addition of 50 ju.1 of glacial acetic acid. A 100
jul aliquot of the reaction mixture was spotted
onto a strip of Whatman No. 1 filter paper
and dried under the hood. The sample was
counted in a Packard Tri-Carb (Model 3320)
liquid scintillation counter in toluene scintil-
lation fluid. The counted samples were cor-
rected for machine efficiency and quenching
June 1980
Andersen, Brotherson: Cold Acclimation
123
and the values converted to disintegrations
per minute (dpm).
Determination of specific activity of RuD-
PCase: RuDPCase has been identified with a
large, rapidly moving boundary observed in
the Spinco Model E ultracentrifuge known as
fraction I protein. The area of the Schlieren
boundary curve corresponding to fraction I
protein provides a means to determine the
relative concentration of RuDPCase present.
An estimate of specific activity per unit of
enzyme can then be calculated by comparing
enzvme activitv in a given extract with the
area of the corresponding fraction I protein
peak (Andersen et al. 1970).
Results
Net photosynthesis, dark respiration, CO2
compensation point: Table 1 shows that cold-
adapted, field-grown mallow exliibited lower
rates of APS than greenliouse-grown mallow
or tomato. However, rates of DR were sim-
ilar in all cases. This resulted in an APS to
DR ratio for field-grown mallow of one-half
that for the greenhouse-grown plants. The
CP shows a significant increase for field-
grown Mallow over values for the green-
house-grown plants.
Reaction velocities for carbonate and ri-
hu\osedip}\osphate substrates: The com-
parative reaction velocities for purified
"cold-adapted" mallow and tomato RuD-
PCase at different carbonate substrate con-
centrations are depicted in Figure 1. The ri-
bulosediphosphate (RuDP) substrate was
maintained at maximum concentration for
both enzymes. The mallow RuDPCase exliib-
ited higher catalytic capacity per unit of en-
zyme than tomato RuDPCase. The turnover
number at V.^^, (4 u moles of carbonate sub-
strate per 200 ul of reaction mixture) for to-
mato (RuDPCase) was calculated at 1036
moles carbonate fixed per mole of enzvme
per minute. The turnover number for puri-
2
V 10
i 8
< 6
O •MALLOW
•■TOMATO
Ofl 1£ 2.4 32 40 4B 56 64
CONCENTRATION HCO," (>i MOLES)
Fig. 1. Dependence of RuDPCase activity in pmified
extracts upon HC0.3 concentrations. RuDPCase was pu-
rified from common mallow and tomato ACE 55 var.
The enzyme activities are based upon total amount of
RuDPCase enzvme present in the reaction mixture as
calculated from the Schlieren curve of the sedimenting
boundaries in a model E ultracentrifin'e.
T.\BLE 1. Rates of net photosynthesis, dark respiration, and the CO2 compensation point of excised plant shoots.
See text for description of plant treatments.
Species
Location
.\pparent
photosynthesis
(ugCo2'dnr^*min"^ )
Mallow
Mallow
Tomato (ACE 55)
Greenhouse
Field
Greenhouse
3.2
1.5
2.6
Dark respiration
(ugCo2'dnr^Mnin"')
L7
1.7
1.3
CC^ compensation
point (uI.LM
64
86
68
*.\pparent photosynthesis {.\?S) was measured bv determining time required to lower closed system COj concentration from 320 ul/I to 280 ul/1 air-1.
^'Dark respiration' (DR) was measured by determining the time required for a darkened closed system to return COj concentration from 280 to 320 ul/1
air.
'^Compensation point was measured by allowing photosynthesizing plants to fix CO2 from a closed atmosphere until no further change in CO2 concentra-
tion could be obser\ ed.
124
Great Basin Naturalist
Vol. 40, No. 2
fied mallow RuDPCase was calculated at
1400 moles of carbonate fixed per mole en-
zyme per minute. The differences in V,„^^ val-
ues for the two enzymes were judged to be
highly significant, based on the student-t test
for measuring differences between paired
variates. The calculated t value exceeded the
0.001 level of significance. Only slight differ-
ences between the corresponding Km values
could be observed.
The reaction velocities of mallow and to-
mato RuDPCase for different ribulosediphos-
phate substrate concentrations are graphed in
Figure 2. The shapes of the reaction velocity
curves for tomato and Mallow RuDPCse are
similar. The mallow enzyme exhibited a sig-
nificantly higher V^,^, value. Both enzymes
showed substrate inhibition at RuDP substate
concentrations higher than 8 u moles per 200
jLil of reaction mixture. The significance of
the differences between the two reaction ve-
locity cmves in Figure 2 was measured by
the student-t test for paired variates. The cal-
culated t value exceeded the 0.01 level of sig-
nificance. The Km values for substrate con-
centration at half maximal velocity were
slightly higher for tomato.
Effect of temperature on reaction velocity
with purified RuDPCase enzyme: Purified to-
mato and mallow RuDPCase enzyme extracts
were compared for catalytic velocities at re-
action temperatures ranging from 4 C to 63
C (Figure 3). Purified mallow RuDPCase had
significantly higher catalytic activity per unit
of enzyme under the temperatine range of 4
to 25 C. The calculated student-t value for
differences between paired variates exceeded
the 0001 level of significance. On the other
hand, purified tomato enzyme showed signifi-
cantly higher catalytic capacity in the 38 to
53 C temperature range. The corresponding
calculated student-t value for paired reaction
rates in the 38 to 53 C temperature range ex-
ceeded the 0.01 significance level. The gener-
al shapes of the temperature curves for the
purified mallow and tomato RuDPCase were
quite similar, however, with heat denatura-
tion for both enzymes occurring near 53 C.
Discussion
Our studies indicate that respiration and
photosynthesis are differentially effected by
COMCCNTRATION OF RuOP (ji MOLES)
Fig. 2. Dependence of RuDPCase activity in purified
extracts from mallow and tomato ACE 55 var. upon
RuDP concentrations. Enzyme activities are normalized
for equal concentrations of RuDPCase.
20 25 30 35 40 45 50 55 60 65
TEMPERATURE °C
Fig. 3. Dependence of RuDPCase activity in purified
extracts from mallow and tomato ACE 55 var. upon
temperatine. Enzyme activities are normalized for equal
concentrations of RuDPCase.
June 1980
Andersen, Brotherson: Cold Acclimation
125
cold acclimation in mallow. The measured
rates of apparent photosynthesis values in
greenhouse-thrown mallow were approx-
imately twice the measured rate of apparent
photosynthesis values of cold-acclimated,
field-grown mallow. No differences were ob-
served for rates of dark respiration (Table 1).
Higher COo compensation points were ob-
.served for field-grown, cold-adapted mallow,
which suggested a depressed efficiency for
CO2 fi.xation. However, because respiration
and photosynthesis measurements were made
at 25 C in the laboratory, it is possible that
relative efficiencies of carbohydrate accumu-
lation would change at lower temperatures.
The cold-acclimated mallow might under
such circumstances become relatively more
efficient. Present evidence, however, in-
dicates that cold acclimation in field-grown
mallow is a matter of maintaining a steady
state of metabolic activity rather than the
rapid acciunulation of carbohydrate reserves.
Further studies are underway to assess inter-
action of lower temperatures and carbohy-
drate accumulation in cold-acclimated mal-
low.
Purified RuDPCase from cold-adapted
Malva neglecta had the highest catalytic ca-
pacity per unit of enzyme. The Vmax values
for carbonate and ribulosediphosphate sub-
strates were highest for mallow RuDPCase
(Figs. 2 and 3). The Km values for tomato
were only slightly higher. The Km and Vmax
values for tomato RuDPCase agree in general
with corresponding reported values (Ander-
sen et al. 1970). Although we cannot yet
compare in vitro RuDPCase activity to in
vivo CO2 fixation without some misgivings,
our studies suggest that mallow RuDPCa.se
would promote slightly more rapid CO2 fix-
ation per unit of enzyme in vivo. The lower
Km value for CO2 substrate of RuDPCase
from the cold-adapted mallow would suggest
a higher photosynthetic efficiency for the in-
tact plant. Yet the higher compensation point
of these cold-adapted plants indicates that
photosynthetic efficiency is depressed in the
intact leaf. Recent work by several in-
vestigators clearly implicates RuDPCa.se as a
major contributing factor to the high com-
pensation points of the C3 species (Ogren
and Hunt 1978). Our study would indicate
that the higher compensation point in the
cold-adapted mallow is due to some other
factor in the photosynthetic carbon cycle
than RuDPCase. On the other hand, since we
are using purified enzyme for our studies, it
is likely that control molecules that may af-
fect Km for CO2 fixation of RuDPCa.se could
be removed in our purification process. In
any case, if RuDPCase has a higher Km for
CO2 in vitro, which would result in a higher
compensation point in the cold-adapted mal-
low, the effect does not persist through puri-
fication of the enzyme. Therefore, at least a
change has not been detected on the purified
enzyme that would affect the compensation
point and thus be a basis for lower photo-
synthetic efficiency durmg cold acclimation.
The mallow RuDPCase enzyme showed
higher catalytic capacity than tomato RuD-
PCase under temperature ranges of 0-25 C,
and the tomato RuDPCase enzyme exhibited
higher activity under temperature ranges
40-60 C. This may be indicative of Mallow's
lower-temperature environmental adaptation
and its COo-fixing enzymes. Other studies
have shown that RuDPCase extracts from
plants of different climatic regions exliibit
correspondingly different temperature reac-
tions (Trihame and Cooper 1969). Also, we
have observed that the RuDPCase activities
in ciiide extracts from Mallow were con-
sistently higher than RuDPCase activities in
tomato crude extracts, (on a per-gram fresh
weight basis). These results, along with the
distribution patterns of these two species,
suggest that the temperature interaction of
the enzyme might be related in some way to
the different seasonal adaptations of the two
species.
The results also suggest that the process of
cold acclimation in mallow affects photo-
synthesis and dark respiration differently.
Respiration was not seriously affected, but
photosynthetic capacity per unit of leaf area
and photosynthetic efficiency were signifi-
cantly reduced (Table 1). It may be hypoth-
esized then that the processes of cold accli-
mation in mallow either depresses the in vivo
activity of RuDPCase or alters in some way
other chloroplast functions which affect the
plant's capacity for photosynthesis.
126
Great Basin Naturalist
Vol. 40, No. 2
Acknowledgment
This research was supported in part by the
National Institute of Health, Grant GM
17868-02.
Literature Cited
Andersen, W. R., G. F. Wildner, and R. S. Griddle.
1970. Ribiilose diphosphate carboxylase from mu-
tant tomato plants. .\rch. Biochem. Biophys.
137:84-90.
Brewster, S. F. 1971. The physiological vitality of scar-
let globemallow, Sphaeralcea gwssuhiriaefolia
(Hook. & .\RN.) Rydberg, under drought. Unpub-
lished dissertation, Brigham Young University,
Prove, Utah.
Gerloff, E. D., M. a. Stahmann, and D. Smith. 1967.
Soluble proteins in alfalfa roots as related to cold
hardiness. Plant Physiol. 42:895-899.
Hochachka, P. W., and G. N. Somero. 1968. Adapta-
tions of enzymes to temperatures. Comp. Bio-
chem. Physiol. 27:659-668.
Ogren, W. L., and L. D. Hunt. 1978. Gomparative bio-
chemistry of ribulose-bisphosphate carbo.xylase n
higher plants. Pages 127-1.39 in Siegelman and
Hind, eds., Photosynthetic carbon assimilation.
Basic Life Sciences 11.
Roberts, D. W. A. 1969. A comparison of isozymes of
wheat plants grown at 6 G and 20 G. Ganadian J.
Bot. 47:263-265.
Treharne, K. J., AND J. P. GooPER. 1969. Effect of tem-
perature on the activity of carboxylase in tropical
and temperature graminae. J. Exp. Bot.
20:170-175.
Zeller, O. 1951. Respiration at low temperatures in
cold hardened cereals. Planta. .39:.500.
RECOVERY OF GAMBEL OAK AFTER FIRE IN CENTRAL UTAH
L. M. Kiinzler' and K. T. Harper'
.\bstr.\ct.— The height of oak (Qucrcus gdinbelii Nutt.) stems was measured on several fire scars within the Uinta
National Forest and vicinity and compared with the height of oak stems on adjacent, nonhumed areas. A significant
relationship exists between the recovery rate of oak after fire and elevation, with the recovery rate being greatest at
low elevations. \ trend also exists showing that recoverv tends to be greater on south to westerly exposvires than on
north to easterly exposures.
Gainbel oak {Qucrcus gambelii Nutt.) is an
important species of the deer winter ranges
of central Utah, providing both food and cov-
er for deer (Alhiian 1952, Smith 1949). How-
ever, because of its growth habit, it often
forms impenetrable thickets (Allman 1952,
Baker 1949, Dills 1970, Marquiss 1972,
McKell 1950). This, coupled with its height,
places most of the available browse out of
reach of big game (Plummer et al. 1966,
1970). By treating these dense stands of oak
with chemical herbicides, fire, or machinery
to break them up, the stands can be opened
up and made available to browsing animals
(Anon. 1966, Dills 1970, Hallisey et al. 1976,
Marquiss 1971, 1972, Plummer et al. 1966,
1970, Price 1938). Because oak stands recov-
er rather rapidly after these treatments, it is
necessary to determine a rotation period for
treatment to maintain an optimum amount of
browse for wildlife (McKell 1950, Plummer
et al. 1966, 1970).
Literature Review
In central Utah, Gambel oak has been re-
ported to occur in almost pure stands from
5000 (1525 m) to 8000 (2440 m) feet eleva-
tion along the Wasatch Range (Allman 1952,
Baker 1949, McKell 1950). This area con-
stitutes a large portion of the deer winter
range in the area (Allman 1952, Anon. 1966,
Dills 1970, Hallisey et al. 1976, Plummer et
al. 1966, 1970, Smith 1949).
Treatments of oak using fire, herbicides, or
machinery to destroy the oak canopy result
in prolific sprouting, with several stems re-
placing each preexisting stem. Impenetrable
thickets often result from such treatments
(Allman 1952, Baker 1949, Dills 1970, Mar-
quiss 1972, McKell 1950). Yet, treatments can
be effective in improving deer range. When a
follow-up program is used, such as seeding
with competitive herbs and grasses, the ben-
efits of the treatment can be prolonged for
over 15 years (Anon. 1966, Dills 1970, Hal-
lisey et al. 1976, Marquiss 1971, 1972, Plum-
mer et al. 1966, 1970, Price 1938). By treat-
ing oak, deer use can be increased up to four
times, but deer use declines as the time from
treatment increases (Anon. 1966, Hallisey et
al. 1976, Price 1938).
Methods
The height of oak stems was measured on
several stands within the Uinta National For-
est and vicinity. One half of these stands
were located in oak stands that had burned 3
to 15 years ago. The other half of the stands
were located in nonburned areas adjacent to
each burned stand considered. Unburned
stands were selected so as to have the same
slope, exposure, and elevation as the burned
stand that each was paired with. Mea.sure-
ments were taken along a 100-foot transect at
16-foot intervals, with the oak stem that was
nearest to the point being measured. Slope
varied from 20 to 70 percent and elevation
ranged from 5100 feet (1555 m) to 6800 feet
'Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602.
127
128
Great Basin Naturalist
Vol. 40, No. 2
(2070 m) in elevation among the stand pairs
considered. Exposure also varied, with one-
half of the stands having a south to west ex-
posure and the other half having a north to
east exposure. A percent recovery value was
calculated for each pair by dividing the aver-
age height of the oak in the fire scar by the
average height of the oak in the nonburned
area. A recovery rate was then calculated by
dividing the percent recovery value by the
age of the fire scar (Table 1). To minimize
the variation among recovery rates caused by
the nuisance factors of age, elevation, slope,
and site, recovery rates were also calculated
on a uniform hill within the Wallsburg Burn
(Table 2). Three transects were placed in the
Ijumed area at 500-foot (152 m) intervals in
elevation.
Results
In comparing the recovery rates from the
stands throughout the Uinta National Forest
and vicinity, a significant correlation (power
equation) exists between elevation and oak
recovery rates (r = -.85, .01 < P < .05, Fig.
1). Using data collected from the Wallsburg
Burn, a similar analysis confirms that a signif-
icant correlation (linear equation) exists be-
tween elevation and the recoverv rates of oak
stems (r = .99, P <.01, Fig. 2)'. Both sets of
data show that, as elevation is increased, the
recovery rates decrease. Although there is in-
dication that stands on south to west expo-
sures have faster recovery rates, the differ-
ence between the recovery rates of these
stands and those with north to east exposures
is not significant (Fig. 3).
Discussion
Comparing the results of the two sets of
data, it seems clear that elevation has a
strong influence on oak's rate of recovery fol-
lowing fire. The variation between recoverv
rates in Figure 2 is primarily due to elevation
Table 1. Annual recoverv rates in percent for oak on burns scattered throuij;hout the Uinta National Forest and
nearby areas.
Average
Average
height of
Age of
height of
oak in non-
Recoverv
Stand
Elevation
burn
oak on burn
burned area
Percent
rate
#
(m)
Exposure
(years)
(cm)
(cm)
recovery
(percent)
1
1,740
East
3
77
579
13.3
4.4
2
1,800
North
15
140
320
43.7
2.9
3
1,560
South
3
145
274
52.8
17.6
4
1,770
South
4
83
343
24.2
6,1
5
1,650
East
3
84
290
28.9
9.6
6
1,680
West
3
61
135
45.0
15.0
7
1,680
East
3
83
290
28.5
9.5
8
1.770
West
4
88
399
22.1
5.5
9
1,860
East
6
116
533
21.7
3.6
10
2,070
West
4
37
320
11.4
2.9
T.\BLE 2. Percent recovery values for oak on a unih)rni hill in the Wallsburg Burn. This stand had burned four
years earlier.
Transect
1
1-a
2
2-a
3
3-a
Elevation
Status
Average height
Percent recovery
1,770 m
Burned
88.4 cm
22.2
1,770 m
Unburned
398.8 cm
1,920 m
Burned
59.9 cm
16.8
1,920 m
Unburned
356.6 cm
2.070 m
Burned
■36.6 cm
11.4
2,070 m
Unburned
320.0 cm
June 1980
KuNZLER, Harper: (jAmbel Oak
129
alone. Even with the nuisance variables men-
tioned earlier, it will he noted that elevation
is a significant factor for recovery rates of
oak (Fig. 1).
Possible reasons for the change in oak re-
coverv rates with elevation include the fol-
lowing: (1) the species is approaching its up-
per elevational limit on some of the burns
and, because of this, its growth may be slow-
er; (2) more moisture and nutrients may be
available to plants at the bottom of slopes be-
cause of precipitation's surface nuioff and at-
tendant erosion, nutrient transport, and re-
sultant differences in soil depth at the top
and bottom of the slope; and (3) the shorter
growing season at the higher elevations gives
less time for growth there. There may be oth-
er reasons or a combination of reasons for
this phenomenon. In anv event, the relation-
ship is strong and has management implica-
tions.
In the winter, deer in the Uinta National
Forest and nearby areas primarily use south-
and west-facing slopes at lower elevations
(Bruce Giunta and Jordon Pederson, Utah Di-
vision of Wildlife Resources, and Juan Spil-
lett, Uinta National Forest, pers. comm.).
Such areas coincide with situations where
oak recovery is most rapid. Thus, manage-
ment programs to regenerate oakbrush on
deer winter ranges in our area may be short-
lived. If Gambel oak is to be manipulated to
improve deer range using conventional meth-
ods in this area, a follow-up program that
will retard oak recovery should be used. The
20-1
15
10-
.85
1,530m
1,740 m 1,950 m
Elevation
2,130m
Fig. 1. The relationship between recovery rates of
oak and elevation on various burns throughout the Uinta
National Forest and nearbv areas.
significant increase in time between major
treatments will thus minimize management
costs.
Conclusions
Elevation is a significant factor in affecting
the recovery rate of oak after fire, with high-
er elevation stands recovering more slowly.
Recovery takes from 6 to .35 years in this
area, with a modal recovery time of about 15
years.
25H
20-
15-
10
r = .9998
1,770m
1,920m
Elevation
2,070 m
Fig. 2. The relationship between recovery rate and
elevation for oak in the Wallsburg Burn area.
10-1
0)
4-1
CD
P^
>,
U
>
o
o
0)
Pi
>
<:
5-
s-w n -e
Exposure
Fig. .3. Histograph of the average recovery rate of oak
stands with south to west exposures and north to east ex-
posures on burns throughout the Uinta National Forest
and nearby areas.
130
Great Basin Naturalist
Vol. 40, No. 2
Acknowledgments
The authors wish to thank the Uinta Na-
tional Forest and the Utah Division of Wild-
life Resources for permitting access to the
study areas. Funding for this study was pro-
vided by the Uinta National Forest (Supple-
ment to Cooperative Agreement 12-11-204-
31). The authors also express thanks to Karl
McKnight for help in the statistical analysis
of the data.
Literature Cited
Allman, v. p. 1952. A preliminary study of the vegeta-
tion in an exclosure in the chaparral of the
Wasatch Mountains, Utah. Unpublished thesis,
Brigham Young University, Provo, Utah. 236 pp.
Anon. 1966. Conversion of thicket covered areas to pro-
ductive grazing lands. USDA, Forest Service, Int.
Reg. Range Imp. Notes 11(4): 1-4.
Baker, W. L. 1949. Soil changes associated with recov-
ery of scrub oak (Querctis gambelii) after fire. Un-
published thesis, Univ. of Utah, Salt Lake City,
65 pp.
Brown, H. E. 1958. Gambel oak in west-central Colo-
rado. Ecology .39:317-327.
Dills, G. G. 1970. Effects of prescribed burning on deer
browse. J. Wildl. Manage. 34:540-545.
Eastmond, R. J. 1968. Vegetational changes in a moun-
tain brush commimity of Utah during 18 years.
Unpublished thesis, Brigham Yoiuig Univ., Provo,
Utah, 64 pp.
Hallisey, D. M., and G. W. Wood. 1976. Prescribed
fire in scrub oak habitat in central Pennsylvania.
J. Wildl. Manage. 40:507-516.
Marquiss, R. W. 1971. Controlling Gambel oak on
rangelands of southwest Colorado. Colo. .'Vgri.
Exp. Stat. Progress Report #PR71-9. 2 pp.
1972. Soil moisture, forage, and beef production
from Gambel oak control in Southwest Colorado.
J. Range Manage. 25:146-150.
McKell, C. M. 1950. A study of plant succession in the
oakbnish (Querctis gambelii) zone after fire. Un-
published thesis, Univ. of Utah, Salt Lake City.
74 pp.
Plummer, a. p., D. R. Christensen, and S. B. Monsen.
1966. Highlights, results, and accomplishments of
game range restoration studies. Utah State Dept.
of Fish and Game. Publ. No. 67-4:7-9.
Plummer, A. P., R. Stevens, and K. R. Jorgensen. 1970.
Highlights, results, and accomplishments of game
range restoration studies. Utah State Dept. of
Fish and Game. Publ. No. 70-3:26-30.
Price, R. 1938. Artificial reseeding on oakbrush range in
Central Utah. USDA Circ. No. 458. 19 pp.
Smith, J. G. 1949. Deer forage observations in Utah. J.
Wildl. Manage. 13:314-315.
RELATIONSHIPS AMONG TOTAL DISSOLVED SOLIDS, CONDUCTIVITY,
AND OSMOSITY FOR FIVE ARTEMIA HABITATS (ANOSTRACA: ARTEMIIDAE)
Nicholas C. Collins' and Gray Stirling'
.\bstract.— Graphs allowing interconversion between various physical chemical parameters are presented for five
Aiicmia habitats in the western USA. Both the mean osmosity and its typical yearly range differ greatly among habi-
tats. Consequently, Arteiiiici populations provide an interesting opportimity to study physiological and life history
adaptations to differing degrees of habitat stability.
Populations of Aiiemia, the brine shrimp,
exist in isolated hypersaline environments
throughout most of the world (McCarraher
1972). Their source waters span the entire
natural spectrum of ion ratios (Cole and
Brown 1967) and range from the massive and
relatively permanent Great Salt Lake to tem-
porary ponds 50 m to diameter (e.g. Broch,
1969; Khalaf et al. 1977). Not surprisingly,
the individual populations exhibit morpholo-
gical, physiological, developmental, and gen-
etic differences that indicate they are locally
adapted (e.g. D'Agostino 1965, Clark and
Bowen 1976, Glaus et al. 1977, Collins 1977).
Because the resting cysts of these popu-
lations are easy to collect, transport, store,
and hatch, Artemia populations are excellent
subjects for comparative studies of genetics
(Barigozzi 1974, Clark and Bowen 1976),
physiology of ion regulation (e.g., Geddes
1975a, b, c), and life history tactics (Collins
1977, Glaus et al. 1977). Many such studies
have involved comparisons of the perform-
ance of various strains grown in a common
medium, usually diluted or concentrated sea-
water. An alternate approach, which accom-
modates some strains that will not grow well
in sea water, involves growing them each in
their own source water, but at a common os-
motic pressure. This alternative requires
measurement of the osmotic characteristics
of various dilutions of water from each
source lake, a time-consuming process requir-
ing osmometers that are both expensive and
uncommon. To reduce the necessity for fu-
ture osmometric measurements for studies of
Artemia populations in the western United
States, this paper presents relationships be-
tween osmosity and more easily measured pa-
rameters for source waters of five popu-
lations. Data on pH changes with source
concentration and information on the natural
range of concentrations for each source are
also presented.
Methods
Source waters from Arizona, New Mexico,
Nebraska, and Washington, collected during
1976 and 1977, were filtered and diluted or
concentrated by evaporation. Locations for
each source are specified in the references in
Table 1. Total dissolved solids (TDS) concen-
trations were determined by evaporating five
or ten ml samples to a constant weight at 100
C. Salt scale had to be repeatedly broken to
insure completion of the drying process.
Drying at temperatures higher than 100 G
resulted in steam explosions within salt mas-
ses that scattered the salt and biased the de-
terminations.
Conductivity meter readings were con-
verted to specific conductance at 20 G using
an NaCl calibration curve based on Wolf,
Brown, and Prentiss (1975).
Specific gravity at 20 C was measured
gravimetrically using individually calibrated
50 ml volumetric flasks. Each determination
'Department of Zoolog)' and Erindale College, University of Toronto, 3359 Mississauga Road, Mississauga, Ontario L5L 1C6.
131
132
Great Basin Naturalist
Vol. 40, No. 2
is the average of duplicate measurements.
Osmosity, the molar concentration of NaCl
having the same freezing point or osmotic
pressure as the measured solution, was deter-
mined with a Wescor vapor pressure os-
mometer. The microvoltmeter output was
calibrated with a series of NaCl solutions
whose osmotic properties were assumed to
correspond with Wolf et al. (1975). Two to
five determinations were averaged for each
data point in the figures.
Results
For three of the five sources (Figs. 1-3) os-
mosity is closely, linearly related to TDS.
Where Y is osmosity, X is TDS, n is the num-
ber of measurements, and r is the product-
moment correlation coefficient, the relation-
ships for Penley Lake (Washington), Green
Pond (Arizona), and Lily Lake (Nebraska) are
respectively:
E
u
(/)
o
E
E
TOTAL DISSOLVED SOLIDS
Fig. 1. Relationships among TDS, conductivity (open circles), and osinositv (closed circles) for Penley Lake, near
Oniak, Washington (Broch 1969). Numbers below upper margin are specific gravity measurements and those above
lower axis are pH values for indicated TDS levels.
June 1980
Collins, Stirling: Artemia Habitats
133
Y = .0081 X +
.0176
n = 23
r = .995
Y = .0160 X +
.0141
n = 19
r = .995
Y = .0103 X +
.0439
n = 30
r = .998
The TDS-osmosity relationships for the other
two lakes (Figs. 4, 5) did not appear to be lin-
ear, and were drawn by eye. Similarly, the
curvihncar TDS-conductivity relationships
lor all lakes were drawn by eye.
E
a
«/)
o
"I
E
>-
>
a
o
u
160
40 80
120
160
1 i 1 II 1
' 1 '
1 1 1
-
1.0390-
1.0465 •
1
CD
CO
eo
yv
-
140
-
/ /
"~
-
o/
° /
-
120
-
GREEN POND
o /
/o
/
—
■
jPo /
100
f /•
80
-
/•
-
60
-
•
—
40
-
—
20
- [
—
-/
9.8
1 1 ll 1
9.4
ll 1
1 1 1
—
3.20
- 2.80
- 2.40
- 2.00 _.
o
E
-11.60 r"
- 1.20
- .80
CO
o
C/)
o
0 40 80 120 160 200
TOTAL DISSOLVED SOLIDS ( g/l )
Fig. 2. Relationships among physical-chemical parameters for Green Pond, near St. Johns, .\rizona (Cole and
Whiteside 196.5). Legend as in Figure 1.
134
Great Basin Naturalist
Vol. 40, No. 2
Discussion
Generally, conductivity is the most con-
venient indicator of osmosity or TDS. The
natural concentrations of these soiuce lakes
fall well above the highest scale of most exist-
ing conductivity meters, but Figs. 1-5 will al-
low rough predictions of TDS or osmosity
from conductivity measurements of samples
diluted to within the range of such meters. In
such cases the errors in any prediction based
on the graphs will be multiplied by the dilu-
tion factor. At the highest source concentra-
tions conductivity is not a precise predictor
of TDS or osmosity (Figs. 4, 5). For such solu-
tions an accurate dilution can be made for
conductivity measurement, or TDS can be
measured directly. Inaccuracies in TDS mea-
O
E,
>-
CO
o
C/3
O
40 80 120 160 200
TOTAL DISSOLVED SOLIDS (g/l)
240
Fit;. 3. Kelationships among pliysical-eheniical paraiufteis ior Lilv Lake, near Alliance, Nebraska (McCarraher
1970). Legend as in Figure L
June 1980
Collins, Stirlinc;: Arteml\ Habitats
135
siirements from water inclusions in the crys-
talizing salt mass can be eliminated by using
a micrometer syringe for applying precisely
measured small volumes (< 1 ml) to a pre-
weighed filter paper circle. Such a technique
will also allow much faster determinations
tlian the one we used.
Hydrometer measurements of specific
gravity are often reported as a measure of
concentration. Although this method appears
to be quick, straightforward, and suitable for
field measurements, our experience indicates
even marginally accurate results require
careful control of water temperature, wind,
and cleanliness of the hydrometer that to-
gether preclude most field measurements. In
160
140
^ 120
E
u
o
E
E
100
80
60
40
20 -
50 100 150 200 250 300 350
T — ^ — T] — rpi — I 1 1 1 1 1 1 1 — I — r
2.0
4.0
50 100 150 200 250 300 350 400
TOTAL DISSOLVED SOLIDS (g/l)
CO
o
CO
Fig. 4. Relationships among phvsical-chemical parameters of Jesse Lake, near .Alliance, Nebraska (McCarraher
1970). Legend as in Figure 1.
136
Great Basin Naturalist
Vol. 40, No. 2
the laboratory, TDS measurements or gravi-
metric density determinations are almost as
easy as hydrometer readings, if a con-
ductivity meter is not available.
A summary of natural osmosity and TDS
levels from western Artemia habitats (Table
1) indicates that different populations expe-
rience not only very different ion ratios and
mean osmosities, but also very different sea-
sonal ranges in source osmosity. Western
North American Aiiemiu populations there-
fore provide an interesting opportunity to
E
o
o
"I
E
>-
>
I—
Q
O
50 100 150
200 250
300
225
1
1.0827-=
1
52 / 52-^
1 1
-75 8-
200
ZUNI SALT LAKE
/
/
:
175
■ 1
'
^
150
/ •
/
-
125
/ /
-
100
- / /
-
75
■ /y
-
50
' 1/
-
25
r
-
• 8.7 \
).5
8.1
7.9
1 1 ill 1. . J-
1 1
1 1 li
1 1
4.5
- 4.0
3.5
-13.0 =;
o
H2,5 ^
- 2.0
- 1.5
- 1.0
- .5
CO
o
C/9
O
0 50 100 150 200 250 300 350
TOTAL DISSOLVED SOLIDS (g/l)
Fig. 5. Relationships amonn pliysical-cliemital parameters of Zuiii Salt Lake, near Queniado, New Mexico (Brad-
bury 1971). Legend as in Figure 1.
June 1980
Collins, Stirling: Artemia Habitats
137
identity the genetic, physiological, and life
history characteristics that evolve in response
to variability of environmental conditions,
and in response to substantial differences in
length of the growing season.
Note added in proof: Recent talks with
Nebraska residents revealed that the lake re-
ferred to in this paper and in Collins (1977)
as Lily Lake of McCarraher (1970, 1972) is
actually an imnained smaller lake slightly
northwest of the tnie Lily Lake. The range of
osmosity for this lake is unknown; therefore
the figures for it in Table 1 should be dis-
regarded.
Acknowledgments
The National Geographic Society, the Nat-
ural Sciences and Engineering Research
Council of Canada, and the University of To-
ronto supported this work financially. 1 thank
Mr. Drew Piatt for permission to sample
Green Pond, Gerald Cole, J. P. Bradbiuy, and
Sarane Bowen for information on sampling
sites, and R. W. Cummins and J. Svoboda for
access to the o.smometer. LeeAnne Wilson
and John Stoneman provided excellent tech-
nical assistance.
Liter.\ture Cited
Baricozzi. C. 1974. Aitcinui: a survey of its significance
in genetic prolileins. Evolutionary Biol. 7:
221-252.
Bhadburv, J. P. 1971. l.iniiiology of Zuni Salt Lake, New
Mexico. Geol. Soc. Anier. Bull. 82: .379-398.
Broch. E. S. 1969. The osmotic adaptation of the fairy
slirinip Brcinrhinccta campestris Lynch to salaine
astatic waters. Linmol. Oceanogr. 14: 48.5-492.
Clark, L. S., and S. T. Bowe.n. 1976. The genetics of
Artemia xalina. VIL Reproductive isolation. J.
Hered. 67: 38.5-.388.
Claus, C, F. Benijts, a.nd P. Sorgeloos. 1977. Com-
parative study of different geographical strains of
the brine shrimp, Artemia salimi, pp. 91-105. In:
Jaspers, E. (ed.). European Mariculture Society
special publication No. 2. Bredene, Belgium.
Cole, G. \., a.nd M. C. Whiteside. 1965. Kiatuth-
lanna— a limnological appraisal. II. Chemical fac-
tors and biota. PJateau .38: .36-48.
Cole, G. \. and R. J. Brown. 1967. The chemistry of
Artemia habitats. Ecology' 48: 8.58-861.
Collins, N. C. 1977. Ecological studies of terminal
lakes— their relevance to problems in limnology
and population biology, pp. 411-420. In: D. C.
Greer (ed.). Desertic terminal lakes. Utah Water
Research Laboratory, Logan, Utah.
D".\(;osTiNo, \. 1965. Comparative studies of Artemia
■salina (development and physiology). Unpub-
lished dissertation. New York Univ.
Gedde.s, M. C. 197.5a. Studies on an .Australian brine
shrimp, Parartemia zietziana Sayce (Crustacea:
.\nostraca)— I. Salinity tolerance. Comp. Bio-
chem. Physiol. 51,\: .5.53-5.59.
Table 1. Comparisons of physical and chemical characteristics of six Artemia .sources in the western USA.
Water body
Major salt
Great Salt Lake,
Utah
NaCl
Jesse Lake.
Nebraska
NaCOj
Lily Lake.
.Nebraska
NaC()3-Cl
Green Pond,
NaC03-Cl
.\rizona
Penley Lake,
Na2S()4
Washington
Zuni Salt Lake,
New Mexico
NaCl
Observed^ TDS
range (g/1)
"Typical" annual
osmosity range^
(moles/1) References
130-300
.52-87
12-69
61-112
33-230
17.5-.350
2.-2.25
..57-.90
.16-. 76
.98-1.8
.24-dryness
Handv and Hahl
1966 '
McC:arraher 1970.
1972
.McCarraher 1970,
1972
Cole and
Whiteside 1965
Broch 1969
..35-5.3-1- Bradbury 1971
Range of values recorded in literature and personal observations, .\nnual range is usually smaller.
IncKides only the period during which active Artemia are present. For some lakes, only one year's data are available, so nothing Ls known about be-
tween-vear differences.
138
Great Basin Naturalist
Vol. 40, No. 2
1975b. Shidies on an Australian brine shrimp,
Parartemia zietziana Sayce (Crustacea: Anost-
raca)— II. Osmotic and ionic rei;iilation. Comp.
Biochem. Physiol. Sl.A: 561-571.
1975c. Studies on an Australian brine shrimp,
Parartemia zietziana Sayce (Cnistaceae: .\nost-
raca)— III. The mechanisms of osmotic and ionic
regulation. Comp. Biochem. Physiol. 51A:
573-578.
Handy, A. H., a.nd D. C. Hahl. 1966. Chemistry of the
water in the Great Salt Lake, pp. 135-151. In:
Stokes, VV. L. (ed.). The Great Salt Lake, Guide-
book 20. Utah Geol. Soc., Salt Lake City.
Khal.\f, a. N., M. a. Lattif, H. H. Mangalo, and M.
Salih. 1977. A bioecological study on the brine
.shrimp Artemia salina L. (Anostraca: Branchiop-
oda) in two inland brine temporary ponds in
Iraq. Bull. Biol. Res. Center, Bagdhad. 8: .37-48.
McCarraher, D. B. 1970. Some ecological relations of
fairy shrimps in alkaline habitats of Nebraska.
.\mer. Midi. Nat. 84: .59-68.
1972. A preliminary bibliography and lake index
of the inland mineral waters of the world. F.\0
Fisheries Circular No. 146. .33 pp.
Wolf, A. V., M. G. Brown, and P. G. Pre.ntiss. 197.5.
Concentratiye properties of aqueous solutions:
conversion tables, pp. D218-D273. In: Weast, R.
C. (ed.). Handbook of chemistry and physics,
56th ed.
SPAWNING OF THE LEAST CHUB [lOTICHTHYS PHLEGETHONTIS)
riioiiKis M. Baugh'
.\bstract.— The least tluib, loticlitlujs phlf^clhonlis (Cope), a relict fisli in I'tali, spawned suecesslully under lab-
oratory conditions.
The least chub, lotichthys pJiIcgethontis
(Cope), is a small (ca 5 cm) relict fish (Hubbs
and Miller 1948) found only in a few local-
ities in the western desert region of Utah. In
1973 the Utah Division of Wildlife Resources
classified this species as endangered. There is
little literature on /. phlcgethontis (Crawford
1979, Hubbs and Miller 1948, Pendleton and
Smart 1954, Sigler and Miller 1963), and
there are no reports of this species spawning
under artificial light in closed-svstem aquaria.
On 26 May 1979, I obtained five male and
five female least chub from an open raceway
at the Utah Division of Wildlife Resources fa-
cility, Logan, Utah. The fish had originally
been collected from Leland Harris Spring
and the associated marsh between May 1977
and February 1978 by Crawford (1979). I
placed the fish in a 61 X 41 X 31 cm (72.5
liter) aquarium with commercial aquarium
gravel placed over a subgravel filter to a
depth of about 5 cm. The water was con-
stantly aerated and the tank was densely
planted with simulated, broad-leafed plants.
The fish were fed a mix of TetraMin^ Staple
Food and Tetra KrillflakesR at 0630 and fro-
zen San Francisco Bay Brand'^ brine shrimp
at 1630 each day.
Once each week, for a two-hour period,
the water was filtered through a Vortex Dia-
tom'^ filter. Also once each week, 15 percent
of the aquarium water was drawn off and re-
placed with an equal amount of aged tap wa-
ter. One oimce of Instead Ocean*^ marine salt
mix in solution was added each week.
On 15 October 1979, I added two 29 cm
long strips of Living World^ spawning grass
to the aquarium. This spawning meditun was
examined daily, and on 26 October one
length of the medium contained five mildly
adhesive eggs. This piece of medium was re-
moved from the aquarium and placed in a 3.6
liter glass jar containing water from the aqua-
rium. The water in the jar was mildly agi-
tated with air. On 27 October one and on 28
October three additional eggs were removed
from the aquarium to the jar. The water in
the jar was maintained at the same temper-
ature as that in the aquarium.
Free swimming larvae were first noted on
31 October, and by 2 November all nine eggs
had hatched. The larvae were able to adliere
to glass and plastic. The mechanism of at-
tachment was not studied.
The following conditions existed at the
time of spawning. During the 10 days prior
to the day of last-noted egg deposition, the
water temperature ranged from 17.7 to 18.8
C and averaged 18.2 C. The photoperiod was
14 hours of daylight and 10 hours of darkness.
Water conditions were: pH 7.6, total alka-
linity 84.1, total hardness 186, CI- 683, Ca
48.4, an Mg 45.1. Due to an equipment mal-
fimction, dissolved oxygen was not measured.
Two additional spawnings took place on 5
Noveiuber 1979 and 8 November 1979. Fif-
teen eggs were gathered from the former and
eight eggs from the latter spawning. In addi-
tion, several other spawnings from these fish
occurred from which the eggs were not
taken.
From the above, it appears that /. pfilc^e-
thontis is amenable to culture in closed-sys-
tem aquaria under artificial light.
'1020 Custer .\ venue, Ogden, Utah 84404.
139
140
Great Basin Naturalist
Vol. 40, No. 2
Acknowledgments
I thank the Utah Division of Wildhfe Man-
agement, especially Donald Andriano, for
granting Permit s'CC-SL-919, which made
this work possible; and Kent Miller, Utah Di-
vision of Wildlife Resources, for reviewing
this note.
Literature Cited
Cr.\\vford, M. 1979. Reproductive modes of the least
chub [lotichtltijs pJilcgetJiontis Cope). Unpub-
hshed thesis. Utah State Univ. 79 pp.
HuBBs, C. L. .\ND R. R. Miller. 1948. Correlation be-
tween fish distribution and hvdrographic historv
in the desert basins of the western United States.
Bull. Univ. Utah, Biol. Ser. 19(7): 17-166.
Pe.ndleton, R. C, and E. W. Sm.\rt. 1954. A stud\ of
the food relations of the least chub, lotichthijs
phlegcthontis (Cope), using radioactive phos-
phonis. J. Wildlife Manag. 12(2): 226-228.
SiGLER. W. F.. .\ND R. R. Miller. 1963. Fishes of Utah.
Pages 82-84 in Utah Div. ^\■ildlife Resour. Salt
Lake Citv.
TRANSFERRIN POLYMORPHISM IN BIGHORN SHEEP,
OVIS CAXADEXSIS, IN COLORADO
Patrick W, Roberts . Donald J. Nash . and Robert E. Keiss'
.\bstr\ct.— Senmi transferrins were analyzed by polyacrylamide gel electrophoresis in four populations of Colo-
rado bighorn sheep. Oti.s canadensis canadensis. Transferrin was found to be polymorphic, with two alleles. Tf D
and Tf E. being represented in each of the four populations. Within herds the phenot) pic ratios confonned to values
predicted h\- the Hardv- Weinberg equilibrium, .\niong populations, significant differences were seen with respect to
phenotvpic frequencies.
Transferrin polymorphisms have been de-
scribed in a number of breeds of domestic
sheep and in different species of wild sheep
including Ovis canadensis, O. dalli, and O.
mouflon (Nadler et al. 1971). They reported
three transferrin alleles, Tf B+, Tf D, and Tf
E, in two subspecies of bighorn sheep, O. c.
canadensis and O. c. mexicana. In 14 speci-
mens of O. c. canadensis from Montana, 13
had the Tf DE phenotype and one sheep was
B^D. Two specimens of O. c. mexicana from
.\rizona were of the EE phenotvpe.
In Colorado, bighorn sheep historically
ranged over much of the central and western
parts of the state, but the distribution has
been fragmented in recent times (Armstrong
1972), and there are now more tlian 30 dis-
junct bands occurring in the less accessible
parts of the higher mountains. A study was
undertaken to characterize electro-
phoretically demonstrable genetic variation
in several sennu proteins and in hemoglobin
of several disjimct herds to determine the de-
gree of genetic similarity or dissimilarity
among and within the bands sampled. The
present stud\ is a report of transferrins ob-
served in several herds.
Blood samples were collected from four
different herds in Colorado. The designation
of the herds and their centers of distribution
are as follows: (1) Poudre— north slope of
Poudre Canvon. Larimer Co.. (2) Tarrvall—
Tarryall and Kenosha Mts., Park Co., (3)
Chalk Creek— Chafee Co., and (4) Gunnison-
Gunnison Co.
Transferrins were analyzed by poly-
acrvlamide disc gel electrophoresis using the
techniques described by Smith (1968). Gels
were prepared at 7 percent (w/v) concentra-
tion. Senuu samples were prepared by mak-
ing serum with 50 percent sucrose containing
0.25 percent brom phenol blue as a tracking
dve. Electrophoresis was carried out in tris-
givcine buffer at pH 9.5. Twelve senmi sam-
ples were electrophoresed for 26 minutes at 3
milliamps per gel at 10 C.
Samples of domestic sheep blood of known
transferrin tvpe were obtained from the Sero-
log\" Laboratorv of Dr. Stormont of the Uni-
versitv of California at Davis and were used
as reference sera.
All populations were polymorphic for
transferrin phenotypes (Table I). Two herds.
Chalk Creek and Poudre. had three pheno-
t\pes and two herds. Tarr\ all and Gunnison,
each had two phenot>pes. The phenotypes
Table 1. Transferrin phenotypic frequencies of big-
horn sheep, Ovis canadensis canadensis, in Colorado
(uiunbers of observations in parentheses).
Herd
Tf DD
TfDE
TfEE
Gunnison (7)
0.000
0.857
0.143
Chalk Creek (16)
0.313
0.374
0.313
Tarrvall (26)
0.577
0.423
0.000
Poudre (18)
0.316
0.526
0.158
'Department of Zoologj- and Entomolog\-, Colorado State University . Fort Collins, Colorado 80523.
-Colorado Di\ision of Wildlife, Fort Collins, Colorado 80521.
141
142
Great Basin Naturalist
Vol. 40, No. 2
were determined to correspond to those pro-
duced by two alleles, Tf D and Tf E. Allelic
frequencies ranged from 0.43 to 0.79 for Tf
D and from 0.21 to 0.57 for Tf E. Significant
differences among herds were observed for
the distribution of phenotypes. Within herds
the transferrin frequencies followed a Hardy-
Weinberg distribution. The proportion of
heterozygotes was high in all populations,
with the lowest value of 0.375 being ob-
served in the Tarryall herd.
Although surveys of isozymes in natural
populations of small mammals have indicated
considerable genetic variability, relatively
few biochemical studies have been done on
large mammals. Bonnell and Selander (1974)
found no polymorphisms in 24 presumptive
loci in northern elephant seals. Hetero-
zygosites of 0.04 have been reported for elk
(Cameron and Vyse 1978), 0.04 for moose
(Ryman et al. 1977), and 0.32 for white-tailed
deer (Manlove et al. 1976). These species
were monomorphic at the transferrin locus
except for white-tailed deer, which had 23
percent heterozygosity.
It is of interest that the bands of bighorn
sheep in Colorado retain such a high degree
of polymorphism, at least at the transferrin
locus, although the populations have been
relativelv isolated and have had relatively
small population numbers. Some recent esti-
mates of population size include Poudre,
65-75, Tarryall, 100, and Chalk Creek,
90-100. Results at the transferrin locus in-
dicate that inbreeding within the herds may
not be a major problem, although surveys of
additional genetic loci should be undertaken.
Literature Cited
Armstrong, D. 1972. Distribution of mammals in Colo-
rado. University Kansas Mas. Nat. Hist., Monogr.
3:1-415.
Bonnell, M. L.. and R. K. Selander. 1974. Elephant
seals: Genetic variation and near extinction. Sci-
ence 184: 908-909.
Cameron, D. C, .a.nd E. R. Vyse. 1978. Heterozygosity
in Yellowstone Park elk, Cervus canadensis. Bio-
cheni. Genet. 16:651-657.
Manlove, M. N., J. C. Avise, H. O. Hillestad, P. R.
Ramsey, M. H. Smith, and D. O. Straney. 1975.
Starch gel electrophoresis for the study of popu-
lation genetics in white-tailed deer. Proc. 19th
Ann. Conf. S. E.. Game and Fish Comm.
29:.392-403.
Xadler, C. F., .\. WooLF, AND K. E. Harris. 1971. The
transferrins and hemoglobins of bighorn sheep
{Otis canadensis), Dall sheep (Oris dalli) and
mouflon (Otis niusimon). Comp. Biochem. Phvs-
iol. 40B: 567-570.
Ryma.n, N., G. Beckman, G. Briun-Petersen, and C.
Reuterwall. 1977. Variability of red cell en-
zymes and genetic implications of management
policies in Scandinavian moose (Alces alces). He-
reditas 85:157-162.
Smith, I. (Ed.). 1968. Chromatographic and electro-
phoretic techniques. Vol. 2. Zone— electro-
phoresis. Wiley, New York, 524 pp.
THE GENUS ERIOGONUM MICHX. (POLYGONACEAE) AND MICHEL GANDOGER
James L. Reveal'
Abstract.— Michel Gandoger, a notorious "splitter," proposed several new entities in the plant genus Eriogonum
(Polvgonaceae) in a 1906 paper published in Belgium. Because he used the term species at two different ranks, in
violation of the International Code of Botanical Nomenclature, manv of his names are invalid. Unlike his papers
published in France, this one was apparentl\- edited so that some names were validly published and some are invalid.
A review of the 1906 Eriogonum paper shows that a majority of both specific and infraspecific entities proposed are
valid, but some names, long in use and assinned to be valid, are, in fact, invalid. Even so, most of his names are
sviionyms. Each name proposed by Gandoger is reviewed and a nomenclatural and taxonomic disposition made. Two
new combinations are made within E. hitcolum Greene, var. caninum (Greene) Reveal and var. pedunrulatum (S.
Stokes) Reveal.
Michel Gandoger (1850-1926), in the
words of Keck (1958), was a "French abbe;
author of 'Flora Europae' (27 vols.); volumi-
nous writer; amasser of a huge herbarium
now at Lyon; a 'splitter' who named thou-
sands of unacceptable species." Gandoger is
also mentioned in the International Code of
Botanical Nomenclature (Stafleu et al. 1978)
as an example of Art. 33.4. This article deals
with the problem of misplaced ranks such as
the use of the term species as a rank within a
species. As noted in the code, Gandoger ap-
plied the term species and used binary no-
menclature for two categories of taxa of con-
secutive rank, the higher rank being
equivalent to that of species in contemporary
literature, while he misapplied the same term
to a lower rank. These latter terms are not
validly published. In his 1906 paper, Gando-
ger used three sets of ranks, the species, spe-
cies of the second order ("speciebus secundi
ordinis"), and a rank which has come to be
identified with the rank of variety (Heller
1907), although Gandoger causally refers to
this latter rank as forms ("formis," "formas
secernendas, " "formae memorabiles," "mem-
orantur sequentes formae," "varieas formas,"
"modo formae," etc.). He used the terms
"varians" and "variabilis" to allude to the
same category. As for the species of the sec-
ond order, in addition to terming them "spe-
ciebus secimdi ordinis," he also used the term
siibspeciebus or subspecies to refer to entities
he then proceeded to treat as binomials and
to designate by the expression sp. n.
The following year. Heller (1907) present-
ed a "compilation" of the Gandoger paper,
noting that he was presenting only "the new
forms described in this paper." Heller himself
had some difficulties with the names. He
mentioned, after Eriogonum aspalathoides (as
it was called bv Heller), that this name was
"perhaps intended as a variety of E. fasci-
ciilatiim, but the way the name is printed
.should indicate a species." Only once does
Heller include Gandoger's terminology allud-
ing to species of a lower rank, this being for
the phrases published under E. sphoerocepha-
him, "Inter formas varias duae sequentes, ut
subspecies, praecipue distingui possunt."
It must be noted that Heller was merely
presenting Gandoger's results, and he cannot
be assumed to have validated any of the oth-
erwise invalid Gandoger names.
Normally, Gandoger published his papers
in the Bulletin de la Societe Botaniqiie de
France, but for Eriogonum his choice was the
Bulletin de la Societe Roijale de Botanique de
Belgique, and, because of this, some of the
names may be validly described. Possibly
'Department of Botany, University ot Maryland. College Park. Maryland 20742, and National Museum of Natural History. Smitfisonian Institution, Wash-
ington, D.C. 20560. Research supported by National Science Foundation Grant BMS75-13063. This is Scientific Article .\2714. Contribution No. 5761 of the
Maryland .\gricultural Experiment Station, Department of Botany.
143
144
Great Basin Naturalist
Vol. 40, No. 2
they may because of the way the editor had
the paper set, thereby removing from the
original manuscript some of Gandoger's ec-
centricities. In reviewing Gandoger's papers
in the French journal, it is clear that the ma-
jority of his names are invalidly published. In
reviewing a few of his articles containing
species from the United States, I failed to
note a single instance when a species name
was validly published. This little known fact,
at least in the United States and perhaps else-
where, means that all the Gandoger names
must be carefully checked before they can be
accepted. This is particularly true for floristic
workers.
Finally, it is sometimes difficult to know
exactly where the various sections of Gando-
ger's paper start and end so that one can de-
termine if the name is validly published or, in
fact, is not because of a sentence presented
several pages before. I have tried to follow
the intent of Gandoger's paper, and hope I
have interpreted each segment correctly.
The following treatment indicates which
names are valid, which are not, and the tax-
onomic status of each name. The full author
citation of each name is given, and the num-
ber following the name is the page on which
it was proposed by Gandoger.
Erio^onum abertianum Torr. in Emory var.
rubenimiim Gandoger, 185, valid, a synonym
of var. abertianum.
Eriogonum aberiianum var. neomexicanum
Gandoger, 185, valid, a synonym of var.
abertianum.
Eriogonum arizonicum Gandoger, non
Stokes ex Jones, 186, valid, a synonym of E.
pharnaceoides Torr. in Emory var. pharna-
ceoides.
Eriogonum alattim Torr. in Sitgr. var. mac-
douglasii (Tandoger, 186, valid, a synonym of
var. mogollonense Stokes ex Jones.
Eriogonum alatum var. brevifolium Gando-
ger, 186, valid, a synonym of var. alatum.
Eriogonum anemophiUim (as anemophijl-
luni) Greene var. cusickii Gandoger, 186,
valid, a synonym of E. cusickii M. E. Jones.
Eriogonum angulosum Benth. var. rectipes
Gandoger, 186, valid, a synonym of E. ma-
cukitum Heller.
Eriogonum angulosum var. patens Gando-
ger, 187, valid, a synonvm of E. maculatum
Heller.
Eriogonum angulosum var. pauciflorum
Gandoger, 187, valid, a synonym of E. ma-
culatum Heller.
Eriogonum angulosum var. flabellatum
Gandoger, 187, valid, a synonym of E. ma-
culatum Heller.
Eriogonum annuum Nutt. var. pauciflorum
Gandoger, 187, valid, a synonym of E. an-
nuum.
Eriogonum hitchcockii Gandoger, 187,
valid, a synonym of E. annuum.
Eriogonum juncinellum Gandoger, 187,
valid, a synonym of E. davidsonii Greene.
Eriogonum salicorniodes Gandoger, 187,
valid, a good species restricted to clay slopes
in southwestern Idaho and adjacent south-
western Oregon. It is most closely related to
E. collinum Stokes ex Jones but most often
confused with E. baileyi S. Wats. Synonyms
of this species include E. demissum S. Stokes
(1936) and the var. romanum S. Stokes.
Eriogonum caespitosum Nutt. var. alijs-
soides Gandoger, 188, valid, a synonym of E.
caespitosum.
Eriogonum nevadense Gandoger, 188,
valid, a synonym of E. ochrocephalum S.
Wats. var. ochrocephalum.
Eriogonum elatum Dougl. ex Benth. var.
limonifolium Gandoger, 188, valid, a syn-
onym of £. elatum var. elatum.
Eriogonum elatum var. erianthum Gando-
ger, 188, valid, a synonym of var. elatum.
Eriogonum fasciculatum Benth. var. oleifo-
lium Gandoger, 189, valid, a synonym of E.
fasciculatum var. fasciculatum.
Eriogonum fasciculatum Benth. var. as-
palathoides Gandoger, 189, valid, a synonym
of var. fasciculatum. As noted by Heller
(1907), there is some problem with this name;
that is, there is an E. prior to the epithet.
However, based upon the rest of the text, this
is clearly as printer's error and should be list-
ed as a variety and not as a species, as was
done by Heller in his "compilation. "
Eriogonum flavum Nutt. in Fras. var. folia-
tum Gandoger, 189, valid, a synonym of E.
jamesii Benth. in DC. va.r. flavescens S. Wats.
Eriogoni4m flavum var. linguifolium Gan-
doger, 189, valid, a synonym of var. flavum.
Eriogonum leucocladum Gandoger, 189,
valid, a synonym of E. baileyi S. Wats. var.
divaricatum (Gandoger) Reveal.
ErioEonum heracleoides Nutt. var. micran-
June 1980
Reveal: Eriogonum Nomenclature
145
thum Gandoger, 189, valid, a synonym of £.
herocleoides var. angustifolium (Niitt.) Torr.
& Gray.
Eriogonum hcracleoides var. viride Gando-
ger, 190, valid, a synonym of £. umhellatum
Torr. var. nevadense Gandoger.
Eriogonum hcracleoides var. multiccps
Gandoger, 190, valid, a synonym of var. hcra-
cleoides.
Eriogonum hcracleoides var. utahensis
Gandoger, 190, valid, a synonvm of var. hcra-
cleoides.
Eriogonum hcracleoides var. rydbergii Gan-
doger, 190, valid, a synonym of var. hcra-
cleoides.
Eriogonum jamesii Benth. in DC. var. sim-
plex Gandoger, 190, valid, a good variety of-
ten included within var. jamesii; this phase of
the species is restricted to southwestern Kan-
sas.
Eriogonum jamesii var. neomexicanum
Gandoger, 190, valid, a synonym of var.
jamesii.
Eriogonum longifolium Nutt. var. long-
idcns Gandoger, 190, valid, a synonym of E.
longifolium var. gnaphalifolium Gandoger.
Eriogonum longifolium var. gnaphalifo-
lium Gandoger, 190, valid, that phase of the
species restricted to Florida; often called E.
floridanum Small.
Eriogonum longifolium var. floridanum (as
floridana) Gandoger, 190, valid, a synonym
of £. longifolium var. gnaphalifolium Gando-
ger.
Eriogonum longifolium var. lindheimeri
Gandoger, 190, valid, the western phase of
the species now best considered a synonvm of
var. longifolium.
Eriogonum longifolium var. caput-fclis
Gandoger, 190, valid, a synonym of var. lon-
gifolium.
Under the heading, Eriogonum micro-
thccum Nutt., Gandoger states "inter quas se-
quentes altem pro subspeciebus habueris"
and refers to the following six entities all
published with binary names.
Eriogonum macdougalii Gandoger, 191, in-
valid, a synonym of E. microthecum Nutt.
var. foliosum (Torr. & Gray) Reveal. The
name was used by Stokes (1936) at the varia-
tal rank within E. microthecum, where she
validated the name as E. microthecum var.
macdougalii S. Stokes.
Eriogonum myrianthum Gandoger, 191, in-
valid, a synonym of E. cffusum Nutt. var. cf-
fusum.
Eriogonum sputhulare Gandoger, 191, in-
valid, a synonym of E. microthecum Nutt.
var. laxiflorum Hook. The name was used by
Stokes (1936) at the variatal rank within E.
microthecum, where she validated the name
as E. microthecum var. spathulure S. Stokes.
Eriogonum intricatum Gandoger, 191, in-
valid, a synonym of E. microthecum Nutt.
var. laxiflorum Hook.
Eriogonum Iwlichrysoides Gandoger, 192,
invalid, a synonym of E. cffusum Nutt. var.
rosmarinioides Benth. in DC. The name was
validly published by Rydberg (1931) and the
citation of the name should be E. helichry-
soides Rydb., Brittonia 1: 87. 1931, without
reference to Gandoger. Rydberg was in-
correct that Gandoger had proposed a varie-
ty, so the combination attributed to Gando-
ger by Rydberg in synonymy, E. nucrothecum
var. helichrysoides, must be attributed to
Rydberg as well.
Eriogonum sarothriforme Gandoger, 192,
invalid. The type of this name, collected by
Osterhout at Glenwood Springs, Garfield Co.,
Colorado, may represent a distinct taxon. It
belongs to the E. brevicaule Nutt. complex
and is seemingly a part of the polymorphic
species E. lonchopln/llum Torr. & Gray, a
taxon that morphologically bridges the E.
brevicaule complex, a group of herbaceous
perennials, with those species typified bv E.
corymbosum Benth. in DC, a series of shrubs
or subshnibs. The Garfield Co. plants tend to
be more slender than typical E. lonchophyl-
lum of southern Colorado and adjacent
northern New Mexico. These plants also tend
to resemble some of the more robust, but yel-
low-flowered, forms of E. brevicaule found in
Rio Blanco Co., Colorado. Although well
known to me, I am still uncertain what to do
with the Glenwood Springs plants.
Eriogonum niveum Dougl. ex Benth. var.
suksdorfii Gandoger, 192, valid, a synonym of
E. niveum.
Eriogonum niveuni var. candelabrum Gan-
doger, 192, valid, a svnonvm of E. niveum.
Eriogonum ochrolcucum Small var. macro-
podum Gandoger, 192, valid, the basionym
for E. ovalifolium Nutt. var. macropodum
(Gandoger) Reveal.
146
Great Basin Naturalist
Vol. 40. No. 2
Eriogonum ochroleucum var. decahans
Gandoger, 192, valid a synonym of E. ocali-
folium Nutt. var. macropodum (Gandoger)
Reveal.
Under the heading of Eriogonum ovalifo-
lium Nutt., Gandoger proposed a series of
names in two ranks, all of which he refers to
by "formas quarum non paucas saltern pro
speciebus secimdi ordinis haberi possimt,"
which I believe he wished to apply only to
those names he indicated by the designation
oi sp. n.
Eriogonum flavissitnum Gandoger, 193, in-
valid, a svTionym of E. ovalifolium Nutt. var.
anserinum (Greene) R. J. Davis. The name
was used by Stokes (1936) as a subspecies of
E. ovalifoliwn, where the name was validated
as E. ovalifolium Nutt. ssp. anserinum S.
Stokes.
Eriogonum cusickii Gandoger, non M. E.
Jones (1903), 193, invalid, a synonym of E.
strictum var. proliferum (Torr. & Gray) Re-
veal. The name was used at the variatal rank
by Stokes (1936) within E. strictum, where
the name was validated as E. strictum Benth.
var. cusickii S. Stokes.
Eriogonum cusickii Gandoger var. califor-
nicum Gandoger, 193, invalid, a synonvm of
E. strictum var. proliferum (Torr. 6c Grav)
Reveal.
Eriogonum ovalifolium Nutt. var. neva-
dense Gandoger, 193, valid, a good varietv of
the species, this being the yellow-flowered,
early flowering expression which I have
called var. multiscapum Gandoger (see be-
low).
Eriogonum ovalifolium var. deltoideum
Gandoger, valid, 193, a synonym of £. ovali-
folium var. nevadense Gandoger.
Eriogonum dichroanthum Gandoger, 193,
invalid, a synonym of E. ovalifolium Nutt.
var. nevadense Gandoger (see discussion un-
der var. multiscapum below).
Eriogonum ovalifolium Nutt. var. utahense
Gandoger, 194, valid, a synonym of var. ova-
lifolium.
Eriogonum ovalifolium Nutt. var. multi-
scapum Gandoger, 194, valid, a synonym of
var. ovalifolium. For several years I have
misapplied this name to the yellow-flowered
phase of the species (Reveal & Munz 196S,
Reveal 1973, 1976). Gandoger based this
name on plants gathered by Nelson (4658) at
Cokeville, Uinta Co., Wyoming, upon which
he also based the name E. dicJiroanthum. An
examination of this collection b\ me in 1966,
and then by my brother, Jon A. Reveal, in
1973, was inconclusive in that it could not be
fully determined if the flowers of var. multi-
scapum were truly yellow. Gandoger divided
the collection and moimted each on a sepa-
rate sheet, with E. dichroanthum having
"flores fructusque flavissimi." and the flowers
of var. midtiscapum, to him. were "'ochro-
leuci." The same collection at KSC is strictly
yellow flowered, but the specimen at RM is a
mixture of a bright-yellowed specimens
matching E. dichroanthum and a whitish or.
at best, pale yellow-flowered specimen that
matches var. multiscapuryi. I now believe that
the yellow-flowered, early spring flowering
phase of the species should be called var.
nevadense and the type of var. lyiultiscapum
assigned to var. ovalifolium.
Eriogonum ovalifolium Nutt. var. cijclo-
phijllum Gandoger, 194, valid, a synonym of
var. macropodum (Gandoger) Reveal. A reex-
amination of the types of var. cyclophyllum
and var. cerastoides (see below), as part of
this study, clearly shows that these names
must be referred to what I (Reveal 1968) had
earlier called var. macropodum. making a
new combination for this name by transfer-
ring it from E. ochroleucum Small to E. ovali-
folium. Previously, these two names have
been referred to var. ovalifolium (Hitchcock
1964). The var. cijclophyUum is close to var.
ovalifolium and may represent one of the
many intermediate expressions between the
two varieties. I retain the usage of var.
macropodum.
Eriogonum ovalifolium Nutt. var. ceras-
toides Gandoger, 194, valid, a synonym of
var. macropodum (Gandoger) Reveal (see the
discussion above).
Eriogonum ruhidum Gandoger. 194, in-
valid, a synonym of E. ovalifolium Nutt. var.
depressimi Blankinship.
Eriogonum ruhidum var. frigidum Gando-
ger, 194, invalid, a synonym of E. ovalifolium
Nutt. var. depressum Blankinship.
Eriogonum roseiflorum Gandoger. 194, in-
valid, a svnonvm of E. ovalifolium Nutt. var.
ovalifolium.
Eriogonum piperi Greene var. ochrocepha-
lum Gandoger, 195, valid, a synonym of E.
June 1980
Reveal: Eriogonum Nomenclature
14'
flaviim \utt. in Fras. var. piperi (Greene) M.
E. Jones.
Eriogonum piperi var. longifloruni Gando-
ger, 195, valid, a synonym of E. flavum Xutt.
in Fras. var. piperi (Greene) M. E. Jones.
Under Eriogonum pohjanthum Benth., now
better known as E. umbellatum Torr. var.
pohjanthum (Benth. in DC.) M. E. Jones,
Gandoger introduces two validly described
species simply stating that "species duae se-
quentes huic sunt affines."
Eriogonum marginale Gandoger, 195,
valid, a synonym of E. umbellatum Torr. var.
aureum (Gandoger) Reveal.
Eriogonum glaherrimum Gandoger, 195.
valid, the basionym of E. umbellatum Torr.
var. glaberrimum (Gandoger) Reveal.
Eriogonum glaberrimum var. aureum Gan-
doger, 195, valid, the basionym of E. um-
bellatum Torr. var. aureum (Gandoger) Re-
veal.
Eriogonum pohjcladon Benth. var. mexi-
canum Gandoger, 196, valid, a synonym of E.
pohjcladon.
Eriogonum polycladon Benth. var. crispum
Gandoger, 196, valid, a synonym of E. poly-
cladon.
Eriogonum racemosum Xutt. var. sagitta-
tum Gandoger, 196, valid, a synonym of E.
racemosum.
Eriogonum racemosum Xutt. var. cordi-
folium Gandoger, 196, valid, a svnonvm of E.
racemosum.
Eriogonum reniforme Torr. & Frem. var.
asarifolium Gandoger, 196, valid, a svnonvm
of E. pusillum Torr. 6f Gray.
Eriogonum praebens Gandoger, 196, valid,
a s\nonym of E. bailcyi S. Wats. var. diva-
ricatum (Gandoger) Reveal.
Eriogonum praebens var. diiaricatum Gan-
doger, 196, valid, the basionym of E. baileyi
S. Wats. var. divaricatum (Gandoger) Reveal.
Under the heading Eriogonum sphaero-
cephalum Dougl. ex Benth.. Gandoger states
"inter formas varias duae sequentes. ut sub-
species, praecipue distingui possunt."
Eriogonum cupreum Gandoger, 196. in-
valid, a synonym of E. umbellatum Torr. var.
umbellatum.
Eriogonum halimioides Gandoger, 197, in-
valid. This name was validated by Stokes
(1936) as E. sphaerocephalum var. hali-
mioides S. Stokes without reference to Gan-
doger.
Eriogonuin subalpinum Greene var. arach-
noideum Gandoger, 197. valid, a synonym of
E. umbellatum Torr. var. dichrocephalum
Gandoger.
Eriogonum subalpinum var. vulcanicum
Gandoger, 197, valid, a synonymy of £. um-
bellatuin Torr. var. majus Hook.
Eriogonum subalpinum var. stenophyllum
Gandoger, 197, valid, a synonym of E. um-
bellatum Torr. var. majus Hook.
Eriogonum subalpinum var. subnivale
Gandoger, 197, valid, a synonym of E. um-
bellatum Torr. var. majus Hook.
Eriogonum tenellum Torr. var. grandi-
florum Gandoger, 197, valid, a synonym of E.
microthecum Nutt. var. laxiflorum Hook.
Eriogonum tenelhw} var. sessiiflorum Gan-
doger, 198, valid, a synonym of E. micro-
thecum Xutt. var. laxiflorum Hook.
Eriogonum tenellum var. erianthum Gan-
doger, 198, valid, a synonym of E. micro-
thecum Xutt. var. ambiguum (M. E. Jones)
Reveal in Mimz.
Eriogonum thurberi Torr. var. parishii Gan-
doger. 198, valid, a svnonvm of £. thurberi.
Eriogonum thurberi var. acutangulum Gan-
doger, 198. valid, a synonvm of E. macula-
turn Heller.
Eriogonum tlnjmoides Benth. in DC. var.
pallens Gandoger, 198, valid, a synonym of
£. thymoides.
Eriogonum umbellatum Torr. var. cran-
dallii Gandoger, 198, valid, a synonym of var.
umbellatum.
Eriogonum umbellatum var. chrysanthum
Gandoger, 198, valid, a synonym of E. um-
bellatum var. stellatum (Benth.) M. E. Jones.
Eriogonum umbellatum var. nevadense
Gandoger, 198, valid, a good variety referr-
ing to that phase of the species found in the
Sierra Xevada of California northward into
Oregon and eastward into uestern Xevada
that has been routinelv called var. umbella-
tum (Reveal & Munz 1968; see Howell 1976).
Eriogonum umbellatum var. cladophorum
Gandoger, 198, valid, a synonym of var. um-
bellatum.
Eriogonum umbellatum var. dichro-
cephalum Gandoger, 199, valid, a good varie-
ty applied to that phase previously called E.
148
Great Basin Naturalist
Vol. 40, No. 2
umbeUatum var. aridum (Greene) C. L.
Hitchc. (Hitchcock 1964).
Eriogonwn umhellatum var. californicuni
Gandoger, 199, valid, a synonym of E. um-
beUatum var. nevadense Gandoger.
Eriogonum vimineiim Dougl. ex Benth.
var. rigescens Gandoger, 199, valid, a synon-
ym of E. vimeneum.
Eriogonum vimineum Dougl. ex Benth.
var. califomicum Gandoger, 199, valid, a syn-
onym of E. luteohim Greene var. caninum
(Greene) Reveal, comb, nov., based on E. vi-
mineum var. caninum Greene, Fl. Francisc.
150. 1891. The type of var. califomicum is
somewhat intermediate between var. luteo-
lum and var. caninum, being closer to the
latter than the former. I have long recog-
nized the caninum phase (Reveal & Munz
1968) as distinct from E. vimineum, but J. T.
Howell, who has considered the expression
only as a variant, has pointed out in our con-
versation that he had observed a large series
of intermediate populations, as had I, which
held the Mt. Tamalpais plant, var. caninum,
well within the boundaries of what he called
E. vimineum. This latter expression, however,
proved to be E. luteohim rather than E. vimi-
neum, and I am now following Howell's
(1970) taxonomic disposition of this local en-
demic. In addition to this variant of E. luteo-
lum, the Sierra Nevada plant I recognized as
a distinct species previously (Reveal 1970)
should be included within this species as well.
Thus, I propose E. luteohim var. peduncula-
tum. (S. Stokes) Reveal, stat. & comb, nov.,
based on E. pedunculatum S. Stokes, Leafl.
W. Bot. 2: 48. 1937.
Eriogonum vimineum var. oregonense Gan-
doger, 199, valid, a synonym of E. vimineum.
Eriogonum restioioidcs Gandoger, 199,
valid, a synonym of E. baileyi S. Wats. var.
bailey i.
Literature Cited
Gandoger, M. 1906. Le genre Eriogonum (Poly-
gonaceae). Bull. Soc. Roy. Bot. Belgique 42:
18.3-200.
Heller, A. A. 1907. Compilations. Miihlenbergia 3:
a3-96.
Hitchcock, C. L. 1964. Eriogonum. Univ. Wash. Publ.
Biol. 17(2): 104-138.
Howell, J. T. 1970. Marin flora. 2d ed. Berkeley: Univ.
California Press. 366 pp.
1976. Eriogonum notes VII. Mentzelia 1: 17-22.
Keck, D. D. 1959. "Abbreviations of authors' names."
Pages 1551-1576 in P. A. Munz, & D. D. Keck,
eds. A California flora. Berkeley: Univ. California
Press.
Reveal, J. L. 1968. Some nomenclatural changes in
Eriogonum (Polygonaceae). Taxon 17: 531-533.
1973. Eriogonum (Polvgonaceae) of Utah. Phvto-
logia 25: 169-217.
1976. Eriogonum (Polvgonaceae) of Arizona and
New Mexico. Phytologia 34: 409-484.
Reveal, J. L., and P. A. Munz. 1968. "Eriogonum."
Pages 33-72 in P. A. Munz, ed. Supplement to A
California flora. Berkeley: Univ. California Press.
Rydberg, p. a. 19.31. Taxonomic notes on the flora of
the prairies and plains of central North .\merica.
Brittonia 1: 79-104.
Stafleu, F. a., et al. 1978. International code of bot-
anical nomenclatme. Regnum Veg. 97: 1-457.
Stokes, S. G. 1936. The genus Eriogonum, a preliminary
studv based on geographic distribution. San Fran-
cisco: J. B. Nebiett. 124 pp.
PARASITES FROM TWO SPECIES OF SUCKERS (CATOSTOMIDAE)
FROM SOUTHERN UTAH
J. Craig Biieiiholt' and Richard A. Heckmann
.\bstr\c:t.— Twenty Ciitostoiims latipiiiuis and 50 Catostomiis lUscohohs from La Verkin Creek and the Fremont
River in southern Utah were collected and surveyed tor parasites. Data from the survey indicated that 83 percent of
the fish were infected with at least one parasite, with the fish from La Verkin Creek harboring more parasites.
Twelve genera and 12 species of parasites were identified from these fish. .\ monogenetic trematode, G\jwd(trtiilus
dedans, which was found in 90 percent of the fish, was the most common parasite. Comments are included on habi-
tat and host variations for the parasitofauna from suckers taken from the two locations.
A survey of the parasites of the cato,sto-
mids, Catostomiis latipinnis and C. discoboUs,
was conducted at La Verkin Creek, southern
Utah, and the Fremont River near Hanks-
ville, Utah. The objectives of this survey were
to provide a hst of parasites for C. discoboUs
and C. latipinnis in La Verkin Creek and
Fremont River and to correlate water param-
eters and benthos from these streams with
parasite loads. Both streams contain well-e.s-
tablished populations of the listed suckers. An
exhaustive survey of parasites can explain the
source or reservoir of serious pathogens for
endangered species and commercially impor-
tant fish.
Catostomids are found exclusively in North
America, excluding two or three Asiatic spe-
cies (Pflieger 1975). Catostomiis discoboUs is
found in Idaho, Utah, and Nevada in the fol-
lowing drainages: Colorado River above the
Grand Canyon, upper Snake River, Bear Riv-
er, and Weber Lake outflows. Catostomiis
latipinnis is unique to the Colorado River
drainage (Eddy 1959). Information con-
cerning the life history of these two .species is
limited. Catostomid levels in both study areas
for this project are maintained by resident
sucker populations.
Both of the streams selected in this survey
are unstable desert streams. Much of the sub-
strate is sand which shifts and prevents deep
pools from forming. Flash floods can disrupt
and completely change the nature of the
streams and change the macroinvertebrate
population. Because of this, fish species, e.g.,
salmonids and centrarchids, that cannot with-
stand the instability of the stream and the
consequent change in macroinvertebrate food
source are not found extensively in these two
streams. The ichthyofauna found in the study
area of the Fremont River are: Longnose
dace, RJiinichthys cataractae; speckled dace,
R. osciiliis; leatherside chub, Gila copei; blue-
head sucker, C. discoboUs; and flannelmouth
sucker, C. latipinnis (Heckmann 1976). The
speckled dace and leatherside chub are omni-
vores that feed on aquatic plants, insects, and
cmstaceans. The flannelmouth suckers are
herbivores which feed on algae, diatoms,
parts of higher plants, and seeds. The blue-
head sucker is a bottom feeder which scrapes
algae and other organisms from rocks (Sigler
and Miller 1963). The largest fish found in
the Fremont River is the flannelmouth sucker
and the smallest is the speckled dace. None
of the fish found in the sample area are pisci-
vorous. All fish feed either on aquatic in-
vertebrates or plant material. Fi,sh predators
mav include birds and small mammals.
Fish .species inhabiting the survey site at
La Verkin Creek are: speckled dace, R. os-
cuUis; Virgin River spinedace, Lcpidomeda
mollispinis; red shiner, \otropiis liitrcnsis;
woundfin minnow, Plagopterus argentissimus;
'Department of Zoolog\', Brigham Young University, Provo, Utah 84602.
149
150
Great Basln Naturalist
Vol. 40, No. 2
flannelmouth sucker, C. kitipinnis; bluehead
sucker, C. discoholis or desert sucker, C.
clarki; and rainbow trout, Salmo gairdneri
(Winget and Baumann 1977). La Verkin
Creek, in comparison to the Fremont River,
is more stable and less turbid, resulting in the
presence of riffles, pools, and some holes
1-1.5 m deep. Because of the difference in
stream conditions, rainbow trout are planted
by the Utah Division of Wildlife Resources in
small numbers. Two species of fish, L. moUis-
pinis and P. argentissimus, are considered en-
dangered. The woimdfin minnow, rainbow
trout, and Virgin River spinedace are consid-
ered carnivores feeding mainly on in-
vertebrates. Catostoriius discoholis and N. lut-
rensis are considered bottom-dredging
detritovores. Catostoinus kitipinnis and R. os-
culus are selective omnivores (Winget and
Baumann, 1977). The top carnivore in a tro-
phic scheme would be S. gairdneri because it
may feed on smaller fish. Direct competition
"s virtually eliminated because those species
vith similar feeding habits have different
habitat preferences or specific food prefer-
ences (Winget and Baumann 1977).
Parasites of catostomids other than C. lati-
pinnis and C. discoholis have been studied by
researchers in the United States and Canada.
Hoffman (1967) lists known parasites for 12
species of catostomids. Other surveys have
been conducted bv Voth and Larson (1968),
Amin (1969), Threlfall and Hanek (1970),
Amin (1974), White (1974), Mackiewicz
(1963), Price and Arai (1967), Dechtiar
(1969), Daly and De Giusti (1971), Clifford
and Facciani (1972), Hatha wav and Herlev-
ich (1973), Schell (1974), andHayunga and
Grey (1976). The most widely studied ca-
tostomids are white suckers, C. cornmersoni,
and longnose suckers, C. catostomiis. These
surveys deal primarily with metazoan para-
sites, and little information concerning the
protozoan parasites is included.
M.\TERIALS AND METHODS
Through the use of electrofishing, 18
flannelmouth suckers and 40 i)luehead suck-
ers were collected from La Verkin Creek
near the Toquerville cemeterv, southern
Utah. Two flannel-mouth suckers and 10
bluehead suckers were collected from the
Fremont River one mile west of Capitol Reef
National Park, near Hanksville, Utah. The
fish were transported to Brigham Young Uni-
versity in iced holding tanks. Limited num-
bers of fish were obtained due to collecting
restrictions.
Each fish was checked for parasites. The
suckers were euthanized by a blow to the
head before being weighed and measured
(Table 1). Following macroscopic exam-
ination, scrapings of the surface, gills, medial
area of the opercula, and eyes, were exam-
ined for parasites. Because the blood vessels
were niptiued, gill scrapings were used to
check for hemoflagellates. Intestine, liver,
and gall bladder were excised and examined
for endoparasites. The presence of metacer-
cariae inhabiting the liver was checked by
pressing a piece of the organ between two
glass slides and examining it without magnifi-
cation.
Protozoans were either air dried or pre-
served in 10 percent formalin. Permanent
preparations of monogenetic trematodes
were made with Turtox mounting and stain-
ing medium (nonresinous stain mountant
CMC-S). Leeches were also fixed in formalin
and all were identified tlirough the use of
keys listed in Hoffman (1967).
Cestodes were placed directly into AFA
fixative to prevent total relaxation. Digenetic
trematodes were placed in 95 C water to
promote relaxation and then placed in AFA
fixative. Cestodes and digenetic trematodes
were stained with Semichon's carmine for 12
hours and then destained in changes of acid
alcohol to improve color contrast. After des-
taining, the specimens were dehydrated in 95
percent and 100 percent ethyl alcohol for
one hoiu' each. Once dehydrated, specimens
were cleared in xylene and then mounted
with Permount on glass slides. Morphological
characteristics given in Hoffman (1967) were
used for identification of trematodes and
nematodes. Preliminary identification of the
caryophyllid tapeworms was confirmed by
John S. Mackiewicz (State University of New
York at Albany).
Water chemistry and macroinvertebrate
data were obtained from studies by Heck-
mann (1976), Winget and Reichert (1976),
and Winget and Baumann (1977).
June 1980
Breinholt, Heckmann: Fish Parasites
151
Results
Data from the examination of 40 bluehead
and 18 flannelmouth suckers from La Verkin
Creek in southern Utah indicated that 55
suckers harbored at least one species of para-
site. Thirty-seven of 40 bluehead suckers
were infected and all 18 flannelmouth suck-
ers harbored parasites (Table 2). Postmortem
examination of 12 suckers from the Fremont
River revealed one of 10 bluehead suckers
and 2 of 2 flannelmouth suckers were para-
sitized (Table 2).
Twelve genera and 12 species of parasites
were identified from fish from La Verkin
Creek. The most frequently encountered par-
asite was Gyrodactylus elegans, a mon-
ogenetic trematode which was found in 52
Table 1. Weights and measurements of fish exam-
ined.
Species
Weight
Length
No.
offish
Stream
(gm)
(TL) (cm)
1
C. discobolis
La Verkin Creek
45
17
2
43
14
3
38
16
4
33
15
5
33
14
6
55
18
7
65
19
8
50
16
9
40
16
10
25
13
11
35
16
12
50
16
13
60
18
14
85
21
15
90
21
16
45
17
17
20
13
18
40
16
19
40
16
20
40
17
21
15
11
22
45
17
23
30
14
24
35
15
25
25
13
26
55
17
27
20
13
28
35
Ifi
29
30
Hi
30
45
17
31
20
13
.32
25
14
33
25
14
34
30
15
35
25
15
(90 percent) of the fish. Other monogenetic
trematodes recovered were Octomacrwn lan-
ceatinn, found in one (2 percent) fish, and
PcUucidhaptor alahauius, foimd in six (10
percent) of the fish. Metacercariae of two
digenetic trematodes were also recovered.
Neascus sp. was found in 25 (43 percent) fish
and Clinostomum marginatum was found in
two (3 percent) fish. Cystidicola sp., a nema-
tode, was found in one (2 percent) fish and
Monobothrium hunteri and Isoglaridacris hex-
acotyle, both caryophyllid cestodes, were
found in 29 (50 percent) and 28 (48 percent)
fish, respectively. Three protozoans were re-
covered from the fish. Myxosoma sp. was
found in 11 (19 percent) fish, Myxidiiim sp. in
four (7 percent), and Tridiondina sp. was
found in 20 (34 percent) of the fish examined
(Table 3).
Table 1 continued.
No.
Species
of fish
Weight
Stream (gm)
Length
(TL) (cm)
36 C. discobolis
37
38
.39
40
41 C. latipinnis
42
43
44
45
46
47
48
49
.50
51
52
53
54
55
56
57 "
58
59 C discobolis
60
61
62
63
64
65
66
67
68
69 C. latipinni.s
70
La Verkin Creek 30 15
45 17
15 11
20 13
20 13
115 24
180 27
185 27
230 30
205 29
260 32
210 30
125 25
185 29
260 31
225 24
110 24
195 23
130 25
170 28
125 24
105 24
85 22
Fremont Hiver 20 14
10 11
35 16
35 17
20 14
30 16
5 10
25 15
10 11
30 15
5.30 40
350 37
152
Great Basin Naturalist
Vol. 40, No. 2
Differences in protozoan parasite load
from the two species of fish taken from La
Verkin Creek are as follows: Myxosoma sp..
25 percent bluehead suckers and 6 percent
flannelmouth suckers; Myxidium sp., 0 per-
cent bluehead suckers and 22 percent
flannelmouth suckers; Trichodina sp., 80 per-
cent bluehead suckers and 22 percent
flannelmouth suckers were infected. For the
metazoan parasites, G. elegans was fomid in
90 percent of the bluehead suckers and 89
percent of the flannelmouth suckers, Octoma-
cntm lanceatum and P. alahamus were found
exclusively on bluehead suckers, 3 percent
and 6 percent, respectively. Neascus sp. was
found on 28 percent of the bluehead suckers
and 78 percent of the flannelmouth suckers,
but C. marginatum was found only in 11 per-
cent of the flannelmouth suckers. The
flannelmouth suckers have a higher incidence
of both species of caryophyllid cestodes.
MonobotJirium hunteri was found in 38 per-
cent of the bluehead suckers and 78 percent
of the flannelmouth suckers, and /. hexacotyJe
was found in 33 percent of the bluehead
suckers and 83 percent of the flannelmouth
suckers. The nematode Cystidicola sp. and
the leech Piscicola sp. were symbiotic exclusi-
vely to the bluehead suckers. Three percent
of the fish were infected with each of these
two parasites.
One species of parasite was recovered from
T.\BLE 2. Number and percentage of bluehead and flannelmouth suckers parasitized from La Verkin Creek and
the Fremont River.
Host species
Total fish
sampled
Total fish
parasitized
La Verkin Creek
fish parasitized*
37 ( 93%)
18 (100%)
.55 ( 95%)
Fremont River
fish parasitized^
Bluehead sucker
Flannelmouth sucker
Total
50
20
38 ( 76%)
20 (100%)
58 ( 83%)
1 ( 10%)
2 (100%)
3 ( 2,5%)
*58 fish examined from La Verkin Creek (40 bluehead and 18 flannelmouth suckers).
''12 fish examined from the Fremont River 1 10 bluehead and 2 flannelmouth suckers).
Table 3. Parasites identified from 58 suckers from La Verkin Creek.
Parasite species
Number and
percentage of
Species
of fish
fish positive
Bluehead
Flannelmouth
11 (19)
10 (25%)
1 ( 6%)
4( 7)
0 ( 0%)
4 (22%)
20 (34)
16 (80%)
4 (22%)
52 (90)
.36 (90%)
16 (89%)
1( 2)
1 ( .3%)
0 ( 0%)
6(10)
6 (15%)
0( 0%)
25 (43)
11 (28%)
14 (78%)
2 ( 3)
0( 0%)
2(11%)
29(50)
15 (38%)
14 (78%)
28 (48)
13 (3.3%)
15 (8,3%)
1( 2)
1 ( 3%)
1 ( 0%)
1( 2)
1 ( 3%)
0 ( 0%)
Protozoans
Myxosoma sp.
Myxiditnn sp.
Trichodina sp.
Trematodes
Gijrodactijlufi elegans
Octomacrum lanceatutn
Pellucidhaptor alahamus
Postodiphstom am mit^ im iim
Clinostomiim marginatum
Cestodes
Monobothritim hunteri
Isoglaridacris hexacotyle
Nematodes
Cystidicola sp.
Leeches
Piscicola sp.
June 1980
Breinholt, Heckmann: Fish Parasites
153
the fish examined from the Fremont River
(Table 4). Gyrodactyhis elegans was identi-
fied in 25 percent of the fish examined. Ten
percent of the bluehead suckers were in-
fected with this inonogentic trematode, and
100 percent of the flannelmouth suckers
were infected.
Table 5 lists the preferred tissue in the host
for each parasite. These parasites were found
in onlv five areas of the fish. Eight of the par-
asite species were fomid on the external sur-
face and only four were found in more than
one area.
Discussion
There are habitat and host variations for
the parasitofauna from suckers taken from La
V'erkin Creek and Fremont River. The ca-
tostomids from La V'erkin Creek were more
heavily parasitized, both in the number of
fish infected and in the number of species en-
countered, than were the fish from Fremont
River. Explanation for these differences may
be attributed to many factors, such as water
qualitv and macroinvertebrates. Water chem-
istr\ for the two streams was found to be sim-
ilar except during spring rvmoff.
Oligocheates, which usually act as the in-
termediate hosts for caryophyllid tapeworms
(Mackiewicz 1972), occur in both streams.
Recent studies listed 14,203 (Winget and
Baumann 1977) and 882 (Heckmann 1976)
oligocheates per m^ in La Verkin Creek and
Fremont River, respectivelv, where the fish
for this studv were obtained. The number of
oligocheates should not have caused the dif-
ference in parasite load because infected
worms would have been ingested by fish
from both streams. Milbrink (1975) correlates
Table 4. Parasites identified from 12 suckers from the Fremont River.
Number and
percentage of
fish positive
Species
offish
Parasite species
Bluehead
suckers
Flannelmouth
suckers
Trematodes
Gyrodactyhis elegans
3 (25)
1 (10%)
2 (100%)
Table .5. Location in host of parasites found in fish from La Verkin Creek and the Fremont River.
Parasite species
Surface
Gills
Operculimi Intestine Gall bladder
Protozoans
Mijxosotna sp.
MyxidiuDi sp.
Trichodiud sp.
Trematodes
Gyrodactyhis ch-gans
Octomacrum hmccatiiiu
PcUucidJiaptor ahihamus
Chnostomuin mart^inatiiui
Cestodes
Monohothriuin hunteri
Isogla ridac ris h cxacoty Ic
Nematodes
Cystidicola sp.
Leeches
Piscicola sp.
= present
o = not present
154
Great Basin Naturalist
Vol. 40, No. 2
the caryophyllid worm burden of fish with
the number of infective oligocheates con-
sumed. If the Fremont River contained in-
fected oHgocheates, some of the fish sampled
should have been infected.
The geographical location of the two
streams may have caused the difference in
parasite load. Parasites can be found in one
area but not in another even though both
have similar aquatic characteristics. Myxo-
soma cerebralis, a myxosporidan parasite
which caused whirling disease in trout, has
been reported in eight states (American Fish-
eries Society, 1974). Whirling disease has not
spread to the other states even though suit-
able habitats exist. Diplostomum spathacewn,
the eye fluke of fish, has been reported in
some areas of Utah but not others (Palmieri,
Heckmann, and Evans 1976).
Most parasites have some effect on the
health of the host (Olsen 1974). The fish sam-
pled from the Fremont River were infected
with only one species of parasite, Gijro-
dactylus elegans, and the incidence of that
parasite was low in comparison to infected
fish from La Verkin Creek. The most com-
mon parasite found on the fish from La Ver-
kin Creek is G. elegans. This organism was on
the surface and occasionally in gill scrapings.
Large numbers of G. elegans can cause dam-
age to the fish by physical blockage of the
gill surface, thus interfering with the gas ex-
change area (Hoffman 1967). Other mon-
ogenetic trematodes, Octomacrum lanceatum
and Pellitcidhaptor alahamiis, and the pro-
tozoan, Trichodina sp., are capable of causing
similar problems in the fish. These parasites
were not foimd in great enough quantities to
pose a threat at the present time. The two
myxosporidans, Myxosonia sp. and Myxidium
sp., are capable of encysting and destroying
tissue. However, no cysts were found. Thus,
it is assumed that these myxosporidans are
causing little damage to the fish.
The fact that no hemoflagellates were
found dining the course of this study does not
disprove their existence in these fish, because
some of these parasites have seasonal fluctua-
tion.
Only one nematode, Cystidicola sp., was
recovered from all fish examined in the sur-
vey. This round worm is not detrimental to
the fish unless it is found in high numbers
(Hoffman 1967). The other intestinal hel-
minths, Monobothrium hunteri and Isoglari-
dacris hexacotyle, are adult cestodes that usu-
ally cause little damage to the definitive host.
High numbers (200 plus) result in mechanical
blockage or cause nutritional deficiencies
(Mackiewicz 1972). The adult worms adhere
to the intestinal lining by means of suckers.
There is little intestinal damage by individual
cestodes because the scolex is unarmed.
The metacercariae of Neascus sp. and Cli-
nostomwn marginatum were recovered from
suckers from La Verkin Creek. For these
trematodes, the metacercarial stage is usually
encysted in the second intermediate host and
does not cause damage through migration
(Hoffman 1967). Unless the metacercaria is
encysted in vital organs, such as a parasite in
the eye lens (D. spathaceum), it does not
present a pathogenic health problem to the
fish. (Neascus sp. was observed encysted in
the fins and gills. The cysts found in the gills
were not numerous and did not appear to in-
terfere with gas exchange for the fish.
The leech, Piscicola sp., did not present a
current problem to the fishing that di^ly one
specimen was recovered. Leeches are period-
ic feeders and should not attach permanently
to the host. The major problem with leeches
on fish is due to large numbers on one host or
tlie transmitting by hemoflagellates (l:)lood
parasites) (Hoffman 1967). No blood parasites
were found during this survey.
None of the fish expired during the journey
from their natural habitat, approximately 400
km, to holding tanks. Thus, it is assumed that
the effects of all the parasites on the suckers
were not evident when the fish were placed
under stress of capture and transportation.
Limited host specificity is demonstrated by
the parasites recovered in this survey. Most
of the parasites encountered have been re-
ported in other species of fish (Hoffman
1967). Species of Myxidium, Myxosoma,
Trichodina, Cystidicola, and Piscicola have all
been reported in trout. The parasitic species
found in salmonids may be different than
those found in the suckers. Gyrodactylus ele-
gans and P. minimum have also been report-
ed in salmonids. Octonuurum lanceatum has
been reported in the catostomids, Catostomus
teres, C. commersoni, C. macrocheilus, and
Erimyzon secetta, the cyprinids, Mylocheilus
June 1980
Breinholt, Heckmann: Fish Pahasites
155
caurinus (peamouth), and Notropis corntitits
(common shiner). FclhicitUuiptor dldluiiniis
has been reported in Ictiohus htihalus, the
smalhnoiith buffalo (Chien and Rogers 1970).
The caryophyllid tapeworms. A/, hunteri and
/. hcxncoti/lc, common to C. discobolis and C.
Uitipinnis, have both been reported from oth-
er catostomids (Hoffman 1967). Because the
parasites recovered in this study have been
reported in other species of fish, it is possible
that these parasites may infect game fish or
commerciallv cultured fish. In the case of the
listed digenetic trematodes, infected birds
can fly from one body of water to another
and "seed" other streams and ponds. Also,
currents can carrv infected fish and other in-
termediate hosts downstream to contaminate
the lower drainage system. Thus, potential
infections of other fish in the same stream
could threaten endangered species such as
the woundfin minnow and the Virgin River
spinedace, which are also found in La Verkin
Creek.
The identity of one of the hosts from La
Verkin Creek is doubtful. Originally, it was
classified as a chiselmouth sucker (Sigler and
Miller 1963). Later studies considered this ca-
tostomid a desert sucker (C. clarki), which is
still a valid species (Bailey et al. 1970). Then,
with the taxonomic revision of some of the
members of the catostomid family, Pan-
tosteus delphinius, the bluehead sucker, and
P. virescens, the green sucker, were combined
to form C. discobolis (Bailey et al. 1970). Af-
ter this revision some investigators have con-
sidered the fish as C. discobolis. Because of
the anatomical similarities between C. dis-
cobolis and C. clarki and the activities and
feeding habits, the two could be considered
similar. It may also be concluded that they
could harbor similar parasites even if they
are two distinct species.
Of the two suckers, C. latipinnis is more
selective in its feeding habits than is C. dis-
cobolis. Winget and Baumann (1977) report-
ed stomach contents of the flannelmouth
sucker contained seeds, identifiable plant
matter, and dipteran larvae; stomachs of the
bluehead sucker contained detritus, uniden-
tifiable plant matter, and very few macroin-
vertebrates. The difference in feeding habits
of these two fish is probably the reason for
the difference in resident parasite species.
The parasites that these two fish have in
common may be due to the ingestion of a
common intermediate host. Even though C.
latipinnis is more selective in its feeding
habits, it still would ingest detritus and other
material due to its feeding technique.
The parasites identified in this survey are
not unexpected. Although no parasitic sur-
veys of C. discobolis and C. latipinnis have
been reported, Mi/xosorna sp., Myxidiian sp.,
Trichodina sp., G. elegans, O. lanceatum, P.
alaJianius, P. mininiitm, C. marginatum, I.
Jiexacotyle, M. hunteri, Cystidicola sp., and
Piscicola sp. have all been reported from
suckers (Hoffman 1967).
Literature Cited
American Fisheries Society. 1974. Suggested procedures
for the detection of certain infectious diseases of
fishes. U.S. Department of the Interior, Washing-
ton. D.C.
Amin, O. M. 1969. Helminth fauna of suckers (Catosto-
midae) of the Gila River system, Arizona. II. Five
parasites from Ccito.'itomus spp. Amer. Midi. \at.
82(2): 429-443.
1974. Intestinal helminths of the white sucker,
Catostomus coninicrsoni (lacepede) in southeast
Wisconsin. Proc. Helminthol. Soc. Wash.
41(l):81-88.
Bailey, R. M., J. E. Fitch, E. S. Herald, E. A.
Lachner, C. C Lindsey, C. R. Robins, and W.
B. Scott. 1970. .\ list of common and scientific
names of fishes from the United States and Clan-
ada. .\merican Fisheries Society Special Pub-
lication No. 6. Washington, D.C.
Chien, S. M., and W. Rogers. 1970. Four new species
of monogenetic trematodes. genus PelliicicUuiptor.
ftoin fishes of the southeast United States. J. Par-
asitol. 56(3):480-485.
Clifford, T. S., and S. Facciani. 1972. Philoinetra
nodulosu in Wvoming white suckers. Prog. Fi.sh-
Cult. .34(4):23.5-2.36. '
Daly, J. J., a.nd D. L. De(;ivsti. 1971. Tn/pdnosoiiia ca-
tosiomi n. sp. from the white sucker Cdtostoiiiiis
cominersoni. J. Protozool. 18(.3):414-417.
Dechtiar, .\. O. 1969. Two new species of monogenetic
trematodes (Treniatoda: Monogcnca) from nasal
cavities of catostomid fishes. J. Fish. Wvs. Board
Can. 26(4):86.5-869.
Eddy, S. 1969. The freshwater fishes. Win. C Brown C.
Puhl., Dubuque, Iowa.
IL\THAWAY, R. p., and J. C. Herlevich. 1973. Gyro-
dactyhis stahlcri sp. n. with new host and locality
records for species of Giircddiliihis. J. Parasitol.
.59(2):801-802.
Hayunga, E. C, and .\. J. Grey. 1976. Cystobracluis
meyeri sp. n. (Hirudinea: Piscicolidae) from Ca-
tostomtis commersoni Lacepede in North Ameri-
ca. J. Parasitol. 62(4):621-627.
156
Great Basin Naturalist
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Heckmann. R. a. 1976. Aquatic habitat evaluation of
the Fremont, Muddy, and Dirt\ Devil rivers and
Caine Springs and Pleasant Creek. Westinghouse
Corporation contract .\o. P. O. ESD- 151-76.
HoFFM.w. G. L. 1967. Parasites of North American
freshwater fishes. University of California Press,
Berkeley and Los .\ngeles.
Mackiewicz, j. S. 1963. Monobothhum httntcri sp. n.
(Cestoidea: Carvoph\llaeidae) from Catustomus
commersoni (Lacepede) (Pisces: Catostomidae) in
North America. J. Parasitol. 49(.5):723-7.30.
1972. Caryophvllidea (Cestoidea): \ review. Exp.
Parasitol. 31(3)':417-512.
MiLBRi.NK, G. 1975. Population biolog> of the cestode
Canjophyllaeus laticeps (Pallas) in bream. Ahra-
mis brama (L.), and the feeding of fish on oligo-
cheates. Institute of Freshwater Research Report
No. 54:36-51.
Olsen, O. W. 1974. .\nimal parasites: Their life cycles
and ecologN . University Park Press, Baltimore.
Palmieri, J. R., R. .\. Heckm.\.xn. and R. S. Evans. 1976.
Life cvcle and incidence of Diplostomum spatho-
ceiim Rudolphi (1819) (Trematoda: Diplostoma-
tidae) in Utah. Great Basin Nat. 36(l):86-96.
Pfliecer. W. L. 1975. The fishes of .Missouri. Missouri
Department of Conservation. Jefferson City.
Price, C. E., .and H. P. .\r.\i. 1967. The monogenean
parasites of Canadian freshwater fishes. Canadian
J. Zool. 45(6/2)123.5-1245.
ScHELL, S. C. 1974. Two new genera and three new^ spe-
cies of allocreadiidae trematodes (Digenea; .\1-
locreadiidae) from freshwater fishes. J. Parasitol.
60(2):24.3-246.
Sigler, W. F., .a.\d R. R. Miller. 1963. Fishes of Utah.
Utah State Department of Fish and Game. Salt
Lake City.
Threlfall, W., a.nd G. Hanek. 1970. Metazoan para-
sites, excluding monogenea, from longnose and
white suckers. J. Fish. Res. Board Can.
27(7): 1317-1319.
N'oTH, D. R., .\ND O. R. Larson. 1968. Metazoan para-
sites of some fi.shes from Goose River, North Da-
kota. Amer. Midi. Nat. 79(l):216-224.
White. G. E. 1974. Parasites of the common white suck-
er (Catastomiis commersoni) from the Kentuckv
River drainage. Trans. Amer. .Micros. Soc.
93(2):280-282.
WiNGET, R. N., AND R. W. Bal'm.a.nn. 1977. Virgin River,
Utah-.\rizona-Nevada, aquatic habitat, fisheries,
and macroinvertebrate studies. Center for Health
and Environmental Studies, Brigham Young Uni-
versitv, L'tah.
WiNGET, R. -M., A.ND .M. K. Reichert. 1976. .\quatic
habitat inventory in the hot desert eis area, Utah.
U.S. Bur. Land Man., Contract No. YA-512-CT6-
77.
SOIL WATER WITHDRWVAL AND ROOT DISTRIBUTION UNDER
GRUBBED, SPRAYED, AND UNDISTURBED BIG SAGEBRUSH VEGETATION
David L. Stiirges'
.Vbstract.— Seasonal depletion by vegetation where sagebrush was selecti\elv removed bv gnibbing and where
sagebrush was sprayed with 2,4-D was 33 and 12 percent less, respectively, than that for undisturbed big sagebrush
vegetation in the surface 122 cm of soil. Differences were located primarilv below 61 cm in vegetation gnibbcd the
previous fall and below 91 cm in vegetation sprayed three years previously. Total root weights under gnibbed and
sprayed vegetation were 29 and 16 percent less, respectively, than for unclisturbed big sagebrush vegetation. Total
herbaceous production by grubbed and sprayed vegetation was 69 and 43 percent less, respectiveh, than production
l)\ undisturbed vegetation.
Big sagebrush [Artemisia trident at a) is
commonly controlled with herbicides, me-
chanical methods, or fire to increase livestock
forage production. Pheno.xy herbicides such
as 2,4-D damage forbs as well as sagebrush,
so that the net effect of spraying is to favor
grass productivity.- Burning or mechanical
sagebrush control techniques, however, do
not selectively favor grasses. Herbaceous pro-
duction commonly doubles or triples bv the
.second or third year after sagebnish removal.
The shift from a shrub to a herbaceous-
dominated vegetation produces other ecolog-
ic and hydrologic changes. This studv was
made to quantify differences in the soil water
regime and in root biomass between undis-
turbed big sagebnish vegetation and (a) her-
baceous vegetation three years after spraying
with 2,4-D and (b) herbaceous vegetation
from which only big sagebnish was removed
by mechanical means the previous fall. Infor-
mation about herbaceous productivity was
also collected.
LlTER.\TL RE ReVIEW
Ghanges in the soil water regime after
sagebrush control are strongly influenced by
rooting characteristics of sagebnish and her-
baceous species. Roots of basin big sagebnish
(A. t. sub. tridcntata) and mountain big sage-
brush (A. t. vaseyana) commonly extend
about 2 m deep and have a maximum lateral
spread from the trunk of 1.5 m (Goodwin
1956, Cook and Lewis 1963, Tabler 1964,
Hull and Klomp 1974, Sturges and Trlica
1978). Most roots are in surface soil where
maximum spread occurs. About 60 percent of
total root length (Tabler 1964) and 85 per-
cent of total root system weight were present
in the surface 61 cm of soil, with only about
4 percent in soil below 91 cm (Sturges and
Trlica 1978).
The principal soil water reservoir utilized
by isolated mountain big sagebnish plants ex-
tended 0.9 m laterally from the trunks and
0.9 m deep (Sturges 1977b). Tlie plants utiliz-
ed water from surface soil adjacent to the
trunk early in the growing season, but use-
zones shifted outward and downward later in
the summer as water adjacent to the trunk
was depleted. .Appreciable water uptake was
detectable until early in .\ugust.
Tabler (1968) and Sturges (1977a) found
that seasonal soil water withdrawal was re-
duced after spraying sagebnish vegetation
'Rocky Mountain Forest and Range Experiment Station, Laramie. Wyoming 82070. Central headquarters is at Fort Collins in cooperation with Colorado
State University; research reported here was conducted at the station's Research Work Unit at Laramie, in cooperation with the University of Wyoming.
Portions of the research were supported by the Bureau of l^nd Management. U.S. Department of the Interior.
This article reports research involving pesticides. It neither contains recommendations for their use nor implies that the uses discussed here have been
registered. .\ll uses of pesticides must be registered by appropriate state and/or federal agencies before they can be recommended. Use all pesticides selecti-
vely and carefullv, read and follow the directions on the label.
157
158
Great Basin Naturalist
Vol. 40, No. 2
with 2,4-D on sites with deep soils that were
fully recharged by snowmelt. This reduction
was located almost entirely below 91 cm as
depletion -of surface soil water by sprayed
vegetation sometimes exceeded depletion by
untreated vegetation. Water depletion in sur-
face soil increased the first few years after
treatment as herbaceous vegetation respond-
ed to release from sagebrush competition.
Other studies also detected an increasing
moisture draft from surface soil with time
(Hyder and Sneva 1956, Cook and Lewis
1963, Shown et al. 1972).
Herbaceous production was measured in
most soil moisture studies. Grass production
doubled the year after spraying mountain big
sagebrush and was 2.6 times higher than un-
treated vegetation three years after treat-
ment (Sturges 1977a). Shown et al. (1972)
found that usable forage production in-
creased 300 percent compared to pre-
treatment conditions once a planted grass
stand became established. Hyder and Sneva
(1956) found the increase in grass production
to be the same whether big sagebrush was
controlled by spraying or by grubbing. Total
herbaceous production increased the most
where sagebrush was grubbed, because forbs
were damaged by the spray.
Study Area
The study was performed at the Stratton
Sagebnish Hydrology Study area 29 km west
of Saratoga in south-central Wyoming. The
experimental site is at an elevation of 2,225
m and lies on a north-facing slope in a mod-
erate snow catchment zone. Annual precipi-
tation is about 500 mm, with two-thirds of
the total falling as snow. Precipitation be-
tween 1 June and 30 September averages 114
mm. Sufficient snow usually accumulates to
completely recharge the soil mantle. Soils de-
veloped in place from sandstone and belong
to the Argic Cryoboroll great soil subgroup.
A dense stand of mountain big sagebrush, un-
derlain by a productive understory of bunch
grasses-primarily Idaho fescue (Festuca ida-
Jwensis), bluegrass (Poa spp.) and needle-
grasses {Stipa spp.)-was present before study
initiation. The site had been grazed by sheep,
but no grazing occurred during the study.
Methods
Work began in 1968 with a study that uti-
lized 14 0.4-ha experimental units arranged
in seven blocks to determine how the soil wa-
ter regime would be affected by spraying big
sagebnish (Sturges 1977a). One experimental
unit within each block was sprayed with 2,4-
D in 1970; the other unit remained un-
treated. Experimental units from three of the
seven blocks were used in the current study.
In October 1972, smaller plots 23 m long and
10 m wide were established on either side of
the common border between sprayed and un-
sprayed vegetation (Fig. 1). These plots were
used to obtain soil cores and to create the
grubbed sagebrush vegetative condition.
Sagebnish was grubbed from four circular
areas 6.1 m in diameter by cutting plants at
or slightly below the ground surface. Grubb-
ing was done in the fall of 1972, when vege-
tation was dormant, thereby minimizing
damage to residual herbaceous vegetation
and insuring that herbaceous vegetation
would be as comparable as possible to that
within the undisturbed sagebnish stand when
study measurements began the following
spring.
Soil Water Measurements
Soil water content was measured with a
neutron-scattering soil moisture meter at four
randomly located access tubes on each exper-
imental unit. Access tubes within grubbed
vegetation were installed at the center of
each cleared circle in October 1972. If one
assumes that big sagebnish has a maximum
lateral root spread of 1.5 m, these tubes were
surrounded by a volume of soil at least 1.5 m
in radius devoid of live sagebrush roots.
Moisture measurements began 31 May
1973 upon completion of snowmelt and con-
tinued at biweekly intervals until 19 Septem-
ber 1973, when vegetation was dormant.
Measurements were taken at eight depths:
15, 30, 46, 61, 76, 91, 107, and 122 cm. The
manufacturer-supplied calibration curve re-
lating field neutron count (expressed as a per-
centage of shield count) to volume moisture
content was applied to all data except that
collected at 15 cm. Here, a correction was
made for escape of neutrons into the atmo-
June 1980
Sturges: Big Sagebrush
159
sphere using a polyethylene shield technique
similar to Pierpoint's (1966).
Root Weights
Soil cores for sampling root weight were
obtained at four random locations within the
small plots that straddled the common border
between sprayed and midisturbed sagebrush
vegetation. The cores were collected in Sep-
tember following the final soil water mea-
surement (Fig. 2). Each core was 7.6 cm in
diameter and 122 cm long. The cores were
collected in 15-cm increments using the de-
vice described by Brown and Thilenius
(1977). Each sample site in grubbed vegeta-
tion was located within 2.4 m of an access
tube, a minimum of 0.6 m from the surround-
ing sagebrush cover. Soil cores were placed
in plastic bags and frozen on the day of col-
lection. After thawing, core segments were
individually washed in a core-washing ma-
chine (Brown and Thilenius 1976) to isolate
root matter from soil. Roots were oven dried
for 24 hours at 70 C and weighed on an ana-
lytical balance. It was not possible to dis-
tinguish between live and dead roots, but
woody sagebrush roots from cores taken
within grubbed vegetation were discarded
before samples were weighed.
Herbaceous Production
Above-ground herbaceous productivity
was measured by clipping 12 randomly lo-
cated plots within each experimental unit as
grasses matured in mid-July. In grubbed veg-
etation, three production plots were placed
Undisturbed sagebrush
vegetation
Sprayed sagebrush
vegetation
10 m
23nn
64 nn
64 nn
X Access tube on 0.4 ha plot
fj Circular area where sagebrush grubbed
+ Access tube in grubbed vegetation
Fig. 1. The experimental design for one block showing the 0.4-ha experimental units of undisturbed and sprayed
sagebrush vegetation and smaller plots where sagebrush was grubbed. Soil moisture data, soil cores, and herbaceous
productivity information for the grubbed treatment were obtained on the small plot.
160
Great Basin Naturalist
Vol. 40, No. 2
at random within 2.4 m of access tubes. Veg-
etation was harvested to a 1 cm stubble
height from plots 30.5 cm wide and 61 cm
long. Vegetative matter was separated into
grass, forb, or sagebnish components and
placed in paper bags when harvested. Only
leaves and herbaceous stem material were in-
cluded with sagebnish herbage. Vegetation
samples were subsequently dried at 105 C for
24 hours and weighed.
Selected big sagebrush and productivity
characteristics were measvired in 1969 on the
0.4-ha experimental units, the year before
spraying (Table 1). No statistically significant
differences before treatment were present.
Big sagebrush contributed 76 percent of
aboveground herbaceous production while
grasses contributed 20 percent and forbs 4
percent. About one-third of the area was cov-
ered by the live, leafy portion of the sage-
bnish canopy. Sagebrush plants had an aver-
age height of 34 cm and an average crown
area of 7 dm-.
Statistical Analysis
Soil water withdrawal and root weight dif-
ferences among the three vegetative condi-
tions were tested for statistical significance
by variance analysis utilizing a split-plot de-
sign. Experimental units (whole units) were
arranged in three randomized 'blocks, and the
eight measurement depths served as subunits.
Analyses were based on average plot values
determined from the four replicated mea-
surements on the plot. Variables analyzed
were the change in soil water content be-
tween successive sampling dates, the seasonal
change in soil water content, and root
weight. Herbaceous productivity data were
analyzed with a randomized block design.
Results
Soil Water Depletion
Soil under undisturbed and sprayed vegeta-
tion was completely recharged by snowmelt
on the first measurement date, but only to 61
cm under grubbed vegetation (Fig. 3). At the
end of summer, water content in the surface
46 cm of .soil was similar for all treatments.
Below 46 cm, progressively more water re-
mained in soil under grubbed vegetation
compared to undisturbed sagebnish vegeta-
tion, but appreciable differences between
sprayed and undisturbed vegetation were
present only below 91 cm.
Seasonal water withdrawal by undisturbed,
sprayed, and grubbed vegetation was 24.3,
21.4, and 16.2 cm of water, respectively, in
the surface 122 cm of soil. These differences
Table 1. Characteristics of vegetation on plots as-
signed to sprav and undisturbed treatments in 1969, one
vear before 2,4-D was applied.
Sagebrush
Undisturbed
Sprayed
Height (cm)
30°
37
Canopy area (dm')
6
8
Canopy intercept (%)
31
32
Density (number/ha)
57,(X)0
52,000
Hebbac;eous production
(kg /ha)
Sagebrush
926
1095
Grass
249
290
Forb
53
60
Total
1228
1445
"Differences between treatment means were not significant for any mea-
surement parameter at the 0.05 level of probability.
tig. 2. Root samples were obtained using a core sam-
pler driven into the soil to a 122-cm depth by 15-cm in-
crements.
June 1980
Sturges: Big Sagebrush
161
were significant at the 0.01 probability level.
Treatment differences did not accrue uni-
formly through the soil mantle, but were con-
centrated at deeper soil depths (Fig. 4). Be-
tween 91 and 122 cm, depletion by grubbed
and sprayed vegetation was 31 and 66 per-
cent, respectively, of depletion by undis-
turbed sagebnish vegetation.
Treatment soil water withdrawal differ-
ences between consecutive measurement
dates were significant (p<0.05) only between
25 June and 10 July. The treatment x depth
interaction term was significant during five
of the eight measurement intervals, though,
indicating that the three vegetative condi-
tions were utilizing water differently from
within the soil. For example, most of the dif-
ference in depletion below 91 cm between
1 20
CL
.^ 0
D
Q
Undisturbed
Sprayed
Grubbed
76 cm
91cm
46 cm
107cm
61cm
122 cm
-J I 1 i_
ilLlJI IlL
15 31 15 30
li .J .1
I nil. I
-Id L
5 31 15 31 15 30 15 31 15 30
I III.. I .1 I 1. L
5 31 15 31 15 30
May June July Aug. Sept. May June July Aug. Sept.
Fig. .3. Daily precipitation and soil water content in the surface 122 cm of soil for undisturbed, sprayed, and
grubbed .sagebnish vegetation in the 197.3 growing season.
162
Great Basin Naturalist
Vol. 40, No. 2
sprayed and undisturbed vegetation devel-
oped after 25 July. Sagebrush remained phys-
iologically active through the summer and
flowered about 1 September, so that appre-
ciable water usage continued all summer.
Most grass and forb species had matured and
set seed by early August, thus reducing the
need for water by grubbed and sprayed vege-
tation.
Root Weights
The average weight of roots obtained from
soil cores extending 122 cm deep was 12.2,
10.2, and 8.7 g under undisturbed, sprayed,
and grubbed vegetation, respectively. Nei-
ther the treatment, nor the depth x treatment
interaction term was statistically significant.
Varying quantities of dead but undecayed
root matter and other organic debris were in-
cluded with sample material and could not
be separated from live roots. Inclusion of ex-
traneous matter probably accounted, in part,
for the low statistical sensitivity of root mea-
surements.
Most of the weight of roots was located in
surface soil (Fig. 4). Material from the surface
15 cm of soil ranged from 36 percent of total
root weight in imdisturbed sagebRish vegeta-
tion to 54 percent of total root weight in
sprayed vegetation. Conversely, only 1 to 2
percent of root weight for each treatment
came from the deepest sampling depth.
Herbaceous Production
Herbaceous production of undisturbed
sagebrvish vegetation was about a third great-
er in 1973 than in 1969, but composition of
vegetation was similar both years. Treatment
differences within sagebrush, grass, and total
production herbage classes were highly sig-
nificant (Table 2). The response by sprayed
vegetation the third year after treatment was
typical to that reported from other locations.
Grass production was 2.6 times greater than
production in imdisturbed sagebmsh vegeta-
tion, but forb production was still depressed
below pretreatment levels. Total herbaceous
production by sprayed vegetation was only
57 percent as large as production by undis-
turbed sagebnish vegetation, the increase in
grass production not compensating for loss of
sagebrush.
Grass production increased 27 percent
where sagebrush was grubbed the previous
fall, but the increase was not statistically sig-
nificant (Table 2). Total production was 31
percent as high as that by undisturbed vege-
tation because of the loss of sagebrush.
Discussion and Conclusions
This study indicates the soil water regime
in the surface 91 cm of soil is unaffected by
sagebrush control once herbaceous vegeta-
tion responds to release from sagebrush com-
petition. However, below 91 cm, substantial
reductions in seasonal withdrawal can occur
as reported by Tabler (1968) and Sturges
(1977a). The overall reduction in soil water
depletion caused by grubbing sagebrush com-
pares closely with that detected on the same
0.4-ha experimental units in 1970 when sage-
bnish was sprayed. Grubbing decreased sea-
sonal water withdrawal 33 percent in this
study, and spraying reduced withdrawal from
the surface 137 cm of soil 37 percent (from
the spray date on 22 June through 30 Sep-
tember). The year after spraying, a 17 per-
cent difference in seasonal withdrawal was
observed with grass production doubling in
response to sagebrush removal.
Reductions in moisture withdrawal are re-
lated to decreased aboveground herbaceous
productivity of treated vegetation. Produc-
tivity in grubbed and sprayed vegetation was
31 and 57 percent as large, respectively, as
that of undisturbed vegetation. Development
of vegetation in years immediately following
sagebrush control also influenced water with-
drawal patterns. Seasonal depletion under
gnibbed vegetation was less than that of un-
disturbed sagebnish vegetation at all depths,
but appreciable differences existed only be-
Table 2. Aboveground herbaceous production
(kg/ha) by undisturbed, sprayed, and grubbed vegeta-
tion in 1973.
Treatment Sagebnish
Grass
Forb
Total
Undisturbed °120P
Spraved 1^
C; rubbed (1*^
347*
918^
442''
86*
Ifif'
67*
1634*
93^
509^"
"Treatment means having different letters within a column are signifi-
cantly different at the 0.05 level o^probability.
June 1980
Sturges: Big Sagebrush
163
o
o
cr
0-15 15
,;yyw^.i...»,.i„.iiiiii„„„„„„„„y,^.. .
^;M. npot weight •mi Depletion Undisturbed
_| Sprayed
t i
Grubbed
15-30 30
30-46 46
E 46-61 61
o
IP
Q.
Q
61-76 76
76-91 91
91-107 107
107-122 122
I 2 3
Root weight (g)
Seasonal water depletion/15 cm soiKcnn]
Fig. 4. Seasonal water depletion and weight of roots in soil cores 7.6 cm in diameter and 122 cm long unde
undisturbed, sprayed, and grubbed sagebrush vegetation.
164
Great Basin Naturalist
Vol. 40. No. 2
low 61 cm. Sprayed vegetation, unlike
grubbed vegetation, had fnlh- responded to
release from sagebrush competition and
depletion did not become appreciabK less
tlian that of imdistiu-bed vegetation mitil a
91-cm depth was reached. Reductions in
treatment effect through time within soil
60-90 cm deep were described by Hyder and
Sneva (1955), Cook and Lewis (1963), and
Shown et al. (1972).
The reduction in seasonal water use and in
root weight caused b\' treatments are similar
when e.vpressed as a percentage of values for
undisturbed vegetation. Seasonal depletion
was 33 and 12 percent less for grubbed and
spraved vegetation, respectively, and root
weights were 29 and 16 percent smaller on
these same treatments. Similar agreement be-
tween depletion and root weight did not exist
for individual measurement depths (Fig. 4).
Thus, root weight measurements do not veri-
f\' or refute the hypothesis that root devel-
opment by herbaceous species in the surface
90 cm of soil subsequent to sagebrush remov-
al accoimts for increases in moisture use from
this zone. Measurement of root length, rather
than root weight, probably would have pro-
vided a better measure of potential moisture
draft because of the differences in morpho-
logy of grass and sagebrush roots.
Comparisons of seasonal moisture change
and root weight with depth does indicate
that deep roots are extremely important in
extracting soil water, even though they com-
prised a small part of root weight in soil
cores (Fig. 4). Summer precipitation is usual-
ly ineffective in replenishing soil water levels
in the sagebrush zone, so that deeper soil be-
comes an important water reservoir when
surface soil dries. A progressive, downward
shift of major water use zones in August was
especially evident for undisturbed sagebrush
vegetation (Fig. 3).
Results of this and other soil water deple-
tion studies indicate that control of big sage-
bru.sh with methods that do not destroy all
vegetation on lands with an adequate popu-
lation of herbaceous species has a relatively
small effect upon the soil water regime.
Changes in the soil water regime can, at
most, result in small increases of streamflow.
This response will onl\ t)ccur on lauds where
soils are deeper than 90 cm and soil water
recharge exceeds that retiuired to fulK wet
the soil mantle. The maxinuuu reduction in
depletion will usually occur in the treatment
\ear because of productivity increases bv
herbaceous species in xears immediatelv after
treatment. Consecjuentlv. justification for big
sagebiiish control must rest on the benefits
derived from shifting site resources to species
more desirable than sagebrush from a given
land management perspective.
Literature Cited
Brown, G. R., and J. F. Thilenius. 1976. A low-cost
machine for separation of roots from soil mate-
rial. J. R;mge Manage. 29;506-507.
1977. .\ tool and method for extracting plant-
root-soil cores on remote sites. J. Range Manage.
30:72-74.
Cook, C. W., .\nd C. E. Lewis. 1963. Competition be-
tween big sagebrush and seeded grasses on foot-
hill ranges in Utah. J. Range Manage.
16:245-250.
Goodwin. D. L. 1956. .\iitecological studies oi Artemisia
tridentata Nutt. Unpublished dissertation. Wash-
ington State Univ., Pullman. 79 pp.
Hi LL, A. C, Jr., .and G. J. Klomp. 1974. Yield of crested
wheatgrass under four densities of big sagebnish
in southern Idaho. U.S. Dept. Agric. Tech. Bull.
No. 1483. 38 pp.
Hyder, D. N., a.nd F. A. Sneva. 1956. Herbage response
to sagebrush spraying. J. Range Manage. 9:34-.38.
Pierpoi.nt, G. 1966. Measuring surface soil moisture
with the neutron depth probe and a surface
shield. Soil Sci. 101:189-192.
Shown. L. M., G. C. Lusby, and F. A. Branso.n. 1972.
Soil-moisture effects of conversion of sagebrush
cover to bunchgrass cover. Water Resour. Bull.
8:1265-1272.
Sti rc;es, D. L. 1977a. Soil moisture response to spraying
big sagebnish: A seven-year study and literature
interpretation. USD.\ For. Serv. Res. Pap. R.\i-
188. 12 pp. Rocky Mt. For. and Range Expt. Stn..
Fort Collins, Colo.
19771). Soil water withdrawal and root character-
istics of big sagebrush. Am. Midi. Nat.
98:257-274.
SriRCES, D. L., AND .M. J. Trlica. 1978. Root weights
and carbohydrate reserves of big sagebrush. Ecol-
ogy 59:1282-1285.
Tabi.er, R. D. 1964. The root system oi Arteniisia triden-
tata at 9,500 feet in Wyoming. Ecology'
45:633-636.
. 1968. .Soil moisture response to spraying big sage-
brush with 2,4-D. J. Range Manage. 21:12-15.
SWARMING OF THE WESTERN HARVESTER ANT.
POGOXOMYRMEX OCCIDEXTALIS
Dorald M. Allred'
,\bstract.— The swarming and mating of harvester ants was observed in Utah in July 1979. Workers groom the
alate forms outside the mound before swarming occurs and are highly aggressive in protecting them. Mating pairs
apparently are not disturbed by other ants. Fertile females likely use moving vehicles that extend their dispersal.
Few detailed accounts of the swarming ac-
tivities of harvester ants are existent in the
literature. Wheeler (1910) reported the
swarming of harvester ants in the desert
along the Colorado River. Michener (1942,
1948) noted swarming and mating of Pogono-
mymiex califomicus and P. barbatus. Strandt-
mann (1942) recorded the mating activities of
P. Comanche, and Chapman (1957) reported
elevational swarming of P. occidentalis on
mountain tops in five states.
During the latter part of June and all of
July in 1979, I traveled extensively over Utah
collecting ants. In most cases when harvester
ants were taken, I partially excavated each
mound from which I collected to determine
the presence of immature and winged forms.
Although winged males and females were
present in the majority of the moimds during
this period, swarming was not seen until the
latter part of July.
On July 23 I stopped at about 11:30 a.m.
(Mountain Daylight Time) to collect from an
area of abimdant. large mounds one mile
west of Elberta. Utah County, Utah, along-
side highway US6 at an elevation of 5400 ft
in a sagebrush-rabbitbmsh habitat [Artemisia
tridentata-Chrysothamnus nauseosus). -\s I
approached a large moiuid on that wanii,
sunny day, an area of one-half square foot
around two enlarged, south-facing openings
was literally red with a mixture of \\orkers,
winged males and females. I would have had
difficulty finding an open space within tlie
mass of ants where I could ha\'e touched the
ground with a pencil. The winged forms
were relatively inactive, none in flight, and
the workers seemed to be grooming and at-
tending them. As I approached the moumd to
aspirate a sample of ants, the majority of the
winged females and some males quicklv en-
tered the nest openings. Some of the males,
however, remained immobile outside the
mound as though mesmerized bv the sToom-
ing activities of the workers. The workers on
the fringe of the mass immediatelv began ag-
gressive tactics toward me in much more of a
frenzied movement than I had heretofore en-
countered with workers when winged forms
were not outside the nest.
After I had taken my sample. I returned to
the car to record the data. Five minutes later
I again visited the mound to see if the
winged forms had left the burrow. The air
was filled with fl\ing ants. At the mound the
workers were no longer congregated around
the openings, but were scurrvin^ around,
near, and over the mound. .\ few winded
forms were crawling around on the mound. I
was quickly deluged with flNing ants, and a
sting on my leg. presimiablv from a winged
female, stimulated m\" hastv retreat to the
car. where I quickly closed all windows. The
outside of the car was soon covered with
winged ants that were mating. Females
seemed to be much more abundant than
males. This was consistent \\ith m\ findings
wherever I had excavated mounds through-
out the state during Jime and July.
In mating, the male moimted the female
'Department of Zcxilog), Bricham Vount; I'niversit) . Privo, I'tah S46Q2.
165
166
Great Basin Naturalist
Vol. 40, No. 2
dorsally, clasped her around the thorax with
his legs, and bent his abdomen strongly
downward to contact her gentitalia. She si-
multaneously bent her abdomen slightly up-
wards to facilitate contact. Once joined, the
partners sometimes assumed different posi-
tions than described above, frequently both
establishing leg contact with the substrate on
which they were resting, although maintain-
ing abdominal junction. Other crawling ants
frequently came in contact with a mating
pair, but the contact was brief. At no time
did I observe other males remaining with
mating pairs as described by Strandtmann
(1942) for P. Comanche. Copulation lasted for
perhaps 20 to 40 seconds, whereupon the two
sexes immediately separated, with the male
the first to fly away. No case was observed
where either the male or female used its
mandibles to grasp or chew on the other as
described by Strandtmann (1942) for P. Com-
anche, and by Michener (1948) for P. har-
batus.
About 12:.30 p.m., after I had observed the
mating activities of numerous pairs and the
numbers of ants crawling on the car and fly-
ing in the area had considerably diminished, I
returned to the mound. Few flying ants re-
mained in the air in its vicinity, but a few
winged forms were crawling around the
mound. Most of the workers had reentered
the entrances, and those that remained out-
side had apparently resumed their normal,
slower speed of routine activity.
I moved to another area of several mounds
situated about 25 yards north of the site of
my previous observations. The ants of several
of these mounds were in various stages of
swarming activities. By periodical rotation
between these, and with some excavations, I
was able to summarize the overall activities
associated with swarming of this species.
When the males and females prepare to
swarm, the workers enlarge the openings
leading from the mounds, the winged ants
and many workers leave the nest and congre-
gate around its openings. The winged males
and females are attended by myriads of
workers who groom them as their bodies
warm in the sun, and at the same time act in
a protective capacity in relation to any
would-be predators. Once the mating flight
begins, the workers disperse and crawl
around the mound for a few minutes, then re-
enter the burrow. Normal worker activity
outside the mound is resumed within a few
minutes after the majority of the winged
forms have departed. A few winged forms
seem to linger around the mound, apparently
hesitant to leave. Some also delay leaving the
burrow to assume their flight of destiny.
Swarming males and females seek some high
point to mate, but many pairs mate on the
ground.
After mating, females migrate in all direc-
tions to locate sites for establishing new colo-
nies. Undoubtedly some of them hitch rides
on moving conveyances such as cars, trucks,
and trains. When I arrived at my home in
Provo, 30 miles from and several hours after
observing the swarming activity, a live
winged female dropped from my car onto the
driveway and crawled into the vegetation.
Such methods of conveyance likely extend
the dispersal of females over relatively great
distances.
Literature Cited
Chapman, J. A. 1957. A further consideration of summit
ant swarms. Canadian Ent. 89:389-95.
Michener, C. D. 1942. The history and behavior of a
colony of harvester ants. Scientific Monthly
55:248-58.
1948. Observations on the mating behavior of
harvester ants. J. New York Ent. Soc. 56:239-42.
Str.\ndtmann, R. W. 1942. On the marriage flight of Po-
go(io»H/n)U'.v comanclic Wheeler. Ann. Ent. Soc.
America .35:140.
Wheeler, W. M. 1910. Ants: Their structure, devel-
opment, and behavior. Columbia Univ. Press,
New York, 663 pp.
RELATIONSHIP BETWEEN ENVIRONMENTAL AND VEGETATIONAL
PARAMETERS FOR UNDERSTORY AND OPEN-AREA COMMUNITIES
William E. Evensoii', Jack D. Brothersoir, and Richard B. Wilcox'
.XbstR'VCT.— Ten individnals from each of four tree species were selected, and their associated understory and ad-
jacent open-area communities were sampled for both environmental and vegetational parameters, including light
intensity, pH, litter depth, soil depth, and percentages of exposed rock, litter cover, living cover, shrubs, forbs,
grasses, and annuals. The four tree species were ponderosa pine. Rocky Mountain juniper, Gambel oak, and snow-
linish ceanothus. The studv site was in the lower Uinta Mountains about 10 miles east of Kamas, Utah. Correlations
among the various biotic and abiotic parameters were examined. The interplay of these factors in differentiating the
understory and open-area communities is discussed.
Understanding the relationship of vegeta-
tional patterns to environment is a primary
goal of community ecology. One aspect of
such relationships is the effect of overstory
trees and shRibs on their associated under-
story communities. In a previous report (Wil-
cox, Brotherson, and Evenson 1981), we ex-
amined the influence of four canopy species
on their associated imderstory plant commu-
nities in comparison to neighboring commu-
nities in open areas outside the canopy in-
fluence. The four canopy species were
ponderosa pine {Piniis ponderosa Dougl.),
Rocky Mountain juniper {Junipenis scopulo-
nim Sarg.), Gambel oak {Quercus gambelii
Nutt.), and snowbrush ceanothus (Ceanothus
vehitinus Dougl.).
Many previous studies have reported envi-
ronmental and vegetational differences be-
tween understories and open areas. Light in-
tensity (del Moral 1972, Cline 1966,
Blackman 1956) and spectral distribution
(Federer and Tanner 1966) are known to
strongly differentiate understory and open-
area plant communities. Soil moisture and
thickness of litter layer are also important
factors (Anderson 1969, McQueen 1973), as is
soil improvement due to nitrogen fixation by
such common understorv plants as bitter-
bmsh {Pursliia tridentata (Pursh) DC.) and
snowbrush ceanothus (Wollam and Young-
berg 1964, Rusel and Evans 1966, Webster,
Youngberg, and Wollam 1967).
Because these and other environmental
modifications are influenced by the canopy
species, cover (Anderson 1969, McQueen
1973, McConnell and Smith 1970) and diver-
sity (Auclair and Goff 1971) of understory
communities are strongly dependent on the
canopy tree or large hebaceous species with
which they are associated (Gordon 1962,
Smith and Cottam 1967).
The present study examines detailed rela-
tionships between the various environmental
and vegetational parameters measured under
the canopies of fovir tree species and in
nearby open areas.
Study Area
The study site is about 10 miles east of
Kamas, Utah, along the Yellow Pine branch
of Beaver Creek (Fig. 1). This area was cho-
sen because of the homogeneity of the under-
lying parent material (an alluvial outwash
gravel bed) throughout the site, its constant
slope and exposure, and its easy accessibility.
The study site is an area of "zone jumbling"
(Cottam 1930) and contains plant representa-
tives from all life zones except lower sonoran.
It is an area of highlv mixed vegetation,
varying from Douglas fir {Pseudotsuga men-
'Department of Physics. Brigham Young I'niversity, Provo, Utah 84602.
'Department of Botany and Range Science. Brigham Young University, Provo, Utah 84602.
'Land Specialist, State of Utah, Room 440, Empire Buildnig, 231 East 400 South, Salt Lake City, Utah 84111.
167
168
Great Basin Naturalist
Vol. 40, No. 2
ziesii (Mirb.) Franco.), white fir (Abies con-
color Lindl.), and ponderosa pine to clumps
of Gambel oak and snowbnish ceanothus.
Also interspersed throughout the area are in-
dividuals of lodgepole pine (Pinus contorta
Dougl. ex Loud.), Rocky Mountain juniper,
quaking aspen (Populus tretnuloides Michx.),
and various other plant species. All can be
found at the same elevation and in fairly
close proximity.
Because of its apparent uniformity, this site
is especially well suited to measure the rela-
tionship of environmental and vegetational
parameters associated with understories of
different tree species and nearby open areas.
In such an area, the likelihood of factors oth-
er than tree overstory affecting such relation-
ships in a major way is small.
"^76 00
y7(.SS
Sfe
iSrW
Yellow Pine
Campground
Fig. 1. Map showing location of stiidv site.
June 1980
EVENSON ET .\L.: PlANT EcOLOCY
169
Methods
Ten individuals of each of four tree or
shrub species (i.e., ponderosa pine, Rocky
Mountain juniper, Gambel oak, and snow-
bnish ceanothus) were chosen at random in
the study area. Eight quadrats (0.25 m^) were
placed aroimd each individual tree or .shrub.
Four of these quadrats were placed inside the
cylinder of the canopy, and four were placed
outside the influence of the canopv. To elimi-
nate bias, quadrats were consistently placed
one at each direction of the compass. Quad-
rats were subdivided into four equal units for
species frequency measurements. Sample
trees and quadrat sites were marked for relo-
cation.
Presence or absence of individual plant
species in the understory was determined for
all four subunits of each quadrat. All species
rooted in the quadrat were recorded. Fre-
quency of each plant species was determined
by dividing the number of quadrat sub-
divisions in which a species occurred by the
number of subdivisions sampled. Total living
plant cover and composition of plant cover
by life form were measured at each quadrat
using an ocular estimate method (Daviben-
mire 1959).
Light intensity was measured in foot-can-
dles at each quadrat, and averages were com-
puted for the understory and open-area quad-
rats associated with each tree species. All
readings were taken between 1200 and 1400
hours on cloud-free days, the last two days of
the study.
Soil pH was measured by the colormetric
method in the field to avoid pH changes
which can occur when soil is stored moist.
Litter depth was measured at the center of
all quadrats taken, and soil depth was deter-
mined by the average of five penetrometer
readings in each quadrat (one at each corner
and one in the center). Correlations of all
variables with each other were nm.
Results .4nd Discussion
Characteristics of the environment and the
vegetation types associated with understory
and open-area communities are summarized
in Table 1 for the four canopy species. As ex-
pected, light intensity values are consistently
lower for the imderstory communities. Un-
derstory communities have consistently high-
er pH (more basic), except for ponderosa
pine which shows an understory tending to
be slightly more acidic than the open area.
Both litter depth and soil depth are far great-
er in tlie imderstory communities than out-
side the canopies. Percent cover of litter is
Table 1. Average measured values of environmental and vegetation parameters for understorv and open area
comnumities ± their standard deviations.
Pintis
Ceanothus
Juniper us
Quercus
ponderosa
vchttintts
scopulortim
gamhelii
Under-
Open
Under-
Open
Under- Open
Under-
Open
story
areas
story
areas
story areas
story
areas
Light intensity
(foot candles)
117 ±52
225 ±65
123 ±29
.3.39 ±50
168 ±77 295 ±67
76 ±.32
242 ±67
Soil pH
6.3 ±0.2
6.4 ±0.1
6.6 ±0.1
6.4 ± 0. 1
7.4 ±0.6 6.4 ±0.2
6.6 ± 0.3
6.4 ±0.2
Litter depth (cm)
6.0±2.7
0.6 ±0.3
2.1 ±0.7
0.1±0.1
2.5 ±0.3 0.3 ±0.2
2.3 ± 0.6
0.2 ±0.2
Soil depth (cm)
10.5 ±2.6
5.3±1.1
6.7±1.7
4.4 ±1.2
6.1 ±.3.2 4.0 ±1.6
8.5 ±2.3
.5.7 ±0.8
% Living cover
20.2 ±6.0
25.8 ±6.8
20.0 ±4.7
31.5 ±7.6
28.0 ±19.1 .32.0 ±8.4
27.5 ±5.2
23.4 ±5.3
% Litter cover
83.6 ±6.0
39.0 ±11.6 45.6± 10.0 9.1 ±7.5
49.3 ±20.3 13.1 ±9.0
72.6 ±5.9
19.3 ±10.4
% Exposed rock
10.8 ±5.7
26.3 ±6.4
24.9 ±6.8
43.4 ±11.1
14.8±11.1 .32.4±8.1
14.6 ±9.9
28.5 ±4.7
% Shrubs
26.8 ± 1.3.0 27.1 ± 17.0 .37.0 ± 16.8 2.3.9 ± 1.3.2
48.0±21.9 22.9±17.8 27.7±11.9 18.4± 1,3.6
% Perennial forbs
5.2 ±7.4
11.4 ±11.3
18.9±9.7
.36.5± 12.9 8.0± 15.3 31.2± 14.9
15.6 ±9.2
27.4 ±12.8
% Perennial grasses
11.5 ±3.4
4.0 ±.3.1
13.9 ±8.5
7.1 ±5.9
16.9 ±1.3.9 10.4 ±9.0
22.8 ±10.2
14.0 ±9.3
% .\nnuals
56.6 ±11.9
57.5 ± 14.5
.30.2 ±12.5
.32.4 ±10.1
27.1 ±2.3.9 .3.5.6 ±2.5.5
.3.3.9 ±13.8 40.2 ±13.1
Average number of
species/tree
10.5
11.9
14.2
14.9
10.9 14.3
12.7
14.0
Average number
of species/
quadrat /tree
4.6
6.2
7.1
6.9
5.4 6.3
7.4
8.4
170
Great Basin Naturalist
Vol. 40, No. 2
much greater in the understory, while per-
cent of exposed rock is greater in the open-
area communities. Percent living cover is
higher in the open-area communities except
for oak.
Species preferences for understory and
open areas were obtained by taking the dif-
ference of total subquadrat occiu-rences for a
species in canopy-covered and open stands
and normalizing by dividing by the total oc-
currences of that species in all stands. The re-
sulting index nms from -1 to 1. Those species
with the highest positive values are found
most often under the canopy while those spe-
cies with the greatest negative values are
found most often in open areas. Table 2
shows species preference indices and total
subquadrat occurrences for each species ob-
served in the study (species are listed alpha-
betically). Frequency is obtained by dividing
the number of occurrences by 160, the total
number of subquadrats sampled for each tree
species within each type (canopy or open
area).
The species preference index was broken
down into components relating to each of the
four tree species as shown in Table 3. Each
component was obtained by taking the differ-
ence of the subquadrat occurrences for a spe-
cies in canopy-covered and open-area stands
Table 2. Number of ocxurrences of each species observed for each of the four canopy species; 160 subquadrats
sampled in each category. Life form codes are f=forb, g = grass, c = cool-season or spring ephemeral but perennial
(so cf = cool-season forb), sh = shrub, e = evergreen, a = annual, t = tree.
Life
Preference
Understory
stands
Open-area
stands
Species
form
index
Ceve
Pipo
Jusc
Quga
Ceve
Pipo
Jusc
Quga
Achillea millefolium
Agoseris glauca
Agropyron spicatum
Agropyron suhsecundum
Agropyron tnichycaulum
Allium acuuiinatum
Amelanchier alni folia
f
f
g
g
g
cf
sh
0.4.3
-0.31
0.00
0.09
0.56
-0.30
-0.50
3
13
0
0
0
1
0
4
9
0
2
12
3
1
11
3
2
2
2
3
0
32
10
0
2
6
0
2
22
0
0
0
8
0
3
23
0
0
0
9
1
9
6
2
4
0
6
2
6
15
0
1
6
1
0
Antennario luzuloides
f
1.00
0
0
0
1
0
0
0
0
Antennaria rosea
f
-1.00
0
0
0
0
0
0
1
0
Apocynum androsaemifolium
Arctostapliylos uva-ursi
f
esh
-0.78
1.00
2
0
0
0
0
4
0
0
9
0
0
0
0
0
0
Artemisia tridentata
esh
-1.00
0
0
0
0
0
0
1
0
Aster chilensis var. adscendens
f
-0.28
6
0
0
3
10
0
0
6
Bromus ciliatus
g
0.32
11
11
4
9
2
5
6
5
Bromus tectorum
g
-1.00
0
0
0
0
0
0
0
1
Carex geyeri
Carex rossii
Ceanothus velutinus
eg
eg
esh
0.36
0.31
0.09
6
23
0
9
0
0
35
1
6
43
18
0
1
14
0
15
0
0
22
5
6
1
0
Chenopodium amhrosioides
Chenopodium fremontii
Chrysopsis tillosa
Cirsium undulatum
a
a
f
f
1.00
-0.,32
-0.79
-0..33
0
9
8
1
0
1
0
0
0
0
4
0
3
5
0
0
0
20
33
0
0
0
17
0
0
3
42
2
0
6
10
0
CoUinsia pnrviflora
Collomia linearis
Comandra umbellata
a
a
cf
-0.13
-0.19
0.07
7
20
11
.33
35
2
39
36
4
19
28
5
IS
8
41
66
1
36
39
8
.33
37
2
Cryptantha circttmscissa
Eriogonum heracleoides
a
esh
-0.42
0.07
6
1
1
14
0
0
0
16
1
0
0
0
9
1
9
Erysimum asperum
Euphorbia rohusta
Galium horeale
f
f
f
-1.00
1.00
-1.00
0
1
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
Gayophytum ramosissimum
Uydrophyllum capitatum
Ipompsis agoregata
Juniperus communis
Lomatium grayi
Mahonia repens
a
cf
f
esh
f
esh
-0.91
-0.02
-1.00
1.00
-0.05
0.08
1
4
0
0
7
70
2
1
0
0
9
40
1
5
0
4
2
60
1
10
0
0
0
48
11
2
0
0
15
47
5
5
1
0
2
43
30
4
0
0
3
44
59
10
0
0
0
50
June 1980
EvENSON ET AL.: PlANT EcOLOGY
171
associated with one of the four canopv spe-
cies. This difference was then normahzed by
dividing by the total occurrence of that spe-
cies in all stands. The four components thus
sum to the species preference index discussed
above (within round-off error). The com-
ponents of the species preference index clari-
fy how a species preference for understories
or open areas is associated with a particular
canopy species. Some luiderstorv species are
highly associated with a particular canopy
tree species, and others are not. For example,
Carex geycri's preference for the understorv
is strongly associated with Gambel oak. Yet,
Stellaria jamesiana prefers the understory
much more evenly for three of the four cano-
py species.
Correlation analysis was performed to
study the relationship of the environmental
and vegetational parameters which were
measured. Table 4 shows the significant posi-
tive and negative correlations.
Light intensity correlates significantly with
all variables except percent annuals. The
negative correlation of light intensity with
pH, Itter depth, soil depth, and percent litter
cover is to be expected because of the gener-
ally higher values of these parameters under
the canopies. Similarly, there is more exposed
rock and living cover in the open areas. The
correlations of light intensity with life forms
reflect the preference of shrul>s and grasses
for understory areas and forbs and annuals for
open areas (Wilcox, Brotherson, and Evenson
1981).
The significant correlations of pH with
Table 2 continued.
Understory
Open
area
Life
Preference
stands
stands
Species
form
index
Ceve
Pipo
Jusc
Quga
Ceve
Pipo
Jusc
Quga
Melica bulbosa
g
1.00
0
2
0
0
0
0
0
0
Merten.mi hrcvistijle
f
-l.(X)
0
0
0
0
2
0
0
0
Osniorhizci obtttsa
f
-l.(K)
0
0
0
0
0
0
0
1
Pdcliystima nujisiiiites
esh
0.19
3
2
10
4
2
3
6
2
Penstcmon sp.
f
-1.00
0
0
0
0
0
0
2
0
Piiuis coutorta
at
l.(X)
0
0
0
1
0
0
0
0
Pintis pondcnmi
at
0.00
0
I
0
0
0
1
0
0
Poa curt a
g
-0.36
0
0
0
-
0
0
3
12
Poa fcndlcriana
g
-1.00
0
0
0
0
3
0
0
0
Poa pratensis
g
0.11
0
5
0
0
0
0
4
0
Pohjgonuin cloui:,hi.sii
a
-0.51
65
17
29
28
129
92
87
119
Potcntilla olanduhmi
f
-0. 1 1
3
0
0
1
0
0
1
4
Pntniis viroiniana
sh
0.54
1
1
6
2
0
2
1
0
Purshid tridcntatci
sh
l.(K)
1
0
0
1
0
0
0
0
{hicrru.s ^ambelii
t
l.(K)
0
0
1
0
0
0
0
0
Rosa ivoodsii
sh
0.20
5
5
6
S
6
4
2
4
Seduiii stenopctaluni
f
-0.36
6
4
15
3
20
11
21
(
Scnccio u in tah ensis
f
-0.64
2
0
0
0
4
0
1
4
Silcnc menziesii
ef
0.33
0
2
0
0
1
0
0
0
Solidaoo tnissouricnsis
f
-0.69
5
2
1
0
25
9
9
0
Solida^o multiradiata
f
0.05
9
17
4
2
4
17
6
2
SoUdaop sparsiflora
f
-0.53
0
0
0
11
0
13
0
23
Stellaria janwsiana
cf
0.51
55
94
27
105
11
36
16
27
y,tipa cohiiubiaiia
g
-l.(K)
0
0
0
0
0
0
0
5
Stipa k'ttcrmanii
g
0.09
5
1
5
1
5
3
2
0
Sijmphoricarpos oreophilus
sh
0.20
8
2
12
5
4
3
9
2
Taraxacum officinale
f
0,(K)
4
6
2
2
2
4
3
5
Tlialictrum fendleri
f
l.(K)
0
0
6
0
0
0
0
0
Thlaspi montanutn
f
-0.20
.53
16
28
40
48
55
25
76
Tragopooon dubius
a
0.25
5
0
0
0
1
0
2
0
Viguicra midtijlora
f
0.00
9
0
2
2
5
6
0
2
Viola nuttallii
cf
0.(X)
0
0
0
1
0
0
0
1
Viola purpurea
f
0.52
11
0
0
5
1
2
0
2
Total number of species
39
34
36
37
38
32
41
.39
172 Great Basin Naturalist Vol. 40, No. 2
other variables again reflect the tendency to- nated communities with relatively low total
ward higher pH under the canopies. Sim- living cover and significant amounts of litter
ilarly, for litter depth, soil depth, percent ex- cover.
posed rock, percent litter cover, and percent By contrast, shrubs and forbs are positively
living cover the correlations generally reflect correlated with percent living cover, but
the relationship of these parameters to the negatively correlated with each other. Shrubs
canopy-covered or open-area condition. provided a significant proportion of living
Correlations of these parameters with per- cover in any quadrat in which they occur just
cent annuals are not so easily interpreted, because of their size. This fact accounts for
however. Annuals tend shghtly to prefer the their positive correlation with living cover,
open areas, hence the negative correlation However, shrubs tend to prefer understory
with pH. Their positive correlation with per- habitats and forbs prefer the open areas. So
cent litter cover, however, is better under- forbs are positively correlated with living
stood by observing the negative correlation cover due to the greater cover outside the
with all other life forms and percent living canopies, while being negatively correlated
cover. Annuals tend to grow in annual-domi- with shrubs.
Table 3. Preference index components by tree species. Life form codes are as in Table 2.
Life
form
Preference _
index
Preference index
components
Species
Ceve
Pipo
Jusc
Quga
Antennaria luzuloides
f
LOO
0.00
0.00
0.00
1.00
Arctostaphylos uva-ursi
ash
LOO
0.00
0.00
1.00
0.00
Chenopodium ambrosioides
a
LOO
0.00
0.00
0.00
LOO
Euphorbia robusta
f
LOO
1.00
0.00
0.00
0.00
Jttniperus conimiinis
esh
LOO
0.00
0.00
1.00
0.00
Melica bulbosa
g
LOO
0.00
1.00
0.00
0.00
Pinus contorta
at
LOO
0.00
0.00
0.00
1.00
Purshia tridentata
sh
1.00
0.50
0.00
0.00
0.50
Querciis gambelii
t
1.00
0.00
0.00
1.00
0.00
Tlialictrwn fendleri
f
1.00
0.00
0.00
1.00
0.00
Agropyron trachycaitlum
g
0.56
0.00
0.44
0.07
0.04
Pruntis virginiana
sh
0.54
0.08
-0.08
0.38
0.15
Viola purpurea
f
0.52
0.48
-0.10
0.00
0.14
Stellaria jamesiana
cf
0.51
0.12
0.16
0.03
0.21
Adiillea millefolium
f
0.43
0.01
0.01
0.03
0.37
Carex geyeri
eg
0.36
0.04
-0.04
0.09
0.27
Silene menziesii
af
0.33
-0.33
0.67
0.00
0.00
Bromus ciliatus
g
0.32
0.17
0.11
-0.04
0.08
Carex rossii
eg
0.31
0.14
0.00
-0.09
0.27
Tragopogon dubius
a
0.25
0.50
0.00
-0.25
0.00
Rosa woodsii
sh
0.20
-0.02
0.02
0.10
0.10
Symphoricarpos oreophilus
sh
0.20
0.09
-0.02
0.07
0.07
Pachystima myrsinites
esh
0.19
0.03
-0.03
0.12
0.06
Poa pratensis
g
0.11
0.00
0.56
-0.44
0.00
Agropyron subsecundum
g
0.09
0.00
0.18
-0.18
0.09
Ceanothus celutinm
esh
0.09
0.00
0.00
0.09
0.00
Stipa lettennanii
g
0.09
0.00
-0.09
0.14
0.05
Mahonia repens
ash
0.08
0.06
-0.01
0.04
-0.00
Comandra umhellata
af
0.07
0.07
0.02
-0.10
0.07
Eriogonum heracleoides
ash
0.07
0.00
0.34
-0.05
-0.22
Solidago multiradiata
f
0.05
0.08
0.00
-0.03
0.00
Agropyron spicatum
g
0.00
0.00
0.00
0.00
0.00
Pinus ponderosa
at
0.00
0.00
0.00
0.00
0.00
Taraxacum officinale
f
0.00
0.07
0.07
-0.04
-0.11
Viguiera multiflora
f
0.00
0.15
-0.23
0.08
0.00
June 1980
EVENSON ET AL.: PlANT EcOLOGY
173
Summary and Conclusions
Canopy tree species clearly influence both
vegetation and environment in their under-
stories (Wilcox, Brotherson, and Evenson
1981). The effects of this influence on plant
distributions are shown clearly for individual
species in Tables 2 and 3.
The correlations of the environmental and
vegetational parameters among themselves
can be imderstood on the basis of a few im-
portant concepts.
(1) The environmental parameters (light in-
tensity, pH, litter depth, soil depth, percent-
age of exposed rock, and percentage of litter
cover) are directly influenced by the pres-
ence or absence of canopy cover. All correla-
tions among these parameters are as expected
on that basis.
(2) The vegetational correlations follow
primarily from the facts that there is more
living cover in open areas than in the under-
stories; shnibs and grasses tend to prefer the
understories; and forbs and annuals tend to
prefer open areas.
(3) Shrubs follow the pattern inferred from
their tendency to prefer understory areas ex-
cept for their positive correlation with the
percentage of living cover. This is because
shrubs themselves provide a large fraction of
the living cover that is found in understory
quadrats.
(4) Annuals are different. They apparently
tend to grow in annual-dominated patches
with low total living cover and relatively
high litter cover.
Table 3 continued.
Life
form
Preference
index
Preference index
components
Species
Ceve
Pipo
Jusc
Quga
Viola ntittallii
cf
0.00
0.00
0.00
0.00
0.00
Hydrophylltiiii capitatum
cf
-0.02
0.05
-0.10
0.02
0.(X)
Lomatiwn gnnji
f
-0.05
-0.21
0.18
-0.03
0.(X)
Potentilla glandulosa
f
-0.11
0..33
0.00
-0.11
-0.33
CoUinsia paniflora
a
-0.13
-0.05
-0.04
0.01
-0.06
Colloniki linearis
a
-0.19
-0.04
-0.11
-0.01
-0.03
TliUispi montanwn
f
-0.20
0.01
-0.11
0.01
-0.11
Aster chilensis tar. adscendens
f
-0.28
-0.16
0.00
0.00
-0.12
Allium acuminatutn
cf
-0.30
-0.19
-0.16
-0.08
0.14
Agoseris oUnica
f
-0.31
-0.09
-0.14
-0.03
-0.05
Chenopodiutn fremontii
a
-0.32
-0.25
0.02
-0.07
-0.02
CirsitiDi iindiilatuui
f
-0.33
0..33
().(X)
-0.67
0.00
Poo eurta
g
-0.36
0.00
0.00
-0.14
-0.23
Sedtim stenopetahnti
f
-0.36
-0.16
-0.08
-0.07
-0.05
Cnjptantha circuinscissa
a
-0.42
-0.42
0.04
0.00
-0.04
Amelanchier alnifolia
sh
-0.50
0.00
0.(X)
-0.50
0.00
Polygonum dotiglasii
a
-0.51
-0.11
-0.13
-0.10
-0.16
Solidago sparsi flora
f
-0.53
0.00
-0.28
0.00
-0.26
Senecio uintahensis
f
-0.64
-0.18
0.00
-0.09
-()..36
Solidago m isso u riensis
f
-0.69
-0.39
-0.14
-0.16
0.(X)
Apocynum androsaemifolium
f
-0.78
-0.39
-0..39
0.00
0.00
Chrysopsis villosa
f
-0.79
-0.22
-0.15
-0..33
-0.09
Cayophytum ramosissimum
a
-0.91
-0.09
-0.03
-0.26
-0..53
Antennaria rosea
f
-1.00
0.00
().(X)
-l.(X)
0.(X)
Artemisia tridentiita
esh
-l.(X)
0.00
0.(X)
-l.(X)
O.tX)
Bromus tectorum
g
-l.(X)
0.00
0.(X)
().(X)
-1.00
Erysimum aspcrum
f
-1.00
-1.00
0.00
0.(X)
0.(X)
Galium horeale
f
-1.00
0.(X)
0.(X)
0.(X)
-l.(X)
Ipompsis aggregata
f
-l.(K)
0.00
-1.00
0.(X)
().(X)
Me rten s ia b re f istyla
f
-l.(K)
-1.00
0.(X)
0.(X)
0.00
Osmorhiza obtusa
f
-l.(X)
().(X)
().(X)
0.00
-l.(X)
Penstemon sp.
f
-l.(X)
0.(X)
().(X)
-1.00
0.(X)
Poa fendleriana
g
-1.00
-1.00
0.00
0.00
().(X)
Stipa Columbiana
g
-1.00
0.00
0.00
0.00
-l.(X)
174
Great Basin Naturalist
Vol. 40, No. 2
Table 4. Significance levels for correlations of life form types, cover, and measured environmental parameters
for all stands studied.
Variables
7
8
10
11
1. Light intensity
2. pH
3. Litter depth
4. Soil depth
5. % exposed rock
6. % litter cover
7. % living cover
8. % shrubs
9. %forbs
10. % grasses
11. % annuals
.001 -.001 +.001 -.001 +.1 -.05 +.001 -.001 NS
NS NS -.05 NS NS + .05 -.01 NS -.05
+ .(K)1 -.001 +.001 NS +.1 -.001 NS NS
-.001 +.fK)l -.05 NS -.01 +.1 NS
-.(K)l +.1 -.1 +.001 -.01 NS
-.001 NS -.001 +.001 +.05
+ .01 +.1 NS -.001
-.001 NS -.001
NS -.05
NS -.05
-.01
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Wollum, A. G. II, and C. T. Youngberg. 1964. The in-
fluence of nitrogen fixation by non-leguminous
woody plants on the growth of pine seedlings.
Journal of Forestry 62:216-321.
SEASONAL ACTIVITY PATTERN OF COLUMBIAN GROUND SQUIRRELS
IN THE IDAHO PRIMITIVE AREA
Charles L. Elliott' and jerran T. Flinders'
.\bstr.\ct.— Data were gathered concerning the seasonal activity pattern of a population of Columbian ground
scjuirrels (Spermophihis colttmbianus) in the Idaho Primitive Area. Adult females were significantly more active in
|une of all vears than were adult males. \ relationship between ground squirrel activity and temperature is postulat-
ed in which the squirrels alter their activity so as to avoid high temperatures and possible heat stress.
Members of the genus Spennophilus are
characterized bv a seasonally short period of
aboveground activity and a prolonged period
of hibernation. During this time of surface
activity ground squirrels must establish terri-
tories, breed, reproduce, and gain .sufficient
weight to survive the inactive season. The an-
nual cycle of activity for various species of
ground squirrels has been reported (Skryja
and Clark 1970, Michener 1974, Loehr and
Ris.ser 1977), but these data as they apply to
populations of Columbian ground squirrels
{Spennophilus cohimbianiis) are limited.
Methods
The study was conducted at Cold Mead-
ows, an 87 ha meadow (elev. 2010 m) located
in the northeastern portion of the Big Creek
Ranger District, Idaho Primitive Area. A de-
scription of the Big Creek area has appeared
elsewhere (Wing 1969, Hornocker 1970, Sei-
densticker et al. 1973). Ground squirrels were
trapped from 12-19 June, 17-24 July, and
14-21 August, 1976-1978. Field work prior
to 12 June was impractical because of bad
weather and the inaccessibility of the study
area. A 90 x 90 m grid with 36 trapping sta-
tions 15 m apart was establi.shed on the cen-
tral portion of the meadow. One live trap (15
X 15 X 48 cm) was placed at each trapping
station. Traps were baited with carrot and
checked every hour. Captured squirrels were
marked using the toe clipping sequence of
Melchior and Iwen (1965), sexed, measured.
weighed, time-of-capture recorded, and re-
leased back onto the grid. Vegetation was
collected using the procedure outlined by
Tadmor et al. (1975). All plant samples were
weighed to the nearest gram in the field and
then brought back to the laboratory, where
they were oven dried at 64 C for three days.
The dried specimens were then weighed to
the nearest gram and percent moisture con-
tent calculated. Daily ambient temperatures
were obtained using a Taylor Maximum-Min-
imum thermometer.
Ground squirrel activity in this study was
equated with the animals presence in the
traps. Bias due to 'trap-shy' or 'trap-happy'
squirrels may have occurred, but attempts to
conduct hourly visual censuses proved unre-
liable during the latter months due to the in-
crease in vegetation height.
Results and Discussion
The number of Columbian ground squirrels
captured, including recaptures, is depicted in
Table 1. Adult female squirrels were signifi-
cantly more active in June of each year than
were adult males (Kolmogorov-Smirnov Two
Sample Test, P <0.05). Activity for July and
August was not significantly different be-
tween sexes of adult or young squirrels.
Males are territorial during the breeding
season (Steiner 1970a), exliibiting extreme ag-
gression toward other males and occasionallv
raiding the nesting ground and colonies of
adjacent males (Steiner 1970a, 1970b, 1971,
'Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602.
175
176
Great Basin Naturalist
Vol. 40, No. 2
1972). This aggressive behavior and Lam-
beth's (1977) findings that Columbian ground
squirrels utilize a core area or arena of activi-
ty between 21 and 40 m in size may account
for the greater number of females captured
in June. Dominant males would have been
excluding other male squirrels from the
trapping grid.
The hibernation entry sequence described
by Manville (1959) and Michener (1974) for
S. columbianus was not apparent for the
Cold Meadows colony. This disparity may be
artificial because adult groimd squirrels dur-
ing August become very lethargic and in-
active and were extremely difficult to trap.
Shaw (1925) has suggested that early
spring activities of Columbian ground squir-
rels are largely controlled by temperature
alone, whereas estivation is induced by the
drying of the vegetation. Howell (1938) felt
that "the date of beginning estivation was de-
termined chiefly by the ripening of the vege-
tation and consequent reduction of the mois-
ture content in their (ground squirrels) food,
and in part also by the accumulation of fat in
the body." Nansel and Knoche (1972) also ob-
served Columbian ground squirrels and pos-
tulated that hibernation was a response to
drought prior to a decrease in temperature.
Cold Meadows plant moisture content de-
clined as the season progressed (Table 2), as
did the squirrels' aboveground activity (Table
Peak daily activity of ground squirrels at
Cold Meadows appears to be determined by
temperature. Table 3 shows the monthly time
interval exhibiting the greatest percentage of
adult ground squirrel activity. The mean
maximum temperature for June 1976 (Table
4) was significantly greater than June 1977
and June 1978. The maximum percent of ac-
tivity for June 1976 occurs later in the day
than for June 1977 or June 1978. This same
type of activity shift in relation to significant-
ly greater maximum temperatures is evident
for July; July 1978 is significantly greater
Table 1. Live trapping results by age and sex for Columbian ground squirrels at Cold Meadows, Idaho Primitive
Area, 1976-1978.
June
July
August
Adult
Adult
Juvenile
Adult
Juvenile
Male
: Female
Male
Femal
e
Male
Female
Male
Female Male
Female
1976
12
25
15
32
15
13
5
5
13
5
1977
15
39
7
12
11
8
3
2
2
4
1978
24
32
18
13
6
7
2
6
/
6
Table 2. Mean percent moisture content ( ± standard deviation) of plant species exhibiting the highest frequency
of occurrence at Cold Meadows, Idaho Primitive Area, 1976-1978.
1976
1977
1978
Species'
July August June
July
August June
July
August
Achillea millefolium 66 ±1 72 ±4 72 ±1 72 ±5 58 ±1 82 ±3 77 ±2 60 ±4
Carex aquatilis 65 ±5 63 ±6 61 ± 7 51 ± 4 30 ± 1 67 ±2 58 ±3 45 ±5
Fragaria virginiana 67 ± 8 66 ± 2 65 ± 8 65 ± 7 52 ± 1 76 ± 1 67 ± 2 55 ± 1
Penstemon procerus 68 ±6 65 ±2 72 ±3 66 ±5 47 ±1 73 ± 1 69 ±2 63 ±2
Phleum alpimim 63 ±5 57 ±7 57 ±1 49 ±8 27 ±1 72 ±2 59 ±4 30 ±1
l:n = 10 samples/species/month.
Table 3. Peak activity time intervals for adult Columbian ground squirrels at Cold Meadows, Idaho Primitive
Area, 1976-1978.
1976
1977
1978
June 12-19
July 17-24
August 14-21
1700-1800 hours
13(K)-14(X)
1600-17(X)
1400-1500
1200-1300
1200-1300
1400-1500
1000-1100
13(K)-1400
June 1980
Elliott, Flinders: Columbian Ground Squirrels
177
Table 4. Monthly mean maximum temperatures
(± standard deviation) at Cold Meadows, Idaho Primi-
tive Area, 1976-1978.
1976
1977
1978
June 12-19 22±2C' 20±2l 20*2^
July 17-24 22±l2 22±li-2 24±l''2
August 14-21 17 ±2''^ 25±1^ 17±2'-2
1; Means tested for significance using unpaired t-test at the 0.05 level.
2: Means tested for significance using Cochran and Cox test at the 0.05
level (Ferguson 1959:143).
than July 1976 and July 1977. August 1977
mean daily maximum temperature is signifi-
cantly greater than August 1976 and August
1978; yet the expected shift in peak activity
does not occur. This may be due to the small
activity sample size for August 1977.
The relationship of temperature to activity
indicates that the higher the average daily
maximum temperature, the earlier or later in
the day peak activity will occur. If this hy-
pothesis is valid, then, for those sampling pe-
riods where the average maximum temper-
atures were not significantly different, the
time interval of peak activity should be sim-
ilar. This relationship is observed for June
1977 and June 1978, and July 1976 and July
1977 (Table 3).
Betts (1976) observed a lower elevation
(1360 m) colony of S. columhianus in western
Montana and reported that, with the increase
in temperatures during lactation and post-
lactation periods, there was an increase in
morning and late afternoon activity and a de-
crease in midday activity. Betts postulated
that temperature or solar radiation may limit
the amount of consecutive time Columbian
ground squirrels can spent aboveground.
The observations of Betts (1976) and data
reported here indicate that the scheduling of
surface activity for these squirrels is an ap-
parent behavioral response designed to es-
cape heat stress.
We thank the University of Idaho for per-
mission to use the facilities at the Taylor
Ranch Field Station, Idaho Primitive Area.
Literature Cited
Betts, B. J. 1976. Behaviour in a population of Colum-
bian ground scjuirrels, Spernu)philus coliuiibianus
columhianus. .^nini. Behav. 24:652-680.
Ferguson, C. \. 1959. Statistical analysis in psychology
and education. McGraw-Hill Book Co., Inc., New
York. .347 pp.
HoRNocKER, M. G. 1970. An analysis of mountain lion
predation upon mule deer and elk in the Idaho
Primitive Area. Wildlife Monogr. 21:l-.39.
Howell, \. H. 1938. Revision of the North American
ground squirrels. North American Fauna
56:85-90.
Lambeth, R. E. 1977. The Coliunhian groimd squirrel in
subalpine forest openings in the Idaho Batholith.
Unpublished thesis, Univ. Idaho, Moscow. 1 1.3
pp.
Loehr, K. a., and a. C. Risser, Jr. 1977. Daily and sea-
sonal activity patterns of the Belding ground
squirrel in the Sierra Nevada. J. Mammal.
58:445-448.
Manville, R. H. 1959. The Columbian ground squirrel
in northwestern Montana. J. Mammal. 40:26-45.
Melchior, H. R., and F. A. Iwen. 1965. Trapping, re-
straining and marking .\rctic ground squirrels for
behavioral observations. J. Wildlife Manage.
29:671-679.
Michener, D. R. 1974. .\nnual cycle of activity and
weight changes in Richardson's ground squirrel,
Spermopliilus richardsonii. Can. Field-Nat.
88:409-413.
Nansel, D., and L. Knoche. 1972. Blood changes in tor-
pid and non-torpid Columbian ground squirrels,
Spermophilus cohimbianus. Comp. Biochem.
Physiol. 41A: 175-179.
Seidensticker, J. C IV, M. G. Hor.nocker, W. V.
Wiles, a.nd J. P. Messick. 1973. .Moimtain lion
.social organization in the Idaho Primitive Area.
Wildlife Monogr. .35:1-60.
Shaw, W. T. 1925. Duration of the aestivation and hi-
bernation of the Columbian ground squirrel.
Ecology 6:75-81.
Skryja, D. D., and T. W. Clark. 1970. Reproduction,
seasonal changes in body weight, fat deposition,
spleen and adrenal gland weight of the golden-
mantled ground squirrel Spennophihis lateralis
lateralis, (Sciuridae) in the Laramie Mountains,
Wyoming. Southwestern Nat. 15:201-208.
Stelner, a. L. 1970a. Etude descriptive de quelques ac-
tivities et comportements de base de Spermo-
philus columhianus columhianus (Ord). I. Loco-
motion, soins du corps curiosite et alarme,
reproduction. Rev. Comp. Animal 4:3-21.
1970b. Etude descriptive de quelques activities et
comportements de base de Spennophilus colum-
hianus coltimhianus (Ord). II. Vie de groups. Rev.
Comp. .Animal 4:23-42.
1971. Plav activitv of Columbian ground squir-
rels. Z. Tierpsychoi. 28:247-261.
1972. Mortality resulting from intraspecific fight-
ing in some ground squirrel populations. J. .Mam-
mal. 53:601-603.
Tadmor, N. H., a. Briegher, I. Noy-Meir, R. W.
Benjamin, and E. Eyal. 1975. .^n evaluation of
the calibrated weight-estimate method for mea-
suring production in annual vegetation. J. Range
Manage. 28:65-69.
Wlng, L. D. 1969. Ecology and herbivore use of five
mountain meadows in the Idaho Primitive ,\rea.
Unpublished thesis, Univ. Idaho, Moscow. 215
pp.
HABITAT AND PLANT DISTRIBUTIONS IN HANGING GARDENS
OF THE NARROWS, ZION NATIONAL PARK, UTAH
George P. Malanson'
,\bstract.— Hanging gardens are insular plant comnuinities of the Colorado Plateau. This study examines hanging
gardens in the Narrows, Zion National Park, Utah. The floristic similarity of gardens and the presence of species in
classes of variables which characterize the habitat are disclosed. Although the gardens are isolated and dissimilar, the
individual species are not restricted in the range of habitat found. Floristic dissimilarity cannot be attributed to
differences in habitat. These results are compared to studies of hanging gardens in eastern Utah.
Hanging gardens are plant communities
growing at seeps on the canyon walls of the
Colorado Plateau. The hanging garden envi-
ronment is characterized by shallow soils at a
seep from bedrock. Seeps occur where water
has percolated through a porous formation
until meeting a less permeable layer of rock.
Then the water flows laterally until a canyon
intersects this plane. The narrow canyons of-
ten shade the hanging gardens. Compared to
other environments of the Colorado Plateau,
the hanging gardens are cool and moist.
The Narrows of the North Fork of the Vir-
gin River in Zion National Park is an arch-
etypal hanging garden locale. In an 8 km sec-
tion there are about 60 gardens, varying in
size from a few square centimeters to over
100 m-. Most of these are at permanent seeps
with small discharges of water. The hanging
gardens of the Narrows assume a variety of
shapes, but in general they occupy a place
where erosion has modified the steepness of
the canyon wall. Often these places are hori-
zontal bands. Other gardens occur where ver-
tical jointing has concentrated the seepage.
Some gardens occupy remnants of potholes,
and others are on bulges of travertine. A few
are in alcoves. Most hanging gardens in the
Narrows are close to the level of the river,
where they may be vulnerable to flash floods.
This study examines the relationship be-
tween the plant species and the habitat in the
hanging gardens of the Narrows. The con-
cepts of Ramensky (1924) and Gleason (1926)
are the basis for hypothesizing that the pres-
ence and importance of species at sites are
determined by their individual tolerances and
requirements in relation to the habitat. This
idea can be evaluated by examining the flo-
ristic similarity between sites and the in-
cidence of species across a range of variables
that characterize the biotopes. The hypoth-
esis leads one to expect a positive relation-
ship between floristic similarity and sim-
ilarity of biotope (i.e., if sites represent a
single habitat).
A few authors have investigated the vege-
tation of hanging gardens. In a general eco-
logical stvidy of Zion National Park, Wood-
bury (1933) outlined the stages of primary
succession that occur at seeps. Welsh and
Toft (1976) described a variety of garden
types in Glen Canyon, Utah, based on the
form created by the erosion of the rock, and
traced the geographical affinities of the spe-
cies they found. They called hanging gardens
"relictual refugia" because the gardens pro-
vided sites for species from other southwest
locations, boreal forests, and earlier epochs.
Welsh and Wood (1976) concluded that
hanging gardens have a stable structure, attri-
buting change in species importance mea-
-sured over a one-year interval to measure-
ment error. Wood and Welsh (1976) found
productivity to be relatively high for this
'Department of Geography, University of Utah, Salt Lake City. Utah 84112. Pre.scnt address: Department of Geography, University of California, Los
Angeles, CaWfomia 90024.
178
June 1980
Malanson: Hanging Gardens
179
type of vegetation, and presumed it to be
steady.
Nebeker et al. (1977) believed that the flo-
ristic dissimihiritv found in hanging gardens
of eastern Utah indicated ''random assort-
ments of individuals capable of exploiting the
environments of individual sites." They con-
clude that over 25 percent of the species
were ecological specialists of hanging garden
habitats, within the Colorado Plateau region.
A few studies briefly mention hanging gar-
dens. Clover and Jotter (1944), Flowers
(1959), and Loope (1976) describe hanging
gardens in general ecological studies of the
Colorado River area. Raines (1976) noted
that hanging gardens can be critical habitats
for small mammals in drought years. All
these studies emphasize the importance of
habitat and support the hypothesis of this
study.
Methods
I sampled 29 hanging gardens in an 8 km
section of the Narrows and in 0.75 km of a
tributary (Orderville Canyon) between June
and September 1977. Sites were chosen infor-
mally on the basis of access, but a representa-
tive range of garden sizes was sought. To es-
timate percent foliar cover of species, I
recorded the number of decimeters inter-
cepted by a species along line transects
spaced at 2 m intervals and perpendicular to
the long axis of each garden. In gardens nimi-
bered 1, 2, and 3, I placed a 2500 cm- quad-
rat every 2 m on transects spaced at 2 m in-
tervals and recorded the number of 100 cm-
grids occupied bv each species. This method
proved to be impractical, although in com-
parison tests the results were not significantly
different.
From the values of relative cover for each
garden, I calculated the similarity of each
pair of hanging gardens according to Ru-
zicka's (1958) index:
SI = (2A/2B)(100)
where A is the smaller and B the larger value
of a species in two sites, considering all spe-
cies present in either site.
I measured 11 variables to characterize the
biotope of each hanging garden. Soil and wa-
ter pH at each site was measured color-
imetrically (Microessential Laboratory 0.2
unit paper). I gathered grab samples of soil
from the surface where there was any depth,
not taking any litter. I took 125 ml samples
of water from the seeps. In a few cases sam-
ples could not be taken because of a paucity
of soil or water. The USDA Soils Testing
Laboratory at Logan, Utah, measured soil
and water conductivity as a surrogate of sali-
nity, and the total phosphorous content of
the soil. I measured the slope of each garden
by taking the height:depth ratio and calcu-
lating a percent slope. I sampled soil depth
every 4 m across the center of each garden
with a wire rod to obtain a single average soil
depth measure; in smaller gardens, one or
two evenly spaced measures were taken.
Direct solar radiation was derived by a
computer program of Williams et al. (1972),
which accounts for slope, azimuth, and lati-
tude of the site, and intervening topography
that shades the site. The program computes
the calories per square centimeter for any
one day, and I summed the 120th, 180th, and
240th days of the year to bracket the grow-
ing season. The scale of resolution does not
account for all possible variations in micro-
relief. In locations with particular sunblock-
ing features, such as alcoves, I reduced the
computed value 10 percent. At one garden
where the three-day sum was zero, although
I observed direct radiation at the site, I arbi-
trarily assigned a value of 50 cal/cm-.
I measured three spatial variables: area,
isolation, and distance to the Gateway to the
Narrows Trail. I derived the area of each gar-
den from the grid formed by the species in-
tercept transects. Isolation is defined here as
the sum of the distances from each garden to
its nearest three neighbors. These distances
were measured on a topographic map
(ZNHA 1977). The Gateway to the Narrows
Trail is the scene of much pedestrian tourist
traffic, but above the terminus human use de-
clines rapidly. The distance to the trail, also
measured on the topographic map (ZNHA
1977), may affect invasion by nonnative spe-
cies.
I analyzed the relationships between spe-
cies presence and absence and the 11 envi-
ronmental variables. These variables were
considered separately because there was low
correlation between any two. I followed
180
Great Basin Naturalist
Vol. 40, No. 2
Strahler's (1978) use of the Gh statistic to dis-
close significant differences between species
presence and absence in ordinal categories of
the environmental variables. Although the
data are ratio scaled and the categories are
ordinal, the presence of many zeros in the
species matrix and the number of tie scores
among the environmental variables make
more powerful tests less dependable. Only
the 13 most common species occur frequent-
ly enough to be tested. I divided each of the
11 environmental variables into four or five
ordinal categories (Table 1). The number of
categories and the ranges are arbitrary, ex-
cept that each category represents at least
one garden. I attempted to categorize the
variables so that the results of the Gh tests
would represent the relationships between
species presence and a continuous change in
the variable.
I also calculated a species-area curve of
the gardens by a regression of log number of
species on log area. Following the Whitehead
and Jones (1969) treatment of island floras, I
deleted the smallest garden on successive cal-
culations to find the point below which the
area detracted from the fit of the curve.
Results
Forty-eight species were coimted in the 29
hanging gardens. Nine taxa were identified
only to genus or family, and four rare plants
could not be identified (Table 2). Richness
ranged from 2 to 20. The frequency of occur-
rence ranged from 1 to 17. Although no spe-
cies occurred in all gardens, the 13 species
occupying five or more gardens are used here
as diagnostic species, because species of mod-
erate constancy are good indicators of varia-
tion in the environment (Mueller-Dombois
and Ellenberg 1974). The many rare species
(35 occupying less than five gardens) are sim-
ply not common enough to give useable cor-
relations with garden environments. For
complete tables of data on species cover, flo-
ristic similarity, and the environmental varia-
bles, see Malanson (1978).
The floristic similarity of the hanging gar-
dens is low, ranging from 0 to 77, and aver-
aging 10.23 for all gardens. This value of sim-
ilarity is close to the results of Nebeker et al.
(1977).
All hanging gardens provide a mesic to
hygric biotope. For water and soil, values of
pH ranged from 5.9 to 7.2 and 6.3 to 7.4, and
values of salinity from 0.3 to 2.5 jumhos/cm
and 230 to 1420 mhos/cm, respectively. In
this study, average soil phosphorous content
ranges from 2.1 to 84 ppm. Direct solar radi-
ation varies from 0 to 683 cal/cm* for the
three-day sum. Average soil depth ranges
from 0.1 to 48.4 cm, but exceeds 10 cm in
only three gardens. Slopes average between
40 percent and vertical. Area ranges from 2
to 100 m- among samples. Most values of iso-
lation are low, 24 gardens are less than 300 m
from the nearest three neighbors. All but four
distances from the Gateway to the Narrows
Table 1. Classes of the environmental variables.
Classes
Variables
1
2
3
4
5
Area (m-)
0-10
10-25
25-50
50
Distance from
trail (m)
50O-ia30
1631-2760
2761-3890
3891-5020
5021-6150
Isolation (m)
0-80
81-160
161-240
241-320
,320
Phosphorous
(ppm)
0-5.0
5.1-15
16-36
37-87
Slope (percent)
1-50
51-125
126-275
276-525
526
Soil depth (cm)
0-1
1-2
2-4
4-8
8
Soil pH
6.3-6.4
6..5-6.6
6.7-6.8
6.9-7.4
Soil salinity
(mhos/cm)
0.1-0.4
0.5-0.8
0.9-1.2
1.3-1.8
1.9-2.5
Solar radiation
(cal/cm-)
0-150
151-300
.301-4,50
451-600
601-750
Water pH
5.9-6.2
6.3-6.5
6.6-6.8
6.9-7.2
Water salinity
(mhos/cm)
0-300
301-600
601-900
900
June 1980
Malanson: Hanging Gardens
181
Trail are clustered bimodally between 500
and 2000 m and between 3000 and 5000 ni.
The results of the species presence tests in-
dicate that there are few relationships be-
tween gradients of the environmental varia-
bles and the presence and absence of
important species in the sampled gardens
(Table 3). Of the 143 tests, only 15 show a
significant difference at p = 0.05, and one
Table 2. Hanging garden plant species.
Species
Frequency
Abies concolor
Acer negiindo
A did utti in cap i7/«.s- veneris
Adiantuni pcdatum
Amaranth tts c^raecizans
Amiphalis margaritaceae
Apocyniim cannahinum
Aquiligia spp.
Aralia racemosa
A rteniisia ludovicia na
Aster eatonii
Berheris rcpens
Brickelia grandiflora
Bromus ciliattts
Calaniagrostis scoptilorum
Cirsiinn arizonicuin
Cystoptcris fragiUs
Dodecatheon pulchellum
Dryopteris filix-nias
Eleocharis sp.
Epipactis gigantea
Equisetttm hyemale
Fraxintis vehitina
Galium aparine
Hepaticae
Heuchera versicolor
Junctts sp.
Lobelia cardinalis
Mimiilus cardinalis
Mimtdtis guttatus
Muhlenbergia andina
Middenbergia mexicana
Nasturtium officinale
Pea nevadensis
Rhus radicans
Rubus leucodennis
Riimex sp.
Salix sp.
Smilacina stellata
Sphagnum sp.
Sphagnaceae
Taraxacum officinalis
Thalictrum fendleri
Viola spp.
unidentified # 1
unidentified #2
unidentified #3
unidentified #4
1
3
15
6
1
4
1
13
12
1
8
3
1
3
5
1
17
9
2
3
3
1
3
4
10
2
2
3
13
1
1
2
2
2
2
3
1
1
7
14
9
4
2
3
1
1
2
1
would expect 7 of the tests to prove signifi-
cant by chance alone. Only solar radiation
consistently returns significant results.
The hanging gardens are smaller isolates
than those described by Whitehead and Jones
(1969) (1.25 to 8.65 ha); yet the changing spe-
cies-area curves are similar. The exclusion of
the four smallest gardens (6 m-) improves the
regression coefficient from 0.55 to 0.67, and
changes the slope from 0.28 to 0.48.
Discussion
The biotopes of the hanging gardens were
delimited in this study by the environmental
variables of soil and water pH and salinity,
soil phosphorous, soil depth, slope, direct so-
lar radiation, area, isolation, and distance to
the Gateway to the Narrows Trail. In gener-
al, the biotopes are within the habitat of the
species, and a particular species composition
is not maintained by the environment. The
distribution of species among the hanging
gardens in the Narrows is not strongly affect-
ed by their tolerances and requirements.
High values of solar radiation seem to mete
against the mosses and ferns, but these values
occur in only 13 percent of the gardens sam-
pled. The great dissimilarity between hang-
ing gardens cannot be attributed to dissimilar
habitats, although they are insular commu-
nities.
In this regard, the hanging gardens of the
Narrows may be very different from those of
the Arches and Canyonlands area. There the
habitat differences found over a wide geo-
graphic area are more likely to be significant
in affecting species presence and garden sim-
ilarity. In the Narrows, the proximity of
many gardens and their probable suscepti-
bility to flash floods prevents a strict com-
parison with the research in eastern Utah.
The Narrows presents a case in which we
must look beyond the structure of the habitat
to find an explanation of plant distributions.
Malanson and Kay (in preparation) consider
disturbance frequencies a likely alternative.
Acknowledgments
I am thankful for the encouragement and
advice of Jeanne Kay, Kimball T. Harper,
and Paul A. Kay on all aspects of this re-
182
Great Basin Naturalist
Vol. 40. No. 2
search. Kezia M. Nielsen, Garry F. Rogers,
Lois A. Amow, and Anne M. Travis helped in
the gathering and preparation of the data.
This research was funded in part by grants
from the University of Utah Student Re-
search Grants in Geography, the Zion Natu-
ral Historv Association, and Sigma Xi, the
Scientific Research Society, and an equip-
ment loan bv Robert W. Austin.
Literature Cited
Clover, E. U., and L. Jotter. 1944. Floristic studies in
the canvon of the Colorado and tributaries.
.\merican Midland Naturalist 32:591-642.
Flow'ers, S. 1959. \egetation of Glen Canyon. Pages
21-61 in A. M. \Voodbur\-. ed. Ecological studies
of flora and fauna in Glen Canyon. University of
Utah .\nthropological Papers 40.
GuE.\so.\, H. .\. 1926. The individualistic concept of
plant association. Bulletin of the Torrey Bot-
anical Club 5.3:7-26.
LooPE, W. E. 1977. Relations of vegetation to environ-
ment in Canyonlands National Park. Unpub-
lished dissertation. Utah State University. Logan.
M.\L.\NSON, G. P. 1978. Distribution of plant species in
hanging gardens of the Narrows, Zion National
Park. Utah. Unpublished thesis. University of
Utah, Salt Lake City.
McELLER-DoMBOis. D.. AND H. Ellenberg. 1974. Aims
and methods of vegetation ecolog) . John Wiley 6c
Sons, New York.
Nebeker, G. T., K. T. 1L\rper. J. D. Brotherson, .\nd S.
L. Welsh. 1977. Characteristics of plants of com-
mon occurrence in hanging gardens of the Colo-
rado Plateau. Utah. Unpublished manuscript.
Brigham Yoimg University. Provo.
R\LNES, J. .\. 1976. Modeling studies of small mammal
trapping, phenolog)', and plant succession in the
Kaiparowits region. Kane County, Utah. Unpub-
lished dissertation. Brigham Young University,
Provo.
R\MENsia-, L. G. 1924. The basic lawfulness in the struc-
ture of the vegetation cover. [In Russian.] \'estnik
opy tnogo dela Sredne-Chemoz. Obi.. Voronezh.
Pages 37-73 excerpted in E. J. Kormondv, ed.
Readings in ecologw Prentice-Hall. Englewood
Cliffs, N.J.
RuzicKA, M. 1958. Anwendung mathematisch— statistis-
cher methoden in der geobotanik, (s\nthetische
bearbeitung von aufnahmen). Biologia. Bratislava
13:&47-661.
Str.\hler, a. H. 1978. Binary discriminant analvsis: a
new method for investigating species-environ-
ment relationships. Ecolog\- 59:108-116.
Welsh, S. L., and C. .\. Toft. 1976. Biotic conununities
of hanging gardens in southeastern Utah in J. R.
Murdock,^S^ L. Welsh, and B. W. Wood, eds.
Navaho-Kaiparowits environmental baseline
studies 1971-1974. Center for Health and Envi-
ronmental Studies. Brigham Young Universitv.
Provo.
Welsh, S. L., .a.nd B. W. Wood. 1976. Structure of a se-
lected hanging garden in Glen Canvon of the
Colorado River drainage in J. R. Murdock. S. L.
Welsh, and B. W. Wood. eds. Navaho-Kaiparo-
wits environmental baseline studies 1971-1974.
Center for Health and Environmental Studies.
Brigham Young University, Provo.
\\ hitehead. D. R.. and C. E. Jones. 1969. Small islands
and the equilibrium theory of insular biogeo-
graphy. Evolution 23:171-179.
Williams, L. D., R. G. B.arry, and J. T. .\ndrews. 1972.
-Application of computed global radiation for
areas of high relief. Journal of Applied Mete-
orology 11:526-5.33.
Wood. B. W., and S. L. Welsh. 1976. Productivit\ of
hanging gardens in J. R. Murdock, S. L. Welsh,
and B. W. Wood, eds. Navaho-Kaiparowits envi-
ronmental baseline studies 1971-1974. Center for
Health and Environmental Studies, Brigham
Young Universitv. Provo.
WooDBiRY. .\. M. 19.33. Biotic relationships of Zion
Canyon. Utah, with special reference to succes-
sion. Ecological .Monographs 3:147-245.
ZNH.\. 1977. Topographic map of Zion National Park
and vicinity. Zion Natural Historv .\ssociation.
Springdale, Utah.
Table 3. Class of environmental variable in which a species was most significanth limited (classes from Table 1).
Variables
Species
Soil Soil Solar Water
Distance Isolation Slope Depth Salinitv Radiation Salinity
Adiantum capillus-veneris
Adiantum pedatum
Aster eatonii
CaUimagrostis seoptilorum
Cijstupteris fragilis
Dodeea th eon p tt Ich ellu m
Mimtdus cardinalis
Smilacina stellata
Sphagnimi sp.
Sphagnaceae sp.
4-5
1-2
I
4-5
3-5
3-5
5
4-5
5
3
1&5
3-4
1-2
SHORT-TERM EFFECTS OF LOGGING ON RED-BACKED VOLES AND DEER MICE
Thomas M. Cainpbcll HI and Tim VV. Clark-
Abstract.— Clearcutting and selective logging effects on red-backed voles (Clethrionomtjs gapperi) and deer mice
(Peromysciis maniculatus) were studied (September-November, 1975; June-October, 1976) in Bridger-Teton Nation-
al Forest, Wvoming. Five selective cuts (total 137 ha) removed 57 percent (range 34-74 percent) of the trees. One
clearcut (9.6 ha) eliminate 84 percent of the trees. Soils remained mesic in selective cuts, but became xeric in the
clearcut. Snap-trapping indicated that voles were most abundant on the imlogged and selectively cut mesic sites (76
percent of 408 captures), whereas deer mice were more common on the xeric clearcut (80 percent of 60 captures).
Species composition remained unchanged on selective cuts following logging (77 percent voles of 256 captures), but
changed from predominantlv voles to mostly deer mice (80 percent of 60 captures) in the clearcut. Intraspecific age
and sex ratios, litter sizes, and morphological measurements were compared between logged and unlogged areas.
The short-term logging effects on the
structure and dynamics (i.e., habitat, num-
bers, and morphological and reproductive
characteristics) of red-backed voles {Cleth-
rionomys gapperi) and deer mice {Peromysciis
maniculatus) were examined on a 646 ha
study area in the Bridger-Teton National
Forest, about 40 km north and 8 km east of
Jackson, Wyoming. It is on the backslope of
an escarpment that nms west from Mt. Leidy
to a point southwest of Toppings Lakes. This
backslope is a series of benches with L5 to 40
percent slopes (Brady 1974) at elevations of
2300 to 2700 m. Soils include loams at the
surface, with siltv clay to clav loam subsoils
(Knight 1973).
Climate is characterized by long, cold win-
ters with deep snow, a short growing season
(average 60 days), and a low mean annual
temperature of 1 C. Snowfall averages 345
cm annually and can occur any month. Mean
annual precipitation is 69 cm, predominantly
snow (Department of Commerce 1975).
Jackson Hole vegetation has been de-
scribed by Read (1952), and Beetle (1961). A
climax spruce-fir {Picea engelmannii-Abies
lasiocarpa) forest covers the study area ex-
cept on recent bums (last 80 years) and at
lower elevations, where lodgepole pine
{Pinus contorta) is dominant, and in areas of
intermixed coniferous forest and meadows
where Douglas fir (Pseudotsuga menziesii)
also occurs. Limber pine {Pinus flexilis) is
sparsely scattered throughout. Understory is
dominated by highbrush huckleberry
{Vaccinium globuhre), grouse whortleberry
(V. scoparium), mountain ash {Sorbus scopu-
lina), and sedges {Carex spp.).
Methods
The density of all trees (dbh > 15 cm),
saplings (dbh 7.5-15 cm), and seedlings (dbh
< 7.5 cm regardless of height) by species was
measured on 10 randomly placed quadrats
(0.004 ha each) in each of the six harvest
blocks before and after logging. Soil beneath
the ground litter was classified as mesic if it
felt damp or xeric if it felt dry.
Small mammals were snap-trapped on
each harvest block just prior to and immedi-
ately after logging and on each harvested
block and adjacent undisturbed sites for up to
one vear thereafter at monthly intervals.
Each sample consisted of 60 traps in three
lines with 10 trap stations 16 m apart per line
and 2 traps 3 m apart per station. Traps were
baited with peanut butter and checked daily
for 3 consecutive days for a total of 180 trap
davs (TD) per sample (one trap set for 24
hours equals one trap day). The species; sex;
age (juvenile or adult); length of body, tail.
'Department of Fishery- and Wildlife Biology, Colorado State University. Fort Collins, Colorado 80522. Reprints: Box 2705, Jackson. Wyoming 83001.
■Department of Biology, Idaho State I'niversity. Pocatello. Idaho 83209.
183
184
Great Basin Naturalist
Vol. 40, No. 2
right hind foot, and right ear; and reproduc-
tive status were recorded for all individuals
trapped. Testes length for males and numbers
of embryos or placental scars for females
were also recorded. Age was based on pelage,
size, and reproductive status.
Statistical tests were based on the Chi
square method of analysis imless otherwise
indicated.
Results and Discussion
Effects of Logging on Vegetation
Collectively, logging altered approx-
imately 24 percent of the study area. Table 1
shows the harvest schedule. Selective cutting
removed a mean of 52, 52, and 77 percent of
the trees, saplings, and seedlings, respective-
ly, and mesic ground conditions persisted for
one year on these sites (Table 2). Clear-
cutting removed 88, 70, and 79 percent of
the trees, saplings, and seedlings, respective-
ly, but ground conditions changed from mes-
ic to xeric within 9 months after harvest. Al-
though not quantitatively measured, logging
and skidding drastically disturbed understory
vegetation and litter in both clear and selec-
tive cuts.
Effects of Logging on Small Mammals
Five species were captured: red-backed
voles, deer mice, western jumping mice
{Zapiis princeps), yellow pine chipmunk (£«-
tamius amoenus), and masked shrews {Sorex
cinereus). Ninety-eight percent of 478 cap-
tures were voles and deer mice; only these
two species are discussed here.
Abundance: Voles were significantly
(P<0.01) more abundant on mesic soils (76
percent of 408 captures. Fig. 1) throughout
the study, but deer mice were significantly
(P<0.01) more numerous on the one xeric
site sampled 10 months after clearcutting (80
Table 1. Logging schedule for the six timber harvest blocks. Toppings Lakes Study Area, Bridger-Teton National
Forest, Wyoming.
Size
Harvest
Harvest schedule
Block
Interval
No.
(ha)
method
Cut
Skidded
(days)
1
19.4
Selective cut
8/19/76
8/21/76
2
2
9.6
Clearcut
9/08/75
9/10/75
2
3
75.0
Selective cut
9/15/76
10/31/76
46
^
8.1
Selective cut
9/10/76
9/10/76
0
5
9.6
Selective cut
9/09/75
8/20/76
315
6
36.4
Selective cut
9/13/76
9/23/76
10
^Logging was suspended after 4 ha were logged aiid it became apparent
that the expected timber voKimes were not there.
Table 2. The changes in tree overstory densities for clear and selective cuts on the Toppings Lakes Study Area,
Bridger-Teton National Forest, Wyoming.
Mean size
of harvest
block (\mf
Mean number per hectare*
before
logging
after
logging
Selective cuts
Trees
Saplings
Seedlings
Clearcut
Trees
Saplings
Seedlings
29.7 ± 27.7 SD
9.6
1064 ± 233
866 ± 84
11027 ± 4124
1213
743
17203
507 ± 47
416 ± 196
2592 ± 545
150
230
3770
"Means and standard deviations for selective cuts (N = 5).
June 1980
Campbell, Clark: Rodent Ecology
185
percent of 60 captures). Koehler et al. (1975)
found similar results in Idaho on undisturbed
niesic sites and on sites xerified by forest fire.
Red-backed voles apparently require a heavy
cover of vegetation or logs (Gashwiler 1959,
LaBue and Darnell 1959, Hooven 1969,
Krefting and Ahlgren 1974) and reside pri-
marily in cool, damp forests (Townsend 1935,
Bailey 1936). The greater diversity of under-
story plants of mesic sites (Daubenmire and
Daubenmire 1968) apparently provide food
and cover for red-backed voles, but deer
mice prefer xeric habitats.
Species composition on logged and un-
logged mesic sites (selective cut #5) did not
differ significantly (P>0.10) one year after
logging (Fig. 2). Conversely, the disturbed
xeric site (clearcut 2) had a highly significant
(P<0.01) change in species composition fol-
lowing harvest (Fig. 2). Preharvest and imme-
diate postharvest data (September and Octo-
ber 1975, respectively) showed a community
composed primarily of red-backed voles (73
percent of 113 captures). Nine to 12 months
after harvest (June to September, following
winter inaccessibility) composition had
changed to 80 percent deer mice of 60 ro-
dent captures. These deer mouse capture
rates indicate a larger population than that
on the original, undisturbed forest. This is at-
tributed to the xerification of this site as a re-
sult of logging. Similar increases of deer mice
were observed by Tevis (1956), Gashwiler
(1959, 1970), Koehler et al. (1975), and Hoov-
en and Black (1976) in forests altered by tim-
ber harvest and forest fires. Clearcuts may be
more attractive to deer mice (Gashwiler
1959) because they tend to move into dis-
turbed, depopulated areas (Stickel 1946).
Age ratios: On newly logged selective
cuts, juveniles outnumbered adults 4.1 : 1.0
for red-backed voles and 5.0 : 1.0 for deer
mice, but 9 to 12 months later juveniles had
decreased significantly (P<0.01) to 0.8 : 1.0
for voles and 0.6 : 1.0 for deer mice. Juveniles
also outnumbered adults on the newly clear-
cut site 1.6 : 1.0 for voles and 1.8 : 1.0 for
deer mice.
90 -■
80
70
60
50 -■
40
30
20
10
RED-BACKED VOLES
DEER MICE
LOGGED
(mesic )
UN LOGGED
(mesic)
LOGGED
(mesic)
UNLOGGED
(xeric)
SELECTIVE CUTS
CLEAR CUTS
Fig. 1. Percent captures of red-backed voles and deer mice on logged and unlogged areas. Bridger-Teton National
Forest, Wyoming. Figures at the top of the bars equal the number of animals trapped.
186
Great Basin Naturalist
Vol. 40, No. 2
Juveniles also comprised 75 percent of all
deer mice captured in a recent burn (Sims
and Buckner 1972). Powell (1972) found
three times as many red-backed vole juve-
niles in a recently blown down forest than in
an undisturbed forest. He concluded that
standing forests were preferred red-backed
vole habitat and adults drove juveniles into
the less preferred, disturbed habitat (presum-
ably via aggressive behavior).
Sex ratios: The overall sex ratio for red-
backed voles was 1.2M:1.0F, comparable to
1.3M:1.0F found in an Oregon study (Gash-
wiler 1959). Adults had an even sex ratio, but
juvenile males significantly (P<0.05) out-
numbered juvenile females (1.5M:1.0F). Vole
sex ratios (adults, juveniles, and total) did not
differ significantly (P>0.05) in unlogged
mesic, logged mesic, or logged xeric sites.
Sex ratios for all deer mice captured (1.0
M:1.0 F); for adults (1.6 M:1.0 F); for juve-
niles (0.6 M:1.0 F); and for logged mesic, un-
70
50 ■-
40
30 --
20
10
y
70
40
30
20 --
10
Block 2 - Clearcut
Sept Jun Jul Aug Sept
1975 1976
Block 5 - Selective Cut
^
# # Total Captures
#— — —• Red-backed Voles
# # Deer Mice
-\ 1 1 H
Block 2 - Unlogged
y
—I 1 1 1 H
Sept Jun Jul Aug Sept
1975 1976
Block 5 - Unlogged
Fig. 2. Numbers of red-backed voles and deer mice trapped on logged and unlogged sites, Bridger-Teton National
Forest, Wyoming (September 1975, Jvme-September 1976).
June 1980
Campbell, Clark: Rodent Ecology
187
logged mesic, or logged xeric sites did not
differ significantly (P>0.05) from 1.0 M : 1.0
F.
Reproduction: Reproductive data for
June-September 1976 showed that most re-
production ceased after August (Tables 3 and
4). Males of both species with scrotal testes
were captured through August but not in
September. Packard (1968) and Clark (1973)
used male rodents exhibiting scrotal testes as
an indication of population breeding. Fe-
males of both species carried embryos and
were lactating from June to September, but
September embryos were near term (Table
4).
Mean litter size and SD based on embryo
and placental scar counts was 6.0 ± 0.9
(range 4-9) for 44 red-backed voles and 6.1
± 0.3 (range 6-7) for 10 deer mice. These
were similar to other studies of red-backed
voles in Crand Teton National Park (Clark
1973) and of deer mice in northwestern
Wyoming (Clark 1975, Long 1964). Repro-
ductive timing or litter sizes for either spe-
cies did not differ between logged and im-
logged areas.
Body measurements: Linear measure-
ments for red-backed voles and deer mice are
shown in Tables 5 and 6, respectively. Adult
females of both species were insignificantly
larger than males. No significant (P>0.05,
AOV) intraspecific age differences in mea-
surement of either sex were observed be-
tween logged and unlogged areas 9 to 12
months after harvest. Male red-backed voles
from this study and nearby Grand Teton Na-
tional Park (N' = 7, Clark 1973) did not differ
significantly (P>0.10, t-test).
Conclusions
Both selective cutting that removed about
57 percent of the trees and clearcutting of
spruce-fir forest resulted in an increase of
juvenile red-backed voles and deer mice two
weeks to a month after logging that was not
apparent 9 to 12 months later. Clearcutting
changed species composition 9 to 12 months
later from predominantly red-backed voles
on unlogged areas to predominantly deer
mice. The change was attributed to soil xeri-
Table 3. Male reproductive condition of red-liacked voles and deer mice, Bridger-Teton National Forest. Wyo-
ming (June-September 1976).
Number
of
Number of
red-hacked voles
deer mice
Condition
June
J"iy
Aug.
Sept.
Total
June
July
Aug. Sept.
Total
Scrotal
9
7
12
0
28
4
9
1
0
14
Nonscrotal
0
0
11
9
20
0
1
13
6
20
Total
9
7
2.3
9
48
4
10
14
6
34
Table 4. Reproductive condition of red-backed vole and deer mouse adult females. Bridger-Teton National For-
est, Wyoming (June-September 1976).
N
umber
of
Number
of
red-backed voles
deer mice
Condition
June
July
Aug.
Sept.
Total
June
July
Aug.
Sept.
Total
Nimiber of females examined
8
5
13
18
44
1
6
10
1
18
Percent pregnant
50
80
23
72
54
100
67
20
100
44
Niunber of embrvos
Mean
6.3
6.8
6
5.9
6.1
6
6.5
6
6
6.3
Range
6-7
5-9
6
4-9
4-9
6
6-7
6
6
6-7
Percent with placental scars
50
40
-"-
28
48
0
.33
80
0
.56
Number of placental scars
Mean
5.5
6
6
.5.6
5.8
0
6
6
0
6
Range
5-6
6
6
4-6
4-6
0
6
6
0
0-6
Number of females lactating
4
6
13
17
40
1
0
10
1
17
188
Great Basin Naturalist
Vol. 40, No. 2
fication on the clearcut. Voles continued to
predominate on selectively cut sites, and soils
remained mesic there. For both species adults
consistently outnumbered juveniles in June,
July, and August in both logged and unlogged
areas, but juveniles outnumbered adults in
areas newly logged in September. Sex ratios,
timing of reproduction, litter size, and body
measurements for both species did not differ
significantly between unlogged, selectively
cut, or clearcut areas 9 to 12 months after
logging.
Acknowledgments
This study was supported by the Bridger-
Teton National Forest, Jackson, Wyoming.
Special thanks is extended to Mr. G. GreuU,
Wildlife Biologist, U.S. Forest Service. Mr.
W. Barmore offered valuable criticism of the
manuscript.
Table 5. Body measurements of 258 red-backed voles from unlogged, clearcut, and selectively cut areas, Bridger-
Teton National Forest, Wyoming (June-September 1976).
Sex and age*
Mean length, SD, and (range) in mm
Body
Tail
Ear
Hind Foot
Control
M2+ (n=18)
F2+ (n = 23)
Ml-(n = 40)
Fl-(n = 26)
139.4 ±8.8(1 19-152)
143.4 ±10.9(118-158)
115.2 ±12.1(88-137)
112.2 ±10.5(90-132)
39.1 ±2.6(34-45)
40.5 ±4.4(26-47)
,33.0 ±4.5(26-43)
.33.2 ±3.7(25-42)
15.8 ±1.8(1,3-21)
16.5± 1.1(14-19)
14.9± 1.8(10-17)
14.9± 1.7(10-18)
18.4 ±0.9(17-19)
18.0 ±1.0(16-19)
16.9 ±1.2(15-18)
17.0± 0.7(15-18)
Clearcut
M2+ (n = 3)
F2+ (n = 4)
Ml-(n = 4)
Fl-(n = l)
147.7 ±6.4(144-155) 41.0 ±3.5(39-45) 14.3 ±0.6(14-15) 18.7 ±0.6(18-19)
139.6 ±10.4(125-147) 40.3 ±2.9(37-44) 17.0± 0.8(16-18) 17.8 ± 1.0(17-19)
113.3 ±13.8(100-130) 33.5 ±6.2(27-40) 13.0± 2.0(12-16) 16.8 ±0.5(16-17)
117.0- - .35.0- - 17.0- - 18.0- -
Selective cut
M2+ (n = 27)
F2+ (n = 17)
Ml-(n = 59)
Fl-(n = 36)
138.9± 10.1 (115-160) 40.8 ±3.5(26-48) 15.9± 1.7(12-18) 18.4 ± 1.0(16-19)
142.7 ±11.2(122-159) 40.6 ±4.7(37-47) 15.9± 1.8(11-18) 17.9± 0.8(16-19)
121.9 ±8.5(96-1.37) ,35.6 ±.3.5(26-42) 15.6± 1.6(11-19) 17.3 ±0.7(16-19)
115.0± 10.7(93-138) .3,3.5 ±,3.6(25-40) 15.4 ± 1.9(10-18) 17.2 ±0.7(16-19)
*2+ = adult; 1- = juvenile.
Table 6. Body measurements of 118 deer mice from unlogged, clearcut, and selectively cut areas, Bridger-Teton
National Forest, Wyoming (June-September 1976).
Mean length, SD, and (range) in mm
Sex and age°
Body
Tail
Ear
Hind foot
Control
M2-I- (n=ll)
F2+ (n = 5)
Ml-(n = 8)
Fl-(n = 9)
Clearcut
M2+ (n = 12)
F2+ (n = 10)
Ml-(n=9)
Fl-(n=17)
Selective cut
M2+ (n = 12)
F2+ (n = 4)
Ml-(n = 6)
Fl-(n=15)
'2+ = adult; 1- = juvenile.
157.5± 11.1(140-17,5)
156.6± 10.1(146-170)
139.4 ±5.0(133-146)
146.0±6.,3(137-155)
68.4 ±4.8(62-77)
69.2±4..3(63-7.3)
6.3.9 ±2.0(61-66)
67.0 ±3.4(62-72)
158.7± 11.1(148-175) 69.8 ±5.0(6,3-77)
166.3 ± 14..3(144-188) 7.3.4 ± 7.1(62-85)
146.4 ±7.2(133-156) 67.4 ±3.6(61-74)
142.5 ± 7.6(130-155) 63.4 ± 5.2(55-72)
154.1 ± 10.1(141-174) 67.4 ±5.2(61-76)
160.1 ± 17.8(145-183) 69.8 ± 8.3(62-80)
146.8 ±6.6(139-157) 66.5 ±3.7(64-74)
143.2 ±9.6(128-160) 65.0 ±5.0(57-74)
18.5 ±0.8(1.3-20)
18.8 ±0.5(1.3-19)
18.3 ±1.0(17-20)
18.8 ±0.8(17-20)
18.0 ±1.7(15-20)
18.3 ±1.3(17-20)
18.7 ±1.0(18-20)
17.6 ±1.8(14-20)
18.3 ±1.4(15-20)
18.5± 1.0(17-19)
18.2 ±0.8(17-19)
17.9 ±1.5(16-20)
18.4 ±0.9(19-20)
19.6 ±0.9(19-21)
19.1 ±0.6(18-20)
18.6± 1.0(17-20)
19.4 ±0.7(19-21)
19.3 ±0.7(18-20)
19.1 ±0.9(18-20)
18.5 ±1.0(17-20)
19.1 ±0.,5( 18-20)
19.3 ±1.0(18-20)
19.2 ±1.0(18-20)
18.8 ±0.8(17-20)
June 1980
Campbell, Clark: Rodent Ecology
189
Literature Cited
Bailey, V. 1936. The mammals and life zones of Oregon.
North American Fauna 55. 416 pp.
Beetle, A. A. 1961. Range survey of Teton County,
Wyoming. Part 1: Ecology of range resources.
Univ. Wyoming Agr. Exp. Stn. Bull. No.
376R:l-42.
Brady, S. 1974. Environmental impact study of Top-
pings Lake timber sale. Bridger-Teton National
Forest, Jackson, Wyoming. Mimeo. 20 pp.
Clark, T. W. 1973. Local distributions and interspecies
interactions in microtines. Grand Teton National
Park, Wyoming. Great Ba.sin Nat. 33:205-217.
1975. Ecological notes on deer mice in Grand Te-
ton National Park, Wvoming. Northwest Sci.
49(1): 14-16.
Clark, T. W., and T. M. Campbell. 1976. Population
organization and regulatory mechanisms of pine
martens in Grand Teton National Park, Wyo-
ming. Paper presented at Conference on Re-
search in National Parks, Nov., 1976, New Or-
leans, Louisiana.
Daube.nmire, R., and J. B. Daubenmire. 1968. Forest
vegetation of eastern Washington and northern
Oregon. Washington .\gr. Exp. Stn. Tech. Bull.
No. 22, 60 pp.
Department of Commerce. 1975. Climatological data
of Wvoming. Natl. Ocean, and Atmos. Admin.,
.\sheville, N.C. Nos. 1-12.
Gashwiler, J. S. 1959. Small mammal study in west-
central Oregon. J. Mammal. 40(1): 128-1.39.
1970. Plant and mammal changes on a clearcut in
west-central Oregon. Ecology 51(6): 1018-1026.
Hoffman, G. R. 1960. The small mammal components
of six climax plant associations in eastern Wash-
ington and northern Idaho. Ecology
41(.3):571-572.
HoovEN, E. F. 1969. The influence of forest succession
on populations of small animals in western Ore-
gon in H. E. Black, ed. Wildlife and reforestation
in the Pacific Northwest. Oregon State Univ.,
Corvallis.
HoovEN, E. F., and H. C. Blac:k. 1976. Effects of some
clear-cutting practices on small-mammal popu-
lations in western Oregon. Northwest Sci.
50(4): 189-208.
Knight, C. A. 1973. Soil resource inventory manual.
Bridger-Teton National Forest, Jackson, Wyo-
ming.
KoEHLER, G. M., W. R. Moore, and A. R. Taylor.
1975. Preserving the pine marten: management
guidelines for western forests. Western Wildlands
2(3) :3 1-36.
Kreftinc, L. W., and C. E. .\hlgren. 1974. Small mam-
mals and revegetation changes after fire in a
mixed conifer-hardwood forest. Ecology
55(6): 1391-1398.
LaBue, J., and R. M. Darnell. 1959. Effect of habitat
disturbance on a small mammal population. J.
Mammal. 40(3):425-437.
Long, C. A. 1964. Comments on reproduction in the
deer mouse of Wyoming. Trans. Kansas Acad.
Sci. 67:149-153.
Love, J. D., and J. C. Reed, Jr. 1968. Creation of the
Teton landscape. Grant Teton Nat Hist, .\ssoc.
Publ., Moose, Wyo. 120 pp.
Packard, R. L. 1968. An ecological study of the fulvous
harvest mouse in eastern Texas. Amer. Midi. Nat.
79:68-88.
Powell, R. A. 1972. A comparison of populations of bo-
real red-backed voles {Cletlirionomys gapperi) in
tornado blowndown and standing forest. Can.
Field Nat. 86(4):377-379.
Reed, J. F. 1952. The vegetation of the Jackson Hole
Wildlife Park, Wyoming. Amer. Midi. Nat.
48(3): 700-729.
Sims, P., and C. H. Buckner. 1972. The effects of clear-
cutting and burning of Pinus barksiana forests on
the populations of small mammals in south-east-
ern Manitoba. Amer. Midi. Nat. 90(1):228-231.
Stickel, L. F. 1946. The source of animals moving into
a depopulated area. J. Mammal. 27(4):302-307.
Tevis, L. 1956. Response of small mammal populations
to logging of Douglas fir. J. Mammal.
37(2): 189-196.
Townsend, N. T. 1935. Studies on small mammals of
central New York. Roosevelt Wildl. .\nnals.
4:6-120.
TERMINAL BUD FORMATION IN LIMBER PINE
Ronald M. Lanner' and James A. Bryan'
,\bstr.'\ct.— The progress of bud development was studied in limber pines growing in the mountains of north-
eastern Utah. Initiation of new bud scales began in mid-June, several weeks after elongation of the current shoot had
begim. Needle primordia first appeared in September and continued to form through the winter, until all were pres-
ent in May. This winter activity is believed to be fostered by surface temperatures on the terminal buds considerably
higher than ambient air temperatures.
The annual shoot of limber pine (Pinus
flexilis James), a five-needled species com-
mon in the Rocky Mountains, consists of a
monocyclic spring shoot formed by the elon-
gation of a winter bud. This is the most fa-
miliar shoot development pattern in northern
pines and has been classified as the Resinosa
pattern (Lanner 1976). But even among spe-
cies of this habit, there is diversity in the de-
velopmental schedule of bud morphogenesis.
For example, in some species formation of
the new winter bud begins while the old one
is still elongating, but in others bud formation
is delayed until the cessation of current-sea-
son shoot growth. Further, the timing of
.short-shoot and needle morphogenesis is also
subject to variation. In this report we de-
scribe the annual cycle of development of
limber pine terminal buds to resolve the
questions of when the short shoots and nee-
dles are formed, both in terms of calendar
date and in regard to the growth stage of the
spring shoot.
The trees studied grow at an elevation of
2130 m on a steep southeast slope in Logan
Canyon, northeastern Utah. They vary from
12 to 27 cm DBH and 4 to 7 m in height. At
each of 18 collection dates during 1978, 2
terminal buds of vigorous first-order branches
in the upper crown were harvested from
each of at least 2 of the 18 study trees. Buds
were stored in formaldehyde-acetic acid-eth-
yl alcohol (FAA), dissected with standard mi-
cro-dissecting tools, and examined at 12-lOOx
with a Wild M-5 stereomicroscope. Two
shoots had steel pins inserted at the base of
tlie bud as a datum for observations of shoot
elongation.
On each of the harvested buds we dis-
sected at least two short-shoot budlets from
the proximal (basal) end of the bud, and two
from the distal (apical) end. We counted the
budlet scales (future fascicle sheath scales)
and needle primordia, if any. The study took
place during a single calendar year, so we ac-
tually observed the late development and
elongation of the 1978 spring shoot and the
early development of the 1979 spring shoot.
Ideally, a study of this kind should begin with
early morphogenesis of a bud and end with
the maturation of the resulting shoot.
Results
One of the marked shoots started to elon-
gate during the interval 20-30 May, and the
other during the interval 30 May-5 June.
These shoots completed their elongation
growth prior to 8 July and 13 July, respec-
tively. Final lengths of these shoots were 30.5
and 21.5 mm. Pollen was shed during the pe-
riod 9-13 July.
Initiation of the primary bud scales (cata-
phvlls) of the newlv forming terminal bud be-
gan between 11 and 18 June. After cataphylls
formed, meristems appeared in the axils of
most of them. These axillary meristems be-
came the apical meristems of the short-shoot
'Department of Forestry and Outdoor Recreation, Utah State University, Logan, Utah 84322.
190
June 1980
Lanner, Bryan: Limber Pine
191
budlets that would later become needle fas-
cicles. Scales formed on these budlets are fu-
ture fascicle sheath scales. Sheath scale ini-
tiation began in proximal budlets in early
July, and in distal budlets in early August.
Scale production continued in both types of
budlets luitil about mid-Januarv.
The earliest needle primordia started to
appear in proximal budlets in late August and
continued to appear over a period of three
months. Needle primordia in distal budlets
did not begin to form until about mid-No-
vember, but continued appearing up to early
May, about 5.5 months later.
When bud scale initiation began, about
two-thirds of the elongation growth of the
marked shoots had been completed. Bud
scale initiation probably continued through-
out the remainder of the elongation period.
Sheath scale initiation began in proximal
budlets about the time shoot elongation was
ceasing, and it began in distal budlets after
the cessation of shoot growth. When needle
primordia started to form, shoot growth had
been inactive for several weeks. The needle
primordia formed in the spring developed
prior to the onset of shoot elongation in late
May.
When the first buds were harvested 22 Jan-
uary 1978, short-shoot budlets from proximal
parts of the terminal bud contained 10-12
scales (x = 11.2) and all had their full com-
plement of 5 needle primordia. But budlets
located at the distal end of those terminal
buds tended to have fewer scales (8-11; x =
10.0) and averaged only 1.7 needle primordia
(Table 1). The difference in scale number was
maintained even beyond 8 May, when, for
the first time, all the distal budlets contained
their full complement of 5 needle primordia
(Table 1).
Discussion and Conclusions
The initiation of bud scales was first noted
on 18 June, when the 1978 shoot had attained
68 percent of its final length, showing that
bud morphogenesis began during the period
Table 1. Progress of sheath scale and needle initiation in proximal and distal short shoot budlets of first-order
terminal buds of Limber Pine, and of shoot elongation.
Proximal budlets
Needle
Distal budlets
Needle
Length of
1978
number
rimordium
primordium
bud/shoot as
Scale
ninnber
Scale
number
number
percent of
final length
1978 Date
X
range
X
X
range
X
1978 annual
shoot
22 Jannary
11.2
10-12
5.0
10.0
8-11
1.7
31.5
21 March'
10.0
9-12
5.0
9.6
9-10
2.5
31.5
12 April
10.3
9-12
5.0
9.5
9-10
3.3
31.5
8 May
10.0
9-11
5.0
10.6
10-11
5.0
31.5
20 May
9.5
8-11
5.0
9.5
9-10
5.0
31.5
4 Jnne
10.0
10
5.0
9.5
9-10
5.0
45.0
1 1 Jnne
13.0
13
5.0
1979 annual
11.0
shoot
11
5.0
55.5
1 1 Jnne
0
0
0
0
0
0
55.5
IS June
0
0
0
0
0
0
68.0
2Jnly
0
0
0
0
0
0
95.0
13 July
2.0
2
0
0
0
0
100
23 July
4.5
3-6
0
0
0
0
1(X)
2 August
5.0
5
0
0
0
0
1(K)
16 August
6.0
6
0
0.75
0-2
0
100
26 August
6.0
.5-7
0
2.3
2-3
0
1(X)
15 September
8.0
8
5.0
7.3
6-9
0
100
5 October
8.3
8-9
3.0
7.3
6-9
0
100
15 November
8.8
8-10
2.2
7.0
6-8
0
100
9 December
8.0
8
5.0
8.0
8
1.8
100
192
Great Basin Naturalist
Vol. 40, No. 2
of shoot elongation. In this regard limber
pine resembles P. strobus L. and P. lamber-
tiana Dougl. (Lanner 1976). It provides fur-
ther evidence that shoot elongation does not
inhibit the initiation of lateral structures on
the shoot apical meristem. Bud morphoge-
nesis in limber pine is delayed, however, in
comparison to that of lodgepole pine (P. con-
torta Dougl.), one of its associates in this area
(Van Den Berg and Lanner 1971). In lodge-
pole, initiation of the new bud and elonga-
tion of the shoot began almost simultaneously
early in May.
A given stage of development— i.e., attain-
ing a certain number of sheath scales or a
certain number of needle primordia— is
reached earlier in proximal than in distal
short shoots. Thus, in the 1978 winter buds,
the full complement of 5 needle primordia
was present in proximal short shoots in Janu-
ary collections, but was not found in distal
short shoots until May. This is another case of
the developmental gradient in short-shoot
maturation described in the more complex
buds of lodgepole pine (Van Den Berg and
Lanner 1971), the much larger buds of slash
pine, P. elliottii Engelm. (Lanner 1978), and
in eastern white pine (Owston 1969).
Perhaps the most unusual finding reported
here is that morphogenetic activity contin-
ued in wintering limber pine buds. Buds col-
lected early in 1978 showed periodic increas-
es in needle primordium number in distal
budlets. Buds collected late in 1978 showed
consistent increases in needle primordium
number in both proximal and distal budlets,
and in sheath scale number in distal budlets
(Table 1). Yet temperatures at a nearby tem-
porary weather station at a comparable ele-
vation fell as low as -21 C in January and
-12 C in November and December (pers.
comm., S. A. Loomis). In lodgepole pine
studied just a few miles from this site, budlets
overwintered with less than their full com-
plements of sheath scales and needle pri-
mordia, but no changes were noted during
the winter (Van Den Berg and Lanner 1971).
Budlets of slash pine actively initiated scales
and needle primordia during the winter
months, but this was in the much milder cli-
mate of Florida (Lanner 1978).
Seed cone primordia have also been re-
ported as morphogenetically active during
the winter. Duff and Nolan (1958) observed
changes in ovulate strobili of red pine (P. resi-
nosa Ait.) between October and January in
the cold climate of Chalk River, Ontario. Gif-
ford and Mirov (1960) also reported female
strobilus development in ponderosa pine (P.
ponderosa Laws.), but in the milder climate
of the Sierra Nevada west slope.
Such meristematic activity may be permit-
ted by surface temperatures considerably
higher than those of the ambient atmosphere.
For example, Tranquillini and Turner (1961)
have reported maximum monthly temper-
atures of Swiss stone pine (P. cernbra L.) nee-
dles 2.7 and 7.4 C higher than air temper-
atures during November and March,
respectively, in the Austrian Alps. In March,
needles reached a maximum of 18.4 C,
though the mean air temperature for that
month was only 0.3 C. At our study site, even
in January, the coldest month of the year, air
temperature on three occasions attained al-
most 6 C. Bud surface temperatures may
have reached as high as 12 C or more, well
above the apparent threshold for meristemat-
ic activity.
Literature Cited
Duff, C. H., and N. J. Nolan. 1958. Growth and mor-
phogenesis in the Canadian forest species. III.
The time scale of morphogenesis at the stem
apex of Pintis resinosa Ait. Canadian J. Bot.
36:687-706, 4 pis.
GiFFORD, E. M., Jr., and N. T. Mirov. 1960. Initiation
and ontogeny of the ovulate strobilus in ponde-
rosa pine. Forest Sci. 6:19-25.
Lanner, R. M. 1976. Patterns of shoot development in
Pinus and their relationship to growth potential.
Pages 223-243 in M. G. R. Cannell and F. T.
Last, eds. Tree physiology and yield improve-
ment. Academic Press, New York and London.
1978. Development of the terminal bud shoot of
slash pine saplings. Forest Sci. 24:167-179.
Owston, P. W. 1968. The shoot apex in eastern white
pine: its stnicture, seasonal development, and
variation within the crown. Canadian J. Bot.
47:1181-1188.
Tranquillini, W., and H. Turner. 1961. Untersuchu-
ngen iiber die Pflanzentemperaturen in der sub-
alpinen Stufe mit besonderer Beriicksichtung der
Nadeltemperaturen der Zirbe. Mitteil. Forstlich.
Bundes-Versuchs. Mariabrunn 59(1): 127-151.
Van Den Berg, D. A., and R. M. Lanner. 1971. Bud de-
velopment in lodgepole pine. Forest Sci.
17:479-486.
STINGER UTILIZATION AND PREDATION
IN THE SCORPION PARUROCTONUS BOREUS
Bnice S. Gushing' and Anne Matherne'
.\bstr.\ct.— The iitihzution of the stinger and the predatory technique of the scorpion, Paruroctanus boretis, was
studied under laboratory conditions. During the study, 83 feedings were observed. Age of the scorpions and the per-
centage of prey stung by them were used to classify the scorpions into groups. The scorpions aged 1.3-61 days always
stung prey, .\fter 62 days the scorpions began to selectively utilize the stinger. Utilization declined until it reached
.30 percent in the adult stage. The stinger is apparently necessary for prey capture only in the earlv life stages.
The role of the stinger in scorpion behav-
ior has never really been studied or estab-
lished. The available information is vague
and inconsistent. Alexander (1959) reported
that different groups of scorpions utilize the
stinger differentially. Pocock (1893) and
Stahnke (1966) stated that scorpions paralyze
prey only when it does not submit to passive
consumption. Finally Hadley and Williams
(1968) reported that Vepvis confusus and
Paruroctonus mesaensis and P. baergi stung
prey (Williams 1972). These reports leave the
adaptive significance of this potent device in
doubt. In this study, we attempted to estab-
lish the use of the stinger through controlled
experimentation and observation.
Methods and Materials
Scorpions were collected from south-
western Oregon on the Malheur National
Wildlife Refuge and adjacent region. They
were kept in 10-gallon terrariums containing
three inches of soil and flat rocks from the
natural habitat. In most ca.ses, two scorpions
were placed in each tank to induce in-
traspecific responses. A large population of
grasshoppers and a small number of beetles,
crickets, and termites were found in the area,
and the.se were selected in relative propor-
tions as the prey.
To maintain natural conditions, the terra-
riums remained outdoors except during peri-
ods of observation. Observations were con-
ducted at night using red light. Red light
provided visibility for us, but apparently did
not affect the scorpions, which possess vision
in the blue and ultraviolet wavelengths
(Machan 1968).
Results
Over a two-year period several different
groups of Paruroctonus boreus were ob-
served. Predation techniques were the same
for scorpions of all ages and sizes. Emergence
from cover occurred between 2130 and 2300
hours. If emergence did not occur by 2.300
the scorpions did not forage that night. After
emergence, contact with prey was estab-
lished through random encounter or active
stalking. When actively stalking, the scor-
pions traveled with the pedipalps extended
forward and held apart at a distance approx-
imately equal to the maximum width of the
abdomen. The telson was arched over the ab-
domen with the caudal vesicle above the
midabdomen. When a potential prey was de-
tected the scorpions rushed it.
Upon contact, the scorpions used their
pedipalps to grasp the prey by one or more
appendages. If stinging occurred at all, it oc-
curred at this time. The telson was arched
over the abdomen and at the same time the
abdomen was quickly raised. This imparted a
downward stabbing motion which allowed
'Department of Wildlife Biology, University of Montana. Missoula, Montana 59812.
"Department of Zoology, Louisiana State University. Baton Rouge, Louisiana 7080.3.
193
194
Great Basin Naturalist
Vol. 40, No. 2
the stinger to penetrate the prey's abdomen.
Resistance by the prey subsided within one
minute after it was stung. Whether or not the
prey was paralyzed, it was held motionless in
the pedipalps for 10 to 30 minutes and then
transferred to the chelicerae. Upon leaving
the site of capture the prey was slung, ven-
tral side up, onto the cephalothorax.The prey
was carried about the terrarium in this posi-
tion for up to several hours. When walking,
the scorpions waved their pedipalps in front
of their path in a "blindman" fashion, a slow
exploratory touching.
Except when scorpions were with young,
prey was taken beneath cover for con-
sumption. Consumption time varied between
2 and 48 hours. Feeding began at the head of
the prey and continued until the prey was
consumed. The hard exoskeleton of beetles
was left as an empty husk. Several of these
husks were found with scorpions in the field.
All scorpions used the same feeding tech-
niques. However, the scorpions also under-
went regular periods of nonfeeding which
lasted up to five months.
The stinger was removed from two adult
scorpions. These scorpions fed six times and
utilized the same techniques as unimpaired
individuals. However, they never attempted
to sting any of the prey.
During intraspecific aggression or canni-
balism, the method of capture was as de-
scribed above with minor modifications. If
there was a significant difference in size, the
smaller scorpion attempted to avoid conflict,
but the larger one often pursued. When ag-
gression occurred, the scorpions grasped each
other by the pedipalps and repeatedly at-
tempted to sting. A size difference always re-
sulted in the death of the smaller scorpion.
Consumption proceeded normally after im-
mobilization. In two instances of the scor-
pions being the same size, both animals were
killed.
Immature scorpions did not capture prey
until they were 13 days old. Prior to this time
they consumed their casting left at birth and
their first exuvium. After the juveniles dis-
persed at 9 to 11 days, the female began
feeding with an alteration in feeding tech-
nique. The adult female consumed prey in
the open and its young congregated about
the adult's cephlathorax. On day 14 the
young began to capture prey. Table 1 sum-
marizes the percentages of prey stung by the
early instars and all other age groups.
Discussion and Analysis
No age group beyond 84 days was ob-
served due to our inability to keep juveniles
alive. This resulted mainly from a high de-
gree of cannibalism and mishandling of a few
remaining scorpions. Data on cannibalism
were excluded from Table 1 because of the
bias it would introduce inasmuch as in-
traspecific aggression always elicited stinger
utilization. Two scorpions per tank and the
limited dispersal range of the young led to an
unnatural increase in incidents of cannibal-
ism. Data on the six feedings by the scorpions
with stinger removed were also excluded due
to their inability to sting.
During active stalking, the scorpions
rushed prey. Some stimulus must have been
present which alerted the scorpions. Paruroc-
tonus boreus, like other desert scorpions, may
be able to detect and utilize Rayleigh waves
for prey location (Brownell 1977). A Rayleigh
wave is a slow-moving secondary vibration
created by movement and propagated
through sand.
The stinger was not essential for feeding by
the adult scorpions. A low percentage of prey
was paralyzed, and stinger-impaired individ-
uals were able to feed without difficulty.
Scorpions are also capable of surviving pro-
longed periods without food (Stahnke 1966).
Considering these factors, we suggest that the
amount of food which would be lost to an
adult incapable of stinging would not have a
significant or deleterious effect upon its sur-
vival.
If stinging occurs, it is triggered by two
stimuli. One, as stated by Pocock (1893) and
Table 1. Relationship between age and stinger utili-
zation.
Age group
Number of
Number
Percent
in days
feedings
stung
stung
01-12°
0
0
00.0
13-61
40
40
100.0
62-84
13
9
69.2
Adult
30
10
30.0
"Fed on exuviae and adult pellets.
June 1980
CusHiNx;, Matherne: Scorpion Predation
195
Stahnke (1966), is an attempt by the prev to
resist capture. This is not the onlv stiuiuhis.
Struggling hard-bodied or powerful prey,
such as grasshoppers, were stung. Termites
and other soft-bodied prey were held in the
pedipalps until resistance subsided. The fac-
tors that elicited a sting were resistance in
combination with the characteristics of the
prey species. This strongly suggests a form of
selective stinger utilization.
Tlie selection process develops over time.
The first instars feed on the exuviae and pos-
sibly on small pellets dropped by the adult
during feeding (Stahnke 1966). Utilization of
pellets for food suggests the reason for the
change in feeding technique by the adult fe-
male, with the young gathered about her
cephalothorax. The next age group, 13-61
days, paralyzed all prey (Table 1). The cause
of this may be that the pedipalps were not
sufficientlv developed at this point to hold
prey against struggle. Therefore, in order to
insure the maximimi number of feedings and
promote growth and development, the juve-
niles must sting prey at first contact. As de-
velopment occurs the pedipalps strengthen,
and certain prey types, such as small ter-
mites, no longer must be paralyzed. This
would reduce the use of toxin and be energy
efficient by reducing the manufacture of
more toxin.
Stinger utilization drops from 100 to 30
percent in the adults (Table 1). This decrease
began about the second month and continued
until the adult stage. The actual percentage
utilization in the group aged 62-84 days may
have been biased in that this group was not
fed a representative class of prey, but instead
was fed whatever small insects and arachnids
happened to be available. Nevertheless, this
group still demonstrates the beginning of the
process of differential selection for stinger
utilization in that not all prey was stung.
In conclusion, the stinger fimctions as a
necessary device for prey capture by the
early instars. As physical development oc-
curs, the pedipalps can hold certain types of
prey and there is a reduction in the use of the
stinger. This decline continues until the adult
stage, where only a small percentage of prey
is stung and these are not essential for survi-
val. However, the stinger is still utilized for
intraspecific aggression and possibly for de-
fense.
Acknowledgment
We thank Dr. John Mates, University of
California, Davis, for his early help in this
project, and Dr. Andrew Sheldon, Depart-
ment of Zoology, University of Montana, for
his critical review of this manuscript.
Literature Cited
Alex.\nder, a. J. 1959. A survey of the l)iologv of scor-
pions of South Africa. Afr. Wildl. 13(2):99-106.
Brownell, p. H. 1977. Conipressional and surface
waves in sand: used by desert scorpions to locate
prey. Science 197:479-481.
RIDLEY, N. F., AND S. C. WiLLiAMs. 1968. North .Ameri-
can scorpions in relation to feeding. Ecology
49(4):726-734.
Macha.n, L. 1968. Spectral sensitivity of scorpions' eyes
and possible role of shielding pigment effect. J.
Exp. Biol. 49(1):95-105.
PococK, R. I. 189.3. Notes upon the habits of some living
scorpions. Nature. 48:1()4-I()V.
Stahnke, H. L. 1966. Some aspects of scorpion behavior.
Southern Calif. Acad. Sci. Bull. 62(2)65-80.
W'illia.ms, S. C. 1972. Four new scorpion species be-
longing to the genus Paruroctonus. Occasional
Papers^Calif. Acad. Sci., No. 94.
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TABLE OF CONTENTS
Feeding ecology of Gila boraxobius (Osteichthyes: Cyprinidae) endemic to a thermal
lake in southeastern Oregon. Jack E. Williams and Cynthia D. Williams 101
First record of the pallid hat (Antrozous palliclus) from Montana. Jeff Shryer and
Dennis L. Flath '. 115
A Cliiivcantliiuiti spider bite. Dorald M. ,\llred 116
Identity of narrow-leaved Chitisotliamnits visciclifloni.s (Asteraceae). Loran C. .\nder-
ll'i
Ribulose diphosphate carboxylase activities in cold-resistant common mallow, Malva
neglecta Wallr. and a cold-sensitive tomato, Lycopersicon esculentum L., Ace
55 var. William R. Andersen and Jack D. Brotherson 121
Recovery of Gambel oak after fire in central Utah. L. M. Kunzler and K. T. Harper .. 127
Relationships among total dissolved solids, conductivity, and osmosity for five A;-
temia habitats (Anostraca: Artemiidae). Nicholas C. Collins and Gray Stirling 1.31
Spawning of the least chub (loticJtthijs phlegt'thuntis). Thomas M. Baiigh 1.39
Transferrin polymorphism in bighorn sheep, Otis canaclcn.^is, in Colorado. Patrick
W. Roberts, Donald J. Nash, and Robert E. Keiss 141
The genus Eriogoniim Michx. (Polygonaceae) and Michel Gandoger. James L. Reveal 143
Parasites from two species of suckers (Catostomidae) from southern Utah. J. Craig
Brienholt and Richard A. Heckmann 149
Soil water withdrawal and root distribution under grubbed, sprayed, and undis-
turbed big sagebni.sh vegetation. David L. Sturges 157
Swarming of the western harvester ant, Pogononnjnnex occiclentalis. Dorald M.
Allred 165
Relationship between environmental and vegetational parameters for understory and
open-area communities. William E. Evenson, Jack D. Brotherson, and Rich-
ard B. Wilcox 167
Seasonal activity pattern of Columbian ground sqviirrels in the Idaho primitive area.
Charles L. Elliott and Jerran T. Flinders 175
Habitat and plant distributions in hanging gardens of the narrows, Zion National
Park, Utah. George P. Malanson 178
Short-term effects of logging on red-backed voles and deer mice. Thomas M. Camp-
bell III and Tim W. Clark 183
Terminal bud formation in limber pine. Ronald M. Lanner and James A. Bryan 190
Stinger utilization and predation in the scorpion Paiuwctoniis boreits. Bruce S. Gush-
ing and -•Vnne Matherne 193
HE GREAT BASIN NATURALIST
lume 40 No. 3
September 30, 1980
Brigham Young University
MUS. COMP. ZOOl_.
MAY 14'^ '
HARVAl-vO
UMfVERSI-TY
GREAT BASIN NATURALIST
Editor. Stephen L. Wood, Department of Zoology, Brigham Young University, Provo, Utah
84602.
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Stanley L. Welsh, Botany; Clayton M. White, Zoology.
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tural Sciences; Ernest L. Olson, Director, Brigham Young University Press, University
Editor.
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U-80 650 48558
ISSN 0017-3614
The Great Basin Naturalist
Published at Provo, Utah, by
Brigham Young University
ISSN 0017-3614
Volume 40
September 30, 1980
No. 3
SPATIOTEMPORAL VARIATION IN PHENOLOGY AND ABUNDANCE OF
FLORAL RESOURCES ON SHORTGRASS PRAIRIE
V. J. Tepedino''^ and N. L. Stanton^
Abstr.\c:t.— Phenolog\- and abinidance of the floral resources used by bees and other flower-visiting insects were
recorded weekly from permanent quadrats for two years on two shortgrass prairie sites in the Laramie Basin. Wyom-
ing. Each site was composed of three distinct plant communities. Residts suggest considerable spatiotemporal varia-
bilit\ in floral resources. Specifically, most species at both sites showed the following temporal variation: (1) bloom
times between one and two weeks earlier in the second year; (2) differences of approximately 1-2 weeks in bloom
span between years; (3) substantial differences in abundance of flowers between years. Species blooming during the
last half of the census period were significantly more variable in flower abundance between years than those bloom-
ing earlier. Spatial variation was shown both by differences between sites and between plant communities within
sites in the direction and magnitude of between-year changes in floral abundance. For example, floral production at
Boulder Ridge in 1976 was much lower than in 1975, but at The Dirt Farm between-year differences were less pro-
nounced and depended upon season. Similarity measures and cluster analysis .suggest differences in the structure of
the bloom season at both sites between years, and a relatively rapid turnover of floral composition within years such
that bees face a very different flora over the latter part of their flight seasons relative to that encountered initially.
Evidence from other reports support the hypothesis of spatiotemporal unpredictability of floral resources.
Spatial and temporal heterogeneity in re-
source supply is receiving increasing atten-
tion in ecological hypotheses concerning
both species diversity and consumer foraging
strategies (see reviews in Wiens 1976, Levin
1976). An important component of such re-
source heterogeneity is its predictability
(Wiens 1976). When resources are unpredic-
table in time or space, generalists are hypoth-
esized to dominate the consumer faima be-
cause they are less vulnerable than specialists
to fluctuations in the supply of particular re-
sources (Pianka 1966, 1970, Levin 1968,
Cody 1974, Moldenke 1975). Though this hy-
pothesis is intuitively attractive, existent evi-
dence is equivocal (Futuvma 1976, Rabenold
1978).
In addition, the resource predictability hy-
pothesis has been extended to explain latitu-
dinal diversity gradients by proposing that
high tropical diversity is the result of clo.se
species packing of specialists in a predictable
environment (Pianka 1966, 1970). However,
the assertion of large differences in predict-
ability between temperate and tropical re-
gions has recentlv been challenged bv Wolda
(1978), who has shown that precipitation pat-
terns appear no less variable in the tropics
and that insect populations in the seasonal
tropics fluctuate as much as their temperate
counterparts.
The impression that high .spatiotemporal
variability in resources is more common in
certain regions is primarily intuitive or rests
on anecdotal evidence; convincing documen-
tation is lacking (Ricklefs 1973). Clearly, if
resource predictability and .spatiotemporal
heterogeneity are to play more than a hypo-
thetical role in ecology, more quantitative
field studies are needed that measure changes
in resource levels and their use across space
and time.
Department of Zoology and Physiolog)'. University Station Box 3166, University of Wyoming, Laramie, Wyoming 82071.
Present address: USD.VSE.\-Ali-\VR, Bee Biology- and Systematics Laboratory, Utah Stale University, UMC 53, Logan, Utah 84322.
197
198
Great Basin Naturalist
Vol. 40, No. 3
For many consumer guilds, it is difficult to
distinguish between what is and what is not a
resource (Haigh and Maynard Smith 1972).
In contrast, flowers, the trophic resources of
bees and other animals, are easy to delimit
and quantify. In this report we present two
years' data on variation in phenology and
abimdance of entomophilous flowers on two
shortgrass prairie sites in SE Wyoming. Fu-
ture papers will relate these data to the struc-
ture of the bee community (Tepedino and
Stanton, in preparation).
The Great Plains are subject to the wide
and unpredictable fluctuations in temper-
ature and moisture availability which typify
interior, temperate climates. Shortgrass
prairie experiences the most unpredictable
fluctuations in precipitation of all North
American grassland biomes (Wiens 1974).
For example, two of every five years may be
expected to deviate by a minimum of 25 per-
cent from mean yearly precipitation and one
of 12 deviates by at least 50 percent; this var-
iation is temporally impredictable (Wiens
1974). In addition, precipitation frequently
occurs in localized patches and is spatially
unpredictable (Coupland 1958).
Low climatic predictability prompted the
following hypotheses for flower production:
1. Floral phenology and abundance exhibit
wide year-to-year variations at a given
site, both at the level of the species and,
more generally, for the whole commu-
nity.
2. If climatic variation is spatially local-
ized, then within-year differences be-
tween sites in floral phenology and
abundance should be evident.
3. Between-year variability is modified by
seasonal effects. In particular, the
spring flora exhibits greater year-to-
year variability than the summer flora
(Leopold and Jones 1947).
4. Within-year predictability, i.e., the
probability of encountering similar flor-
al composition for several consecutive
weeks, is greatest during the summer
blooming season.
Methods and Study Sites
Study Sites
The study was conducted in the southern
part of the Laramie Basin, a semiarid inter-
mountain valley in Albany County, Wyom-
ing. In general, soils are shallow, rocky, and
poorly developed. Precipitation in nearby
Laramie averages 25.6 cm per year, with 70
percent falling from April through Septem-
ber. The growing season is short, varying be-
tween 80 to 100 days, with killing frosts com-
mon in June and early September. With few
exceptions, the flora is composed of perennial
species.
The Dirt Farm
Located approximately 16 km southeast of
Laramie, The Dirt Farm site is 1.6 ha in area
at an altitude of 2250 m. The vegetation is
divided into three contiguous communities. A
cushion plant commimity covers 0.77 ha and
is located on windswept shallow soils with
frequent bedrock exposure. Abundant species
include Phlox bryoides. Astragalus serico-
leucus, A. spatidatiis, Arenaria Jiookeri, and
Paronychia sessiliflora, all of the caespitose,
herbaceous growth form. Adjoining the cush-
ion plant community is a level area of 0.33
ha dominated by the shrub, Cercocarpus
montanus, mountain mahogany. Associated
species include Allium textile, A. cernuum,
and the half-shrub Chrysothamnus vis-
cidiflorus. Soils are very shallow or nonex-
istent here also, with the roots of the shrubs
penetrating cracks in the bedrock. Extending
south from the mountain mahogany commu-
nity is a 0.5 ha section of typical shortgrass
prairie. Soils are deeper here than in the
other two communities.
Boulder Ridge
The Boulder Ridge site covers 1 ha and is
located approximately 38 km southwest of
Laramie (22 km southwest of The Dirt Farm)
at an altitude of 2425 m. The vegetation is
foothill scrub (Porter 1962) and is divided
into three communities. A central section of
0.35 ha is dominated by the shrub Cerco-
carpus montanus with a few individuals of
the shrubs Primus virginiana, Amelanchier al-
nifolia, and Ribes cereum. Abundant associ-
ated herbs are Allium textile, A. geyeri, A.
cernuum, Ranunctdus ranuncuUnus, and Ce-
rastium arvense. The western part of the
communitv is level, but the eastern end
September 1980
Tepedino, Stanton: Bee Ecology
199
slopes at an angle of approximately 30 de-
grees. At slope bottom movmtain mahogany
gives Way to a community of 0.28 ha domi-
nated by sagebrush {Arteinisia tridentato).
The most abimdant associates are Castilleja
flava, Astmoalus flexuosiis, CoUinsia parii-
folia, Orthoccirpus luteus, and Chrysopsis vil-
losa. Bordering the study site at its eastern
and western ends is a heterogeneous commu-
nity of 0.37 ha that includes representatives
of the cushion plant, shortgrass prairie, and
sagebrush communities. Nomenclature is that
of Harrington (1954), Porter (1965), and We-
ber (1967).
Methods
Censusing.— Floral phenology and abun-
dance of species with entomophilous flowers
were estimated for two years at each site by
weekly censuses of the number of flowers by
species in permanent m^ quadrats. Censusing
extended from the last week in May through
the last week of August, except for the Boul-
der Ridge site in 1975, which began one
week later because of a spring snow storm.
Quadrats were chosen by using a stratified
random sampling technique to insure that
each community was sampled in proportion
to its percentage of the entire study area. Ap-
proximately 1.6 percent of the total area of
each site was sampled. Individual flowers
were counted in all cases except for most
Compositae, where heads were counted, and
the Umbelliferae and Polygonaceae, where
umbels were counted.
Analysis.— To avoid the assumptions of
normality and homoscedasticity, non-
parametric statistics were used. The Sign
Test (Conover 1971) was used to test for be-
tween-year differences in total floral abun-
dance at each site by comparing the total
number of flowers in each permanent quad-
rat for each set of paired sampling dates
(Table 1). Comparisons were made for each
site as a whole and by vegetation tvpe.
To provide a measure of the similarity be-
tween sampling dates both within and be-
tween years for each site we used the Czeka-
nowski measure (also known as the Brav-
Curtis Index: Goodall 1973) to generate sim-
ilarity matrices, which were then subjected
to cluster analysis. The Czekanowski Index is
written
PS = 22 min (x.y.)/ 2 (x. + y.),
1=1 i= 1
where Xi and y^ are the number of flowers of
species i on dates x and y.
The matrices were analyzed by single,
complete, and average linkage clustering al-
gorithms using the BMDP computer package
(Dixon 1975) and the best grouping method,
decided by calculating the cophenetic corre-
lation coefficient of Sokal and Rohlf (Sneath
and Sokal 1973). The average linkage method
provided the best results, and only these are
reported. Matrices for each site for all census
date comparisons as well as separate within-
site, within-year matrices were clustered.
Only the four within-site, within-year clusters
are reported here because we consider them
most informative.
Table 1. Actual floral census dates and their corresponding census code numbers for each study site.
Census
Dirt Farm
Boulder
Ridge
Code
1975
1976
197,5
1976
1
26 May
24 Mav
No census
27 May
2
2 June
31 May
5 June
3 June
3
9 June
7 June
13 June
10 June
4
16 June
16 June
20 Jiuie
18 June
5
23 June
21 June
27 June
24 June
6
30 June
28 June
4 July
1 July
7
7 July
6 Julv
11 July
7 Julv
8
14 Julv
12 Julv
18 July
18 Julv
9
21 Julv
21 Julv
25 July
25 Julv
10
28 July
28 July
31 July
31 July
11
6 August
5 August
7 August
8 .\ugiist
12
13 August
11 August
14 August
14 August
13
19 August
18 August
23 August
19 August
14
26 August
28 August
29 August
28 August
200
Great Basin Naturalist
Vol. 40, No. 3
Results
Precipitation and temperature.— Precipi-
tation patterns over the two years of study
were quite different (Data from weather sta-
tion at Laramie Airport). Rainfall in spring
and early summer 1975 was much heavier
than normal (Fig. la), and total precipitation
for the year was 6.1 percent above normal.
Conversely, 1976 was a dry year with below
normal rainfall for every month from March
through June. By the end of June precipi-
tation was 32.0 percent below normal. July
and August received greater than average
rainfall, and by the end of the study precipi-
tation was only 17.0 percent below normal
for the January through August 1976 period.
In general, temperatures were warmer in
1976 (Fig. lb). In particular 1976 was warm-
er from April through July, a period which
was also (July excluded) much drier than nor-
mal (Fig. la). The frost-free period in 1975
extended from 16 June to 5 September (79
days) and in 1976 from 25 June to termi-
nation of the study (29 August) (63 days).
JFMAMJJASOND
6-
I
a
E,.
u
c
«
o"-
,
/\
—
,\
1
x^\
.«-
0]
/ ^'
,
J
/"'/ ''\
**"* 3'
\
Q.
1/
K^
U
y
1 ^'v
f
/
a;2
A. /
/ ' \ /
I /\
/
T
■ -i
1
"7
/i \j
~^v'V>
y^
/
1975
MAM
1976
Fig. 1. Monthly precipitation (a) and temperature (b)
records from the Laramie .\irport (Brees Field); a) solid
line = 1975, 1976, dashed line = normal; b) solid line
= 1975, dashed line = 1976.
Species Composition.— We recorded 63
and 73 entomophilous plant species during
the two years at The Dirt Farm and Boulder
Ridge, respectively (Appendix A). Total flow-
ers by species and year are also shown in Ap-
pendix A. The family Compositae was repre-
sented by the largest number of species at
both sites, followed by the Cruciferae at The
Dirt Farm and the Scrophulariaceae at Boul-
der Ridge.
Dirt Farm
Phenology.— 'The flowering phenology of
selected species is shown in Figure 2. Phe-
nological variation between years took two
forms: differences in first bloom and in bloom
span. Evidence for variability in first bloom
comes from several sources. First, when total
floral abundance is graphed by date for each
year, it is clear that both June and August
peaks were advanced in 1976 (Fig. 3). It is
worthwhile to note, however, that the ad-
vancement in each peak is not due to similar
responses in each vegetation type. The early
peak is advanced due to responses of the
cushion plant and mountain mahogany com-
munities, and the shortgrass and mountain
mahogany communities account for advances
in the late peak (Fig. 3). Earlier flowering in
1976 was due to a warmer spring and sum-
mer relative to 1975.
Phenological advancement in 1976 is also
seen when first bloom dates are compared by
species. Analysis shows that 27 of 38 species
differed by at least one week in anthesis.
Eleven species were excluded because they
were in bloom when censusing began, and 14
others were eliminated because they flower-
ed in only one year. Of the 27 species that
differed in phenology, 24 were earlier by an
average of 10 days in 1976 (X2 = 16.33,
P<0.001).
Are there seasonal differences between
early- and late-blooming species in phenolo-
gical predictability? lo test this the census
period was halved and species grouped ac-
cording to the half in which they began
blooming (Fig. 3). Twelve of 18 first-half spe-
cies showed a mean difference of one week in
beginning bloom, and 15 of 20 second-half
species showed mean advance of 12.1 days.
Species blooming during the last half of the
September 1980
Tepedino, Stanton: Bee Ecology
201
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
CO 20
021
U22
d) 23
CL24
00 25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
Census week
^456789
10 11
12
13 14
-J I
Fig. 2. Bloom spans of selected species at The Dirt Farm for 1975 (solid) and 1976 (dashed); stars = did not flow-
. Clensiis date numbers refer to Table 1. species numbers to Appendix A.
202
Great Basin Naturalist
Vol. 40, No. 3
MOUNTAIN MAHOGANY
SHORTGRASS
200r
120 -
en
UJ
Q
tr lOOr
UJ
CD
:^
3
I 2 3 4 5 6 7 8 9 10 II 12 13 14
JUNE h- JULY -\ AUG
2 3 4 5 6 7 8 9 10 II 12 1314
JUNE — +- JULY H — AUG
CENSUS WEEK
Fig. 3. Total floral abundance per ni^, irrespective of species, at The Dirt Farm for each census date in 1975 and
1976. Census date numbers refer to Table 1. Asterisks mark significant differences (P<.05) between years for paired
dates.
September 1980
Tepedino, Stanton: Bee Ecology
203
season showed significantly greater phenolo-
gical differences than those blooming during
the first half (Mann-Whitney U-Test,
P<0.05).
Between-year comparisons of bloom spans
are a second indicator of phenological varia-
bility. Differences of at least one week in
bloom span were shown by 22 of 38 species.
Average difference in bloom span for the 22
species was two weeks with a range of one to
seven weeks. Ten species had longer spans in
1975 and 12 had longer spans in 1976. A
comparison by seasonal grouping of bloom
span data into early and later blooming spe-
cies shows that longer bloom spans during
the last half of the season occurred mostly in
1976, and 1975 had more longer blooming
species during the first half (X^ = 2.76, P =
0.097).
Floral Abundance.— Differences in floral
abundance between years is first shown by
DIRT FARM
comparing total number of flowers by date
(Fig. 3). Total abundance comparisons show
the early peak to be higher but the late peak
lower in 1975. The late peak difference is
due largely to profuse flowering of Eriogo-
nitm effusum. Exclusion of this species re-
sults in much closer agreement of abundances
from late July to mid-August.
When total floral abundance is partitioned
into component communities it is again evi-
dent that there is no typical, overall site re-
sponse (Fig. 3). For example, the mountain
mahogany community shows nine significant
between-year differences in abundance, with
1976 having more flowers on eight dates. In
contrast, between-year differences in the
cushion plant community show 1975 with
more flowers for seven or nine significant
comparisons and the shortgrass community
with more flowers in 1975 for five of six
dates. A cold period during the week begin-
BOULDER RIDGE
60r
50 -
40 -
iij
o 30h
UJ
Q.
20 ■
10 -
0
z
<
X
a:
<
X
o
o
llJ
r
1.00- 2.00- >3.00
1.99 2.99
<
X
o
o
UJ
(A)
F
<
X
CO
T
1.00- 2.00- >l3.00
1.99 2.99
RHL
Fig. 4. Frequency distril)iition of the ratio of the number of flowers in the most abundant year to the number of
flowers in the least abundant year for each species (RHL) at both sites.
204
Great Basin Naturalist
Vol. 40, No. 3
ning 16 Jiine 1976 reduced floral production
appreciably on the mountain mahogany com-
munity and probably affected the shortgrass
community as well. During this period tem-
peratures were below freezing for three
nights and snow and sleet fell twice. The ef-
fect of such periods upon organisms in the
Rocky Mountains has been described by
Ehrlichetal. (1972).
Floral abundance was also compared by
species between years. An expression of dif-
ferences in abundance is the ratio of total
number of flowers observed in the year of
highest production divided by total number
of flowers in the year of lowest production
(RHL). We eliminated from this analysis
those species that either flowered in only one
year or failed to produce at least 50 flowers
in either vear. For the remaining 44 species
mean RHL = 5.39 (SD = 8.56, range =
1.04-48.60), suggesting that the abundance of
flowers of most species may show significant
bet ween-y ear- variation. The data are shown
as histograms in Figure 4. Of the 44 species,
26 were more abundant in 1975 and 18 were
more abundant in 1976. Thus, no year effect
was observed.
Do RHL values display a seasonal pattern?
Again, species were grouped according to the
half of the season in which they began flow-
ering and the Mann- Whitney U-Test used to
test for differences in RHL between groups.
The comparisons show that plants blooming
during the latter half of the season were sig-
nificantly more variable (P = 0.05), i.e., had
larger RHL values than those blooming ear-
lier. No year effect was evident since both
years had an almost equal proportion of spe-
cies with highest RHL values in each half of
the year.
Similarity and Cluster Analysis.— We used
cluster analysis to elucidate differences be-
tween census dates within and between years
and to illustrate seasonal groupings. High
similarity values for paired between-year
sampling dates were expected; however, the
data do not support this hypothesis. Mean
similarity for paired census dates was only
0.519 (SD = 0.196, range 0.204-0.854).
Mean similarity was highest (0.593 [SD =
DIRT FARM
BOULDER RIDGE
0.8 r
23456789 10 II
I JUNE 1— JULY H
0.8 r
0.7
0.6
0.5
0.4
0.3
0.2
0.1
0.0
/
/
/|976
\
J I I L
J I I I I
23456789 10
JUNE \— JULY — I
CENSUS WEEK
Fig. 5. Czekanowski similarity measures between floral ahiindance for each census date and the third subsequent
week at both sites. Census date numbers refer to Table I.
September 1980
Tepedino, Stanton: Bee Ecology
205
0.149, range = 0.265-0.772]) when each
1975 date was compared with the week prior
to that date in 1976. Thus, the similarity data
provide additional evidence for phenological
advancement in 1976.
Within-year similarity comparisons were
also quite variable. We reasoned that, since
bees are the predominant pollinators on
shortgrass prairie and because females of
most species of solitary bees fly for a mini-
mum of four weeks (Linsley 1958), a conserv-
ative estimate of within-year resource varia-
bility for a bee would be the similarity
between the week of emergence and three
weeks later. This measure is conservative be-
cause we used similarity measures between
dates that are one week less than the mini-
mum flight span. In Figure 5 we graph the
results for each year. Except for the last
month of the census period, values are very
low. Any species emerging during the first
two-thirds of the blooming season would face
a very different flora during the latter part of
its flight season relative to that encountered
initially.
Cluster analysis aids in depicting seasonal
groupings and transitional periods within the
blooming season (Fig. 6). The number of clus-
ters formed at a value of .50 differs between
years as does the number of unclustered
dates, suggesting that the "structure" of the
blooming season may differ from year to
year. For example, five clusters plus one un-
clustered date form in 1975, but four clusters
and four unclustered dates form in 1976.
Within both years late season dates cluster
strongly, again indicating higher within-year
predictability for late summer bees. Almost
all other clusters are composed of only two
consecutive census dates, indicating a high
rate of turnover in floral composition for the
first two-thirds of the blooming sason.
Boulder Ridge
Phenohgij.— Flowering also began earlier
in 1976 at Boulder Ridge (Fig. 7).^ Of the 73
species censused, 33 were either in bloom
when censusing began of flowered in only
one year and were eliminated from this anal-
ysis. Of the remaining 40 species, 32 showed
phenological differences of at least one week.
Twenty-six of the 32 species were earlier an
average of 10 days in 1976 (X^ = 12.50,
P< 0.001). The seasonal differences in ad-
vancement between first- and second-half
DIRT FARM
0.0 r
0.
0.2
0.3
> 0.4
q:
< 0.5
1
'^ 0.6
0.7
0.8
0.9
1.0
1975
1976
a
I 2 3 4 5 6 7 8 9 10 II 12 13 14
I JUNE 1— JULY —I AUG
I 2 3 4 5 6 7 8 9 10 II 12 13 14
I JUNE (— JULY — I AUG
CENSUS WEEK
Fig. 6. Dendrograms of floral similarity between census dates for each year at The Dirt Farm. Census date niim-
hers refer to Table 1 .
206
Great Basin Naturalist
Vol. 40, No. 3
CO
a;
u
(D
Cl
00
Census week
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
3
I
4 5 6
7
I
8
9
I
10 11
_J 1_
12 13 14
_i I I
Fig. 7. Bloom spans of selected species at Boulder Ridge for 1975 (solid) and 1976 (dashed); stars = did not flower.
Census date numbers refer to Table 1, species numbers to Appendix A.
species observed at The Dirt Farm were not
detected here.
Most species differed in bloom span be-
tween years. After eliminating species that
began blooming prior to censusing, 29 of the
remaining 40 differed by at least one week.
Average difference in bloom span for the 29
species was 17 days with a range of one to
five weeks. Chi-square analyses for sea.sonal
patterns were insignificant. Year effects,
however, were present; longer bloom spans
were concentrated in 1975 (20 of 29, X2 =
4.17,F<0.05).
Abundance.— Differences in total floral
abundance between years were remarkable
(Fig. 8). Twelve of the 13 comparisons
September 1980
Tepedino, Stanton: Bee Ecology
207
showeci significantly greater abundance in
1975. There was a marked midyear peak in
1975 that is only suggested in 1976.
In contrast to The Dirt Farm, floral abun-
dance for all vegetation types at Boulder
Ridge was higher in 1975 (Fig. 8). Between-
year differences on the heterogeneous area
were primarily responsible for the total flow-
er differences. The last 10 dates showed sig-
nificantly more flowers on this section in
1975 (Fig. 8). The second peak in the total
abundance curve in 1975 (Fig. 8), absent in
1976, was due mainly to the heterogeneous
area. Differences observed on the other sec-
tions showed floral abundances in 1976 to be
consistently below those of 1975, although
many of these comparisons were not signifi-
cant.
The RHL ratio was used to compare abun-
dance by species between years. We elimi-
nated all species with less than 50 flowers in
either years and those that flowered in only
one year. For the remaining 38 species, mean
RHL = 5.47 (SD = 6.56, range =
BOULDER RIDGE
c/)
q:
UJ
o
UJ
CD
60
50
40
30
20
10
0
I 2 3 4 5 6 7 8 9 10 II 12 13 14
28i- TOTAL
24 h
'
HETERO-
GENEOUS
r-
,1975
"
A
i
1
v^
-
>y V
V
*
y
\ __^ \
\*y
/
^
^
jy
^
\
y
^ \
1976
\
1111
-L
— L
1 1 1
-L.
8 -
4 -
0
\^\
.1976
-- \
MOUNTAIN MAHOGANY
J I L
J.
J_J l_l L-J I
20
0
0
28
24
20
16
12
8
4
0
2 3 4 5 6 7 8 9 10 II 12 13 14
SAGEBRUSH
2 3 4 5 6 7 8 9 10 II 12 13 14 I 2 3 4 5 6 7 8 9 10 II 12 13 14
|— JUNE I JULY— I AUG |— JUNE — | JULY 1 — AUG
CENSUS WEEK
Fig. 8. Total abundance per m^, irrespective of species, at Boulder Ridge for each census date in 1975 and 1976.
Census date numbers refer to Table 1. Asterisks mark significant differences (P<.05 between years for paired dates.
208
Great Basin Naturalist
Vol. 40, No. 3
1.07-27.50; Fig. 4). Again, there were large
between-year differences in abundance for
most species on the site.
We tested for seasonal differences in RHL
values using the same methods as for The
Dirt Farm. Again, RHL values for the latter
part of the blooming season are significantly
higher (P<0.05). However, in contrast to
The Dirt Farm, each year is not equally rep-
resented by species with high RHL values.
Twenty-eight of the 38 species were more
abundant in 1975 (X^ = 8.53, P< 0.005). This
phenomenon is particularly striking for the
latter part of the blooming season, when only
one of 16 species was more abundant in 1976.
Similarity and Cluster Analysis.— As with
The Dirt Farm data, the expected high sim-
ilarity for paired between-year comparisons
was not evident. Mean similarity for paired
census dates was only 0.424 (SD = 0.200,
range = 0.184-0.896). Again, highest mean
similarity was between 1975 dates and the
week prior to the identical date in 1976 (x =
0.518, SD = 0.161, range = 0.309-0.869).
Within-year similarities [between each
census date and the census taken three weeks
later] were as low as at The Dirt Farm (Fig.
5). The graph for 1975 is uniformly low with-
out the appreciable rise la^te in the season
present in The Dirt Farm and Boulder Ridge
1976 data.
Cluster analysis again suggests "structural"
differences between the blooming seasons
(Fig. 9). Four clusters plus three unclustered
dates form in 1975, and five clusters and
three unaffiliated dates are found in 1976. As
at The Dirt Farm, late season dates cluster
most densely and all but one other cluster is
composed of only two consecutive dates at
the .50 level.
Discussion
Plant species varied substantially between
years in the onset and length of the blooming
period and in the number of flowers pro-
duced. Comparison of phenology and abun-
dance, both between sites and among vegeta-
tion types within sites, shows other important
differences. Though phenological advance-
ment in 1976 was a uniform occurrence at
both sites, changes in the direction and mag-
nitude of floral abundance were not. A com-
parison of total floral abundance between
The Dirt Farm and Boulder Ridge shows that
the two sites behaved quite differently. Floral
production at Boulder Ridge in 1976 was
consistently well below that of 1975. In con-
0.0 -
0.1 -
0.2 -
0.3 I-
>- 0.4
o:
2 0.5 h
^ 0.6
0.7
0.8
0.9
1.0
1975
BOULDER RIDGE
1976
Q
2 3 4 5 6 7 8 9 10 II 12 13 14
I— JUNE —I JULY \ — AUG
CENSUS WEEK
I 2 3 4 5 6 7 8 9 10 II 12 13 14
I — JUNE —\ JULY 1 AUG
Fij;. 9. Dciidrograiii.s of floral similarity between census dates for each year at Boulder Ridge. Census date num-
bers refer to Table 1 .
I
September 1980
Tepedino, Stanton: Bee Ecology
209
trast, floral production was significantly high-
er at The Dirt Farm in 1976 over the latter
part of the blooming season. Evidently, the
spring and early summer drought was either
more severe at Boulder Ridge or the plant
communities at Boulder Ridge were more
susceptible than those at The Dirt Farm.
Between-site differences appear due to
varying responses to weather by each vegeta-
tion type within each site. At Boulder Ridge
all commmiities displayed consistently lower
floral productivity in 1976, but at The Dirt
Farm each community responded indepen-
dently. Indeed, at The Dirt Farm each spe-
cies seemed to display an independent re-
sponse as shown by the lack of any year
effects on the distribution of RHL values ei-
ther in between-year comparisons or be-
tween-season comparisons. These observa-
tions suggest that variation in floral
production is expressed as spatially localized
patches of high or low abundance that
change from year to year.
Because we collected data for only two
years, it is necessary to ask how representa-
tive of routine variability these results are.
Schemske et al. (1978), in a study of seven
spring herbs, found the onset of flowering to
range, by species, from 8 to 22 days over
three years. More importantly, peak flower-
ing did not usually coincide with optimal
pollinator conditions. Long-term bloom re-
cords for several regions in North America
are available for analysis. In several cases
data are available for period of up to 30
years in the same area (Lindsey and Newman
1956-Indiana; Smith 1915-Ohio; Hulbert
1963-Kansas; Hodson 1971-Minnesota). In
examining these data we have used varia-
bility in first flowering as an indication of re-
source predictability since this phenophase is
common to all studies. In brief, we find that
almost all variability in the date of first
bloom is accounted for with 10 years of ob-
servations and that the range of first bloom is
between four and five weeks for most species
(Tepedino and Stanton, impublished manu-
script). Other support for phenological varia-
bility exists. Recently West and Gasto (1978)
reported that the onset of bloom of two arid
land shrubs in northwestern Utah varied over
seven years by 44 and 39 days. Thus, the sub-
stantial phenological variability recorded in
our study over two years is low relative to
what can be expected over a 10-year period.
Long-term studies of floral abundance are
few. Tamm (1948, 1956, 1972a, b) counted
flowers of several species in permanent quad-
rats in forest and meadow in mid-Sweden for
14 to 29 years. All species showed large ir-
regularities in year-to-year floral abundances
from no flowers in some years to profuse
abundance in others.
Short-term studies are more numerous.
Ackerman and Bamberg (1974) reported
large variation in floral abundance over a
three-year period for Lijciinn andersonii in
Nevada. Bykov (1974), in a general review of
vegetation dynamics of the arid Turanian
Plain, reported wide variation in floral abun-
dance of both ephemerals and perennials. Sa-
rukhan (1974) supplied floral abundance data
for three species of Ranunculus for two years
from permanent plots, with all species pro-
ducing many more flowers in the first year.
Holway and Ward (1965), studying the alpine
plant community in northern Colorado over
two years, noted that floral production in the
second year was much lower. Davies (1976)
used the same five trees of each of two spe-
cies and recorded the number of individuals
flowering over an eight-year period in west-
ern Australia. Combining data for both spe-
cies {Acacia pruinocarpa, Hakea lorea; Davies
1976; Table 6) showed that in three of eight
years the number of individuals flowering
was 40 percent or less. Data on fruit crops
also were presented for 10 species of shrubs
and trees for 10 years. If we can assume that
fruit crop bears at least a partial relation to
floral production (Grubb 1977), floral pro-
duction was irregular in 9 of the 10 species.
Schemske (1977, 1978) has shown that the
number of flowers of two woodland herbs
censused in 78 permanent m^ quadrats varied
considerably between years. Moldenke (1976)
noted that floral production varies widely be-
tween years in California grasslands. Treshow
(1979), in a six-year study of the pinyon-
juniper community in Utah, has shown that
forb cover in almost every year differed sig-
nificantly from each other year.
Year-to-year variation in floral abundance
is not restricted to "unpredictable" temper-
ate zone communities (Federov 1966). Mass
flowering via synchronization of all members
210
Great Basin Naturalist
Vol. 40, No. 3
of a particular species or many species in a
community in the tropics at periodic inter-
vals is well known (Whitmore 1975). Med-
way (1972) and McClure (1966) provided
data showing widespread irregularity in flow-
ering for 46 species of tropical rain forest
trees in Malaya. Although most of the obser-
vations were recorded on very few individ-
uals, it is enlightening to learn that the per-
cent of species flowering each year ranged
from 44 to 88 over the period from 1963 to
1968, with an average of 58 percent. Of 42
species observed for the entire six-year peri-
od, only 11 (26.2 percent) flowered every
year, and 24 (57.1 percent) flowered in three
or fewer years. In a study of flowering
phenology in Ceylon, Koelmeyer (1959) re-
ported: "There was no regularity in the se-
quence of years of flowering and years in
which there was no flowering in the individ-
ual trees. The result is the absence of a defi-
nite cycle of flowering."
The data seem clear. Where data on year-
to-year floral abundances have been record-
ed, large variations in floral production are
the rule rather than the exception.
Variability in floral resources may also be
modified by seasonal influences. First, some
parts of the blooming season may exhibit
more year-to-year variability than others.
Leopold and Jones (1947) hypothesized that
early blooming species are more "turbulent"
in first bloom than those which bloom later
in the year. We reexamined the phenology
data of Leopold and Jones (1947) for Wiscon-
sin using multiple regression analysis and
found that their Sauk County data do show a
significant inverse correlation between range
of flrst bloom and average first bloom date.
Though the Dane County data show the
same pattern, it is not significant (Tepedino
and Stanton, unpublished manuscript).
The Wyoming data do not support the hy-
pothesis of greater year-to-year "turbulence"
in the spring flora. At The Dirt Farm, be-
tween-year differences in flrst bloom were
signiflcantly greater for plants that bloomed
over the second half of the census period. At
Boulder Ridge no significant difference be-
tween flrst- and second-half plant species was
detectable. In addition, between-year differ-
ences in abundance (as judged by RHL ratios)
were significantly greater for the last half of
the blooming season at both sites. Whether
these differences were due to only two years'
data from Wyoming or to conditions that are
site specific is not clear.
The second way in which floral resources
may vary seasonally is in predictability of
subsequent resource abundance and composi-
tion. From this perspective the spring flora
is, indeed, more turbulent; predictability, as
judged by floral similarity values calculated
at three-week intervals (Fig. 5), was low rela-
tive to average flight time for bees until the
latter third of the bloom season when com-
posites became dominant. This result may be
somewhat misleading however, because more
species begin bloom in spring than in late
summer and low spring similarity values are
due in large part to species additions.
Low year-to-year predictability in floral
phenology and abundance must exert strong
selection on flower-visiting insects. This is
particularly true for bees because every stage
in their life cycle is obligately dependent
upon floral resources for food. When re-
sources are unpredictable in time and/or
space, selection should favor generalized con
sumers. Alternatively, specialization would
require precise synchronization between bee
emergence and anthesis of the host plant,
particularly when the host has a brief bloom
span. It is unclear how such precise synchro-
nization might be affected. In most plant spe-
cies studied photoperiodic stimuli initiate
flower formation, but subsequent devel-
opment and anthesis is profoundly modified
by diverse factors such as moisture and nutri-
ent availability and temperature (Evans 1969,
Schwabe 1972). Our knowledge of the stimuli
used by bees to cue emergence in a natural
setting is scanty (Linsley 1958), but in the
laboratory temperature alone is a reliable
stimulus for several species {Megochile rotiin-
data (Fabricius), Osmia lignaria Say, Hylaetis
bisintiatiis Forster, Nomia melanderi Cock-
erell, and several others; G. E. Bohart, F. D.
Parker, P. F. Torchio, pers. comm., pers.
obs.). Thus, though anthesis is determined by
a complex of factors, bee emergence may be
primarily responsive to temperature. Because
of these differences in potential stimuli used
by the two groups, synchronization may be
rare. In this regard, Linsley (1958) noted that
September 1980
Tepedino, Stanton: Bee Ecology
211
studies of oligolectic bees frequently reveal consistent with documented fluctuations in
poor synchronization.
Even if specialized bees could achieve
close synchronization with host plant an-
thesis, the problem of year-to-year variation
in resource quantity still remains. Attempting
to track specific floral resources that vary
widely could cause large fluctuations in the
populations of bee specialists, thereby in-
creasing the probability of local extinction
(Tepedino 1979). The frequently expressed
view that most temperate bees are special-
ized (van der Pijl 1966, Faegri and van der
Pijl 1971, Heinrich 1976, Raw 1976,
Heithaus 1979) will probably require modifi-
cation because such specialization seems in-
floral
resources.
Acknowledgments
We thank J. M. Loar and T. M. Root for
providing meticulous assistance in the field, J.
Meyer for drawing the figures, and Dr. L. L.
McDonald for advice on cluster analysis. The
manuscript was improved by comments by
M. S. Boyce, K. T. Harper, D. H. Knight, P.
Lincoln, M. D. Marcus, and A. R. Moldenke.
The study could not have been conducted
without support from NSF Grant BMS75-
14044. "Doc" and Peggy Wollbrinck of Lar-
amie and the proprietors of The Dirt Farm,
Inc., graciously allowed use of private land.
Appendix A. The number of flowers recorded in permanent m^ quadrats at The Dirt Farm and Boulder Ridge in
197.5 and 1976. Nomenclature: Harrington (1954), Weber (1967), Porter (1965). Numbers in the far left column refer
to Figures 2, 7; the first number to Figure 2 (Dirt Farm), the second to Figure 7 (Boulder Ridge). A zero signifies
nonrepresentation.
Species
Dirt Farm
1975 1976
Boulder Ridge
1975 1976
Berberidaceae
(0,8)
Berberis repens Lindl.
Boraginaceae
(19, 0)
Crijptantha flovoculata (A. Nels
(23, 0)
Cijnoglossitm officinalis L.
(14, 0)
Hackelia florihwida (Lehm.)
Lappulo redoicskii (Horneni.)
(10, 0)
Lithospermtim incisum Lehm.
(0,5)
Mertetisia Iwinilis Rydb.
(38, 0)
(0, 23)
(24, 19)
(0,3)
(22, 20)
(40, 34)
(35, 28)
(39, 0)
(42, 0)
(0, 24)
Cactaceae
Opiintia pohjacantha Haw.
Capparidaceae
Cleorne serrtilata Pursh
Caryophyllaceae
Arenoria fendlcri A. Gray
Arcnaria hookcri Nutt.
Cerastitim arvense L.
Paronychia sessiliflora Nutt.
Stellaria media (L.)
Compositae
Acliillea miUefoliuin L.
Antennaria microphyUa Rydb.
Antennaria rosea Greene
Artemisia frigida Willd.
Aster rtibrotincttis Blake
Chnjsopsis villosa (Pursh)
Chrysothamntts nauseosus (Pallas)
Chrysothamntis viscidiflorus (Hook.)
Cirsitim tmdulatum (Nutt.)
Erigeron canus A. Gray
101
52
68
575
60
-
32
50
47
—
—
67
109
—
—
_
—
5
2
133
16
9
9
12
5
16
42
_
_
122
58
1165
628
1577
474
_
—
.332
50
3489
.3000
718
40
—
—
9
—
382
19
_
.3.343
959
_
83
-
311
114
.393
-
,393
73
87
18
262
116
2426
876
1882
73
-
-
1906
4216
11
3
17
4
2
—
-
2
817
314
212
Appendix A continued.
Great Basin Naturalist
Vol. 40, No. 3
(12, 2)
(41, 35)
(34, 29)
(30, 14)
(6,0)
(13, 9)
(0,30)
(0, 18)
(32, 0)
Species
Erigeron nematophyUus Rydb.
Erigeron pumilus Nutt.
GailUirdia aristaia Pursh
Gutierrezia sarothrae (Pursh)
Haplopappus nutiaUii T. & G.
HelkintheUa uniflom (Nutt.)
Hijmenoxijs acaiiUs (Pursh)
Hymenoxijs torreijana (Nutt.)
Senecio catum Hook.
Senecio integerrimus Nutt.
Solidago spathulata DC
Taraxicttm sp. Hall
Townsendid scricea Hook.
Crassulaceae
S«/i("i stenopetalum Pursh
Cniciferae
(4,0)
Arabis fendleri (Wats.)
Arcihis holbocUn Hornem.
(21, 0)
Dcsnirainia sophia L.
Dmha nemorosa L.
(15, 11)
b'.rijsimum capitatum (Dougl.)
Halimolohos virgata (Nutt.)
(8, 10)
LcsqucrcUa ludoviciana (Nutt
(0,7)
Phiisaria amtralis (Payson)
(17, 0)
Smjmbrium ultissimum L.
Sunjmhrium linefolium Nutt.
Euphorbiaceae
Euphorbia sp. L.
Gentianaceae
Swertia radiata (Kellogg)
Labiatae
Hedcoma drtDtitiiondii Benth.
Snitelhirid bhtionii Porter
Legiuuinosae
(25, 0) Astragcdm hisulaitus Hook.
Astragalus crassicarpus Nutt.
(0, 21) Astragalus flexuosus Dougl.
(9, 0) Astragalus sericolettcus Gray
Astragalus shortianus Nutt.
(7, 0) Astragalus spatulatiis Sheld.
(0, .32) Astragalus striatus Nutt.
(0,12) TItcrmopsis rhoinhifolia Nutt.
Liliaceae
(37, 31) Allium cernuum Roth
(0, 16) Ellium getjeri Wats.
(18, 17) .A//ii/m textile Nels. & Macbr.
Caloehorttis nuttallii Torrev
(0, 1) Leticocrintim montanum Nutt.
(11, 0) VAjgadenus venenosus Wats.
Linaceae
Lidi/Hi lewisii Pursh
Dirt Farm
Boulder Ridge
1975
1976
1975
1976
434
406
895
1014
59
—
12
-
_
7
-
2163
3170
1512
735
45
—
—
45
35
50
3
62
81
82
24
342
248
—
—
499
701
21
53
10
11
—
19
227
100
_
—
57
—
4
_
2
—
279
971
125
9
325
75
9
88
1768
366
535
89
166
—
—
—
13
17
128
225
—
-
35
581
207
—
122
-
160
12
—
-
-
1839
332
466
360
48
—
219
84
24
52
-
-
27
—
—
—
187
_
—
4.38
31
21
57
-
-
_
—
18
-
386
329
-
-
_
41
—
-
-
84
43
1771
2693
478
98
_
_
853
607
622
1284
187
200
1
—
—
—
—
73
122
211
314
-
-
25
17
September 1980
Appendix A continued.
Tepedino, Stanton: Bee Ecology
213
Species
Malvaceae
SpluicKiIcea coccinea (Pursh)
Onagraceae
(31, 0)
Gaum coccinea Nutt.
Oenothera cownopifolia T. & G.
Poleinoniaceae
Gilia apicata Nutt.
Mirrostcris htivtilis (Dougl.)
(1,0)
Plilox hnjoides Nutt.
(2,6)
Phlox hoodii Rich.
Polygonaceae
Eriogonum ahitttm Torr.
(36, 0)
Eriogonum effiistini Nutt.
(0, 33)
Eriogonum janicsii Benth.
(28, 22)
Eriogonum umheUatum Torr.
Portulacaceae
Clatjtonia hinceohita Pursh
Primulaceae
Androsace septentrionalis L.
Ranunculaceae
(26, 0)
Delphinium nehoni Greene
(0. 4)
Ranunctihts ranunetdinus (Nutt.)
Rosaceae
(0, 15)
Amelanehier ahiifolia Nutt.
(16. 13)
Ccrcoearpus montonus Raf.
Potcntilla coneinna Richards
PotcntUla fi-ssa Nutt.
(29, 0)
PotentiUa liippiana Lehm.
Santalaceae
Commandra umbellata (L.)
Saxifragaceae
Ribes cereum Dougl.
Scrophulariaceae
(0, 25)
CastUIeja flava Watson
CoUinsia parviflora Dougl.
Oddiocarpus hiteus Nutt.
(20, 0)
Penstemon angustifolius Nutt.
(27, 0)
Penstemon erianthcrus Pursh
(33, 27)
Penstemon laricifohus exHifolius (A. Neis.
(0, 26)
Pemtemon strictus Benth.
Umbelliferae
(5, 0)
Harbouria trachijpleura (A. Grav)
(3, 0)
Violaceae
Viola nuttallii Pursh
Dirt Farm
1975 1976
28
317
41
67
Boulder Ridge
1975 1976
-
-
4
6511
7526
—
91
42
12
192
27
85
28,869
57,115
-
-
-
-
84
298
852
284
282
73
19
22
11
56
13
14
-
-
4284
5509
_
220
8
mi
3752
128
219
-
-
9
1
-
—
6
2
862
375
20
3
243
5
14
98
-
-
3
7
912
.361
—
—
565
402
—
-
571
32.3
67
^
-
-
61
12
4
1
449
220
966
a3
-
-
44
7
667
196
-
8
73
52
4
Species total
Totals (both years)
56 5.5
63
57
60.
214
Great Basin Naturalist
Vol. 40, No. 3
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September 1980
Tepedino, Stanton: Bee Ecology
215
1978. Sexual reproduction in an Illinois popu-
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DOG OWNERS AND HYDATID DISEASE IN SANPETE COUNTY, UTAH'
Peter M. Schantz' and Ferron L. Andersen'
.\bstract.— a questionnaire survey was conducted in Sanpete County, Utah, to determine the knowledge of dog
owners concerning hydatid disease and an identification of some basic sheep management practices there. The
households surveyed included 21 (Group I) that had one or more dogs infected with Echinococciis gmuiilostis tape-
worms at more than one annual field clinic, and 19 others (Group II) that had one or more dogs infected when the
studv first began in 1971-72, but had not had any infected dogs identified at field clinics during subsequent years.
The results showed that 92.5 percent of households knew the cause of the disease and how it is transmitted, and that
9() percent knew of someone who had been operated on for surgical removal of hydatid cysts. There was no signifi-
cant difference in the level of knowledge of the disease between the two groups of respondents, nor in their sheep
management practices. Even though the level of infection of the parasite in dogs has decreased since the project
started, certain sheep management practices persist among respondents in both groups that allow for continued
transmission of the parasite in this region.
Hydatid disease is an infection of people,
sheep, and some other animals that produces
fluid-filled (hydatid) cysts in the liver, lungs.
or other organs (Fig. 1). The cysts are the lar-
val (immature) forms of a tapeworm parasite,
Echinococciis granulosus (Fig. 2), which lives
Fig. 1. Fluid-filled hydatid cysts in the livers and lungs of infected sheep.
'Supported in part by U.S. Public Health Grant AI-10.588.
'Parasitic Diseases Division, Center for Disease Control, Atlanta. Ceorgia 303.33.
'Department of Zoology, Brigham Young University, Provo, Utah 84602.
216
September 1980
ScHANTz, Andersen: Hydatid Disease
217
^ a.
•^tiga^-'Ti'^-
Fisi;. 2. Tlie adult tapeworm. Echinococni.s oranitlosus (approximatelv 5-6 mm in leiiiith). removed from the small
intestine ot an infeeted dog.
as an adult in the .small intestine of dogs.
People and sheep contract the hydatid cysts
when they inadvertently ingest the tapeworm
eggs passed in the stools of infected dogs.
This may occur when people handle dogs
that harbor the parasite, and when sheep
graze on contaminated pa.stures. Dogs be-
come infected with the tapeworm when they
ingest hydatid cy.sts in the viscera of sheep.
The parasite occurs throughout the world
wherever dogs, sheep, and other suitable ani-
mal hosts are kept together. The common
practice among .sheep ranchers of allowing
dogs to eat the uncooked viscera of home-
killed sheep provides optimum conditions for
continued transmission.
In the United States, transmission of Ech-
inococctis granulosus in the dog-sheep cycle
is known to occur in several western states,
including California (Araujo et al. 1975), Ari-
zona and New Mexico (Schantz 1977), and
Utah (Spniance et al. 1974). The most serious
problem is in Utah, where nearly 50 human
cases have been diagnosed since 1944. Sever-
al of these cases were fatal, and most of the
others have required surgical removal of the
hvdatid cvsts. Many of the victims were resi-
dents of Sanpete County, which is in the cen-
tral part of the .state.
Since 1971 hvdatid disease has been stud-
ied and control measures initiated through
the combined efforts of Brigham Young Uni-
versity (Provo, Utah), the Utah State Depart-
ments of Health and Agriculture (Salt Lake
Citv, Utah), and the Center for Disease Con-
trol (Atlanta, Georgia). These measures have
included (1) the development and distribu-
tion of educational displays and brochures on
the life cycle of the hydatid tapeworm, (2)
the development of adequate methods for
disposal of sheep carcasses at community
dumping grounds, (3) the periodic holding of
public health clinics to detect new ca.ses of
human infection, and (4) annual field clinics
to detect new or persistent cases of infected
dogs (Fig. 3). Following the implementation
218
Great Basin Naturalist
Vol. 40, No. 3
Fig. 3. Sheep dogs from Sanpete County restrained at
cocnts gr«nr//o.v(/.v tapeworms.
of these control measures, the number of
dogs found infected at the field clinics has
decreased from 27 percent in 1971 (Loveless
et al. 1978) to 14 percent in 1978 (unpub-
lished ms.). Most sheep ranchers have shown
a cooperative attitude with regard to proper
disposal of sheep carcasses or viscera. Certain
individuals, however, have not been success-
ful in preventing reinfection of their dogs as
evidenced by the fact that some of their dogs
were found repeatedly infected on numerous
occasions. We believed that if the reasons
could be determined why some dog owners
were imable or unwilling to comply with the
recommended preventive measures, it might
be possible to change or modify the recom-
mendations to obtain more cooperation, and
ultimately an improved control program.
Materials and Methods
A questionnaire survey was conducted of
the owners of dogs that had been found to be
field clinic during examination for detection of Echino-
infected in Sanpete County. The survey in-
cluded 40 households, 21 of which had one or
more dogs found infected at more than one
annual clinic (Group I) and 19 others that
had one or more dogs infected only at either
the first or second annual clinic (1971 or
1972), but did not have infected dogs at sub-
sequent clinics (Group II). During the visits,
questions were asked about dog-feeding prac-
tices, dog control, sheep-killing procedures,
and knowledge of the life cycle and control
of hydatid disease.
Results
What emerged from our study may be con-
sidered a general description of the habits
and practices of dog owners that tend to
maintain the cycle of hydatid disease in San-
pete County. Each household selected had
both sheep and dogs. The average number of
dogs per household was 2.5 and the average
flock size was approximately 1000. We found
that nearly everyone was aware of the dis-
September 1980
ScHANTZ, Andersen: Hydatid Disease
219
ease. Persons interviewed in 90 percent of
the households knew of someone who had
been operated on for the disease. This was
usually someone from the same town, and in
10 households (25.0 percent) the victim
known was a member of the nuclear or ex-
tended family. Moreover, persons inter-
viewed in 92.5 percent of households knew
the cause of the disease and how it is trans-
mitted. Specifically, they knew that people
become infected with hydatid cysts by in-
gesting eggs passed in the feces of infected
dogs, and that dogs become infected with the
hvdatid worms by ingesting the cysts in the
lungs and livers of sheep.
More than four-fifths of the households in-
dicated they sometimes killed and butchered
sheep on their premises or in the fields. De-
spite their awareness and understanding of
how hvdatid disease is transmitted, nearly
two-thirds admitted their dogs had access to
the sheep-killing area, and nearly half said
the dogs sometimes ate part of the sheep car-
cass.
The main diet of dogs in more than 85 per-
cent of households was commercial dog food,
and in none was the main diet reported as
sheep muscle or organ meat. Nevertheless, it
was clear that most dogs could possibly eat
sheep at least occasionally, since in two-
thirds of households dogs were allowed to
roam free, and, therefore could scavenge on
sheep carcasses at the town dump or in the
fields. Less than one-third of households
regularly tied or locked up their dogs when
the dogs were not working.
Persons interviewed at more than 80 per-
cent of households indicated they believed
that the recommended control measures were
adequate to break the chain of transmission
and eliminate the infection. Persons at only 6
(15 percent) of households indicated they had
taken no active measures to eliminate the in-
fection. At the 34 households that indicated
they had done something, the most frequent-
ly mentioned steps taken were (1) periodic
treating of dogs for tapeworms, and (2) dis-
carding of viscera from home-killed sheep in
such a way that dogs could not get to it. Four
households indicated they no longer had dogs
because of the potential of contracting hyda-
tid disease. There was a general consensus
(82.5 percent) that government authority
should not make it illegal for dogs to eat
parts of the sheep carcass.
When the households were categorized ac-
cording to whether their dogs had been
found infected at only one of the first clinics
or whether their dogs had been found repeat-
edly infected, there were no obvious differ-
ences that would allow us to conclude whv
the first group of households was apparently
successful in preventing reinfection. There
were no statistically significant differences in
the two groups regarding the number of dogs
or sheep they owned, the frequency that
sheep were butchered for home consumption,
the apparent access of dogs to sheep viscera,
the household members' knowledge and un-
derstanding of hydatid disease, nor willing-
ness to take measures to prevent the infection
in the dogs. In fact, the responses to our ques-
tions appeared to suggest that dog owners
with repeatedly infected dogs were more
likely to have tied their dogs up when not
working and to have taken other deliberate
measures to prevent their dogs from eating
parts of the sheep carcass. This apparent
anomaly is most likely explained by the fact
that. owners of repeatedly infected dogs had
more recently been made aware of what they
should be doing to prevent infection than the
other group of dog owners whose dogs had
been given a "clean bill of health " at the
most recent dog clinics.
In summary, we did not learn from our
study why some dog-owning households were
successful in preventing reinfection of their
dogs and why others were not. What was
clear, however, was that numerous opportu-
nities still existed at these households for dogs
to become infected with hydatid tapeworms.
As a result of health education and other con-
trol activities, virtually all the Sanpete Coun-
ty dog owners interviewed in our survey
knew the basic facts ab^nit hydatid disease;
however, few had actually taken all the nec-
essary steps to insure its elimination. Evi-
dence obtained from the survey suggests that
manv dog owners apparently believe that pe-
riodic treatment of dogs is sufficient to solve
the problem; however, that may be an over-
simplified solution. To effectively break the
chain of transmission, all dogs must be pre-
vented from eating the viscera of infected an-
imals. This means not only that dog owners
220
Great Basin Naturalist
Vol. 40, No. 3
must refrain from feeding such organs to
their dogs, but, since dead sheep are fre-
quently discarded in open pits and are acces-
sible to roving dogs, dogs must be kept under
control at all times. An additional feasible
control measure would be the installation of
large metal pit covers or sturdy fences at the
animal pits in order to prevent ready access
of roving dogs to animal carcasses discarded
at those sites.
From its inception in 1971, the Hydatid
Disease Control Program has been an entire-
ly voluntary campaign. Results of this survey
suggest that some additional incentives may
be necessar\' to insure that all dog owners
take the necessar\' steps to stop the transmis-
sion of hydatid disease.
Literature Cited
Aracjo. F. p.. C. W. Schwabe. J. C. Sawder, and ^\^
G. Davis. 1975. Hvdatid disease transmission in
California. A study of the Basque connection.
Am. J. Epidemiol. 102: 291-302.
ScH.\NTZ. P. M. 1977. Echinococcosis in American In-
dians living in Arizona and New Mexico. Review
of recent studies. Am. J. Epidemiol. 106:
.370-.379.
Spru.ance, S. L., L. F. Klock. F. Cha.vg. T. Fikushima.
F. L. .\ndersen, .and 1. G. K.\g.an. 1974. Endemic
hvdatid disease in Utah. A review. Rocky Mtn.
Med. J. 71: 17-23.
Loveless, R. M., F. L. .\ndersen. M. J. Rwis.ay, and R.
K. Hedelius. 1978. Echinococciis granulosus in
dogs and sheep in central Utah. 1971-1976. Am.
J. Vet. Res. 39: 499-502.
NEW GRASS DISTRIBUTION RECORDS FOR ARIZONA, NEW MEXICO, AND TEXAS'
Stephan L. Hatch-
Abstr^ct.— New distribution records are given for seven grass species now found in Arizona, New Mexico, and
Texas.
Recent collections have revealed new dis-
tribution records for seven southwestern grass
species. These records are extensions to the
known distributions of these species as re-
corded in general for the United States bv
Hitchcock (1951), in Arizona and Te.xas by
Gould 1 1951. 1975^. and in the intermountain
area by Cronquist et al. il977>. The checklist
for New Mexico published by Martin and
Castetter (1970) is the basis of distributions
for that state. Voucher specimens for these
new records have been distributed in the
Tracy Herbarium (^TAES).
Eremopyron triticeum (Gaertn.^ Nevski
.\rizona: Coconino Co.: Locallv abundant
as an adventive on disturbed sites. 0.5 km
north of Fredonia, east side of Highway 89A.
on a hard gra\ clav soil at 1600 m elevation,
17 May 1978. Brown 652 (TAES). This in-
troduced grass is a new state record for Ari-
zona. Previous collections have been report-
ed from Oregon, Idaho, Nevada, and Utah by
Cronquist et al. il977'. Montana (Hitchcock
1951k and New Mexico ^Hatch 1977).
Afrna barbota Brot.
New Mexico: Dona Ana Co.: Rare adven-
tive along Interstate 10. 10 km south of Las
Cnices, 3 April 1978. Mocliange 21 (TAES).
This is a new state record for New Mexico.
Previous reports show a distribution from
Washington to Arizona ^Gould 1951).
Bothriochloa ischaemum yh.) Keng.
New Mexico: Colfax Co.: Locally abun-
dant as an adventive. 3.5 km south of Raton.
18 August 1978, Hatch 4072 (TAES). Dona
Ana Co.: Locally abundant. 11 km south of
San Agustin Pass on the north slopes of the
Organ Mountains. 18 September 1977, Dick-
Peddie 55 i^TAES). Grant Co.: .\bundant, 32
km north of Silver City along Highwav 15, 9
October 1977. Hoefler'33 TAES). Sierra Co.:
Five km south of Williamsburg along Inter-
state 25, 26 August 1977, Hatch 2604
I TAES). This is a new state record for New
Mexico. Gould (1975) reported this species as
being introduced in Te.xas as a pasture grass.
Dactyloctenium aegyptium (L.i Beauv.
New Mexico: Dona Ana Co.: Rare as an
adventive in the New Mexico State Univer-
sitv Agronomy Field Laboratory land. Las
Cruces, 10 September 1977, Hatch 2565
TAES). This is a new record for New Mexi-
co. Gould ^195L reported this species from
Arizona and later ^1975) from Te.xas. Gould
(1975) stated that this species was well adapt-
ed to sandy soils of the southern United
States.
Eragrostis superba Peyr.
New Me.xicg: Dona Ana Co.: A relatively
rare introduced grass along Interstate 10. 6
km south of Las Cruces in sandy soil. 15 Oc-
tober 1977, Yehca 3S (T.\ES). This is a new
state record for New Mexico. The plant has
been collected in Texas 22 May 1957. Gould
7550 (TAES), but was not included in Gould's
(1975) Grasses of Texas. This species has been
collected in .\rizona. 3 November 1961. Mat-
tox and White sn. (T.\ES). It is native to
South Africa and was introduced to the
United States as P.I. 185516 (No. 39) 12 De-
cember 1949.
Technical Bulletin T..\. 15803, Texas Agricultural Eipenment Stabon. Teias A & M L'niversit). College Station. Texas . iS*3.
'Department of Range Science. Tesas A & Vt Universitv-. College Station. Teias 77843.
221
222
Great Basin Naturalist
Vol. 40, No. 3
Leptoloma arenicola Swallen
New Mexico: Lea Co.: A rare-abundant
perennial grass on sandy sites, 22 km east-
northeast of Maljamar along Highway 82, 10
August 1976, Elhworth 23 (TAES). This col-
lection is a new state record for New Mexico.
Hitchcock (1951) and Gould (1975) report
this taxon as occurring only in Kenedy Co.,
Texas. This is a rhizoinatous form of the
widespread L. cognatum (Schult.) Chase,
classified by Gould (1975) as L. cognatum
var. arenicola (Swallen) Gould.
Texas: Kent Co.: Locally abundant in
stands of shinnery oak {Querciis havardii
Rydb.), 3.0 km south-southeast of Girard, 9
June 1979, Slosser sn. (TAES). The Slosser
collection is a distribution extension within
Texas of several hundred miles.
Neeragrostis reptans (Michx.) Nicora
New Mexico: Sierra Co.: Locally abun-
dant on the mudflats of Elephant Butte Res-
ervoir, near Alamosa, 10 September 1978, So-
pyn sn. (TAES). This is a new record for New
Mexico. Gould (1975) reported this species as
being abundant on exposed lake beds in the
central United States from Kentucky and
South Dakota to Louisiana, Texas, and Flor-
ida.
Literature Cited
Cronquist, a., B. H. Holmgren, N. H. Holmgren, J. L.
Reveal, and P. L. Holmgren. 1977. Inter-
mountain flora. Vol. 6. Columbia Press. New
York. 584 pp.
Gould, F. W. 1951. Grasses of the southwestern United
States. University of Arizona Press, Tucson. 352
pp.
1975. The grasses of Texas. Texas A & M Univer-
sity Press, College Station. 65.3 pp.
H.\TCH, S. L. 1977. New grass distribution records for
New Mexico and the United States. Great Basin
Nat. 37:530-531.
Hitchcock, A. S. 1951. Manual of the grasses of the
United States. USDA Miscellaneous Publication
200, rev. by Agnes Chase. 1051 pp.
Martin, W. C, and E. F. Castetter. 1970. A checklist
of gymnosperms and angiosperms of New Mexi-
co. Published privately.
A COMPARISON OF EPIPHYTIC DIATOM ASSEMBLAGES ON
LIVING AND DEAD STEMS OF THE COMMON GRASS PHRAGMITES AUSTRALIS
Judith A. Grimes'. Larry L. St. Clair', and Samuel R. Rushforth'
Abstract.- Diatoms epiphytic on Phragmites austmUs (Cav.) Trin. ex Steaded stems were collected from a single
clone at the .southern end of Provo Bay, Utah Lake, Utah. Diatom populations from both living and dead stem sec-
tions were analyzed. Species diversity in each .sample was high, indicating that the stems provide a relatively stable
habitat for diatom epiphytes. Of the 23 genera found, only Gomphonema and \avicula showed significant trends
toward stem preference. The diatoms in this study support the current view that Utah Lake is a slightly saline,
eutrophic system.
The occurrence of diatom assemblages as
epiphytes on httoral, emergent macrophytes
is well documented (Godward 1934, 1937,
Knud.son 1957, Prow.se 1959). Likewise, the
impact of such epiphytes on primary produc-
tivity and community trophic structure has
been examined in several estuarine environ-
ments (Mclntire et al. 1971, Stowe et al.
1971, and Main et al. 1974), but has been
largely ignored in freshwater systems (Wetzel
1964). Tlie epiphytic diatom communities at-
tached to emergents inevitably play a role in
the overall productivity of lakes and es-
tuaries. They also contribute to regulation of
the overall metabolism of such waters by al-
tering the amount and quality of alloch-
thonous organics entering the lake by acting
as physical and metabolic traps or filters. The
attached diatom flora also serves as an au-
tochthonous source of particulate organic
and dissolved organic matter that is readily
available to pelagic animals. The degree of
influence of these epiphytic organisms on the
productivity of standing waters has rarely
been determined. However, Allen (1971) esti-
mated that up to 31.3 percent of the total lit-
toral production could be attributed to
epiphtyic algae, with up to 21.4 percent of
the total lake production being attributable
to such attached communities. In addition, a
comparison between phytoplankton and epi-
phyte production demonstrated that the lat-
ter was equivalent to 75 percent of the
phytoplankton production over the annual
period (Allen 1971).
Even though epiphytic commimities have
been demonstrated to be important, the dis-
tribution patterns of such assemblages on the
basis of variation in ho.st species and host sub-
strate conditions have received little atten-
tion. Likewise, the complex physiological
relationship between the host macrophyte
and the attached diatom species has received
less attention than warranted (Wetzel 1964,
1965, 1969b, Allen 1971, Hough et al. 1975).
The impact of this relationship is fimdamen-
tal to understanding the basic distribution
patterns of epiphytes not only on different
macrophyte host species but also on members
of the same species at different levels of se-
nescence.
The purpose of this study is to illuminate
distribution patterns of diatom epiphytes on
living and dead specimens of a single macro-
phyte host {Phragmites australis (Cav.) Trin.
ex Steaded) in Utah Lake, Utah. The data
from this study will be used as a baseline for
extended research in Utah Lake on epiphyte
distribution patterns and epiphyte impact on
lake productivity and trophic structure.
Methods
Samples were collected 20 September
1978 from a single clone of Phragmites ous-
trcilis located at the southern end of the
mouth of Provo Bay in Utah Lake. Five sam-
ples of living and five of dead Phragmites
australis stems were collected as cut 10 cm
sections, measured from the water level
'Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602.
223
224
Great Basin Naturalist
Vol. 40, No. 3
down. Samples were prepared according to
standard acid-oxidation methods, and per-
manent diatom slides were made using Naph-
rax diatom momitant (St. Clair and Rushforth
1977). Slides were examined and diatom spe-
cies were identified at lOOOX with a Ziess RA
research microscope with bright field and
Nomarski interference phase-contrast acces-
sories.
Quantitative data on the diatom assem-
blages were recorded by counting 250-400
diatoms for each sample. Previous studies
have shown that a statistically valid count
can be obtained within this range (Squires
1977). Each slide was then thoroughly
scanned to record the rare species. The re-
sults were converted into percent relative
density values for all species for each site.
Shannon-Wiener diversity indices were cal-
culated for individual samples (Shannon and
Wiener 1963).
The relative density figures for each
sample were compared to all other samples
and similarity indices were calculated (Ru-
zicka 1958). These indices were clustered
(Sneath and Sokal 1973) to identify unique
community associations within and between
the living and dead Phragmites aiistralis stem
sections.
The most prevalent diatoms encountered
in the study and the diatoms that significant-
ly differed between the living and the dead
specimens of Phragmites australis were deter-
mined.
Results
Twenty-three genera and 114 diatom spe-
cies were found on the 10 Phragmites austral-
is stem sections (Table 1). The most prevalent
diatoms throughout the study were Navicula
graciloides, Nitzschia inconspicua, and Nit-
zschia filiformis (Table 2). Nitzschia dis-
sipata, Stephanodiscus astrea var. minutula,
and Nitzschia palea were also common.
Among the prevalent species, Amphora ve-
neta was found to occur only on the living
stems, whereas Navicula schroeteri var. es-
camhia was essentially restricted to the dead
stems.
Table 1. Alphabetical list of the diatom taxa found
on living and dead Phragmites australis stem sections
from Provo Bay, Utah Lake, and their average relative
density.
Species
Living
Dead
Achnanthes hauckiana"
.07
_
Achnanthes lanceolata var. dtibia"
—
P
Achnanthes lewisiana
.06
P
Achnanthes minutissima
2.20
.57
Achnanthes sp."
P
—
Amphora ovaUs
.31
P
Amphora ovalis var. pedicuhis'
—
.08
Amphora perpusilla
.76
1.79
Amphora veneta
8.78
1.26
Amphora sp.°°
.19
-
Anomoeoneis sphaerophora" °
P
—
AsterioneUa formosa
.09
.08
Colonels fenzlioides' °
.02
—
Cocconeis placenttila var. eughjpta'
—
P
Cocconeis placentula var. lineata
.07
.07
Coscinodiscus sp.
.06
.06
Cyclotella kiitzinghiana'
—
.13
Cijclotella meneghiniana
1.87
1.80
Cijmbella affinis'
—
P
Cymbella miniita"
—
.06
Cijmbella muelleri"
.14
-
Cymbella prostrata
.07
.05
Cymbella sp. ° °
P
—
Diatoma tentie var. elongatum
.13
.23
Diploneis oblongella' °
.02
—
Epithemia adnata var. porcellus
P
P
Epithemia adnata var. proboscidea"
-
P
Fragilaria brevistriata"
—
P
Fragilaria brevistriata var. inflata
.06
.15
Fragilaria construens'
—
P
Fragilaria constriiens var. binodis
.06
P
Fragilaria constrtiens var. venter
.65
.11
Fragilaria crotonensis'
—
.17
Fragilaria lapponica"
.19
-
Fragilaria pinnata"
.37
—
Fragilaria pinnata var. lancettula
.07
.13
Fragilaria similis
.19
.17
Fragilaria vaucheriae
.90
.63
Fragilaria virescens"
P
—
Gomphonema affine"
.13
-
Gomphonema gracile"
.02
—
Gomphonema intricatum'
—
P
Gomphonema olivaceiim
.30
.05
Gomphonema parvulum
.44
.21
Gomphonema subclavatiim
var. commutatum
.08
P
Gomphonema tenellum"
.02
—
Gomphonema ventricosum' °
.44
—
Gomphonema sp. ° °
.06
—
Melosira granulata
.55
.65
Melosira granulata var. angustissima
1.58
1.05
Melosira italica
.21
P
Navicula arvensis'
_
.15
Navicula arenaria'
—
P
Navicula aurora"
.02
-
Navicula capitata var. hungarica'
—
.1
Navicula cincta'
—
.59
Navicula circumtexta'
—
.15
Navicula cryptocephala
.29
.35
September 1980
Grimes et al.: Diatom Assemblages
225
Table 1 continued.
Species
Living
Dead
Navicula cryptocephala var.
veneta
3.45
4.38
Navicuhi cxigua"
—
P
Navicuhi gmciloides
12.35
9.17
Navicula Iwiifleri var. leptocephala
.08
P
Navicuhi lanccolata °
—
.08
Navicula tnitiima
.07
.84
Navicula oblonga
P
.08
Navicula peregrina'
-
.08
Navicula pupula
P
.08
Navicula radiosa var. tcnclla
1.13
2.42
Navicula rh yncocephala °
—
P
Navicula salinarum °
—
.13
Navicula salinarum var. intc
rmedia
.08
.05
Navicula schroetcri var. cscambia
3.06
6.23
Navicula tcnclloidcs
.15
1.27
Navictila tripunctata
P
.33
Navicula tripunctata var. sch
lizonemoides .44
1.51
Navicula sp.°
—
.63
Nitzschid acicularis
.51
.66
Nitzschia amphibia
.37
.25
Nitz-schia apiculata
.06
.18
Nitzschia di.ssipata
6.00
4.95
Nitz-'ichia filiformis
7.10
6.83
Nitzschia frustulum
.80
.53
Nitzschia gracilis"
.23
—
Ni tzsch ia han tzsch ia na
4.94
4.40
Nitzschia holsatica
1.67
2.35
Nitzschia inconspicua
12.40
13.25
Nitzschia linearis
.06
.08
Nitzschia longissima var. clostcrium
.08
.08
Nitzschia ovalis
.18
..32
Nitzschia palea
6.90
13.50
Nitzschia paleaceae
4.63
3.92
Nitzschia punctata'
—
P
Nitzschia sigmoidca °
—
P
Nitzschia stagnorum' "
P
—
Nitzschia s-p. 1
P
.42
Nitzschia sp. 2°°
.21
—
Ophephora martiji'
-
P
Rhoicosphenia curvata
3.96
6.32
Rhopalodia gibba°
-
.05
Rhopalodia gibberuhi var. vanheurckii
.10
P
Stcphanudiscus astrea"
.06
-
Stcphanodiscus astrea var. minutula
5.15
4.34
Stcphanodisctis niagarae °
-
.08
Surirella angustata
P
P
Surirella ovalis var. brightwe
llii°
—
.15
Surirella ovata°°
.08
—
Sijnedra actis
.08
1.60
Sijnedra dclicatissima var. angiistissima
.15
.02
Sijnedra fasciculata var. trunctata'
-
.08
Sijnedra mazamaensis
.15
.08
Sijnedra socio
P
.08
Synedra ulna
P
.11
Synedra ulna var. contractu
.81
.02
°° Species unique to living Phragmites australis stems.
'Species unique to dead Pbragmites australis stems.
P Species not recorded on the transects taken for relative de
ures but found on other sections of the diatom slide.
iity fig-
Species diversity according to number of
species encountered was high, averaging 48
species per sample. However, there were
generally 2 to 4 dominant species ranging be-
tween 10-18 percent relative density in each
sample, which allowed for only moderately
high Shannon-Wiener diversity values (Table
3). Forty-nine percent of the diatom species
was found in 30 percent of all samples, and
34 percent of the diatom species was found
in 50 percent of all samples.
The results of the cluster analysis (Fig. 1)
demonstrate the high degree of similarity en-
countered for all ten .samples. Even so, mar-
ginal separation into .samples from living and
dead stems was obtained.
A similarity matrix comparing all 10 sam-
ples was constructed.
Means of similarity indices for living stems,
dead stems, and between living and dead
stems were computed. T-tests were per-
formed and it was determined that there was
no significant difference in similarity within
or between these samples sets.
Discussion
Each of our 10 samples consistently con-
tained approximately 50 identifiable species.
In general, no one species represented more
than 18 percent of the total population of
any sample. These conditions are indicative
of a diverse flora that is further supported by
our Shannon-Wiener diversity values and the
average number of species per substrate
(Table 3). Such conditions indicate that the
epiphyte flora in Utah Lake is more diverse
than we previously believed. T-tests were
computed comparing the means of the Shan-
non-Wiener diversity indices of both .sub-
strates as well as the average number of spe-
cies from both living and dead stems. No
significant differences between the values in
either compari.son existed.
A total of 23 diatom genera were encoun-
tered during this study. The number of spe-
cies included in these genera was nearly
equally distributed between living and dead
stems (Table 4). However, substratum prefer-
ences were noted in the genera Navicula and
Gomphonema and in individual .species with-
in several other genera. Of the 114 species
found in the study, 22 were unique to living
and 29 were unique to dead stems.
226
Great Basin Naturalist
Vol. 40, No. 3
T\BLE 2. Important species encountered on Phragmites australis stem sections from Provo Bay, Utah Lake, with
their percent relative densities. Important species are those species with a percent relative density greater than .3
percent in any one sample.
Livin
g stems
Dead stems
Species
1
2
3
4
5
6
/
8
9
10
Achnanthes minutissima
3.1
Amphora perpusiUa
3.3
3.2
Amphora ceneta
17.3
5.6
7 2
11.2
Cyclotella meneojiiniaiui
3.0
Melosira granulata
var. angustissima
3.9
S'avicula cnjptocephala
var. veneta
3.4
4.9
4.2
3.7
5.4
3.3
6.1
3.5
Xaiicula graciloides
9.9
13.2
13.5
14.0
11.2
5.5
13.4
14.9
8.5
13.6
Naiicula radiosa
var. tenelki
3.3
3.2
6.2
Saviciila schroeteri
var. escambia
5.7
6.3
6.2
6.2
10.6
\aviciila tripunctata
var. scliizonemoides
.3.3
\itzschia dissipata
6.8
8.7
3.6
7.8
3.1
4.1
4.6
7.9
3.6
4.7
Xitzschia filifonnis
3.1
12.5
5.4
11.1
3.5
6.6
3.2
7.0
5.2
12.2
Xitzschia inconspicua
15.5
23.4
9.5
7.3
12.2
25.7
6.2
12.1
10.1
.Vitast7i ia hatiztsch ia no
3.1
8.7
4.6
5.4
12.3
3.1
Xitzschia holsatica
3.7
3.3
5.4
.3.9
Xitzschia palea
5.0
5.7
5.7
7.8
10.4
17.7
9.4
12.4
16.1
12.0
Xitzschia paleacea
10.6
4.5
6.8
5.0
3.3
7 7
7.4
Rlioicosph en ia cu na ta
7.7
6.2
6.6
9.4
12.2
Stephanodiscus astrea
1
var. miniitida
3.4
5.3
5.2
6.2
5.5
4.6
4.1
3.2
43 1
The most important species in each sample are indicated by boldface type.
Table 3. Shannon-Wiener diversity values for the five living and five dead Phragmites australis stem sections from
Provo Ba\. Utah Lake.
Sample No.
Livins
Dead
Living
Dead
2.96
2.96
2.86
2.96
3.07
2.962
3.12
2.98
3.09
2.85
3.15
3.038
48
50
33
44
47
44.4
52
47
50
38
51
47.6
*.\verage number of species/substrate.
An analysis of the diatom types unique to
the living steins reveals that most were peri-
phvtic stalk formers, whereas those imique to
the dead stems were mostly periphytic mo-
bile forms.
The distribution of species of Gompho-
nema and Navicula on Phragmites stems
showed significant deviation from random.
Thus, of a total of nine Gomphonema species
encoimtered during this study, eight of these
occurred on living stems, five of which were
restricted to hving stems. Conversely, of four
species that occurred on dead stems, only one
was restricted to that habitat. These data sug-
gest that the living stems provide a more
suitable substrate for several Gomphonema
species. Such species tend to be strictly epi-
phytic in distribution, usuallv being attached
by a gelatinous jelly stij)e (Patrick and Rei-
mer 1966). Whether the preference of these
species for living stems is relative to the
availability of nutrients or the physical condi-
tion of the substrate is yet to be determined.
Some interesting distribution patterns were
also observed in the 26 species of Navicula.
Twenty-five of these were found on the dead
stems, of which 11 were restricted to that
substrate. On the other hand, onlv 15 Xav-
September 1980
Grimes et al.: Diatom Assemblages
227
20
30
PERCENT SIMILARITY
40 50
60
70
—I
1—5 Living
6—10 Dead
2
10
4
Fig. 1. Cluster dendrogram showing similarities of diatoms on living and dead Phragmites australis stem sections
from Provo Bav, Utah Lake.
icula species were observed on the living
steins and only one taxon was restricted to
living stems.
We believe the high number of Navicula
species in our samples can be accounted for,
at least in part, by the fact that many are op-
portimistic, occurring on a wide variety of
substrates. These opportunistic Navicula spe-
cies occurred primarily on dead Phragmites
stems except for one or two species that dom-
inated both living and dead stems. The rea-
son for this is open to speculation, but it is
probably related to nutrient interaction, the
physical condition of the substrate, or re-
duced competition on the dead stems.
The hypothesis that condition of the Phrag-
mites stems had no effect on the presence or
absence of Gomphonema and Naviculu was
tested by chi-square analysis using a 2 X 2
contingency table. The results departed sig-
nificantly from random. This supported the
concept that Gomphonema and Navicula
were separated on the basis of habitat type.
Consistent with other Utah Lake studies,
the diatoms in this study reflect the condition
of the lake waters. Most of the prevalent dia-
toms were either alkaphilous or alkabiontic
forms and also indicators of eutrophy. Addi-
tionally, many are known to have the ability
to withstand elevated levels of dissolved salts.
These data, together with the elevated diver-
sity found at Utah Lake, support the current
view that Utah Lake is a saline-eutrophic
ecosystem.
We recognize the preliminary nature of
the present study. Even so, we believe the
differences shown in communities on the liv-
ing versus the dead stems are significant. Fu-
ture studies are planned to expand our data
228
Great Basin \atur.\list
Vol. 40, No. 3
Table 4. Alphabetical list of diatom genera found on
Phragniites australis stem sections from Pro%o Bay. Utah
Lake, and the occurrence of species from those genera
on Hvine and dead substrates.
Genus
Li\ing
Dead m^in
Achnanthes
Amphora
Ationioeneis
Asterionella
Caloneis
Cocconeis
Coscinodisctis
Cyclotella
Cymbella
Diatonia
Diploneis
Epithemia
Fragilana
Goinphonema
Melosira
Savicula
Sitzschia
Ophepbora
Rhocosphenia
Rhoipalodia
Steplianodiscits
Surirella
Synedra
4
4
1
1
1
1
1
1
3
1
1
1
9
8
3
15
18
0
1
1
2
2
6
1
0
2
9
4
3
25
17
1
1
2
2
■7
base to the other species of emergent macro-
ph\1:es in Utah Lake. Furthermore, we plan
studies to answer the following questions: (1)
Are some epiphytes host specific? (2^ What
patterns of seasonal succession are evident in
the epiphytic flora? (3) \Miat impact does the
epiphxtic flora have on productivity and tro-
phic structure of the lake? These questions
take on added significance for future re-
source management in light of proposed
large-scale changes in Utah Lake, such as the
dikino; of Provo and Goshen bavs.
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linois Press. Urbana.
Sneath. R. H. .\., .\ND R. R. SoK-\L. 1963. Numerical tax-
onom%': principles and practice of numerical clas-
sification. \\". H. Freeman Co.. San Francisco.
573 pp.
Snedecor. G. ^^'., .\ND W. CocHR.\N. 1968. Statistical
methods. Iowa State Press. 593 pp.
Squires. L. E., S. R. Rushforth. .\.sd J. D. Brotherso.n.
1979. Algal response to a thermal effluent: study
of a power station on the Provo River. Utah.
U.S.A. Hvdrobiologia 63(1): 17-32.
St. Cl\ir, L. L., .\.nd S. R. Rushforth. 1977. The dia-
tom flora of the Goshen Warm Springs ponds and
wet meadows, Goshen, Utah. U.S..\. Nova Hed-
wigia 28:35.3-425.
Stowe. W. C.. .\nd J. C. GossELiNK. 1971. Community
structure and production of the epiphvtic algae
in the Barataria Bay area of Louisiana. Paper
read at the .34th annual meeting of the .\merican
Society of Limnolog)' and Oceanography, \\inni-
peg. June 14-17.
^^'ETZEL. R. G. 1964. A comparative study of the pri-
mary productivity of higher aquatic plants, per-
iphvton. and phvtoplankton in a large, shallow
lake. Int. Rev. Ges. Hvdrobiol. 49:1-61.
196.5a. Techniques and problems of primary pro-
ductivity measurements in higher aquatic plants
and periphvton. Mem. 1st. Ital. Idrobiol, Suppl.
18:147-165.'
1969b. Factors influencing photos\-nthesis and e.x-
cretion of dissolved organic matter by aquatic
macroph\tes in hardwater lakes. Verb. Int. \'er.
Limnol. 17:72-85.
POISONOUS PLANTS OF UTAH
Jack D. Brotherson.' Lee A. Szyska,' and William E. Evenson-
Abstr\ct.— a list of the major livestock-poisoning plants has been compiled for the state of Utah. Two hundred
fifteen taxa representing 36 families. 119 genera, and 209 species occur within the state. Fortv-one percent are from
two families, the Asteraceae and the Fabaceae. The remaining families of major imjx)rtance are: Poaceae, Ranuncu-
laceae. Solanaceae. Chenopodiaceae. Brassicaceae. .\scelpiadaceae, Liliaceae. and Euphorbiaceae. Sixtv-nine percent
of the genera occur with a single species. Thirty-three percent of the ta.xa are introduced to the state. .Most of the
plants are insect pollinated; 57 percent are herbaceous perennials.
Most livestock poisoning occurs during the spring. This is due both to concentration of toxins in emerging vegeta-
tion and to the absence of more suitable forage on late winter and spring ranges. Green herbage is poisoning in about
80 percent of all taxa, seeds and fruits in about 15 percent, and the remaining 5 percent have toxic compounds
confined to flower heads, sap. tubers, or roots. Disturbed or cultivated habitats and poorlv managed range harbor the
greatest diversity of poisonous plants. Wetlands contain fewer poisonous taxa than do xeric or mesic areas.
The predominating plant toxins are various alkaloids and glycosides. Sixteen percent of the plants have un-
characterized toxins. Cattle and sheep are more susceptible to poisoning than are horses, swine, or poultrw
Records document man's encounters with
poisonous plants since ancient times. Thev
have played both positive and negative roles
in human cult\ire (Dayton 1948). This con-
spicuous dualitv of poisonous plants remains
a major concern for range management.
Kingsbury's (1964) manual on the poisonous
plants of the United States and Canada was
designed to aid veterinarians and ranchers in
recognizing poisonous plants and the svmp-
toms they produce in poisoned livestock. Val-
entine (1978^ prepared an extensive bibliog-
raphv on the poisonous plants of American
rangelands, and numerous works have been
published dealing with local species lists and
descriptions (Evers 1972, Mihalopoulus 1974,
Schmutz et al. 1968. Stoddard et al. 1949.
USDA 1968).
The scope of this paper is twofold: to pro-
\"ide a list of taxa of the major poisonous
plants of Utah, and to present some general
patterns observed among poisonous plants
within the state. It is hoped that this anno-
tated compilation and discussion will prove
useful to range managers and biologists alike.
NL\TERIALS .\.ND METHODS
Data on poisonous taxa were gleaned from
the published literature and by consultation
with specialists in botany and toxicolog}-.
Much of the descriptive literature on poi-
sonous plants is redundant, consequently,
only the more recent works are cited here.
Criteria used in compiling the list of poi-
sonous plants were:
1. The ta.xon had to be sufficientlv abun-
dant I either native or introduced) in
natural ecosystems to constitute a legiti-
mate threat to livestock or wildlife. For
example, some species of the genus As-
tragalus are known to be toxic but are
not abundant enough within the state to
be considered dangerous (Williams and
Bameby 1977).
2. Ornamentals were included only if they
have escaped widelv from cultivation.
Such plants are frequent along ecotones
or in disturbed habitats.
3. Suspicions of toxicity had to be reason-
abh well-founded. The genus Astra-
galus, for example, is represented by
more than 100 species in Utah (Welsh
1978), but only those ta.xa demonstrably
to.xic were included in the present list-
ing.
Additional variables considered for each
taxon were: life history strategy (annual.
biennial, perennial), patchiness of distribu-
Department of Botany and Range Science. Brigham Young University, Provo, l"tah &4602.
Department of Phvsics and .\stronomv. Brigham Young University. Provo, Utah 84602.
229
230
Great Basin Naturalist
Vol. 40, No. 3
tion, occurrence as a cultivar or as a common
range plant, growth form (vine, forb, grass,
shrub, or tree), generalized habitat require-
ments including elevation, soil texture and
acidity, and moisture preference, nature of
the toxin and its localization within the plant,
seasonality of poisoning, animals affected,
specific juvenile mortality and abortifacient
properties. Not all variables could be docu-
mented for each species.
Results
Taxonomy
Conservatively estimated, Utah has at least
215 major toxic taxa of plants representing 36
families, 119 genera, and 209 species. Thirty-
three percent of these taxa are introduced to
the state. Two of these 36 families, the Aste-
raceae and the Fabaceae contain 41 percent
of the total known taxa of poisonous plants
(Table 1). In decreasing order of floral promi-
nence, the eight next important families are:
Poaceae, Ranunculaceae, Solanaceae, Che-
nopodiaceae, Brassicaceae, Asclepiadaceae,
Liliaceae, and Euphorbiaceae. The number
of toxic taxa within a family is not related to
the degree of toxicity of individual taxa: two
of the most deadly plants, rosary pea or pre-
catory bean {Abms precatorious) and poison
hemlock {Conium maculatum) belong to the
Fabaceae and Apiaceae, one major and one
relatively minor family if numbers alone are
considered.
Table 1. Poisonous plant taxa by families.
Family
No. taxa
Percent
of taxa
occurring in
each family
Fabaceae
50
23
Asteraceae
39
18
Poaceae
19
9
Ranunculaceae
13
6
Solanaceae
11
5
Chenopodiaceae
9
4
Brassicaceae
8
4
Ascelepiadaceae
7
3
Liliaceae
7
3
Euphorbiaceae
6
3
26 other families
46
21
Total
215
Sixteen genera (13 percent) occur in the
flora with more than two taxa. Twenty-two
other genera (18 percent) occur with two
taxa (including Cannabis, which has two sub-
species of a single species). Sixty-nine percent
of the genera occur with a single species.
The taxonomy of poisonous plants is not
readily explained. The poisonous flora of the
eastern half of the United States resembles
that of Europe more than it does that of the
western states (Kingsbury 1961), and Day-
ton's paper (1948) on the poisonous plants of
the continental United States gives a differ-
ent listing of major families than that found
for the state of Utah alone.
Table 2. Major toxins found in Utah's poisonous
plant flora.
Toxin
Percent of
taxa
in which
toxin occurs
Alkaloids
22
Glycosides
22
cyanogenetic glycosides
goitrogenic glycosides
irritant oils
coumarm
steroids and triterpenoids
Seleniinn
11
Nitrates
5
Unknown toxins
16
Other toxins
19
amines
oxalates
resins and resinoids
photosensitizing compounds
nutritional deficiencies
Mechanically injurious
4
Plant Toxins
Most poisonous principles are secondary
by-products of the plants' metabolism (Kings-
bury 1964, Levin 1976). In Utah's flora, many
of these toxic metabolites are loosely classed
as alkaloids and glycosides (Table 2). These
two biochemical groups are primarily arti-
ficial constructs, and each consists of numer-
ous toxins (only a few of which have been
identified) having generally similar molecular
structures or modes of activity.
When the impact of poisonous range
plants on livestock mortality is assessed, how-
ever, many of the minor toxins become or
September 1980
Brotherson et al.: Poisonous Plants
231
serious concern to ranchers. Selenium poison-
ing by members of the genus Astragalus and
other species is an important source of mor-
tahty, as is oxalate poisoning by halogeton
Halogeton glomeratus) and fivehook bassia
Bassia hyssopi folia), and loss of sheep due to
photosensitization by plants such as spring
jarsley {Cijmoptcrus icatsonii) and St. Johns-
.vort {Hypericum fowiosuui). Thus, the com-
nonness of a particular plant species or toxin
loes not necessarily imply high mortality.
Seasonality and Specificity
Most livestock losses occur in the early
pring (Table 3) as animals are turned out
into slowly greening range (Evers 1972, Kee-
er 1978, Kingsburv 1964, Kreger and Sharp
978, Merrill and Schuster 1978). A second,
mailer peak in mortality follows in the sum-
ler as the more palatable vegetation withers
^ the heat and toxins are concentrated in
ruits and seeds of poisonous species.
Cattle are susceptible to poisoning by
lore taxa than sheep, with consequently
igher mortality rates and greater economic
)ss (Nielson 1978). Most of the toxic taxa
ommonly found on Utah's rangeland will
'oison all livestock but others are relatively
pecies specific (such as Delphinium poison-
ig in cattle— ranges infected with the vari-
us species of this plant may be safely grazed
y sheep). Other poisonous taxa may be
razed in moderate amounts without harm if
Iternative palatable forage is available, and
inie species, for example, halogeton, can be
tilized by sheep if the animals are in-
oduced gradually to the plant, allowing
leir rumen microflora to adapt to
etoxifying large amounts of calcium oxalate
ameset al. 1976).
Table 3. Livestock inortalitv bv seasons.
•ason
Percent of
reported poisonings
.ring"
65
miner
17
ill
11
inter
6
Mortality especially high in the late winter and earlv spring
The Ecology of Poisonous Plants
The ecology of Utah's poisonous flora is
highly variable, but our data permit a few
general observations. Most of the plants are
insect pollinated, and 57% are herbaceous
perennials. Green herbage is toxic in about
80% of all taxa, .seeds and fruits in about 15%,
and about 5% of the taxa have toxins con-
fined to flower head, sap, tubers and roots.
The relative locations and concentrations of
toxins within a plant may vary through the
growing season, and depend upon the par-
ticular taxon being considered.
Toxic species are likely to be found any-
where in the state, although disturbed habi-
tats and poorly managed rangelands are espe-
cially prone to harbor dangerous species. The
predominance of poisonous plants in these
habitats reflects both a bias in the reported
literature and the very real dangers of over-
grazing in the western states. Dry desert soils
have more taxa of poisonous plants than mes-
ic or hydric ones; however, some of the indi-
vidually most deadly plants occur in wet-
lands. Data on elevation, soil texture and
acidity were insufficiently complete to per-
mit any valid conclusions.
Discussion and Summary
The benefit to the plant of manufacturing
and maintaining toxic substances is unknown.
A few phytotoxins such as abrin and cicuto-
xin appear to function specifically as verte-
brate poisons (Kingsbury 1961, 1964) and
may have evolved in response to herbivore
pressures (Laycock 1978). Selenium accu-
mulators are toxic due to characteristics of
the soil, although primary accumulators may
actually require trace amounts of selenium
for proper growth (Kingsbury 1964).
There does not appear to be any univer-
sally applicable method for managing range-
lands infested with poisonous plants. Species
that occur in dense clumps or thickets may
be individually irradicated by chemical or
mechanical means. More commonly, how-
ever, vast areas will be infested, often as a re-
sult of overgrazing. Wise management will
include aspects of the following practices:
1. Recognition of poisonous plants and an
accurate assessment of their potential
232
Great Basin Naturalist
Vol. 40, No. 3
lethality. Unless forced by hunger, live-
stock generally will ignore poisonous
plants in favor of more palatable for-
age.
2. Removal of susceptible animals.
3. Provision of sufficient alternate forage
if animals must be turned onto ranges in
early spring before palatable plants are
abundant.
4. Scheduling of range use around live-
stock susceptibility patterns.
There are several good reviews of manage-
ment practices (Evers 1972, Keeler 1978,
Knieger and Sharp 1978, Merrill and Schus-
ter 1978), and the range literature abounds in
articles dealing with specific plants and their
effects (see Valentine 1978 for a com-
prehensive listing). The purpose of the pres-
ent paper is not to review management tech-
niques in detail, but primarily to provide an
updated listing of Utah's dangerous flora.
Further research needs to be done not only in
management but in toxicology and pharma-
cology so that control of poisonous plants
will be a matter of understanding instead of
irradication and vast areas of range can again
be utilized and productive.
POISONOUS PLANTS OF UTAH
The present list of plant taxa was gleaned
from published literature and by consultation
with experts in botany and toxicology as
cited above.
Certain plant characteristics are desig-
nated for each taxon in parentheses immedi-
ately following the taxon name. The abbre-
viations are as follows:
p
Perennial
B
Biennial
A
Annual
N
Native
I
Introduced
T
Tree
S
Shnib
F
Forb
G
Crass
R
Rush
The following list of families, genera, spe-
cies, and varieties is arranged in alphabetical
order for ease of reference. Plant synonymy
follows Manual of the Plants of Colorado
(Harrington 1964) and Utah Plants (Welsh
and Moore 1973).
Amaranth ACE AE
Amaranthus retroflexus L. (AIF)
Common name: pigweed, carelessweed,
redroot, redroot amaranth.
Toxin: nitrates, under conditions of over-
fertilization with too little water.
Habitat: common garden and field weed;
waste places.
Animals affected: livestock.
Reference: Brakenridge 1956, Kingsbury
1964.
Anacardiaceae
Toxicodendron rydbergii (Small) Greene
(PNS)
Common name: poison ivy.
Toxin: 3-n-pentadecylcatechol.
Habitat: moist areas at lower elevations.
Animals affected: humans (dermatitis).
Reference: McNair 1923, Kingsbury 1964.
Apiaceae
Cicuta douglasii (DC.) Coult. & Rose (PNF)
Common name: Douglas waterhemlock.
Toxin: alcohol (cicutoxin).
Habitat: swampy or wet habitats along
streams and in marshes.
Animals affected: livestock, humans.
Reference.: Alberts 1931, Kingsbury 1964.
Conium tnaculatum L. (BIF)
Common name: hemlock, poisonous hem-
lock, spotted hemlock, California or Nebras-
ka fern.
Toxin: alkaloids (conine, N-methyl conine,
conhydrine, lambda-coniceine, pseudoconhy-
drine).
Habitat: weed of roadsides, ditches, edges
of cultivated fields and other waste areas.
Animals affected: livestock, humans.
Reference: Penny 1953, Kingsbury 1964.
Daucus carota L. (BIF)
Common name: wild ;parrot. Queen Ann's
lace.
Toxin: nitrates, under conditions of over-
fertilization with too little water.
Habitat: waste places.
Animals affected: horses, cattle.
Reference: Volker 1950, Kingsbury 1964.
Pastinaca sativa L. var. sylvestris DC. (BIF)
Common name: common parsnip.
Toxin: unknown.
September 1980
Brotherson et al.: Poisonous Plants
233
Habitat: widely naturalized weed.
Animals affected: humans (dermatitis).
Reference: S. L. Welsh (personal commu-
nication, 1980).
Slum suave Walt. (PNF)
Common name: water parsnip, hemlock
water parsnip.
Toxin: unknown.
Habitat: marshy lands and wet soils.
Animals affected: hogs, cattle.
Reference: Fyles 1920, Kingsbury 1964.
Apocynaceae
Apocynum androsaemifolium L. (PNF)
Common name: spreading dogbane.
Toxin: resins, glycosides.
Habitat: common weed of open places, in
coarse soils along streams, meadows, and
wooded hillsides.
Animals affected: cats, dogs.
Reference: Moore 1909, Kingsbury 1964.
Note: The closely related species A. me-
dium Greene and A. sibericwn Jacq. niav
show similar effects.
Apocynum cannabinum L. (PNF)
Common name: Indian hemp, dogbane,
hemp dogbane.
Toxin: resins, glycosides.
Habitat: common weed of open places, in
coarse soils along streams.
Animals affected: cats, dogs.
Reference: Finnemore 1909, Kingsbury
1964.
Nerium oleander L. (PIS)
Common name: oleander.
Toxin: glycosides.
Habitat: cultivated greenhouse plant,
street plant in St. George.
Animals affected: livestock, humans.
Reference: West 1957, Kingsbury 1964.
Asclepiadaceae
Asclepias asperula (Decne) Woodson (PNF)
Common name: asper milkweed, spider
antelopehorn.
Toxin: resinoids, glycosides and an alka-
loid.
Habitat: open dry soils, flats, desert swales,
sandy or rocky hillsides with pinyon, juniper
or oak.
Animals affected: sheep, cattle, goats,
horses, poultry.
Reference: Huffman 1956, Kingsbury 1964.
Asclepias fascicularis Decne ex DC. (PNF)
Common name: Mexican whorled milk-
weed.
Toxin: resinoids, glycosides and an alka-
loid.
Habitat: dry hillsides and roadsides; pas-
tures, moist streamsides.
Animals affected: sheep, cattle, goats,
horses, fowl.
Reference: Schmutz et al. 1968.
Asclepias incarnata L. (PNF)
Common name: swamp milkweed.
Toxin: resinoids, glycosides and an alka-
loid.
Habitat: marshes.
Animals affected: sheep, cattle, horses,
poultry.
Reference: Hansen 1924, Kingsbury 1964.
Asclepias labriformis Jones (PNF)
Common name: labriform milkweed.
Toxin: resinoids, glycosides and an alka-
loid.
Habitat: in sandy soils along old stream
beds.
Animals affected: sheep.
Reference: Holmgren 1945, Kingsbury
1964.
Asclepias latifolia (Torr.) Raf. (PNF)
Common name: broadleaf milkweed.
Toxin: resinoids, glycosides and an alka-
loid.
Habitat: dry plains in sandy soils.
Animals affected: sheep.
Reference: Schmutz et al. 1968, Kingsbury
1964, Shrift 1958.
Asclepias speciosa Torr. (PNF)
Common name: showy milkweed.
Toxin: resinoids, glycosides, and an alka-
loid.
Habitat: prairies and open areas.
Animals affected: sheep.
Reference: Fleming 1920, Kingsbury 1964.
Asclepias subverticillata (Gray) Vail (PNF)
Common name: whorled milkweed, west-
ern whorled milkweed.
Toxin: resinoids, glycosides and an alka-
loid.
234
Great Basin Naturalist
Vol. 40. No. 3
Habitat: dry plains and foothills; spreads
rapidlv along waterways and irrigation
canals, forming dense stands; prefers sandy
soils.
.\niinals affected: sheep.
Reference: Glover 1917, Kingsbiir\- 1964.
.\steraceae
Achillea miJlefolium L. iPNF)
Common name: \arrow.
To.xin: alkaloids and glycosides.
Habitat: various.
Animals affected: livestock.
Reference: Hurst 1942, Kingsbury 1964.
Ambrosia tomentosa Nutt. (PNF^
Common name: white ragweed, skeleton
leaf bursage
To.xin: nitrates, imder conditions of over-
fertihzation with too little water.
Habitat: dr\" plains, hills, waste ground and
fields.
Animals affected: livestock.
Reference: Huffman 1956, Kingsbury 1964.
Anthemis cotula L. (AIF)
Common name: dog fennel, mayweed.
ma\Aveed camomile.
Toxin: acrid substance irritating to mucous
membranes.
Habitat: weedy plant of disturbed soils,
fields and waste places; common weed in
hay.
.\nimals affected: poultry".
Reference: Los Angeles Count}- Livestock
Department 1938, Kingsbur>- 1964.
Artemisia fiUfoUa Torr. (PNS^
Common name: sand sagebrush, old man
sagebrush.
Toxin: volatile oils.
Habitat: sandy soils.
Animals affected: horses.
Reference: Beath 1953, Kingsbur>- 1964.
Artemisia spinescens (DC.) Eaton PNS)
Common name: bud sagebrush.
Toxin: volatile oils.
Habitat: dry plains and hills.
Animals affected: hvestock.
Reference: Sampson 1942, Kingsbur\- 1964.
Aster chilensis Nees ssp. adscendens (Lindl.)
Cronq, PNFi
Common name: pacific aster.
Toxin: secondary- selenium accumulator.
Habitat: widely scattered in moist habitats.
Animals affected: livestock.
Reference: Trealease and Beath 1949.
Kingsbury 1964.
Aster glaucodes Blake i^PNF^
Common name: gray aster.
Toxin: secondary- selenium accumulator.
Habitat: moimtains.
Animals affected: sheep.
Reference: Trelease and Beath 1949,
KingsbuT)" 1964.
Aster laevis L. (PNF)
Common name: smooth aster.
Toxin: secondarv selenium accumulator.
Habitat: widely scattered in drv to moist
habitats.
Animals affected: livestock.
Reference: Trelease and Beath 1949.
Kingsbur) 1964.
Aster occidentalis (Nutt.) Torr. 6c Grav
iPNF)
Common name: western aster.
To.xin: secondary- selenium acctunulator.
Habitat: mountain meadows at moderate
elevations.
Animals affected: livestock.
Reference: Trelease and Beath 1949,
Kingsbury 1964.
Aster pauciflorus Nutt. (PNF)
Common name: fewhead aster.
Toxin: secondary- selenium accumulator.
Habitat: widespread in saline soils.
Animals affected: livestock.
Reference: J. D. Brotherson (personal com-
munication. 19801
Bahia oppositifolia (Nutt.) DC. iPNF)
Common name: bahia. plains bahia.
Toxin: cvanogenetic glvcoside.
Habitat: dr)' soils; plains and hills.
Animals affected: cattle, sheep.
Reference: Deem et al. 1939. Kingsburv
1964.
Baileya midtiradiata Har\-. & Gray (BNF)
Common name: desert baileya. cloth of
gold, desert marigold.
Toxin: unknown.
Habitat: sandy and gravelly soils in dr\'
areas.
Animals affected: sheep, goats.
Reference: Mathews 1933, Kingsburv
1964.
September 1980
Brotherson et al.: Poisonous Plants
235
Baileya pleniradiata Har\ . & Gray (ANF)
Common name: desert marigold baileya.
Toxin: unknown.
Habitat: mesas and deserts of southeastern
Utah.
Animals affected: sheep, goats.
Reference: Schmutz et al. 1968.
Centaurea repens L. (PIF)
Common name: Russian knapweed.
Toxin: unknown; produces nigropallidal
encephalomalacia.
Habitat: fields, roadsides and waste places.
^Animals affected: horses.
Reference: Mecke 1979.
Centaurea sohtitialis L. (AIF)
Common name: yellow star thistle, vellow
centaurea.
Toxin: imknown; produces nigropallidal
encephalomalacia; also mechanically in-
jurious.
Habitat: waste places, fields and roadsides.
Animals affected: horses.
Reference: Mettler and Stem 1963, Kings-
bur>- 1964.
Chrysothamnus nauseosus (Pall.) Britton
(PXS)
Common name: rubber rabbitbrush.
Toxin: unknown.
Habitat: dr\. open places at moderate and
low elevations.
Animals affected: livestock.
Reference: Sampson 1942, Kingsbury 1964.
Gritidelia squarrosa (Pursh) Dunal (BXF)
Common name: gum weed, gumplane, cur-
lycup gumweed.
To.xin: secondarv selenium accumulator.
Habitat: dr\ open places; prairies, plains,
roadsides and fields.
Animals affected: livestock.
Reference: Trelease and DiSomma 1960,
Kingsbury 1964.
Heleiiium autumnaJe L. (PXF^
Common name: sneezeweed. bitterweed.
Toxin: unknown acrid substance.
Habitat: moist low ground in lowlands and
foothills.
Animals affected: sheep and cattle.
Reference: Kingsbury 1964.
Helenium hoopesii Gray (PNF)
Common name: sneezeweed. orange
sneezeweed.
Toxin: glycoside (dugaldine).
Habitat: high mountain slopes and vallevs,
often forming dense stands in moist, sunnv,
undisturbed localities.
Animals affected: sheep and cattle.
Reference: Marsh et al. 1921, Kingsbury
1964.
Heleomeris longifolia Rob. & Greenm. var.
annua (Jones) Yates (AXF)
Common name: annual goldeneye, resin-
weed, talloweed.
Toxin: unknown.
Habitat: ranges, hills, plains, and river bot-
toms.
Animals affected: cattle.
Reference: Schmutz et al. 1968, Kingsbury
1964.
Hymenoxys richardsonii (Hook.) Cockerel!
iPXF
Common name: pingue, Colorado rubber-
weed, pingue hymeno-xys, rubberweed.
Toxin: unknown; may be associated with
mineral imbalance.
Habitat: dry. rocky or clay soils of plains
and mountain slopes from 1500 to 12,000
feet.
Animals affected: sheep, cattle, goats.
Reference: Aanes 1961, Kingsbury 1964.
Oxytenia acerosa Xutt. PXS)
Common name: copperweed; prickly oxy-
tenia.
Toxin: unknown.
Habitat: alkaline soils in draws or stream-
beds of desert ranges and foothills.
Animals affected: cattle, sheep.
Reference: Throp et al. 1940, Kingsburv
1964.
Psathyrotes annua (Nutt.) Gray i.\XF)
Common name: annual psathyrotes.
Toxin: unknown.
Habitat: dry, sandy, often alkaline soils, es-
pecially of creek beds and dr)- washes.
.\nimals affected: sheep.
Reference: Binns et al. 1962, Kingsbur\
1964.
Psilostrophe sparsiflora (Gray) A. Nels.
(PXF)
Common name: greenstem paperflower.
Toxin: unknown; induces kidney damage.
Habitat: dr\-, open range.
236
Great Basin Naturalist
Vol. 40, No. 3
Animals affected: sheep.
Reference: Huffman 1956, Kingsbury 1964.
Rudbeckia occidentalis Nutt. (PNF)
Common name: western coneflower, nig-
gerheads.
Toxin: unknown.
Habitat: streambanks and woodlands.
Animals affected: generally unpalatable to
livestock; affects hogs and sheep in feeding
trials.
Reference: Pammel 1911, Kingsbury 1964.
Senecio integerrimus Nutt. (PNF)
Common name: groundsel, senecio, lambs-
tongue groimdsel.
Toxin: alkaloids.
Habitat: dry or moist open woods and
slopes, from valleys to near timberline.
x\nimals affected: livestock, humans (?).
Reference: Clawson 1933, Kingsbury 1964.
Senecio longilobus Benth. (PNF)
Common name: wooly groundsel, thread-
leaf groundsel.
Toxin: pyrrolizidine alkaloids.
Habitat: dry slopes, mesas and dry washes.
Animals affected: cattle, horses, sheep,
goats.
Reference: Clawson 1933, Kingsbury 1964.
Senecio spartioides Torr. & Gray (PNF)
Common name: broom groundsel.
Toxin: pyrrolizidine alkaloids.
Habitat: valleys, plains; open areas and
pine forests.
Animals affected: cattle, horses, sheep,
goats, humans(?).
Reference: Clawson 1933, Kingsburv 1964,
Davis 1958.
Senecio vulgaris L. (AIF)
Common name: common groundsel.
Toxin: pyrrolizidine alkaloids.
Habitat: weed of gardens and waste places.
Animals affected: cattle, horses, sheep,
goats, humans(?).
Reference: Steyn 1934, Kingsbury 1964.
Solidago parryi (Gray) Greene (PNF)
Common name: Parry goldenweed.
Toxin: unknown; causes milk-sickness or
trembles.
Habitat: mountains, coniferous forests.
Animals affected: cattle.
Reference: Schmutz et al. 1968, Kingsburv
1964.
Tanacetum vulgare L. (PIF)
Common name: common tansv.
Toxin: abortifacient.
Habitat: weed along roadsides, waste areas,
ditchbanks and other moist areas.
Animals affected: cattle, humans(?).
Reference: Cress 1935, Kingsbury 1964.
Tetradymia canescens DC. (PNS)
Common name: spineless horsebrush, gray
horsebrush.
Toxin: photosensitizing compounds.
Habitat: dry desert and sagebrush ranges.
Animals affected: sheep.
Reference: Kingsburv 1964, Schmutz et al.
1968.
Tetradymia glabrata Gray (PNS).
Common name: littleleaf horsebrush,
spring rabbitbrush, coaloil brush.
Toxin: photosensitizing compounds.
Habitat: dry desert and sagebrush ranges.
Animals affected: sheep.
Reference: Kingsburv 1964, Fleming et al.
1922.
Tetradymia ntittallii T. & G. (PNS)
Common name: Nuttall horsebrush.
Toxin: photosensitizing compounds.
Habitat: dry desert and sagebaish ranges.
Animals affected: sheep.
Reference: Kingsbury 1964.
Tetradymia spinosa T. and G. var. long-
ispina Jones (PNS)
Common name: longspine horsebrush.
Toxin: photosensitizing compounds.
Habitat: dry desert and sagebrush ranges.
Animals affected: sheep.
Reference: S. L. Welsh (personal commu-
nication, 1980).
Tetradymia spinosa T. and G. var. spinosa
(PNS)
Common name: spinv horsebrush.
Toxin: photosensitizing compounds.
Habitat: dry desert and sagebrush ranges.
Animals affected: sheep.
Reference: S. L. Welsh (personal commu-
nication, 1980).
Xanthium strumarium L. (AIF)
Common name: spiny clotbur, spiny cock-
lebur.
September 1980
Brotherson et al.: Poisonous Plants
237
Toxin: hydroquinone.
Habitat: fields and wastelands; along
shores of ponds, rivers and in flood plains.
.\nimals affected: livestock, fowl, hogs, hu-
mans (dermatitis).
Reference: Forrest 1938, Kuzel and Miller
1950, Kingsbury 1964.
Xanthocephalum microcephahim (DC.)
Gray (PNS)
Common name: broomweed, perennial
snakeweed, slinkweed, turpentine weed,
threadleaf snakeweed, matchweed, resin-
weed.
Toxin: saponin.
Habitat: dry stony plains, slopes and mesas.
Animals affected: cattle, sheep, goats,
swine, chicks, rabbits.
Reference: Dollahite 1957, Kingsburv
1964.
Xanthocephalum sarothrae (Pursh) Britt.
and Rushy (A\F)
Common name: broom snakeweed, snake-
weed, matchbnish.
Toxin: saponin.
Habitat: dry stony plains, slopes and mesas.
Animals affected: cattle, sheep, goats,
swine, chicks, rabbits.
Reference: Dollahite 1962, Kingsburv
1964.
BORAGINACEAE
Amsinckia intermedia Fisch, & Mey (AXF)
Common name: tarweed, fiddleneck, fire-
weed fiddleneck.
Toxin: imknown; potentially lethal nitrate
levels, pyrrolizidine alkaloids (?).
Habitat: dry open cultivated ground or
waste areas.
Animals affected: horses, hogs, cattle.
Reference: McCulloch 1940, Kingsburv
1964.
Note: The closelv related species A. tesse-
lata Gray and A. retrorsa Suksd. may show
similar effects.
Cynoglossiim officinale L. (BIF)
Common name: houndstongue.
Toxin: unknown.
Habitat: waste places of plains and hills.
Animals affected: livestock.
Reference: Kingsbury 1964, S. L. Welsh
(personal commimication, 1980).
Brassicace.\e
Brassica hirta Moench. (AIF)
Common name: white mustard.
Toxin: cyanogenetic glycoside.
Habitat: cultivated weed, escaped to waste
areas.
Animals affected: cattle, sheep.
Reference: Eaton 1941, Kingsbury 1964.
Brassica kaber Wheeler (.\1F)
Common name: charlock, wild mustard.
Toxin: cyanogenetic glycoside.
Habitat: common weed of grain crops and
in waste areas.
Animals affected: cattle, hogs, sheep.
Reference: Thomson and Sifton 1922,
Kingsbm-y 1964.
Descurainia pinnata (Walt.) Britt. (ANF)
Common name: tansv mustard, pinnate
tansy mustard.
Toxin: unknown.
Habitat: heavy stands on dry, sandy soils.
Animals affected: cattle.
Reference: Hershev 1935, Kingsburv 1964.
Erysimum cheiranthoides L. (ANF)
Common name: wormweed mustard,
treacle wallflower.
Toxin: cvanogenetic glycoside.
Habitat: weed of cultivation, roadsides,
meadows; moist waste areas in valleys and
canyons.
Animals affected: hogs.
Reference: Thomson and Sifton 1922,
Kingsbur) 1964.
Stanleya integrifolia James (PXS)
Common name: wholeleaf desert prince's
plume.
Toxin: primarv selenium accumulator.
Habitat: dry plains and hills.
Animals affected: Not observed to be eaten
bv livestock.
' Reference: Beath et al. 1953, Kingsbur\'
1964.
Stanleya pinnata (Pursh) Britt. (PNS)
Common name: prince's plume, desert
prince's plume.
Toxin: primary selenium accumulator.
Habitat: desert soils, dry plains and mesas.
Animals affected: experimental; normally
unpalatable.
Reference: Beath et al. 1953, Kingsbury
1964.
238
Great Basin Naturalist
Vol. 40, No. 3
Stanletja viridiflora Nutt. (PNF)
Common name: greenflower prince's
plume.
Toxin: primary selenium accumulator.
Habitat: dry plains and hills.
Animals affected: Not observed to be eaten
by livestock.
Reference: Beath et al. 1953, Kingsbury
1964.
Thlaspi arvense L. (AIF)
Common name: fanweed, field penny-
cress.
Toxin: cyanogenetic glycoside.
Habitat: Common weed of cultivated and
waste places.
Animals affected: livestock.
Reference: Thomson and Sifton 1922,
Kingsbury 1964.
C.wnabinaceae
Cannabis sativa L. ssp. sativa (AIF)
Common name: marijuana, hemp.
Toxin: narcotic element contained in tetra-
hydrocannabinol.
Habitat: waste places.
Animals affected: humans, livestock.
Reference: Steyn 1934, Kingsbury 1964,
Small and Cronquist 1976, Welsh 1980.
Cannabis sativa L. ssp. indica (Lam.) Small
& Cronq. (AIF)
Common name: marijuana, hemp.
Toxin: narcotic element contained in tetra-
hydrocannabinol.
Habitat: waste places.
Animals affected: humans, livestock.
Reference: Small and Cronquist 1976,
Welsh 1980.
Caprifoliaceae
Sambucus coerulea Raf. (PNS)
Common name: blue elderberry.
Toxin: unknown.
Habitat: moist soils of plains and hills.
Animals affected: cattle, children (?)
Reference: Schmutz et al. 1968.
Sambucus racemosa L. (PNS)
Common name: red elder.
Toxin: unknown; concentrated in root.
Habitat: moist forests, 7,500-10,000 ft.
Animals affected: cattle, children(?)
Reference: Schmutz et al. 1968.
Caryophyllaceae
Saponaria officinalis L. (PIF)
Common name: bouncing bet, soapwort.
Toxin: saponin.
Habitat: fields, waste places; cultivated
and escaping.
Animals affected: sheep.
Reference: Kingsbury 1964.
Chenopodiaceae
Atriplex gardneri Moq. (PNS)
Common name: Nuttall saltbush, Gardner
saltbush.
Toxin: secondary selenium accumulator.
Habitat: saline plains and hillsides.
Animals affected: livestock.
Reference: Fleming 1920, Kingsbury 1964.
Bassia hyssopifolia (Pall.) Volk (AIF)
Common name: fivehook bassia, smoth-
erweed.
Toxin: oxalates.
Habitat: dry, saline soils.
Animals affected: sheep.
Reference: Pammel 1911.
Chenopodium ambrosioides L. (AIF)
Common name: wormseed goosefoot.
Toxin: antihelminthic oil.
Habitat: weed of waste places.
Animals affected: geese, humans.
Reference: Bamford 1951, Kingsbury 1964.
Chenopodium album L. (AIF)
Common name: lambsquarter.
Toxin: nitrates, under conditions of over-
fertilization with too little water.
Habitat: weed of waste places.
Animals affected: livestock.
Reference: Case 1957, Kingsbury 1964.
Chenopodium glaucum L. (AIF)
Common name: oakleaf goosefoot.
Toxin: nitrates, under conditions of over-
fertilization with too little water.
Habitat: weed of waste pmces.
Animals affected: livestock.
Reference: Case 1957, Kingsbury 1964.
Halogeton glomeratus (Bieb.) C. A. Mey
(AIF)
Common name: halogeton, barilla.
Toxin: oxalates.
Habitat: dry saline plains and alkaline
soils; roadsides.
September 1980
Brotherson et al.: Poisonous Plants
239
Animals affected: livestock, especially
sheep.
Reference: Cook and Stoddart 1953, Kings-
bury 1964.
Kochia scoparia L. Schrad, (AIF)
Common name: summer cypress, burning
bush, Mexican fireweed. Belvedere summer
cypress.
Toxin: photosensitizing compounds.
Habitat: dry soils, roadsides and waste
places.
Animals affected: cattle, sheep, horses.
Reference: Rottgardt 1944, Kingsbury
1964, Schmutz et al. 1968.
Salsola iberica Sennen & Pau (AIF)
Common name: Russian thistle.
Toxin: nitrates (?), possible oxalates (?). '
Habitat: dry soils of plains and foothills.
Animals affected: livestock.
Reference: Huffman et al. 1956.
Sarcobatus vermiculutus (Hook.) Torr. (PNS)
Common name: greasewood, black grease-
wood.
Toxin: oxalates.
Habitat: dense stands confined to alkaline
flats or saline soils of low and lower middle
elevations.
Animals affected: sheep, sometimes cattle.
Reference: Kouch 1922, Kingsbury 1964.
Cyperaceae
Scirpus pungens Vahl (PNR)
Common name: bulrush, three-square,
American bulrush.
Toxin: unknown; suspected of producing
pulmonary emphysema.
Habitat: wet or moist ground.
Animals affected: cattle.
Reference: Beath et al. 1953, Kingsburv
1964.
Equisetaceae
Equisetum arvense L. (PNF)
Common name: horsetail, foxtail, rush,
marsh horsetail.
Toxin: alkaloids.
Habitat: sandy or gravelly soils along
streams and in moist fields and meadows.
Animals affected: horses.
Reference: Gussow 1912, Kingsbury 1964.
Equisetum laevigatum A. Br. (PNF)
Common name: smooth horsetail.
Toxin: thiaminase.
Habitat: marshes, alluvial thickets, sandy
banks; weed of cultivation.
Animals affected: horses.
Reference: Samp.son and Malmsten 1942,
Kingsbury 1964.
Ericaceae
Kalmia microphylla (Hook.) Heller (PNS)
Common name: pale laurel, bog laurel, al-
pine kalmia.
Toxin: resinoids, andromedotoxin.
Habitat: wet meadows and bogs of high
elevation.
Animals affected: sheep, calves, goats.
Reference: Kingsbury 1964.
Ledum glandulosum Nutt. (PNS)
Common name: western Labrador tea.
Toxin: resinoids, andromedotoxin.
Habitat: wet meadows and bogs of high
elevation.
Animals affected: sheep, cattle.
Reference: Kingsbury 1964.
Euphorbiaceae
Croton longipes Jones (PNF)
Common name: croton.
Toxin: croton oil (caustic).
Habitat: roadsides, fields, and dry stream-
beds; artemisia and pinyon belts.
Animals affected: livestock.
Reference: Schmutz et al. 1968, S. L.
Welsh' (personal communication, 1980).
Croton texensis (Klotz.) Muell. Arg. ex DC.
(ANF)
Common name: Texas croton.
Toxin: croton oil (caustic).
Habitat: roadsides, fields, and dry stream-
beds; artemisia and pinyon belts.
Animals affected: livestock.
Reference: Volker 1950, Kingsbury 1964,
Schmutz et al. 1968.
Euphorbia cyparissias L. (PIF)
Common name: cypress spurge, graveyard
weed.
Toxin: unknown acrid principle.
Habitat: cultivated and escaping to road-
sides and waste places.
240
Great Basin Naturalist
Vol. 40, No. 3
Animals affected: cattle.
Reference: Muenscher 1935, Kingsbury
1964.
Euphorbia esula L. (PIF)
Common name: leafy spurge.
Toxin: unknown acrid principle.
Habitat: field weed of roadsides and waste
places.
Animals affected: horses, sheep.
Reference: Johnston and Peake 1960,
Kingsbury 1964.
Reverchonia arenaria Gray (ANF)
Common name: reverchonia, sand rever-
chonia.
Toxin: unknown.
Habitat: uncommon; plains and hillsides,
sandy areas, Kane Co.
Animals affected: sheep.
Reference: Schmutz et al. 1968, Kingsbury
1964.
Ricinus communis L. (AIF)
Common name: castor bean.
Toxin: ricin (a phytotoxin).
Habitat: cultivated as an ornamental.
Animals affected: livestock, humans.
Reference: Clarke 1947, Kingsbury 1964.
F.\BACEAE
Acacia greggii Gray (PNT)
Common name: catclaw acacia.
Toxin: cyanogenetic glycoside.
Habitat: plains and dry canyons; forms
thickets along Beaver Dam Wash, Washing-
ton Co.
Animals affected: sheep.
Reference: Schmutz et al. 1968, Kingsburv
1964.
Astragalus asclepiadoides Jones (PNF)
Common name: milkweed milkvetch.
Toxin: primary selenium accumulator.
Habitat: saline desert areas.
Animals affected: livestock.
Reference: Welsh 1978.
Astragalus hisulcatus (Hook.) Gray (PNF)
Common name: two-grooved poisonvetch,
two-grooved milkvetch.
Toxin: primary selenium accumulator.
Habitat: plains and bottom lands, sage-
brush zone.
Animals affected: livestock.
Reference: Welsh 1978, Kingsbury 1964.
Astragalus convallarius Greene var. con-
vallarius (PNF)
Common name: timber poisonvetch, lesser
rushy milkvetch.
Toxin: produces locoism.
Habitat: dry hillsides, desert shrub to lower
montane zones.
Animals affected: livestock.
Reference: Muenscher 1951, Kingsbury
1964.
Astragalus drummondii Dougl. ex Hook.
(PNF)
Common name: Drummond milkvetch.
Toxin: produces locoism.
Habitat: plains and hillsides, brushy places.
Animals affected: livestock.
Reference: Welsh 1978, Kingsbury 1964.
Astragalus eastwoodae Jones (PNF)
Common name: Eastwood poisonvetch,
Eastwood milkvetch.
Toxin: primary selenium accumulator.
Habitat: dry hillsides.
Animals affected: livestock.
Reference: Welsh 1978, Kingsbury 1964.
Astragalus flavus Nutt. ex Torr. & Gray var.
argillosus (Jones) Barneby (PNF)
Common name: yellow milkvetch.
Toxin: primary selenium accumulator.
Habitat: dry plains and hillsides, salt desert
areas.
Animals affected: livestock.
Reference: Welsh 1978.
Astragalus flavus Nutt. ex Torr. & Gray var.
candicans Gray (PNF)
Common name: Canada yellow milkvetch.
Toxin: primary selenium accumulator.
Habitat: dry plains and hillsides, shales and
clays of southern Utah.
Animals affected: livestock.
Reference: Welsh 1978.
Astragalus flavus Nutt. ex Torr. & Gray var.
flavus (PNF)
Common name: yellow milkvetch.
Toxin: primary selenium accumulator.
Habitat: dry plains and hillsides, saline silts
and clays in saline desert areas, south-central
Utah.
Animals affected: livestock.
Reference: Welsh 1978, Kingsbury 1964.
September 1980
Brotherson et al.: Poisonous Plants
241
Astragalus iselyi Welsh (PNF)
Common name: Isely milkvetch.
Toxin: primary selenium accumulator.
Habitat: dry hillsides, salt desert areas.
Animals affected: livestock.
Reference: Welsh 1978.
Astragalus lentiginosus Dougl. ex Hook. var.
araneosus (Sheld.) Barneby (PNF)
Common name: spider locoweed, cobweed
milkvetch.
Toxin: produces locoism.
Habitat: dry hillsides in sagebrush.
Animals affected: livestock.
Reference: Welsh 1978, Kingsbury 1964.
Astragalus lentiginosus Dougl. ex Hook. var.
palans (Jones) Jones (PNF)
Common name: straggling milkvetch.
Toxin: produces locoism.
Habitat: salt desert areas, dry hillsides and
canyons, mixed desert shrub communities.
Animals affected: livestock.
Reference: Welsh 1978, S. L. Welsh (per-
sonal communication, 1980), Kingsbury 1964.
Astragalus lentiginosus Dougl. ex Hook. var.
wahweapensis Welsh (PNF)
Common name: Wahweap loco, Wahweap
milkvetch.
Toxin: produces locoism.
Habitat: dry hillsides in sagebrush, sandy
soils, Kane Co.
Animals affected: livestock, esp. horses.
Reference: Welsh 1978, Kingsbury 1964.
Astragalus miser Dougl. ex Hook. var. ob-
longifolius (Rydb.) Cronq. (PNF)
Common name: timber milkvetch, Ryd-
berg weedv milkvetch.
Toxin: miserotoxin.
Habitat: widely scattered in lower mon-
tane zones.
Animals affected: livestock.
Reference: Williams 1969, Welsh 1978.
Astragalus moencoppensis Jones (PNF)
Common name: Moenkopi poisonvetch,
Moenkopi milkvetch.
Toxin: primary selenium accumulator.
Habitat: heavy soils, salt desert through
pinyon-juniper areas.
Animals affected: livestock.
Reference: Welsh 1978, Kingsbury 1964.
Astragalus mollissimus Torr. (PNF)
Common name: Thompson woolly loco-
weed.
Toxin: produces locoism.
Habitat: dry plains and hillsides.
Animals affected: livestock.
Reference: Welsh 1978, Kingsbury 1964.
Astragalus pattersonii Gray ex Brand. (PNF)
Common name: Patterson locoweed, Pat-
terson milkvetch.
Toxin: primary selenium accumulator.
Habitat: dry plains and hillsides.
Animals affected: livestock.
Reference: Welsh 1978, Kingsbury 1964.
Astragalus praelongus Sheldon (PNF)
Common name: stinking milkvetch.
Toxin: primary selenium accumulator.
Habitat: dry plains and hillsides, clay and
seleniferous soils.
Animals affected: livestock.
Reference: Welsh 1978.
Astragalus preussii Gray (PNF)
Common name: Preuss milkvetch.
Toxin: primarv selenium accumulator.
Habitat: dry plains and hillsides, seleni-
ferous clays and silts.
Animals affected: livestock.
Reference: Welsh 1978, Kingsbury 1964.
Astragalus pubentissimus Torr. & Gray
(PNF)
Common name: Green River locoweed.
Green River milkvetch.
Toxin: produces locoism.
Habitat: canyons, mountainsides.
Animals affected: livestock, mainly sheep.
Reference: Buck 1961, Kingsbury 1964.
Astragalus racemosus Pursh var. treleasi Por-
ter (PNF)
Common name: alkali milkvetch.
Toxin: primary selenium accumulator,
causes "alkali disease" or "blind staggers."
Habitat: Uinta and Duchesne River forma-
tions.
Animals affected: cattle.
Reference: Welsh 1978.
Astragalus rafaelensis Jones (PNF)
Common name: San Rafael milkvetch.
Toxin: primary selenium accumulator.
Habitat: seleniferous clays and silts, salt
desert shrub zones, Emery Co.
242
Great Basin Naturalist
Vol. 40, No. 3
Animals affected: livestock.
Reference: Welsh 1978.
Astragalus sabuhsus Jones (PNF)
Common name: straightstem poisonvetch,
Cisco milkvetch.
Toxin: primary selenium accumulator.
Habitat: salt desert shrub zone on shales.
Animals affected: livestock.
Reference: Welsh 1978, Marsh 1935.
Astragalus saurinus Barneby (PNF)
Common name: dinosaur milkvetch.
Toxin: primary selenium accumulator.
Habitat: salt desert shrub and pinyon-juni-
per zone, Uintah Co.
Animals affected: livestock.
Reference: Welsh 1978.
Astragalus tetrapterus Gray (PNF)
Common name: fourwing poisonvetch,
four-wing milkvetch.
Toxin: produces locoism.
Habitat: plains, dry hillsides.
Animals affected: cattle, sheep.
Reference: Marsh 1920, Welsh 1978.
Astragalus toanus Jones (PNF)
Common name: Toano milkvetch.
Toxin: primary selenium accumulator.
Habitat: clay soils, salt desert shrub zone.
Animals affected: livestock.
Reference: Welsh 1978, Kingsbury 1964.
Astragalus woodruffii Jones (PNF)
Common name: Woodruff milkvetch.
Toxin: primary selenium accumulator.
Habitat: desert shnib zone on sandy and
sandy-silt soils.
Animals affected: livestock.
Reference: Welsh 1978.
Lathyrus latifolius L. (PIF)
Common name: perennial peavine, per-
ennial sweet pea.
Toxin: alpha, gamma-diaminobutyric acid.
Habitat: rangeland.
Animals affected: rats (experimental),
sheep (lathyrism).
Reference: Lewis 1949, Kingsbury 1964, S.
L. Welsh (personal commvmication, 1980).
Lathyrus sylvestris L. (PNF)
Common name: everlasting sweetpea, flat
pea.
Toxin: alpha, gamma-diaminobutyric acid.
Habitat: rangeland.
Animals affected: sheep, rats (experimen-
tal).
Reference: Lewis 1948, Kingsbury 1964.
Lupinus alpestris A. Nels. (PNF)
Common name: mountain lupine.
Toxin: alkaloids.
Habitat: mountains.
Animals affected: cattle, sheep.
Reference: Beath et al. 1953, Kingsbury
1964.
Lupinus arbustus Dougl. ex Lindl. var. cal-
caratus (Kellogg) Welsh (PNF)
Common name: Douglas spurred lupine,
spur lupine.
Toxin: alkaloids.
Habitat: hillsides, dry soils.
Animals affected: cattle, horses, sheep.
Reference: Clawson 1931, Kingsbury 1964.
Lupinus argenteus Pursh (PNF)
Common name: silvery lupine.
Toxin: alkaloids.
Habitat: dry flats and slopes in woods,
plains and hills.
Animals affected: sheep, cattle, horses,
goats, hogs, deer.
Reference: Marsh 1916, Kingsbury 1964.
Lupinus caudatus Kell. (PNF)
Common name: Kellogg's spurred lupine,
tailcut lupine.
Toxin: alkaloids.
Habitat: exposed hillsides.
Animals affected: cattle.
Reference: Animal Disease and Parasite
Research Division USDA 1958, Kingsbury
1964.
Lupinus leucophyllus Dougl. (PNF)
Common name: woolly-leaved lupine, vel-
vet lupine.
Toxin: alkaloids.
Habitat: dry soil, foothills.
Animals affected: sheep, cattle, horses,
goats, hogs, deer.
Reference: Marsh 1916, Kingsbury 1964.
Lupinus pusillus Pursh (ANF)
Common name: low lupine, rusty lupine.
Toxin: alkaloids.
Habitat: dry plains, foothills.
Animals affected: sheep.
Reference: Sampson 1942, Kingsbury 1964.
September 1980
Brotherson et al.: Poisonous Plants
243
Lupinus sericeus Pursh (PNF)
Common name: silky lupine.
Toxin: alkaloids.
Habitat: dry hillsides and valleys.
Animals affected: sheep, cattle, horses,
goats, hogs, deer.
Reference: Binns and James 1961, Kings-
bury 1964.
Medicago sativa L. (PIF)
Common name: alfalfa, lucerne.
Toxin: saponin.
Habitat: cultivated and escaping.
Animals affected: cattle, chicks, hogs,
sheep.
Reference: Walter 1954, Kingsbury 1964.
Melilotus alba Desr. (BIF)
Common name: white sweetclover.
Toxin: dicoumarin.
Habitat: waste places and fields, escaped
from cultivation.
Animals affected: cattle.
Reference: Roderick 1931, Kingsbury
1964.
Melilotus officinalis L. Lam. (BIF)
Common name: yellow sweetclover.
Toxin: dicoumarin.
Habitat: waste ground, fields; used for for-
age and fertilizer.
Animals affected: cattle, sheep, horses.
Reference: Roderick 1931, Kingsbury
1964.
Oxytropis lambertii Pursh (PNF)
Common name: white loco, white paint
loco, silky crazyweed.
Toxin: unknown alkaloids; produces
locoism.
Habitat: prairies and mountains, usually in
drier areas, lower to middle elevations.
Animals affected: livestock.
Reference: Couch 1929, Kingsbury 1964.
Oxytropis sericea Nutt. ex Torr. & Gray
(PNF)
Common name: white paint loco, silky cra-
zyweed.
Toxin: alkaloids; produces locoism.
Habitat: open gravelly or well-drained
slopes and hills at lower to middle elevations.
Animals affected: livestock.
Reference: Porter 1951, Kingsbury 1964.
Visum sativum L. (AIF)
Common name: garden pea.
Toxin: unknown.
Habitat: cultivated.
Animals affected: sheep, cattle.
Reference: Whiting et al. 1957, Kingsbury
1964.
Poinciana gilliesii Hook. (PIS)
Common name: bird of paradise.
Toxin: unknown; green seed pods are gas-
trointestinal irritants.
Habitat: cultivated ornamental, small pop-
ulation established in Washington Co.
Animals affected: humans, livestock.
Reference: Cann and Verhulst 1958, Kings-
bury 1964.
Prosopis glandulosa Torr. (PNT)
Common name: mesquite.
Toxin: unknown; may cause a nutritional
deficiency.
Habitat: dry ranges, washes and draws at
low elevations, especially along streams
where the water table is high.
Animals affected: cattle.
Reference: Adler 1949, Kingsbury 1964.
Robinia pseudoacacia L. (PIT)
Common name: black locust.
Toxin: unknown.
Habitat: escaped from cultivation; around
dwellings or along fencerows.
Animals affected: horses, cattle, sheep,
poultry, humans.
Reference: Power 1901, Kingsbury 1964.
Thermopsis montana Nutt. (PNF)
Common name: goldenpea, mountain ther-
mopsis, yellow pea.
Toxin: alkaloids.
Habitat: common in pastures.
Affected animals: cattle.
Reference: Schmutz et al. 1968.
Trifolium hybridum L. (PIF)
Common name: .\lsike clover.
Toxin: photosensitizing compound.
Habitat: cultivated; escaped to roadsides
and meadows.
Animals affected: horses, hogs, sheep,
cattle.
Reference: Fincher and Fuller 1942, Kings-
bury 1964.
244
Great Basin Naturalist
Vol. 40, No. 3
Trifolium praetense L. (PIF)
Common name: red clover.
Toxin: unknown.
Habitat: cultivated and escaping along
roadsides and ditches.
Animals affected: cattle, horses, sheep.
Reference: O'Dell 1959, Kingsbury 1964.
Trifolium repens L. (PIF)
Common name: white clover.
Toxin: cyanogenetic.
Habitat: cultivated and escaping.
Animals affected: newborn pigs.
Reference: Garner 1957, Kingsbury 1964.
Vicia villosa Roth (PIF)
Common name: hairy vetch, winter vetch.
Toxin: photosensitizing compound.
Habitat: cultivated; occasionally escaping.
Animals affected: cattle, horses.
Reference: Claughton and Claughton 1954,
Kingsbury 1964.
Haemodoraceae
Iris missouriensis Nutt. (PNF)
Common name: wild iris, blue flag, fleur-
de-lis, western blue flag.
Toxin: unknown.
Habitat: moist soils along stream banks, in
marshes or moist mountain meadows.
Animals affected: calves, laboratory ani-
mals.
Reference: Beath et al. 1953, Kingsbury
1964.
Hypericaceae
Hypericum formosum H.B.K. (PNF)
Common name: southwestern St. Johns-
wort.
Toxin: photosensitizing compounds.
Habitat: moist soils of plains and hills.
Animals affected: cattle, sheep, horses,
goats.
Reference: Harris 1951.
Fagaceae
Quercus gambelii Nutt. (PNT)
Common name: Gambel oak.
Toxin: tannins.
Habitat: throughout the state, often form-
ing dense thickets.
Animals affected: cattle, sheep, and goats.
Reference: Boughton 194.3, Kingsbury
1964.
FUMARIACEAE
Corydalis aurea Willd. (ANF)
Common name: Golden corydalis.
Toxin: alkaloids.
Habitat: woods and well-shaded mountain
slopes.
Animals affected: sheep, cattle.
Reference: Sperry 1955, Kingsbury 1964.
Gentianaceae
Centaurium calycosum (Buckl.) Fern. (ANF)
Common name: Buckley centaury, moun-
tain pink, arizona centaury.
Toxin: unknown.
Habitat: moist soil, river valleys.
Animals affected: sheep, goats.
Reference: Dollahite and Allen 1962,
Kingsbury 1964.
JUNCAGINACEAE
Triglochin concinna Davy (PNF)
Common name: arrowgrass, goosegrass,
sourgrass, podgrass, Utah arrowgrass.
Toxin: hydrocyanic acid.
Habitat: salty marshes and ponds.
Animals affected: sheep, cattle.
Reference: Schmutz et al. 1968.
Triglochin debilis L. (PNF)
Common name: arrowgrass, weak arrow-
grass.
Toxin: hydrocyanic acid.
Habitat: damp soils, marshes and sloughs;
usually where the soil is alkaline or the water
calcareous or brackish.
Animals affected: sheep, cattle.
Reference: Schmutz et al. 1968, Kingsbury
1964.
Triglochin maritima L. (PNF)
Common name: arrowgrass, seashore ar-
rowgrass, shore arrowgrass.
Toxin: hydrocyanic acid.
Habitat: damp soils, marshes and sloughs;
usually where the soil is alkaline or the water
calcareous or brackish.
Animals affected: sheep, cattle.
Reference: Beath et al. 1933, Kingsbury
1964.
September 1980
Brotherson et al.: Poisonous Plants
245
Lamiaceae
Lamium amplexicaule L. (AIF)
Common name: henbit, dead nettle.
Toxin: unknown.
Habitat: occasional weed of fields and
waste places.
Animals affected: sheep, horses, cattle.
Reference: Hurst 1942, Kingsbury 1964.
LiLIACEAE
Allium schoenoprasm L. (PNF)
Common name: chives.
Toxin: unknown.
Habitat: cultivated; river bars, lake shores,
wet meadows.
Animals affected: horses.
Reference: Kobayashi 1950, Kingsbury
1964.
Asparagus officinalis L. (PIF)
Common name: asparagus.
Toxin: unknown.
Habitat: cultivated and widely escaped.
Animals affected: cattle and dairy cows.
Reference: Los Angeles County Livestock
Department 1938, Kingsbury 1964.
Ornithogalum umbellatum L, (PIF)
Common name: star-of-Bethlehem, snow-
drop.
Toxin: cholchicine alkaloid.
Habitat: weed of grasslands and thickets.
Animals affected: sheep, cattle, children.
Reference: Reynard and Norton 1942,
Kingsbury 1964.
Veratrum californicum Durand (PNF)
Common name: false hellebore, corn-lily,
skunk cabbage.
Toxin: alkaloids.
Habitat: bogs and wet meadows from 7500
to 9500 feet.
Animals affected: cattle, sheep, fowl, hu-
mans.
Reference: Schmutz et al. 1968, Kingsbury
1964.
Yucca L. (PNS)
Note: members of this genus have been re-
ported to contain saponins, salicylic acid, the
alkaloid imperialin, and several resins. None
of the species in which these toxins have been
identified are found in Utah.
Reference: Pammel 1911.
Zigadenus elegans Pursh (PNF)
Common name: death camas, mountain
death camas.
Toxin: alkaloids.
Habitat: prairies, meadows.
Animals affected: cattle, horses, hoes, fowl
humans.
Reference: Marsh et al. 1915, Kingsbury
1964.
Zigadenus paniculatus (Nutt.) Wats. (PNF)
Common name: death camas, foothill
death camas, sandcom.
Toxin: alkaloids.
Habitat: dry soils; hills and plains.
Animals affected: cattle, horses, hogs, hu-
mans.
Reference: Fleming et al. 1921, Kingsbury
1964.
Zigadenus venenosus Wats. (PNF)
Common name: death camas, meadow
death camas.
Toxin: alkaloids.
Habitat: moist, grassy meadows.
Animals affected: sheep, cattle, horses,
hogs, humans.
Reference: Cameron 1952, Kingsbury
1964.
PiNACEAE
Pinus ponderosa Doug, ex Laws (PNT)
Common name: western yellow pine, pon-
derosa pine.
Toxin: unknown.
Habitat: coniferous forest at moderate ele-
vations; dry hillsides, plateaus, slopes, valleys
and mesas.
Animals affected: cattle.
Reference: MacDonald 1952, Kingsbury
1964.
POACEAE
Avena fatua L. (AIG)
Common name: wild oats.
Toxin: mechanically injurious.
Habitat: cultivated land and waste places.
Animals affected: livestock.
Reference: Pammel 1911.
Avena sativa L. (AIG)
Common name: cultivated oats.
Toxin: nitrates, photosensitizing com-
pounds, grass tetany.
246
Great Basin Naturalist
Vol. 40, No. 3
Habitat: open ground, grasslands, waste
places; lawns and golf courses.
Animals affected: cattle, horses, hogs, tur-
keys, goats, sheep and wild ruminants.
Reference: Newsom et al. 1937, Kingsbury
1964.
Bromus rigidus Roth (AIG)
Common name: ripgut brome.
Toxin: mechanical injury from mature
awns.
Habitat: common weed.
Animals affected: cattle and sheep.
Reference: Range Plant Handbook 1937.
Bromus rubens L. (AIG)
Common name: foxtail chess, red brome.
Toxin: mechanical injury from mature
awns.
Habitat: common weed, dry and saline
soils.
Animals affected: cattle and sheep.
Reference: Davis 1952.
Bromus tectorum L. (AIG)
Common name: cheatgrass, downy cheat.
Toxin: mechanical injury from mature
awns; may also be implicated in ergot poi-
soning.
Habitat: common weed, especially in dry
places; plains and foothills.
Animals affected: cattle.
Reference: Pammel 1911.
Cynodon dactylon L. Pers. (PIG)
Common name: Bermuda grass.
Toxin: photosensitizing compound.
Habitat: open ground, grasslands, waste
places; lawns and golf courses.
Animals affected: cattle.
Reference: Gibbons 1953, Kingsbury 1964.
Eragrostis cilianensis (All.) Link (AIG)
Common name: lovegrass, stinkgrass, stick
grass.
Toxin: unknown.
Habitat: cultivated ground, gardens and
waste places; weed in fields and along road-
sides.
Animals affected: horses.
Reference: Gates 1930, Kingsbury 1964.
Festuca arundinacea Schreb. (PIG)
Common name: fescue, tall fescue, alta fes-
cue, goar fescue.
Toxin: alkaloids.
Habitat: unimproved pastures; wet, heavy
soils of high organic content.
Animals affected: cattle.
Reference: Maag and Tobiska 1956, Kings-
bury 1964.
Glyceria striata Lam. Hitch. (PNG)
Common name: fowl mannagrass.
Toxin: cyanogenetic.
Habitat: wet areas.
Animals affected: cattle.
Reference: Reynard and Norton 1942,
Kingsbury 1964.
Hilaria rigida (Thurb.) Benth ex Scribn.
(PNG)
Common name: galleta grass, big galleta,
dixie grass.
Toxin: unknown.
Habitat: dry lands and desert ranges to
4000 feet.
Animals affected: cattle.
Reference: Quortrup and McFarland 1956,
Kingsbury 1964.
Holcus lanatus L. (PIG)
Common name: velvet grass, mesquite
grass, Yorkshire velvet grass.
Toxin: unknown.
Habitat: open ground, meadows and moist
places; occasionally cultivated.
Animals affected: livestock.
Reference: Couch 1932, Kingsbury 1964.
Hordeum jubatum L. (PNG)
Common name: squirreltail grass, foxtail
grass, wild barley.
Toxin: mechanically injurious.
Habitat: weed in open ground, meadows,
prairies, along streams, ditches and waste
places.
Animals affected: sheep, cattle, horses.
Reference: Fleming and Peterson 1919,
Kingsbury 1964.
Hordeum vulgare L. (AIG)
Common name: cultivated barley.
Toxin: mechanically injurious.
Habitat: cultivated for grain and along
road shoulders; sometimes spontaneous in
waste places but not persistent.
Animals affected: hogs, dogs, humans,
poultry.
Reference: Christensen and Kernkamp
1936, Kingsbury 1964.
September 1980
Brotherson et al.: Poisonous Plants
247
Setaria lutescens (Wiegel) Hubb. (AIG)
Common name: yellow bristle grass, foxtail
grass, pigeon grass.
Toxin: mechanically injurious.
Habitat: common weed of cultivated and
waste areas.
Animals affected: livestock.
Reference: Bankowski et al. 1956, Kings-
bury 1964.
Sorghum halpense L. Pers. (PIG)
Common name: Johnson grass.
Toxin: hydrocyanic acid, nitrates.
Habitat: weed of cultivated fields, waste
places and along irrigation ditches and
stream bottoms.
Animals affected: cattle, sheep, horses.
Reference: Slade 1903, Kingsbury 1964.
Sorghum vulgare Pers. (AIG)
Common name: grain sorghum.
Toxin: cyanogenetic glycoside.
Habitat: cultivated.
Animals affected: cattle, sheep, horses.
Reference: Slade 1903, Kingsbury 1964.
Stipa commata Trin. & Rupr. (PNG)
Common name: needle-and-thread grass.
Toxin: mechanically injurious.
Habitat: dry plains and hillsides, sandy soil.
Animals affected: livestock.
Reference: Pammel 1911.
Stipa neomexicana (Thurb.) Scribn. (PNG)
Common name: New Mexican feather-
grass.
Toxin: mechanically injurious.
Habitat: common in dry rocky canyons
and mesas.
Animals affected: livestock.
Reference: Pammel 1911.
Zea mays L. (AIG)
Common name: com, maize.
Toxin: nitrates, under conditions of over-
fertilization with too little water.
Habitat: cultivated for grain, forage or si-
lage.
Animals affected: livestock, humans.
Reference: Brady et al. 1955, Kingsbury
1964.
POLYGONACEAE
Beta vulgaris L. (AIF)
Common name: beet, sugar beet, fodder
beet, mangel-worzel, mangold.
Toxin: oxalates; nitrates, under conditions
of overfertilization with too little water.
Habitat: cultivated.
Animals affected: livestock.
Reference: Baker and Eden 1954, Kings-
bury 1964.
Rheum rhaponticum L. (PIF)
Common name: rhubarb.
Toxin: oxalic acid, oxalates.
Habitat: cultivated and persisting.
Animals affected: livestock, humans.
Reference: Hansen 1930, Kingsbury 1964.
Rumex acetosella L. (PIF)
Common name: sheep sorrel, dock.
Toxin: oxalates.
Habitat: common weed of acid or sterile,
gravelly soils of pastures and meadows; waste
places.
Animals affected: sheep.
Reference: Connor and Adams 1951,
Kingsbury 1964.
Rumex crispus L. (PIF)
Common name: curly dock.
Toxin: oxalates.
Habitat: moist fields and waste places.
Animals affected: sheep.
Reference: Connor and Adams 1951,
Kingsbury 1964.
POLYPODIACEAE
Dryopteris felix-mas (L.) Schrott (PNF)
Common name: male fern.
Toxin: thiaminase.
Habitat: mountains; damp soils, deep
shaded ravines, in cliffs or tallus.
Animals affected: horses.
Reference: Harvey et al. 1944, Kingsbury
1964.
Pteridium aquilinum (L.) Kuhn (PNF)
Common name: Bracken fern.
Toxin: thiaminase and its coenzymes.
Habitat: upland pastures, a.spen zone.
Animals affected: livestock.
Reference: Carpenter 1950, Kingsbury
1964.
PORTULACACEAE
Portulaca oleracea L. (AIF)
Common name: purslane, pusley.
Toxin: oxalates.
248
Great Basin Naturalist
Vol. 40, No. 3
Habitat: common weed of garden and cul-
tivated areas.
Animals affected: sheep.
Reference: Mathams and Sutherland 1952,
Kingsbury 1964.
Ranunculaceae
Aconitum columbianum Nutt. (PNF)
Common name: western monkshood.
Toxin: alkaloids.
Habitat: mountains from 5000 to 10,000
feet; along streams and wet meadows; moist
places and thickets.
Animals affected: livestock, humans.
Reference: Stern 1960, Kingsbury 1964.
Actaea arguta Nutt. (PNF)
Common name: baneberry, western bane-
berry.
Toxin: irritant oil.
Habitat: mountains in rich, moist soil.
Animals affected: livestock, humans.
Reference: Schmutz et al. 1968.
Caltha leptosepala DC. (PNF)
Common name: elkslip marshmarigold.
Toxin: glycoside (protoanemonin).
Habitat: wet mountain soils.
Animals affected: livestock.
Reference: Schmutz et al. 1968.
Delphinium andersonii Gray (PNF)
Common name: Anderson larkspur.
Toxin: alkaloids.
Habitat: subsaline soils of plains and hills.
Animals affected: cattle, sheep.
Reference: Miller 1923, Kingsbury 1964.
Delphinium barbeyi Huth. (PNF)
Common name: barbey, larkspur, tall
larkspur.
Toxin: alkaloids.
Habitat: mountains; meadows and open
woods, summer ranges; common under aspen
and along streams.
Animals affected: cattle, occasionally
sheep.
Reference: Cook 1951, Kingsbury 1964.
Delphinium nuttallianum Fritz. (PNF)
Common name: Nuttall larkspur, low
larkspur. Nelson larkspur.
Toxin: alkaloids.
Habitat: moist soils, hills, foothills, and
sagebrush deserts.
Animals affected: cattle, sheep.
Reference: Ewan 1945, Kingsbury 1964.
Delphinium occidentale Wats. (PNF)
Common name: duncecap larkspur, tall
larkspur.
Toxin: alkaloids.
Habitat: mountain summer ranges, com-
mon under aspen and along streams; moun-
tain meadows.
Animals affected: cattle, occasionally
sheep.
Reference: Couch 1936, Kingsbury 1964.
Ranunculus acris L. (PIF)
Common name: tall field buttercup, tall
buttercup.
Toxin: protoanemonin.
Habitat: common pasture weed.
Animals affected: livestock.
Reference: Tehon et al. 1946, Kingsbury
1964.
Ranunculus cymbalaria Pursh (PNF)
Common name: alkali buttercup; trailing
buttercup.
Toxin: glycosides (protoanemonin).
Habitat: muddy banks along brackish
streams and marshes.
Animals affected: livestock.
Reference: Fleming 1920, Kingsbury 1964.
Ranunculus flammula var. filiformis
(Michx.) Hook. (PNF)
Common name: creeping spearwort but-
tercup.
Toxin: glycosides (protoanemonin).
Habitat: marshy ground of lakes, streams
and ditches.
Animals affected: livestock.
Reference: Hill and van Heyningen 1951,
Kingsbury 1964.
Ranunculus repens L. (PIF)
Common name: creeping buttercup.
Toxin: protoanemonin.
Habitat: meadows and marshes at lower
elevations; wet pastures.
Animals affected: livestock.
Reference: Gilkey 1958, Kingsbury 1964.
Ranunculus scleratus L. (PNF)
Common name: celeryleaf crowfoot.
Toxin: glycosides (protoanemonin).
Habitat: borders of lakes, ponds and
streams.
September 1980
Brotherson et al.: Poisonous Plants
249
Animals affected: livestock.
Reference: Fleming 1920, Kingsbury 1964.
Ranunculus testiculatus Crantz (AIF)
Common name: burbuttercup; testiculate
buttercup.
Toxin: glycosides (protoanemonin).
Habitat: general in the intermountain re-
gion.
Animals affected: livestock.
Reference: Schmutz et al. 1968.
ROSACEAE
Cercocarpus montanus Raf. (PNS)
Common name: mountain mahogany,
birchleaf mountain mahogany, true mountain
mahogany.
Toxin: cyanogenetic glycoside.
Habitat: stony hills and slopes.
Animals affected: livestock.
Reference: Burke 1960, Kingsbury 1964.
Prunus armeniaca L. (PIT)
Common name: apricot.
Toxin: cyanide.
Habitat: cultivated and persisting.
Animals affected: livestock, humans.
Reference: Hurst 1942, Kingsbury 1964.
Prunus persica Batsch. (PNT)
Common name: peach.
Toxin: cyanide.
Habitat: cultivated.
Animals affected: livestock.
Reference: Reynard and Norton 1942,
Kingsbury 1964.
Prunus virginiana L. (PNT)
Common name: choke cherry.
Toxin: cyanogenetic glycoside.
Habitat: common in hills, mountains, along
streams, thickets, fencerows and edges of
woods.
Animals affected: sheep, cattle.
Reference: Pijoan 1942, Kingsbury 1964.
Santalaceae
Comandra umbellata (L.) Nutt. (PNF)
Common name: bastard toadflax.
Toxin: alkaloids, glycosides, secondary se-
lenium accumulator.
Habitat: common weed, found in various
habitats.
Animals affected: livestock.
Reference: Trelease and Beath 1949,
Kingsbury 1964.
SOLANACEAE
Datura meteloides Dunal (ANF)
Common name: datura, stramonium,
thornapple, Jimson weed, Jamestown weed,
apple of Peru, tolgaudia, sacred datura, In-
dian apple.
Toxin: alkaloids (atropine, hyoscvamine,
hyoscine).
Habitat: plains, dry hills and valleys; culti-
vated and escaping.
Animals affected: humans, horses, cattle,
sheep, hogs, mules, chickens.
Reference: Hansen 1924, Kingsbury 1964.
Datura stramonium L. (ANF)
Common name: Jimsonweed, sacred da-
tura.
Toxin: alkaloids (atropine, hyoscyamine,
hyoscine).
Habitat: waste areas, rich soils of barn-
yards, heavily used portions of pastures.
Animals affected: horses, cattle, sheep,
hogs, mules, chickens, humans.
Reference: Hansen 1924, Kingsbury 1964.
Hyoscyamus niger L. (BIF)
Common name: black henbane, henbane.
Toxin: alkaloids (hyoscyamine, hyoscine,
atropine).
Habitat: widespread dry soils of roadsides
and waste areas.
Animals affected: humans, fowl, livestock.
Reference: Long 1917, Kingsbury 1964.
Lycium halmifolium Mill. (PIS)
Common name: matrimony vine, tea vine.
Toxin: unknown.
Habitat: cultivated and escaping around
homesites and cemetaries.
Animals affected: calves, sheep.
Reference: Hansen 1927, Kingsbury 1964.
Nicotiana attenuata Torr. ex S. Wats. (.\NF)
Common name: wild tobacco, coyote to-
bacco.
Toxin: nicotine.
Habitat: dry, sandy stream beds and flats.
Animals affected: horses, pigs, livestock,
humans.
Reference: Marsh et al. 1927, Kingsbury
1964.
250
Great Basin Naturalist
Vol. 40, No. 3
Nicotiana trigonophylla Dunal ex DC.
(ANF)
Common name: wild tobacco, desert to-
bacco.
Toxin: nicotine.
Habitat: dry desert soils.
Animals affected: horses, pigs, livestock,
humans.
Reference: Marsh et al. 1927, Kingsbury
1964.
Solarium dulcamara L. (PIF)
Common name: European bittersweet,
climbing nightshade, bitter nightshade.
Toxin: glycoalkaloids.
Habitat: woods, thickets and waste places;
cultivated and escaping.
Animals affected: cattle, horses, sheep, hu-
mans.
Reference: Craig and Kehoe 1925, Kings-
bury 1964.
Solanum eleagnifolium Cav. (PNF)
Common name: silverleaf nightshade,
white horsenettle, trapillo.
Toxin: glycoalkaloids.
Habitat: serious weed of prairies, open
woods and disturbed soils; dry ground; barn-
yards.
Animals affected: cattle, sheep.
Reference: Buck et al. 1960, Kingsbury
1964.
Solanum nigrum L. (AIF)
Common name: black nightshade.
Toxin: glycoalkaloids.
Habitat: common weed of fields and waste
places.
Animals affected: livestock, humans.
Reference: Carey 1955, Kingsbury 1964.
Solanum rostratum Dunal (ANF)
Common name: buffalo bur, Kansas thistle,
Texas thistle, buffalobur nightshade.
Toxin: glycoalkaloids.
Habitat: plains, roadsides, barnyards.
Animals affected: hogs.
Reference: Simic 1943, Kingsbury 1964.
Solanum triflorum Nutt. (ANF)
Common name: three flowered nightshade,
cutleaf nightshade.
Toxin: glycosides.
Habitat: prairies, fields and waste places;
weed of cultivation and disturbed soils.
Animals affected: horses, cattle.
Reference: Pammel 1921, Kingsbury 1964.
Typhaceae
Typha latifolia L. (PNF)
Common name: cattail.
Toxin: unknown.
Habitat: common in moist soils, marshes
and ponds.
Animals affected: horses.
Reference: Hansen 1930, Kingsbury 1964.
Zygophyllaceae
Tribulus terrestris L. (AIF)
Common name: puncture vine, caltrap.
Toxin: nitrates, photosensitizing com-
pound.
Habitat: dry soils of waste lands, roadsides
and deserts.
Animals affected: sheep.
Reference: Durrell et al. 1952, Kingsbury
1964.
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THE SUCCESSIONAL STATUS OF CUPRESSUS ARIZOMCA
Albert J. Parker
.\bstbact.— Several investigations isize-class analysis, age-determination inquiries, and germination tests"* suggest
that Cupressus arizonica of southeastern .\rizona is a pioneer species. The tree requires disturbance to remove or
reduce soil litter, which other\%-ise inhibits the reproduction of the species. Reduction of light intensity caused by
canopy closure appears to be less important than litter accumulation in restricting C. arizonica reproduction. Fol-
lowing disturbance, successful establishment of seedlings may occur over an e.xtended period ,50 to 100 years I as
Utter graduallv accumulates. The absence of C. arizonica seedlings in present populations suggest that fire suppres-
sion policies on federal lands where C. arizonica occurs have altered fire frequency, and consequently have fostered a
short-term reduction in C. arizonica establishment. Only in floodplain en\ironments. where flooding disturbs the soil
surface, has much reproduction occurred in recent years. The long-term population pattern of C. arizonica appears
stable, due to the great longevity of the species.
Rough-barked Arizona c^'press {Cupressus
arizonicu Greene; all taxonomy after Kearney
and Peebles I960' is a tree species of local
occurrence in the mountain ranges of south-
eastern .\rizona, southwestern New Me.xico,
western Texas, and northern Mexico. Though
it has topically been characterized as a mois-
tiu-e demanding species of riparian associ-
ation iWolf 1948a. \Miittaker and Xiering
1965. BrouTi and Lowe 1974', recent findings
(Parker 1980 > demonstrate that C. arizonica
may occupy relatively xeric landscape posi-
tions—including south-facing slopes, ridge
tops, and convex slope configurations— at
higher elevations laroimd 2000 ml
In the first pubUshed report mentioning
Cupressus arizonica regeneration. Posev and
Goggans - 1967' obser\ed little reproduction
anxAvhere in the Southwest, although they
collected no data to substantiate or in-
vestigate this phenomena. Thev did express
concern that the limited natural range of the
species, coupled with its apparent inability to
reproduce in many natural circtmistances,
could indicate that its existence was threat-
ened. This paper investigates the population
d\Tiamics, reproductive strategy, and role in
species replacement processes of C. arizonica
in the upland forests of southeastern Arizona.
Study Are.a.s
Populations of Cupressus arizonica were
sampled in the Chiricahua Moimtains, the
Department of Geography. Univeran- of Georgia, .Mhens, Georgia 30602.
Blue Range north of Clifton, the Dragoon
Mountains, and the Santa Catalina Moun-
tains, all located in .\rizona (Fig. l\ Cupr-
essus arizonica is most common in the wood-
land zone of these mountain ranges
(1300-1900 m; Wliittaker and Xiering 1965),
a zone characterized by the frequent co-do-
minance of alligator juniper Juniperus dep-
peana\ pinyons [Pinus cembroides and P.
edulisl and a nmnber of oak species, includ-
ing silverleaf oak (Quercus hypoleucoides),
netleaf oak [Q. rugosa), and .\rizona white
oak Q. arizonica"!. Cupressus arizonica is pro-
gressively more restricted to riparian habitats
where the woodland zone grades into the
shrub-dominated desert and semidesert vege-
tation types below. It is also found in stands
at higher elevations (1600-2300 mi with
other conifers, notably Arizona pine (Pinus
ponderosa var. arizonica^ Apache pine {P.
engehiannii), Chihuahua pine P. leiophylla),
and Douglas-fir iPseudotsuga menziesii);
these forested stands often have closed ca-
nopies.
The climate of the woodland zone is sub-
humid and mild, with about 50 cm annual
precipitation and a mean annual temperature
of 12.5 C. The wettest periods of the year are
middle to late summer and midwinter (Bry-
son and Lown.- 1955 1.
The isolated mountain ranges of this region
of basin and range topography are roughly
linearlv aligned, northward trending, and
254
September 1980
P.\rker: Arizona Cypress Succession
iDO
ARIZONA
/^C AT A LIN A ^
MTS.
M3
i"''*"/"„
aV"
•■;/'-$,
f.ii-i/'//^
•'/;
%
''^*i;
i
V"'..
-:. km.
'-/y,...„.? ARIZONA
MEXFCO"
CARTOGRAPH.C LASGRATOfiY. uMVERSiTV Gf W.SGG\S;f4 - V.ADISOM
Fig. 1. Study area uith the species range inset. The location of stands included in this study are noted by number.
Open circles in the inset represent the three extensive areas of nonriparian C. ari:onica occuTTence in southeastern
.\rizona: the Chiricahua Mountains, the Blue Range, and the Santa Catalina Mountains.
256
Great Basin Naturalist
Vol. 40, No. 3
separated by extensive sediment-filled low-
lands. The bedrock core of these ranges in-
clude both igneous and sedimentary rocks
(Fenneman 1931).
Methods
Environmental and vegetational measure-
ments were obtained from 19 internally
homogeneous stands of C. arizonica. Within
each stand, three 4 by 25 m quadrats were
oriented so that their central long axes were
normal to a contour line bisecting the stand,
with intervals of 10 m separating each quad-
rat. The following measurements were re-
corded within each quadrat: the species and
circumference at breast height (1.37 m) of
each tree, the number of saplings of all tree
species, and the number of C. arizonica seed-
lings. Any stem of at least 20 cm circum-
ference at breast height was considered a
tree. A sapling was defined as the stem of any
tree species which exceeded breast height,
but which was less than 20 cm in circum-
ference at that height. All stems of tree spe-
cies less than breast height were counted as
seedlings. The depth of the litter layer in
each stand was measured at 1 m intervals
along the central long axis of each quadrat. A
total of 60 litter depth measurements was
made per stand.
In two stands, both in the Chiricahua
Mountains, all C. arizonica trees with a cir-
cumference at breast height exceeding 30 cm
were cored with an increment borer at breast
height. These two stands were chosen for
their differing size structures. One stand was
characterized by trees few in number but
large in diameter, but the other stand had a
great number of individuals, particularly in
the smaller size-classes. These two stands
were selected as representative of 14 of the
19 stands sampled. The other five stands has
a size structure intermediate in character be-
tween those chosen for coring. In total, 79
trees were cored.
Cores were aged in the laboratory, al-
though the aging was complicated by two
factors. First, several of the C. arizonica trees
possessed rotted xylem tissue that caused
fragmentation of some cores and loss of re-
cord from the damaged segments. Second, C.
arizonica has been shown to produce false an-
nual rings (Bannan 1954). To circumvent the
first difficulty, the length of each core was
measured and the following formula was used
to compute a "tree ring" age for each
sample:
0.95 (^)(g) = t
where: t = extrapolated tree ring age (yr)
c = circumference of the tree
(cm)
g = number of annual rings per
unit length of core (yr*cm-i; this value is
later referred to as "mean time required
per unit of radial growth")
0.95 = a coefficient to adjust for bark
thickness
With respect to the second complication.
Clock and Agerter (1963) were able to utilize
microclimatic records from a plantation of C.
arizonica near Lubbock, Texas (500 km from
the species' natural range) to investigate the
production of false annual rings in this spe-
cies. They reported that multiple false rings
may be produced by an individual in a single
year, and that the number of false rings is not
consistent from tree to tree, or even from
place to place on the same tree. Because of
these difficulties, precise absolute aging was
not possible, and tree ring ages reported in
this study may be as much as two times
larger than the actual tree age. Nevertheless,
the ring counts and extrapolated ages provide
an indication of the relative ages of individ-
uals and thus allow identification and inter-
pretation of the approximate age structures
of the stands.
Cones of C. arizonica were collected for
the purpose of determining the mean number
of seeds per cone, as well as to provide seeds
for use in germination tests. All 250 cones
collected were from the second year foliage
of a number of C. arizonica individuals lo-
cated in the Blue Range. Only closed cones
were collected. Fifty of tht«e cones were ran-
domly selected, and seeds from each cone
were counted. Following this, the other cones
were emptied of seeds, and all seeds were
then sorted by size, the larger seeds being
kept for use in germination tests.
A series of germination tests were per-
formed on lots of 100 seeds selected ran-
domly from the seed source. These tests in-
September 1980
Parker: Arizona Cypress Succession
257
vestigated the relationship of C. arizonica
germination success to the following factors:
freezing, flooding, fire, litter buildup, and
light intensity. The physical conditions in
which the germination tests were performed
followed those of Wolf (1948b) in general
(see Table 4).
Results
Stands were segregated into three groups,
according to the size-class frequency distribu-
tion of individuals of Cupressus arizonica.
The eight stands of the first group display a
generally logarithmatic decrease in the num-
ber of individuals in successively larger size-
classes, with the exception that the seedling
layer is virtually absent (Fig. 2). Populations
with a size stnicture similar to that of the
first group are often characterized as stable
through time, with losses of canopy individ-
uals being balanced by replacements from
the sapling layer (Daubenmire 1968). The six
stands of the third group, in contrast, lack a
logarithmatic trend. Instead, the number of
individuals in successively larger size-classes
is more constant (Fig. 2), suggesting that a
past period of reproduction has ended. The
400-1
350-
Group 1
1 — \ — \ — ] — I — r-1
i—\ — I — \ — I — I T* I '
(DraiDirjinLnioinminTTTTTT ■
ID £ r^
CO
CNJ CD
in to h-
(DO) o>ai^CT)Oio>ojoi a>
Group 2
"I 1 1 ""1 — I I I I I
ID <b 1^ CO a>
ininiDuSininioiT)
*- CNJ to ^ *0 to ^
60-
40-
20-
Group 3
Fig. 2. Composite size-class structure histograms for stands of C. arizonica. See text for a definition of each of the
three groups.
258
Great Basin Naturalist
Vol. 40, No. 3
second group includes five stands inter-
mediate in character between the other
groups.
Even in those cases which appear to pos-
sess a steady-state population above the sap-
ling layer, the almost universal absence of C.
arizonica seedlings in stand understories
(Table 1) is conspicuous, and agrees with the
observation of Posey and Goggans (1967).
Conditions in the two stands which do con-
tain a relatively large number of seedlings
are significant; in both cases, disturbance has
exposed bare mineral soil, in one case by nat-
ural flooding and in the other by logging ac-
tivities that mechanically stripped off the lit-
ter from a portion of the forest floor.
The behavior of other tree species in C.
anzontca -dominated commimities fall pri-
marily into two types, according to stand
size-class information (Table 2). A first group
consists of intolerant pioneer species, which
reproduce episodically in association with
disturbance events, and probably depend on
wide dispersal of seeds to maintain represen-
tation on a given site. These species display
an even aged structure in most C. arizonica
stands. The common upland conifers of the
region, such as Apache pine, Arizona pine,
and Douglas-fir fit this behavior pattern. A
second group consists of tolerant species with
stable populations that reproduce either veg-
etatively of by widespread dispersal from ad-
jacent locales. These display continues repro-
Table 1. The seedling class.
No. seedlings Mean
Recently flooded stand
Recently logged stand
All other stands (17)
Total
20
36
17
73
1.0
3.7
duction in C. arizonica-dominated stands,
with a maximum number of stems occurring
in recent cohorts. This group includes the
typical pinyon/oak woodland dominants-
Mexican pinyon, silverleaf oak, netleaf oak,
and Arizona white oak.
Core analysis indicated that the mean tree
ring age of C. arizonica in the stand with few
but large individuals was 317.1 years, with
ages ranging from 197.6 to 456.7 years (Table
3). The majority of tree ring ages were be-
tween 250 and 375 years. No seedlings, sap-
lings, or trees of less than 30 cm circum-
ference at breast height were found in the
stand. Thus, the tree ring age of the youngest
C. arizonica individual was almost 200 years.
The mean tree ring age of C. arizonica trees
in the stand with the greater density of small-
er trees was 116.9 years, with an age range
from 62.4 to 178.1 years. However, there
were also a large number of saplings and
trees too small to core, and these size-classes
probably extended the age range down to
about 25 years. The conspicuous absence of
C. arizonica seedlings in the imderstory of
this stand (Table 3) suggested that reproduc-
tion has been inhibited during about the last
quarter century.
To determine the degree to which under-
story individuals of C. arizonica were sup-
pressed by the overstory, a linear correlation
and regression analysis was performed on the
mean time required for unit growth (yr*cm-i)
for each tree against tree radius. A negative
correlation coefficient would be predicted in
conditions of understory suppression, because
small trees would require longer periods of
time to produce an amount of radial growth
equal to that produced on the larger, more
rapidly growing trees in the canopy in a
shorter period of time. The results suggest
that suppression of understory individuals
Table 2. Composite size-class structnres for upland conifers and pinyon-oak species. The figures heading each
size-class category are the smallest possible circumference in that class, and size-classes include all stems up to the
next larger value. Upland conifers include Arizona pine, ponderosa pine, Apache pine, and Douglas fir. Pinyons in-
clude Mexican pinyon and two-leafed pinyon. Oaks include silverleaf oak, netleaf oak, and Arizona white oak.
sapl
20
30
40
Size-class
50
(cm
60
circumference)
70 80
90
100
110
120
Upland conifers
3
4
4
7
4
3
8
4
0
4
5
9
Pinyons
Oaks
63
111
27
27
18
18
13
24
4
18
7
8
2
4
2
3
1
0
1
1
0
0
0
1
September 1980
Parker: Arizona Cypress Succession
259
Table 3. Characteristics of cored stands.
First
stand
Second
stand
Tree density (ha"^)
Total basal area
(dm2-ha-i)
Mean basal area of C.
arizonica trees
(dm^'tree"^
Number of C. arizonica
cored
'Mean time required
per unit growth of C.
arizonica (yr-cnr')
Mean extrapolated tree
age of C. arizonica (yr)
Standard deviation of
extrapolated tree age (yr)
Oldest C arizcmica
extrapolated age (yr)
Yoimgest C. arizonica
extrapolated age (yr)
Number of C. arizonica
saplings and trees too
small to core (20-30
cm)
Number of C. arizonica
seedlings
Correlation coefficient
for suppression test
1333
3733
11901
5854
9.51
1.49
21
58
24.2
16.6
317.1
116.9
63.0
26.4
456.7
178.1
197.6
62.4
0
81
0
0
-0.833
-0.673
'Calculated by dividing the number of annual rings per core by the
length of that core, it expresses the number of years required for an individ-
ual to add one cm of radial growth and is the reciprocal of the growth rate.
The great number of seeds produced does
not necessarily insure abundant reproduction.
Sudworth (1915) noted that seeding of C.
arizonica was best on moist, bare mineral
soils. Field observations made during the
course of data collection confirm this sugges-
tion, in that C. arizonica reproduction was
generally restricted to areas within 2 of the
19 stands, and then only on exposed mineral
soil associated with recent floods or logging.
Moreover, the weighted average of litter
depth where C. arizonica seedlings were en-
countered (1.74 cm) was only half the mean
depth of litter for all stands in this study (3.42
cm), indicating that successful reproduction
is associated with reduced litter depth.
Germination tests also indicated that a lit-
ter layer sharply reduced the germination
and survival of seedlings. Onlv 3.7 percent of
the seeds germinated on the litter-covered
substrate, in contrast to 10.0 percent on min-
eral soil controls (Table 4). Of the other fac-
tors tested, freezing of seeds prior to germi-
nation, reduced light intensity, and
immersion of seeds in water did not alter ger-
mination success. Simulated exposure of seed-
bearing cones to canopy fire conditions,
while reducing germination success (6.0 per-
was evident in both stands (Table 3). The de-
gree of suppression was greater in the older
stand, as a consequence of the longer period
of time over which competition was acting.
These results indicate that small C. arizonica
trees may persist beneath a canopy, and sug-
gest that the species is shade tolerant.
Evidence from core aging also .suggests
that C. arizonica possesses potentially great
longevity. Based on the rates of growth de-
termined in this study (Table 3), and account-
ing for multiple annual rings, some of the
larger individuals encountered in canyon bot-
toms (exceeding 1 m dbh) may be 300 to 500
years of age.
Not only does the species achieve old ages,
but also it produces great numbers of seed.
The mean number of seeds per cone was
104.2, with a standard deviation of 19.8. It
was estimated that healthy trees of moderate
size (50 cm circumference) may produce
from 103 to 10* cones per year, therefore re-
sulting in an annual seed crop per tree of
from 10^ to 10'^ seeds.
Table 4. Germination test results. The soil in each
pot was composed of a mixture of 50 percent sand and
.50 percent silt loam. The surface was without litter cov-
er. Pots were supplied with adequate water regularly
and were grown under high levels of simlight. The pots
were 15 cm in diameter and allowed free drainage be-
low. The air temperature was between 20 and 24 C
throughout the duration of the tests.
Composite
Number of
germination
Treatment
test pots
rates (%)
Litter cover over
mineral substrate
6
3.7
Seeds exposed to
freezing before
planting
4
8.8
Seeds immersed in
agitated water
2
10.0
"Seeds exposed to
canopy fire conditions
1
6.0
°° Seeds exposed to
groimd fire conditions
1
0.0
Seeds grown under
reduced light levels
2
8.5
°° "Controls
10
10.0
•—Cones filled with seeds were exposed to 80 C for eight minutes.
■-Cones filled with seeds were exposed to 315 C for eight minutes.
*— Control conditions.
260
Great Basin Naturalist
Vol. 40, No. 3
cent), did not destroy all seeds. In contrast,
exposure to simulated ground fire conditions,
which are much hotter, killed all seeds within
the cones. Thus, C. arizonica seeds enclosed
in cones on branches may remain viable after
fires, and have the capability to germinate on
the mineral seedbeds exposed by burning.
Discussion
The absence of C. arizonica reproduction
under adults of the same species in natural
stands is striking. Two interpretations of this
observed absence of young individuals are
possible. First, successful reproduction may
be temporally sporadic and linked to occa-
sional optimal climatic conditions that foster
waves of C. arizonica reproduction. If such
optimal conditions have not occurred during
the last 20 or 30 years, the paucity of C.
arizonica seedlings would be explained. How-
ever, successful reproduction of C. arizonica
was encountered in two recently disturbed
habitats, and this indicates that recent climat-
ic conditions are not inimical to the tree's re-
production. The environmental conditions as-
sociated with these cases of successful
reproduction do support, however, a second
interpretation of the absence of C. arizonica
seedlings, namely, that disturbance is neces-
sary to create conditions favorable for rege-
neration, and that factors associated with a
closed forest inhibit seeding by the tree.
Stand size-class analyses would seem to
both siipport and contradict this character-
ization of C. arizonica as a species requiring
disturbance for reproduction. Stands of the
first group suggest that the species is tolerant,
able to maintain itself under a closed canopy
through time. Stands of the second and third
size-class groups imply that the species is in-
tolerant, with a period of establishment, fol-
lowed by the cessation of C. arizonica repro-
duction. Therefore, the latter groups support
the hypothesis that the species is opportunis-
tic, capable of invading open habitats, but
not able to compete effectively with other
species as successional processes proceed.
Inspection of core analyses resolve this ap-
parent paradox. The first stand cored, repre-
senting those stands that are dominated by
large individuals and which suggest that the
species is a pioneer, possesses no individuals
with a tree ring age of less than 200 years.
The trees from this stand exhibit a tendency
to clump in the 250- to 375-year tree ring
age range. Lack of successful C. arizonica
regeneration over an extended period, com-
bined with a tendency toward clumping of
ages, suggests that, following a lengthy peri-
od of C. arizonica establishment, reproduc-
tion is virtually eliminated. The second stand
cored, representing those stands with a mixed
size structure and suggesting that the species
is a climax-type, possesses trees ranging in
tree ring age from 25 to 150 years. This stand
possesses, however, a conspicuous absence of
reproduction during the last quarter century.
These data, in the same manner as those from
the first stand, suggest that following a
lengthy period of colonization and reproduc-
tion of C. arizonica, perhaps 50 to 100 years
in length, reproduction subsequently ceases.
The population structures of these two
stands, therefore, differ only in the elapsed
times since disturbance, rather than in some
fundamental difference in the ecological be
havior of the species. In both cases, C. arizo-
nica acts as an intolerant species, requiring
disturbance to create invadable habitats. In
the denser, more youthful C. arizonica popu-
lation of the second stand, disturbance was
more recent than in the older population of
C. arizonica in the first stand. The initial in-
terpretation of stand structures of the first
type— that the species is a climax-type, with a
stable, self-perpetuating population— is
shown to be incorrect. Stands of the first
group attain this pseudoclimax type structure
only temporarily as a consequence of the ex-
tended period following disturbance when C.
arizonica reproduction is possible. Thus, the
three groups identified by differing size
structures may be interpreted as stages of a
temporal continuum, with each stage reflect-
ing a longer period of development since a
disturbance event.
Other results yield evidence favoring the
interpretation of C. arizonica as an intolerant
species. The longevity of C. arizonica is char-
acteristic of many pioneer species, as it in-
creases the allowable length of time between
periodic disturbances, thus enhancing the
probability that perturbation will occur on or
near the site while mature seed trees are
available for colonization. Seed counts and
estimated cone crop sizes indicate that C.
September 1980
Parker: Arizona Cypress Succession
261
arizonico is a prolific seeder, also a character-
istic of the reproductive strategy of a pioneer
species, because it increases the probability
that a germule will be dispersed to an open
habitat. The inverse relationship of litter
depth to C. arizonica seedling number in-
dicates that partial or complete removal of
litter on the forest floor is necessary to pro-
mote reproduction. The inhibition of litter on
germination in test pots further corroborates
the negative effects of litter accumulation on
this tree's regeneration. This need for mineral
seedbeds, created by natural disturbance (i.e.,
fire or flooding) or by certain human activi-
ties (i.e., logging) is typical of pioneer ele-
ments. Finally, the ability of seeds to remain
viable following a canopy fire is an opportu-
nistic character often favored by intolerant
species that must colonize open habitats fol-
lowing such fires.
In contrast to the characteristics suggesting
that C. arizonica is a pioneer species, the
tree's abilities to germinate in low light con-
ditions and to tolerate shaded sites by sup-
pression of growth are characteristics more
commonly associated with climax-type than
with pioneer species. These responses to light
are puzzling and deserve closer inspection in
the future. In general, however, it is con-
cluded that C. arizonica is a pioneer species,
requiring periodic disturbance to open new
sites for colonization.
A unique characteristic of C. arizonica
stand dynamics is the long period, as long as
a century, during which colonization and re-
production remain possible after the disturb-
ance event. Jenny et al. (1949) have demon-
strated that, in ponderosa pine forests at 1220
to 2220 m elevation in California, 100 to 200
years are required to reach a near-equilib-
rium steady-state condition of litter thickness,
in which mineralization of organic matter
balances the addition of litter produced by
the vegetation. Such a long period of litter
accumulation is characteristic of moist, cool
montane conifer forest ecosystems, and fits
well with the observed time scale of C. arizo-
nica establishment on a site. This peculiarly
long period of colonization appears to be re-
lated to the shade-tolerant character of the
species. Most pioneer species, being intoler-
ant, rapidly cease reproduction as light levels
decrease under the species' own canopy. The
ability of C. arizonica to reproduce in shade
allows it to continue reproducing for a much
longer period than most colonizing species.
This period is later terminated by the more
gradual process of litter accumulation.
Thus, C. arizonica may be considered to be
a pioneer species that is intolerant of litter
accumulation under a closed canopv, even
though it is tolerant of low light levels fol-
lowing colonization of disturbed sites. Addi-
tionally, its habit of seed persistence in the
cone is critical to its maintenance on most
sites. It may not depend, then, on seed dis-
persed from distant sources to colonize a giv-
en disturbed area.
Using the Noble and Slatyer (1977) model
of successional processes (from Cattelino et
al. 1979), this study suggests three tvpes of
species that exhibit fundamentally different
responses to disturbance events. First, the up-
land conifers are DI species, characterized bv
widespread dispersal capabilities (D) and in-
tolerance to a closed canopy (I). These trees
depend on colonizing disturbed sites with
seed from an off-site source. Second, the pin-
yon oak group are DT/VT species, which
persist on a disturbed site by colonization
from an off-site source (D— pinyon) or by
vegetative reproduction (V— oaks). These spe-
cies maintain themselves by virtue of their
tolerance of closed canopy conditions (T).
Cupresstis arizonica, the third type, is a CI
(DI) species. As described above, it is intoler-
ant (I) of closed forest conditions and gener-
ally is maintained locally by seed persistence
in cones from the canopy (C), which shelter
viable seeds through disturbance events. Be-
cause of its limited range, C. arizonica is less
commonly maintained by dispersal from adja-
cent poulations (D), except perhaps in ripa-
rian stands, where surface water wash may
supply seeds to wash environments from sur-
rounding upland slopes.
These three types of species, in com-
bination with differential longevity patterns,
yield three possible pathways of community
development (Fig. 3). The initial composition
of each case is represented by at least one
stand sampled in this study. The first and sec-
ond cases diagram multiple successional
pathways in communities which possess rep-
resentatives of the three behavioral types. In
the first case, the model assumes a greater
262
Great Basin Naturalist
Vol. 40, No. 3
Case 1
<b
/
UC, CA, PO
CA, PO
PO
-J
1
I x<a
z'
V
-^_L
UCPO
PO
—^ — TTL "ii^-
X >b
Case 2
' \a<x<b
UC, CA, PO
UCPO
b<x<c
/
-^_lJ
UC, PO
PO
i
I x>_c I
Case 3
N
\ a<x<b
J-
CA, PO
- PO
T
L x_l§ !
L
Fig. 3. Potential multiple pathways in a successional development model. Solid lines indicate developmental
changes in the absence of disturbance. Dashed lines represent disturbance events. Boxes define the constraints on C.
arizonica populations, if they are to avoid local extinction. Symbols:
UC Upland conifers
CA Cupressiis arizcmica
PO Pinyon-oak species
X period of time since last catastrophic disturbance event
a time required by C. arizonica to reach maturity and produce seeds
b longevity of C. arizonica
c longevity of upland conifers
September 1980
Parker: Arizona Cypress Succession
263
life span for C. arizonica than for the upland
conifers, but in the second case the model as-
sumes that upland conifers live longer than
C. arizonica. In both cases, upland conifers
are maintained indefinitely in the landscape
by their wide dispersal of seeds, and pinyons
and oaks are maintained by their tolerance to
a closed forest canopy. Cupressus arizonica,
however, may become locally extinct, depen-
ding on the timing of disturbance events. If
disturbance recurs more frequently than the
time required for C. arizonica maturity, the
species will not be maintained in the commu-
nity because the canopy seed reserve is de-
stroyed. Such local extinction assumes that no
introductions occur by dispersal from non-
local populations, and that the disturbance
was large enough in areal extent to kill the
entire local population of C. arizonica. A sec-
ond mechanism for local extinction in each
case would be for disturbance to be so in-
frequent that the local population dies out
before reinitiation of a successional cycle.
The third case in the model represents a sim-
plified commimity in which the upland con-
ifer element is locally absent, whether by
habitat restriction or chance.
It is apparent from this model of succes-
sion that, assuming habitat factors remain un-
changed, local extinction of C. arizonica
would occur only with excessively frequent
or infrequent catastrophic fire events. Low
net productivity and fuel loadings do not
make frequent catastrophic fire a highly
probable event in montane forests of south-
eastern Arizona. Suppression of fires by hu-
mans may temporarily reduce C. arizonica
reproduction, but, unless the program is
maintained effectively for several himdred
years, it is unlikely to result in extinction of
local populations.
The situation in riparian settings is even
more favorable for C. arizonica perpetuation.
In these habitats, population maintenance is
effected by dispersal from adjacent upland
populations, as well as by local preservation
of seeds in the canopy. Extinction on such
sites would require elimination, not only of
the local population, but also of other C.
arizonica populations higher in the water-
shed. Additionally, flooding assumes an im-
portant role as a disturbance mechanism in
these stands. This is particularly significant
with respect to C. arizonica regeneration, be-
cause, while it may not increase surface light
intensities, it will remove litter and expose
bare mineral soil, which triggers reproduc-
tion, apparently irrespective of light condi-
tions at the forest floor. (Certainly, light in-
tensities influence growth rates and vigor,
but not necessarily establishment potential.)
In effect, C. arizonica appears to be a
stable, terminal element of the restricted
habitats in which it presently occurs. Its life
history characteristics (including its great po-
tential longevity), population maintenance
mechanisms, and colonization patterns com-
bine to make it a persistent species, threat-
ened in a temporal sense only by the remote
probability of either very frequent or in-
frequent catastrophic disturbance events.
Any short-term efforts designed to stimulate
C. arizonica reproduction should be ad-
dressed to the elimination of fire suppression
policies in regions of its occurrence. Long-
term preservation of this locally dispersed,
intolerant species requires that open habitats
continue to be created by catastrophic per
turbations with a frequency of recurrence
greater than the age of first viable seed pro-
duction, and less than the life span of the in-
dividuals in the population.
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Arizonan and Californian species of Cupressus.
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264
Great Basin Naturalist
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Kearney, T. H.. and R. H. Peebles. 1960. Arizona flora.
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Parker, .\. J. 1980. Site preferences and community
characteristics of Cupressus arizonica Greene
(Cupressaceae) in southea.stern Arizona. South-
western Naturalist. In press.
Posey, C. E., and }. F. Gogg.-vns. 1967. Observations on
species of cypress indigenous to the United
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19 pp.
SuDwoRTH, G. B. 1915. The cypress and juniper trees of
the Rocky Mountain region. USD.\ Bull.
207:1-35.
Whittaker, R. H., and W. A. Nierinc. 1965. Vegeta-
tion of the Santa Catalina Mountains, Arizona: a
gradient analysis of the south slope. Ecologv
46:429-452.
Wolf, C. B. 1948a. Taxonomic and distributional stud-
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1948b. Horticultural studies and experiments on
the New World cypresses. El Aliso l:.325-444.
A SELF-POLLINATION EXPERIMENT IN PINUS EDULIS
Ronald M. Lanner'
Abstract.- Controlled pollinations were performed on fonr pinyons (Pinus cdtilis Engelm.) to compare the results
of selfing and outcrossing. Final cone size was the same under both treatments. There was no significant difference
in number of seeds per cone. Filled-seed yields averaged 14.4 percent in selfings and 90.5 percent in outcrossings.
Relative self-fertility averaged about 15 percent, a level comparable with that of other pine species studied.
The breeding system of pinyon {Pinus
edulis Engelm.) has received Httle study. In
several particulars pinyon resembles other
members of Pinus. For example, it is general-
ly monoecious (Lanner 1975) and the sexes
are usually segregated with the upper crown
tending to be more female than male and
vice versa, though there may be broad over-
lap. Pollen shed and conelet receptivity are
limited to a short period in the spring (Lan-
ner 1970). Meiosis leading to microspore for-
mation is temperature sensitive during meta-
phase and anaphase, and temperatures below
4 C may result in inviable polyploid pollen
grains (Chira 1967). Pinyon crosses readily
with singleleaf pinyon {P. monophylla Torr.
& Frem.) and natural hybrids and in-
trogressants are widely distributed where
their ranges overlap (Lanner 1974, 1975).
Though pines are generally considered to
be outcrossing species, many studies have
shown that self-fertilization is usually at least
marginally successful in producing sound
seeds. This paper reports the results of a con-
trolled pollination experiment in which self-
ing was compared to outcrossing.
Methods and Materials
Isolation bags of nonwoven cloth with cel-
luloid windows were placed over female
branches of four pinyon pines of unknown
seed source on the campus of Utah State Uni-
versity 4 June 1976. Pollen extracted the pre-
vious year and stored in a freezer was ap-
plied with a hypodermic syringe 8 June 1976,
shortly after conelets emerged from their
'Department of Forestry and Outdoor Recreation, Utah State University', Logan, Utah 84322.
covers of bud scales. Table 1 summarizes the
crosses made. Pollination treatments were
self-pollen and outcross pollen. Most of the
outcross pollen was a bulked mix from all the
seed trees except tree 3, plus an additional
tree. Tree 3 was outcrossed with trees 9 and
28. Isolation bags were removed 22 June
1976. Cones were inventoried 13 July 1976
and in June 1977; they were harve.sted 25
August 1977 prior to cone dehiscence. Seeds
were extracted in the laboratory as cones de-
hisced at room temperature. Seeds were tal-
lied as filled or empty on the basis of weight
and seed coat color: empty seeds are easilv
detected in handling and by their light-col-
ored seed coats.
Results
1. Cone survival— Within 2 weeks of pol-
lination 12 of the 31 .selfed cones and 2 of the
35 outcrossed cones had aborted. By 15 June
1977, a year after pollination, one more self-
ed cone and 4 more outcrossed cones had
aborted and been shed. No further losses oc-
Table 1. Distribution of pollination treatments
among trees.
Pollination treatment
Tree
Selfed
Outcrossed
Number of conelets
.3
—
16
4
7
5
.36
12
8
58
12
6
Total
31
35
265
266
Great Basin Naturalist
Vol. 40, No. 3
curred prior to maturity. Final survival rates
were 57 percent for selfed cones and 83 per-
cent for outcrossed cones.
2. Cone size— Lengths of 14 mature selfed
cones ranged from 3.3-3.9 cm (x = 3.6 cm).
Lengths of 16 outcrossed cones showed iden-
tical ranges and mean.
3. Seed yield — Yields of seeds per cone
are summarized in Table 2.
In all three trees that were both selfed and
outcrossed the yield of seeds per cone was
slightly higher under selfing, but the differ-
ences were nonsignificant in all cases.
Yield of filled seed and relative self-
fertility are summarized in Table 3. Filled
seed percent averaged 14.4 percent in self-
ings and 90.5 percent in outcrossings. Rela-
tive self-fertility, a statistic designed to neu-
tralize variation not associated with embryo
genotype but which can influence seed yield
(Sorensen 1970), ranged from about 10 to 21
percent and averaged about 15 percent.
Discussion and Conclusions
It is unclear why survival of selfed cones
was exceeded by that of the outcrossed cones.
All losses of selfed cones occurred by 13 July
1976, just five weeks after pollination. Fer-
Table 2. Number of seeds per cone after selfini; and
outcrossing.
Pollination treatment
Tree
Selfed
Outcrossed
3
_
13.9
4
5.8
4.8
.36
10.4
9.4
58
5.2
3.7
Total
7.76
10.52
Table 3. Yield of filled seed by pollination treatment
and relative self-fertility of seed trees.
Pollination treatment
and yield of filled seed
Relative
Tree
Selfed
Outcrossed
self-fertility'
Percent
4
8.7
89.5
9.72
36
15.7
74.2
21.2
.58
15.4
lOO.O
15.4
MEAN
14.4
90.5
15.4
'Relative self-fertility (Sorenson
1970) is computed as:
No. of filled seed/total number of seed in selfings
No. of filled seed/total number of seed in crossings
tilization does not occur in pines until about
one year after pollination and cones are
made up entirely of female parent tissue, so
early cone abortion is not related to such
genetic causes as homozygosity of recessive
genes. The most likely explanation of a lower
average level of cone set among selfed cones
is a lower level of pollination. Pollen of indi-
vidual trees was in shorter supply than
bulked pollen, and may have been used too
sparingly in a few cases, re.sulting in close to
zero pollination of ovules. Sorenson (1970)
has reported 75 percent cone set in out-
crossed cones of ponderosa pine {P. ponderosa
Laws.) and 70 percent in .selfed cones, but the
difference was not statistically significant.
Reduction in filled seed per cone by selfing
as compared to outcrossing is a common find-
ing among pines. Soreason (1970) found that
filled seed yields in ponderosa pines average
23.7 percent for .self and 66.5 percent for out-
crosses; in Scotch pine (P. sijlvestris L.) self-
ing produced 13.4 percent filled .seed and
outcrossing 71.4 percent (Johnsson 1976).
Bramlett and Pepper (1975) reported average
filled seed yields of 16.4 percent and 90.7
percent for selfed and outcrossed cones of
Virginia pine (P. virginiana Mill.). Squillace
and Kraus (1962) found that selfing slash pine
{P. elliottii Engelm.) resulted in 9 sound seeds
per cone, and outcrossing produced 34 seeds
per cone. Snyder (1968), also working with
sla.sh pine, generalized that selfing yields
about 15 percent as many seeds as wind polli-
nation. In this experiment, the filled-seed
yield of selfed pinyon was about 15 percent
that of the outcrosses.
The reduced .sound-.seed yield in selfed
pines and other conifers is due to the en-
hanced homozygosity of embryonic lethal al-
leles. The results of the crosses reported here
suggest that the number of lethals in pinyon
is comparable to that in other species of
Pinus.
Acknowledgments
This work was funded by the Mclntire-
Stennis program and is published as Utah Ag-
ricultural Experiment Station Journal Paper
2549. Mark Buckbee and Michael Jenkins a.s-
sisted in the field and laboratory work.
September 1980
Lanner: Pinyon Pollination
267
Literature Cited
Bramlett, D. L., and \V. D. Pepper. 1974. Seed yield
from a diallel cios.s in Virginia pine. Pages 49-55
in Seed Yield from Southern Pine Seed Orchards
Colloq. Proc., Macon, Ceorgia.
CniRA, E. 1967. Pollen grains of Pinus edulis with more
than the haploid number of chromosomes. Silvae
Genet. 16:14-18.
JonNssoN, H. 1976. Contributions to the genetics of
empty grains in the seed of pine (Pintts silvestris).
Silvae Genet. 25:10-14.
La.nner, R. M. 1970. Origin of the sunmier shoot of pin-
von pines. Canadian ). Bot. .38:1759-1765.
1974. Natural hybridization between Pinus edulis
and Pinus monophylla in the American South-
west. Silvae Genet. 2.3:108-116.
1975. Pinon pines and junipers of the south-
western woodlands. Pages 1-17 in G. F. Gifford
and F. E. Busby, eds., Proc. Symp. on Pinyon-
Juniper Ecosystems. Utah State University, Lo-
gan.
Snyder, E. B. 1968. Seed yield and nurserv performance
of self-pollinated slash pines. Forest Sci.
14:68-74.
SoRENSEN, F. C. 1970. Self-fertility of a central Oregon
source of ponderosa pine. USD.\ Forest Service
Res. Pap. PNW-109, 9 pp.
SyuiLLACE, A. E., and J. F. Krals. 1962. Effects of in-
breeding on seed yield, germination, rate of ger-
mination, and seedling growth in slash pine.
Pages 59-63 in South. Forest. Tree Imp. Comm.
and Soc. Amer. Foresters Tree Imp. Comm. For.
Genet. Workshop Proc.
COMPARATIVE FLORAL BIOLOGY OF PENSTEMON EATONII AND
PENSTEMON CYANANTHUS IN CENTRAL UTAH: A PRELIMINARY STUDY
Lucinda Bateman'
Abstract.— A comparison of the floral visitors of two closely related plant species, Penstemon ctjanonthus and P.
eatonii suggests that flower shape and color may affect the number and type of pollinators, and the ability of the
plant to set fniit. Penstemon cijananthus, which is most attractive to hymenopteran visitors, has a blue flower, large
in diameter, that is positioned as a convenient "landing pad." Although many types of insects visit the flower, the
transport of pollen directly to flowers of another individual of the same species is somewhat inefficient, since fruiting
success is low (66.7 percent). The tubular red flowers of P. eatonii are narrow and droop downward from the stem.
The nectar is accessible to a specific and well-adapted visitor, the hummingbird. This less promiscuous, bird-polli-
nated species sets fruit more successfully (82.4 percent) than P. cijananthus.
Observations of animal visitors to flowers
suggest that the broad range of phenology,
size, structure, color, and odor evident
among flowers of any complex plant commu-
nity is related to the size, morphology, be-
havior, and sensory acuity of the animals vis-
iting the flowers. It has been observed, for
instance, that nocturnal blooming flowers are
specially adapted to night-flying insects or
bats (Faegri and Pijl 1971). Bees appear to be
more influenced by flower shape than color.
Bees also have appendages specialized for
collection and transport of pollen, since pol-
len is an important food item for their off-
spring. Accordingly, flowers visited by bees
may be white, blue, or yellow, but commonly
offer a generous reward of both nectar and
pollen (Proctor and Yeo 1972, Raven, Evert,
and Curtis 1976). Flowers whose most fre-
quent visitors are nonhovering individuals
such as bees are usually so structured as to
provide a "landing pad" near the flower's re-
productive parts and nectar or pollen "re-
ward" (Free 1970).
In view of the fact that few insects are be-
lieved able to distinguish red (Raven, Evert,
and Curtis 1976), it seems signiflcant that red
flowers worldwide are regularly visited by
birds, known to be more stimulated by that
color than any other (Faegri and Pijl 1971). It
is an interesting and probably not imrelated
fact that red flowers, unlike most flowers of
other colors, are essentially odorless (Grant
1966). Significantly, insects have keen olfac-
tory senses, but those faculties in birds tend
to be poorly developed (Proctor and Yeo
1972, Faegri and Pijl 1971). The corolla of
many flowers visited by birds is typically
tubular and narrow and without a landing
platform, excluding all but the smallest insect
intruders (Raven, Evert and Curtis 1976).
Clearly, flowers that are tubular, red, and
odorless should offer minimal attraction to
insects, specifically nonhovering insects, but
should be highly attractive to birds (Boyd and
Brown 1978).
Phenological, structural, color, and odor
differences among the flowers of any particu-
lar complex plant community undoubtedly
increase the degree of fidelity between par-
ticular flower types and specific insects.
Since flowering periods of different species in
a common plant community often overlap,
flower-pollinator fidelity should enhance re-
productive success of plant species having
such flowering overlap. Plants able to selec-
tively entice pollinators should be more suc-
cessful in the distribution of their pollen. The
animal visitor should simply find it more
profitable to visit nonpromisCiious flowers,
since there is a greater probability that such
flowers will yield a reward on any given visit.
Promiscuous pollinators would be particu-
larly detrimental to the reproductive success
of rare to moderately common plants that are
obligate outcrossers, and that flower simulta-
'Department of Botany and Range Science, Brigham Young University, Prove, Utah 84601. Present address: 1212 Ash Avenue, Provo. Utah 84601.
268
September 1980
Bateman: Comparative Floral Biology
269
neously with a variety of other species, since
pollen of a given species could be expected
to be largely dislodged from the body of the
promiscuous pollinator before it encountered
another individual of that plant species (Le-
vin and Anderson 1970).
Evidence suggests that, by chance muta-
tion, plants gradually develop characteristics
attractive to the most consistent pollinators
in the commimity. Surviving individuals of
the species become specialized for visits from
the more efficient pollinators. Tantalizing
odors beckon hungry insects, and brightly
colored corollas are a signal to the pollinators
of the presence of a nectar reward. These
forms of advertisement attract pollinators,
and thereby accomplish a more efficient
spread of pollen.
Statement of the Problem
Few studies have been made of the com-
parative floral biology of two species of the
same genus growing in a common environ-
ment and flowering simultaneously. The pur-
pose of this study is to compare the floral
morphology, insect visitors, and fruit set of
two closely related species, Pensternon eaton-
ii Gray and Penstemon cijananthus Hook. At
the site studied, these species grow in close
proximity, although individuals of P. cya-
nanthiis are approximately twice as numer-
ous. I have tested the following hypotheses:
(a) Corolla size and color affect the number
and types of pollinators, (b) P. eatonii, with a
red, narrow corolla tube, will attract fewer
insects and will be visited by hummingbirds.
(c) P. cijananthus, with a broader, blue co-
rolla, will be more promiscuous, attracting a
variety of insects, (d) The less promiscuous,
bird-pollinated species will set fruit more
successfully.
Materials and Methods
Penstemon eatonii plants produce from
5-10 inflorescences, each approximately 50
cm in length. Narrow tubular corollas hang
downward along the upright stem. The five-
lobed corolla is red. Four fertile stamens lie
within the corolla tube, and one sterile sta-
men protrudes beyond the corolla orifice
(Fig. lA).
Fig. lA. Penstemon cifanantlms flower shape and ar-
rangement of reproductive organs (above); IB. Penste-
mon eatonii flower shape and arrangement of reproduc-
tive organs.
Penstemon cyananthus generally produces
four to five times as many inflorescenses as P.
eatonii, each approximately 45 cm in length.
Each blue, five-lobed corolla is displayed at
an ascending angle from the stem and has
fused petals and five stamens, four fertile and
one sterile. Two stamens are as long as the
petals, but the other two are only two-thirds
that length (Fig. IB).
The data were collected 29 June, and 2
July 1979 on the west face of Mount Tim-
panogos, in Battlecreek Canyon, near Pleas-
ant Grove, Utah. A small stream flows
through the relatively dry study site. Some of
the major contributors to the vegetation of
the area are: sagebrush {Artemisia tridentata
Nutt.), rabbitbrush {Chrysothamnus nau-
seosus [Pall.] Britt.), poison ivy (Tox-
icodendron radicans L.), scrub oak [Quercus
gambelii Nutt.), big tooth maple {Acer gradi-
dentatum Nutt.), chokecherry {Prunus virgi-
270
Great Basin Naturalist
Vol. 40, No. 3
niana L.), squawbush (Rhus trilobata Nutt.),
and various grasses.
Observations of the plants were made daily
in the first half of each hour from 0730 to
1800 hours. On 29 June P. cyananthus was
observed during the first hour, P. eatonii dur-
ing the second, and so on throughout the day.
On 2 July the observation order was reversed.
The two plants observed during these time
periods were within four feet of each other.
Fruit set data were collected 14 July 1979,
about 0.4 km down the canyon from the first
observation site. Individuals of each species
were randomly selected and checked for
height, spent flowers, developing fruit, open
flowers, and buds. Ten stems of each of 8 dif-
ferent P. cyananthus plants were analyzed.
Because of fewer stems per plant, 13 P. ea-
tonii individuals with up to 10 stems per
plant were also tallied. Of the total flowers
that had been produced per inflorescence,
both species showed at least 90 percent spent
flowers, the remainder of the number con-
sisting of open flowers and buds.
Average fruit set was calculated using the
formula:
No. filled fruits
No. spent flowers
Plant nomenclature follows Welsh and
Moore (1973). Insect family names are taken
from Borror and White (1970). Bird identi-
fication is from Bobbins, Brium, and Zim
(1966).
S 30-
7 AM 8 9 10 11 12 1 2 3 4 5 6 PM
Fig. 2. Temperature readings in degrees Celsius for
29 June (dotted line) and 2 July (solid line).
Results
Insect activity was minimal during the
early morning hours, but as air temperature
rose, greater numbers and more types of in-
sects appeared. Temperatures for the two
days of observation were similar (Fig. 2).
Penstemon cyananthus attracted a greater
number of total visitors (153 in two days),
and also more insect families (9) than P. ea-
tonii (Table 1). Not all insect visitors came to
the plant in search of pollen or nectar. Some
coleopterans landed on the showy petals as if
to rest, making no attempt to enter the flow-
er. Still other types of insects crawled among
the stems and leaves. Neither of these types
of visitors were recorded as pollinators. The
most numerous and determined visitors were
hymenopterans in search of pollen (Table 1).
Hymenopterans accounted for 85 percent of
the visitors and over 89 percent of the flow-
ers visited. Thirteen percent of the visitors to
P. cyananthus were lepidopterans and the re-
maining 2 percent were dipterans.
Penstemon eatonii attracted fewer insect
visitors (23) or 15 percent as many as P. cya-
nanthus. The visitors that were observed be-
30
28
•s
/ ^
26
/
"^
24
/
\ .'^
22
I 1
\ ,' »
1
0 \
20
1
\
18
1
1
\
\
16
^
\
14
•
\
r
\
12
*
*
\
10
/
1
1
8
/
I
X .
•
6
^ Av
4
2
^y^^^X.^^^^
0
•• ^ tL^^^>»^ / ^"^
T
Fig. 3. Hourly distribution of individual flower vis-
itors to Penstemon cijaniinthtis (dotted line) and P. ea-
tonii (solid line) throughout the observation period.
Fig. 4. Hourlv distribution of insect families that vis-
ited Penstemon ri/anantlius (dotted line) and P. eatonii
(solid line) throughout the observation period.
September 1980
Bateman: Comparative Floral Biology
271
longed to three hymenopteran families. Dur-
ing tlie heat of the day, when the greater
numbers of insects were actively foraging, 4
or 5 attempted to enter the narrow corolla
tube. Few were successful in their efforts.
Once during the observation period, a broad-
tailed hummingbird visited the plant at 0930,
before the temperature had risen above 24 C
(75 F). The bird moved systematically down
the canyon, stopping at every P. eatonii indi-
vidual within 10 or 20 m of either side of the
path. Upon reaching the observation plant.
the bird sampled five or six flowers on four of
the six stalks, pausing less than one second at
each flower. It hovered in the air .slightly be-
low each flower (Figs. 3 and 4).
Penstemon cyananthu.s averaged 66.7 per-
cent fruit set per plant, and P. eatonii showed
a much higher 82.4 percent (Table 2).
Discussion
The results of this study clearly indicate
that the two penstemons considered have de-
Table 1. Activity and presence of each family throvigho\it the observation period. The first (upper) n\imber in-
dicates the number of individual visitors; the second (lower) indicates the total number of flowers visited.
Hour observed
Family
8
9
10 11 12 1
2
3
4
5
6
Total
Penstenion cyanantlnis
.\pidae'
Chrysididae'
1
2
1
3
3
10
3
12
1
3
3
8
6
22
Halictidae'
Hesperiidae-
Megachilidae'
'Nymphalidae
Pieridae-
Syrphidae'
Vespidae'
Total
1
3
1
6
2
4
2
7
3
9
10
25
2
11
6
17
10
27
14
36
5
4
3
2
2
1
21
6
4
5
6
15
2
3
2
1
1
29
13
25
/
16
10
10
18
2
75
6
4
39
26
15
47
5
183
6
6
2
2
1
1
1
1
11
1
1
3
3
8
2
8
2
3
25
22
6
26
9
15
89
r;
28
26
21
24
7
153
s
(.3
66
46
61
23
370
Penstenwn eatonii
Formicidae'
Halictidae'
Megachilidae
'Trochilidae'
Total
1
15
2
16
2
9
5
17
1
15
2
1
2.3
5
2
54
Key to the orders: 1, Hymenoptera; 2, Lepidoptera^ 3. Diptera; 4, Apodiformes {'Selasphorus platycercus).
272
Great Basin Naturalist
Vol. 40, No. 3
Table 2. Percent fruiting success per plant of Penste-
mon cyananthus and P. eatonii (Number of fruit/spent
flowers/plant). (Difference significant at the .05 level)
Plant
P. cyananthus
P. eatonii
%
%
1
84.7
45.5
2
47.5
93.3
3
59.8
73.1
4
47.6
68.5
5
74.8
89.7
6
75.5
90.0
7
69.7
77.4
8
79.3
9
90.8
10
86.1
11
95.4
12
82.4
13
100.0
Average %
66.7
82.4
veloped different ways of attracting pollina-
tors. The narrow red corolla tube of P. eaton-
ii physically excludes all but a few small in-
.sect visitors, and the absence of odor appears
to minimize attractiveness to insect visitors.
The absence of any sort of a landing pad hin-
ders the ability of nonhovering visitors to
successfully work the flowers. The only ob-
vious diurnal pollinator, a hummingbird, is
le.ss frequent but more systematic and specif-
ic. That the job of pollination is done more
efficiently by such a specific pollinator is sug-
gested by the higher fruit set.
The more promiscuous P. cyananthus also
enjoys a fairly high fruit set. It does this,
however, with larger, more accessible blos-
soms and with no assurance that its pollina-
tors will be species specific. The flowers are,
of necessity, displayed so as to form a con-
venient landing pad for approaching hyme-
nopterans.
Limitations
Although the initial implications are clear,
these data constitute only preliminary results.
Data were collected during daylight hours
late in the flowering season, when flowers of
both species contained little or no nectar.
Only one hummingbird was observed directly
during the study, but my presence may have
frightened usual avian visitors away. No at-
tempt was made to observe early evening or
nocturnal pollinators. The degree to which
either species is capable of self-pollination is
unknown.
Literature Cited
Bond, H. W., and W. Brown. 1978. The exploitation of
floral nectar in Eucalyptus incrassata by honey-
eaters and honeybees. Unpublished manuscript.
BoRROR, D. L., and R. E. white. 1970. A field guide to
the insects. Houghton Mifflin Co., Boston.
Faegri,K., and L. van der Pijl. 1971. The principles of
pollination ecology. 2d ed. Pergamon Press, Ox-
ford, England.
Free, J. B. 1970. Effect of flower shapes and nectar
guides on the behavior of foraging honeybees. Be-
havior 37:269-285.
Grant, K. A. 1966. A hypothesis concerning the preva-
lence of red coloration in Californian humming-
bird flowers. American Nat. 100:85-98.
Levin, D. A. 1969. The effect of corolla color and out-
line on interspecific pollen flow in Phlox. Evolu-
tion 23:444-445.
Proctor, M., and P. Yeo. 1972. The pollination of flow-
ers. Taplinger Publishing, New York.
Raven, P., R. F. Evert, and H. Curtis. 1976. Biology of
plants. 2d ed. Worth Publishers, New York.
Bobbins, C. S., B. Bruun, and H. Zim. 1966. Birds of
North America. Western Publishing Co., New
York.
Welsh, S. L., and G. Moore. 1973. Utah plants:
Tracheophyta. Brighain Young University Press,
Provo, Utah.
DIFFERENTIAL HABITAT UTILIZATION BY THE SEXES OF MULE DEER
Michael M. King' and H. Duane Smith'
Abstract.- Habitat segregation trends have been observed and published for the sexes of mule deer {Odocoilem
hcmionus) based on elevation and slope exposure. Despite these brief descriptions, quantitative studies on habitat
segregation by the sexes of mule deer are lacking. Results of research conducted in central Utah indicated no signifi-
cant difference in elevation positions used by males, but did show significant difference in utilization of studv sites
based on slope exposure, relative percentage forb cover, and relative percentage hiding cover. Males were most com-
mon at sites characterized by low forb abundance and hiding cover, and on south-facing exposures. Females were
most common at sites characterized by high forb abundance and hiding cover, and on north-facing exposures. Pos-
sible advantages of habitat separation to both sexes and management implications are discu.ssed.
Mule deer {Odocoileus hemionus), like
many other ungulates, seem to exhibit habitat
partitioning between sexes (Darling 1937,
Estes 1974, Geist 1974, 1977, Gest and Pe-
tocz 1977, Hirth 1977, Leuthold 1978).
DeVos et al. (1967) indicated that male and
female mule deer are separated throughout
the year with the exception of the breeding
season. Dasmann and Taber (1956) found that
males occupied more open south-facing
slopes and females occupied densely vegeta-
ted north-facing slopes. Several workers have
suggested that males prefer higher altitudes
and ridge tops more than do females (Cowan
1956, Miller 1970). The same trend was ob-
served in a Nevada mule deer herd by Robin-
ette et al. (1977) where subalpine and alpine
conditions were prevalent. Males were found
predominantly above 3000 m elevation,
whereas females were more often below 2500
m elevation. Although habitat separation by
male and female mule deer has been report-
ed, little attempt has been made to quantify
differential habitat use or to describe site dif-
ferences other than to suggest slope exposure
and elevational differences.
The objectives of this study were: (1) to de-
termine quantitatively if male and female
mule deer differentially utilize habitat, (2) to
suggest possible advantages to habitat separa-
tion by sexes of mule deer, and (3) to identify
critical management problems related to dif-
ferential resource utilization between male
and female mule deer.
Study Area
The study was conducted on the Bighorn
Ranch, a privately owned ranch in the Nebo
Range of the Wasatch Mountains, Utah. The
study area was approximately 1130 ha in
size, ranging from 2200 to 2500 m elevation.
Human access is restricted, thus providing a
relatively undisturbed area for observation of
mule deer behavior, distribution, and habitat
utilization. Ridge tops, south-facing slopes,
and other well-drained areas were dominated
by Gambel oak {Quercus gamhelii) and big
sagebrush {Artemisia tridentata) communities
with little herbaceous growth. Drainage bot-
toms, north-facing slopes, and well-watered
areas were dominated by quaking aspen {Pop-
ulus tremuloides). Rocky Mountain maple
(Acer glabrum), and chokecherry {Prttnus vir
giniana) communities, with numerous forb
species in the understory.
Methods
Preliminary observations made in Septem-
ber 1977 to determine deer distribution in
the various watersheds of the ranch indicated
habitat segregation between male and female
mule deer. Based on that survey, the follow-
ing spring nine study sites (Fig. 1) were se-
lected where deer numbers were relatively
high. Other areas had equally as many deer,
but excessive area or distance from access
roads made observation unrealistic.
'Department of Zoology, Brigham Young University. Provo, Utah 84602.
273
274
Great Basin Naturalist
Vol. 40, No. 3
THE BIGHORN RANCH
O Study Sites
— Ranch Border
Nephi and
Fountain Green
Highway 89
N
Highway 89
Fountain Green
Fij;. 1, Map ot the Bit^honi Ranch showing the approximate location of study sites.
September 1980
King, Smith: Deer Habital Utilization
275
Each study site was observed weekly from
1 June 1978 to 1 September 1978. Observa-
tions were made from established observation
points or by vehicle from sunrise until late
morning or from early afternoon until dark.
Ob.servation time for each site was alternated
weekly between morning and evening so ap-
proximately equal observation time was
spent at each site during each time period. A
variable 15-45X spotting scope and 12X bi-
noculars were used for daytime observation,
and a 200,000 candlepower spotlight oper-
ated through the electrical system of the ve-
hicle, along with spotting scope and binocu-
lars, was used for observation at night. Total
observation time for the study exceeded 900
hours.
Observed deer were recorded according to
sex, slope position (Fig. 2), and slope expo-
sure. A 2X4 contingency analysis (Zar 1974)
was performed to determine significant dif-
ferences in utilization of slope positions by
males and females. Total numbers of males
and females recorded at north- and south-fac-
ing exposures were also subjected to contin-
gency analysis to determine slope exposure
usage differences. To characterize study sites
two critical parameters, relative abundance
of forb and hiding cover, were examined at
each site. Forbs were defined as succulent,
low-growing, nonwoody vegetation, and hid-
ing cover as vegetation more than 2 m in
height. Both estimates were determined by a
line-point transect method for determining
relative abundance of vegetation (Kershaw
1973).
Simple correlation procedures relating rel-
ative abundance of forb and hiding cover
with the corresponding male/female ratio for
each site were used to determine if utiliza-
tion of sites by males and females differed
significantly based on forb abundance and
hiding cover (Zar 1974). The maximum prob-
ability accepted for statistical significance
was 0.05; probabilities less than 0.01 were
considered highly significant.
Results and Discussion
Analysis of slope positions used by males
and females (Table 1, Fig. 3) showed no sig-
nificant difference in slope position utiliza-
tion by sex (P = 0.06). The data, though not
significant at the 0.05 level, approach signifi-
SLOPE POSITIONS
^^^^jM-i^-jt-^is^aN.
Fig. 2. Diagram of the four slope positions at each site; slope position 1 = canyon bottom to '4 slope, slope position
2=1/2 slope to '/2 slope, slope position 3 = '/2 slope to % slope, and slope position 4 = \ slope to ridgetop.
276
Great Basin Naturalist
Vol. 40, No. 3
Table 1. Total number of male and female mule deer at each slope position for all study sites. 2X4 contingency
analysis indicates a nonsignificant difference in utilization of slope position by male and female mule deer (X- = 7.42,
df = 3, P = 0.06). Numbers in parentheses are expected values.
Slope position
Sex
1
2
3
4
Total
Males
74
(86.9)
63
(56.4)
42
(39.9)
14
(9.8)
193
Females
157
(144.1)
87
(93.4)
64
(66.1)
12
(16.2)
320
Total
231
150
106
26
513
cance and indicate support for elevational
segregation observed in other areas. Further
categorical analysis to determine usage of in-
dividual slope positions by males and females
shows that considerable differences exist in
utilization of slope positions 1 and 4 by males
and females (Fienberg 1977), with more fe-
males than males at slope position 1, and
more males than females at slope position 4
(position 1 male = -.208, position 1 females
= + .208; position 4 males = -I- .245, posi-
tion 4 females = -.245; positive values in-
dicate most usage at slope position). Analysis
of slope exposure use by males and females
UJ
<
O
111
<
100 -I
75 -
^ 50 -
+ FEMALES
• MALES
Ik
O
O
o
u
85
25
T 1 r
1 2 3
SLOPE POSITION
Fig. 3. Percentage of total males and total females observed at each slope position.
September 1980
King, Smith: Deer Habital Utilization
277
showed a significant difference (P< 0.005),
with males most often at south-facing slopes
and females most often at north-facing slopes
(Table 2, Fig. 4).
Calculated male /female ratios for each site
correlated with corresponding relative per-
centages of forb and hiding cover (Table 3,
Fig. 5) showed a highly significant negative
correlation between forb abundance and
male/female ratios (r = -.89, df = 7,
P< 0.005) and a significant negative correla-
tion between male/female ratios and hiding
cover (r= -.69, df = 7, P<0.05). Therefore,
as forb and hiding cover increased, the
male/female ratios decreased, indicating that
females select areas characterized by rela-
tively high forb and hiding cover densities,
but males select areas characterized by low
forb and hiding cover densites.
Table 2. Total number of male and female mule deer
at each slope exposure for all study sites. 2X2 contin-
gency analysis indicates a significant difference in utili-
zation of slope exposure bv male and female mule deer
(X^ = 29.3, df = 1, P< 0.005). Numbers in parentheses are
expected values.
Slope exposure
Sex
North-facing
South-faciiiu
T..i,,l
Males
64
(93.7)
129
(99.3)
193
Females
185
(155.3)
135
(164.7)
320
Total
249
264
513
To comprehend implications of this pat-
tern of spatial separation, po.ssible advantages
accrued by males and females in their spring-
summer habitats should be examined. During
the spring-summer season perhaps the most
critical events to females are production and
100 -1
3
o
Q.
X
Q.
o
<
<
ui
<
z
<
UJ
<
75 -
50 -
67
33
25 -
NORTH-FACING
EXPOSURE
SOUTH-FACING
EXPOSURE
MALES
FEMALES
Fig. 4. Percentage of total males and total females observed at each slope exposure.
278
Great Basin Naturalist
Vol. 40, No. 3
§
U
flO
O
Ik
>
<
Ul
>
O
o
z
o
z
Ul
>
50 -
40 -
30 -
20 -
10 -
r= -.89
P<0.005
50 -
40 -
= 30 -
20 -
10 -
»'=-.69
P<0.05
T"
1.0
2.0
1^
3.0
"T"
4.0
male/female ratios
Fig. 5. Correlation of relative percentage fort) cover and relative percentage hiding cover with corresponding
male/female ratio for each study site.
September 1980
King, Smith: Deer Habital Utilization
279
Table 3. Relative percentage forb cover, relative percentage hiding cover, total number of male and female mule
deer, and male/female ratios for each study site.
Site No.
Relative
% forb cover
Relative
hiding cover
Males
Females
M/F ratios
40.50
19.00
7
63
0.11
32.25
14.00
6
39
0.15
41.25
19.(X)
12
61
0.20
46.50
22.(X)
2
25
0.08
37.75
16.00
7
46
0.16
16.50
15.00
52
29
1.80
19.00
13.00
32
23
1.40
20.75
8.00
38
15
2.53
19.75
17.00
37
19
1.95
rearing of offspring. This implies that consid-
erable energy is apportioned to gestation,
parturition, and lactation beyond normal
body maintenance requirements (Nelson
1975, Stebbins 1977). If these requirements
are not met through diet resources, body re-
serves are utilized, thus reducing offspring
vigor and survival. Since energy demands for
offspring production exceed normal energy
requirements, it is important that females oc-
cupy areas where nutritious, high-quality for-
age is readily available. Adequate nutrition
insures successful offspring production and
facilitates proper lactation. Research in-
dicates that, during the time period critical
to fawn production, high moisture content,
ease of digestion, and increased nutrition
content (Smith 1952, Short 1966, Short and
Reagor 1970, Boeker et al. 1972) make the
forbs preferred diet items of mule deer
(Smith 1952, Morris and Schwartz 1957, Lo-
vass 1958, Anderson et al. 1965, Crouch
1966, Dasmann et al. 1967, Nelson 1975). It
has also been shown that deer herds having a
variety of succulent forage in their diets have
greater herd productivity and vigor than
those that utilize a greater percentage of
woody vegetation (Biswell 1961, Julander et
al. 1961, Boeker et al. 1972, Nelson 1975, Pe-
derson and Harper 1978). We assume, there-
fore, that forb abundance is related to site
quality and that areas of high forb abundance
are considered high quality, whereas areas of
low forb abundance are low quality. Deer
that select forb-rich areas would have survi-
val advantages because of availability of
choice forage. It follows that females should
select high-quality sites to assure adequate
nutrition and energy for fawn production and
survival. However, it seems strange that
males would occupy areas of low forb abun-
dance when selection of high-forb areas
would more readily insure adequate energy
for increasing body size, accumulating fat re-
serves, and developing antlers. We suggest as
a partial explanation that it is more advanta-
geous for the sexes to be separated to reduce
chances of energy-expensive agonistic expres-
sion between males and females (Geist and
Petocz 1977, McCullough 1979). This allows
energy allocation to gestation, parturition,
and lactation rather than to stresses of haras.s-
ment. Females on ranges uncontested by
males should leave more offspring, and those
behavioral traits responsible for habitat parti-
tioning should be selected to increase the
population. Males that did not compete for
resources necessary for fawn production,
though occupying lower-quality sites, should
likewise leave more offspring to succeeding
generations than males whose behavior bring
them into competition with their offspring
(Wilson 1975, Geist and Petocz 1977,
McCullough 1979).
Differential habitat use patterns could also
confer advantages to males and females by
increasing odds for predator avoidance. Ecol-
ogists working on vmgulates have suggested a
theory of predator avoidance based on the
relative degree of habitat openness and group
size. Ungulates that inhabit densely vegetated
areas usually occur as solitary animals or in
small groups that use hiding as a mechanism
for predator avoidance, whereas ungulates
that occupy open areas are primarily herding
animals that rely on the use of collective
senses and high mobility for predator escape
280
Great Basin Naturalist
Vol. 40, No. 3
(Dasmann and Taber 1956, Kitchen 1974,
Hirth 1977).
Life history studies of mule deer (Linsdale
and Tomich 1953, Robinette et al. 1977),
along with personal observations, give sup-
portive evidence to this hypothesis. Males
and females form different-sized groups in
their preferred habitats. During spring and
summer months females seek isolation in
areas where hiding cover is relatively abun-
dant (Fig. 5) and tolerate few deer other than
their offspring of the year. This partial. soli-
tary existence has advantages from a predator
avoidance standpoint in that females can se-
cret themselves and their offspring in dense
vegetation during periods of high vulnera-
bility to potential danger. After the young
are bom, maternal duties restrict the mobil-
ity and escape efficiency of females. It
would, therefore, be advantageous for fe-
males with fawns to avoid open habitats
where predators can detect and capture them
or their fawns more easily.
In contrast, males on the Bighorn Ranch
during the same time period were often ob-
served to form fraternal groups in areas with
relatively low abundance of hiding cover
(Fig. 5). Male groups that inhabit open areas
of high visibility can collectively monitor
their surroundings and take advantage of rap-
id flight when escape is necessary. This is
better than hiding in dense cover, where ef-
fectiveness of concealment would be reduced
by large groups and would increase chances
of detection by predators.
Furtlier supportive evidence for the pre-
dator avoidance theory is provided by the
differential use of slope positions 1 and 4 by
males and females. More females than males
occupied slope position 1, the lower position
of the slope where the greatest abundance of
hiding cover was located. This suggests the
importance of cover to females. More males
than females used slope position 4, the open
ridges, suggesting preference by males for the
areas of high visibility.
In this study habitat separation by males
and females is primarily based on slope expo-
sure, forb abundance, and hiding cover, with
some evidence, though not significant, for
elevational segregation. We do not detract
from the importance of elevational segrega-
tion as it has been observed frequently in
other areas, but suggest to wildlife managers
that there are several habitat separation pos-
sibilities, depending on characteristics of
mule deer range in a given area. We encour-
age that further research delineating segrega-
tion characteristics, advantages, and mecha-
nisms be initiated so that management
implications can be evaluated.
Knowledge of habitat separation between
the sexes of mule deer will have considerable
influence on several critical management
problems. Three important problems as we
view them are now discussed. First, as deer
populations are being censused in various
areas and habitats, a prime concern is the de-
termination of an accurate sex ratio. If man-
agers are not aware of site-specific habitat
separation by male and female deer, biases
favoring one sex over the other will arise in
calculated ratios depending on the area sam-
pled. Failure to determine accurate sex ratios
will allow faulty plans to be devised and im-
plemented. Second, through recognition of
specific habitat requirements of the sexes, it
is possible that habitat can be manipulated
through appropriate techniques to create
conditions favorable to either sex. This will
allow sex ratio manipulation depending on
management needs. Third, critical areas to
females and offspring as well as males must
be protected from detrimental commercial,
industrial, and recreational development. De-
struction of important fawning areas through
development will force females to occupy
suboptimal habitats and result in reduced
fawn production and survival. Development
in areas occupied predominantly by males
will restrict fall hunting and ultimately re-
duce herd productivity if adequate
male/female ratios are not maintained.
An understanding of how male and female
mule deer partition the habitant and how hab-
itats preferred by females differ from those
most frequented by males will undoubtedly
improve abilities to effectively manage mule
deer habitat. Proper use of knowledge re-
garding differential habitat and resource uti-
lization by the sexes of mule deer can in-
crease the efficiency with which agencies
manage the deer resource.
September 1980
King, Smith: Deer Habital Utilization
281
Literature Cited
Anderson, A. E., W. A. Snyder, and G. W. Brown.
1965. Stomach content analysis related to condi-
tion in mule deer, Guadalupe Mountains, New
Mexico. J. VVildl. Manage. 29: .352-.366.
BiswELL, H. H. 1961. Manipulation of chamise brvi.sh for
deer range improvement. California Fish and
Game 47:128-144.
BoEKER, E. L., V. E. Scott, H. G. Reynolds, a.nd B. A.
Donaldson. 1972. Seasonal food habits of mule
deer in southwestern New Mexico. J. Wildl.
Manage. .36:56-6.3.
Cowan, I. M. 1956. Life and times of the coast black-
tailed deer. Pages 52.3-617 in W. P. Tavlor, ed..
The deer of North America. Harrishurg: Stack-
pole Co. 668 pp.
Crouch, G. L. 1966. Preferences of black-tailed deer for
native forage and Douglas-fir seedlings. J. Wildl.
Manage. .30:471-475.
Darling, F. F. 1937. A herd of red deer. London: Ox-
ford University Press. 215 pp.
Dasmann, R. F., and R. D. Taber. 1956. The behavior
of Columbian black-tailed deer with reference to
population ecology. J. Mammal. 37:143-164.
Dasmann, W., R. Hubb.\rd, VV. G. MacGrecor, and A.
E. Smith. 1967. Evaluation of the wildlife results
from fuel breaks, browsewavs, and tvpe con-
versions. Proc. 7th Tall Timbers Fire Ecol. Conf.
7:179-193.
Devos, a., P. Brokx, and V. Geist. 1967. A review of
social behavior of North American cervids during
the reproductive period. Amer. Midi. Natur.
77:390-417.
Estes, R. D. 1974. Social organization of the .\frican bo-
vids. Pages 166-205 in V. Geist and F. Walther,
eds.. The behavior of ungulates and its relation to
management. Vol. 1. Morges, Switzerland: lUCN
Publ. No. 24. 511 pp.
FiENBERG, S. E. 1977. The analysis of cross-classified cat-
egorical data. Cambridge: MIT Press. 151 pp.
Geist, V. 1974. On the relationship of social evolution
and ecologv in ungulates. .\mer. Zool.
11:205-220. '
1977. A comparison of social adaptations in rela-
tion to ecology in gallinaceous bird and ungulate
societies. Ann. Rev. Ecol. Syst. 8:19.3-207.
Geist, V., and R. G. Petroczy. 1977. Bighorn sheep in
winter: do rams maximize reproductive fitness bv
spatial and habitat segregation from ewes? Can.
J. Zool. ,55:1802-1810.
Hirth, D. H. 1977. Social behavior of white-tailed deer
in relation to habitat. Wildl. Monogr. 53:1-55.
Julander, O., W. L. Robinette, and D. A. Jones. 1961.
Relation of summer range condition to mule deer
herd productivity. J. Wildl. Manage. 25:54-60.
Kershaw, K. A. 1973. Quantitative and dynamic plant
ecology. 2d ed. New York: American Elsevier
Publishing Co. Inc. .308 pp.
Kitchen, D. W. 1974. The social behavior and ecolog>'
of the pronghorn. Wildl. Monogr. ,38:1-96.
Leuthold, W. 1978. .\frican ungulates: a comparative
review of their ethology and behavioral ecology.
New York: Springer- Verlag. .307 pp.
Linsdale, J. M., and p. Q. Tomich. 1953. A herd of
mule deer. Berkeley: University of California
Press. 567 pp.
LovASs, A. L. 1958. Mule deer food habits and range
use. Little Belt Mountains, Montana. J. Wildl.
Manage. 22: 275-283.
McCullough, D. R. 1979. The George Reserve deer
herd. Ann Arbor: University of Michigan Press.
271 pp.
Miller, F. L. 1970. Di.stribution pattern of black-tailed
deer in relation to enviromneut. J. Mammal. .52:
248-259.
Morris, M. S., and J. E. SvHwartz. 1957. Mule deer
and elk food habits on the National Bison Range.
J. Wildl. Manage. 21:18.3-189.
Nelson, J. R. 1975. Forest fire and big game in the Pa-
cific .Northwest. Proc. 15th Tall Timbers Fire
Ecology Conf. 15:85-102.
Pederson, J. C, AND K. T. Harper. 1978. Factors in-
fluencing productivity of two mule deer herds in
Utah. J. Range Manage. 31:105-110.
Robinette, W. L., N. V. Hancock, and D. A. Jones.
1977. The Oak Creek mule deer herd in Utah.
Salt Lake Citv: Utah State Div, Wildl. Res. Publ.
No. 77-15. 148 pp.
Short, H. L. 1966. Effects of cellulose levels on appar-
ent digestability of feeds eaten bv mule deer. J.
Wildl. Manage.' .30: 163- 167.
Short, H. L., and J. G. Reagor. 1970. Cell wall diges-
tability affects forage value of woody twigs. J.
Wildl. Manage .34:964-967.
Smith, A. D. 1952. Digestability of some native forages
for mule deer. J. Wildl. .Manage. 16:.309-,312.
Smith, J. G. 1952. Food habits of mule deer in Utah. J.
Wildl. Manage. 16:148-155.
Stebbins, L. L. 1977. Energy requirements during repro-
duction of Peromyscus tnanictilattts. Canadian J.
Zool. 55:1701-1704.
Wilson, E. O. 1975. ScxiobiologV'. Cambridge: Belknap
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Zar, J. H. 1974. Biostatistical analysis. Englewood Cliffs:
Prentice-Hall Inc. 620 pp.
TEMPORAL ACTIVITY PATTERNS OF A DIPODOMYS ORDII POPULATION
Clive D. Jorgensen,' H. Diiane Smith,', and James R. Garcia'
Abstract.— Temporal activity patterns for Dipodomijs ordii were generally bimodal during the summer, with the
highest peak occurring during early predawn hours when conditions were optimum for water conservation. Removal
of dominant members in the population resulted in a substantial shift in the activity pattern to increased activity
during the evening hours.
Ecologists studying small mammals must
contimially attempt to resolve the problems
of inadequate methods to study daily activity
patterns in natural environments, especially
when studying secretive and/ or nocturnal
species that require trapping procedures.
Usefulness of temporal activity data has been
demonstrated in simulation trapping studies
(Burnham and Overton 1969, Manly 1970,
Jorgensen et al. 1972), population estimator
studies (Scott et al. 1978), and energetics
studies (Kenagy 1973), among others. Al-
though methods for obtaining data under
field conditions have not been well devel-
oped, some have been reported (Jorgensen
and Hayward 1965, Eider 1968, Marten
1973). Generally, their results are deficient in
one or more of the following: (1) numbers of
recorded activity events per day, (2) timing
of the observed or measured activity event,
(3) ability to assign an activity event to a spe-
cific individual, and (4) correlation between
the measured activity under laboratory con-
ditions with analogous activity in natural en-
vironments.
Harling (1971) attempted to relieve some
of the difficulty in recording activity by de-
veloping a trap that could be continually
monitored during a trapping period. His
traps were electrically wired to a central
communications console. Using walkie-talkie
communications between someone on the
grid and another at the console, Harling
(1971) was able to obtain the precise time
when an animal was caught as well as when
it was released.
We extended Harling's (1971) methods to
include an entire grid of traps comparably
wired (Garcia et al. 1974) to study a popu-
'Department of Zoology. Brigham Voting University, Provo, Utah 84602.
lation of Dipodomys ordii under field condi-
tions. Our objectives were to determine the
optimum foraging times for D. ordii in the
salt desert shrub community of west-central
Utah, and illustrate activity patterns that
could assist in interpreting trapping data
from other studies that include this species.
Study Site and Methods
Data were collected from two sites at the
Desert Range Experiment Station, Millard
Co., Utah, from 21 August to 3 September
1971 and 25 June to 27 August 1973. Al-
though D. ordii was the species trapped most
frequently during this study, Perognathus
longimernbris was common and Peromyscus
maniculatus and Onychomys leucogaster
were trapped occasionally. Both sites were
sandy and dominated by Oryzopsis hyme-
noides, Chrysothamnus nauseosus, and Sal-
sola kali, although Ambrosia acanthicarpa.
Astragalus spp., Atriplex canescens, Gilia
hiitchinsifolia, and Hilaria jamesii also were
present.
The trap design and surveillance methods
were described in detail by Garcia et al.
(1974). A 10 X 10 (100 traps) grid was wired
to a central communications console, where
one researcher recorded traps as they were
"set off." He then informed an assistant
working on the grid where trapped animals
were. Animal data were radioed from the as-
sistant to the researcher at the console by
walkie-talkie. Data were then recorded and
the trap reset. Animals seldom were detained
in traps for more than a few minutes. Data
collected in our study included: species, rela-
tive age (juvenile, subadult, adult), sex, repro-
282
September 1980 Jorgensen et al.: Dipodomys Activity Patterns
283
ductive condition, and identification mark
(toe clip). Supportive data, used as the inde-
pendent variables in our analyses and collect-
ed each time an animal was captured, were:
ambient temperature, soil temperature, wind
speed, cloud cover and lunar events (sub-
jectively assessed), light intensity, barometric
pressure, and relative humidity. Absolute hu-
midities (gm/m^) were determined using the
methods described by Piatt and Griffiths
(1964), i.e., X = 217(RH) (^'J/IOOT, where T
is degrees Kelvin and e^ is vapor pressures in
air saturated with water.
The period between sundown and sunrise
was divided into 20 subperiods, which were
used as the time units while plotting activity.
Since activity periods changed slightly as day
lengths changed, it was necessary to establish
a standard set of subperiods before data for
different periods could be pooled. Stepwise
regression methods were used to assess the ef-
fects of independent variables on activity
rates among the pooled data for all activity
subperiods. Differences in activity among
sexes, ages, and reproductive condition class-
es were determined using Chi-square tests of
independence.
An opportunity to assess possible effects of
socially dominant individuals in the D. ordii
population was noted after three weeks of ac-
tivity data had been gathered in 1973. An an-
imal was considered dominant if it was the
only adult captured within the area pre-
scribed when its capture points had been
connected, or if it was the only adult repeat-
edly caught in a specific trap. Eleven domi-
nant individuals among the 42 individuals re-
corded on the grid were removed from the
poulation and activity data gathered for an
additional five days. Temporal activitv pat-
terns of the populations before and after the
selected removals were compared.
Results and Discussion
Activity for the intact population of D. or-
dii was essentially bimodal, with the greatest
activity occurring during the predawn hours
c
0)
u
0)
Q.
9.0 n
so-
ld
T3
•i 70 H
a;
Q.
"I 6.0
CD
0)
5.0-
" 4.0-1
<
3.0
2.0-
1.0 -
Dipodomys ordii (Dominants included)
Dipodomys ordii (Dominants removed )
I I I ■ ■
7 9 11 13 15
Activity Subperiods
17
19
21
Fig. 1. Activity patterns for Dipodomys ordii pooled among 20 subperiods over a 24-hoiir activity period.
284
Great Basin Naturalist
Vol. 40, No. 3
(Fig. 1). These observations agree with Jor-
gensen and Hayward (1965), although our
early evening peak is not as distinct. Since
their data were gathered over the entire year
from trapping rates per hour, we question
whether their results are directly comparable
with ours. Although we have no season-spe-
cific data to demonstrate seasonal variations
in activity patterns, any variation would pro-
duce composite patterns difficult to interpret
and compare.
Precipitation (11.4 cm) was unusually low
in 1971 and rather high (20.5 cm) in 1973.
I
0
Q.
a
o
</)
o
E
'E
<
These differences in moisture were accom-
panied by comparable differences in vegeta-
tive production— low in 1971 and high in
1973 (Jorgensen, unpublished data). Chi-
square analyses resulted in only 6 of a pos-
sible 20 tests being significant (p = .95). All
the 6 cohorts of animal classes that were sig-
nificant included adults (Table 1). The ober-
vations suggest that adults are more sensitive
to seasonal changes in precipitation and veg-
etative production than immature classes
(Fig. 2). The apparent difference was a
change from reduced activity of fewer adults
Sunset
Time
Sunrise
Fig. 2. 1971 and 1973 activity patterns for adult Dipodonujs orclii pooled among 20 subperiods over a 24-hour
activity period.
September 1980 Jorgensen et al.: Dipodomys Activity Patterns
285
Table 1. Significantly independent (p = .05) values of
Chi-square tests for Dipodontys ordii activity patterns
(1971 and 1973).
Variable contrasts
d.f.
X2
1971 vs. 1973 (all classes combined)
19
131.37
1971 vs. 1973 (males)
19
95.66
1971 vs. 1973 (females)
19
.51.41
1971 vs. 1973 (sexuallv active males)
18
32.48
1971 vs. 1973 (sexuallv inactive males)
18
68.29
1971 vs. 1973 (all adults)
19
117.41
around a relatively con.stant level in 1971 to
a higher level of activity for more animals in
the predawn hours of 1973, a pattern that
persisted when all 1973 observations were
pooled (Fig. 1).
The activity pattern with all animals still
in the population (Fig. 1) was then examined
to determine the effects independent vari-
ables may have had on it. Schmidt-Nielson
(1964) reported that Dipodomys merriami
was most active at low ambient temper-
atures, high relative humidity, and high abso-
lute humidity. In 1973, we found that activi-
ty increased as temperature decrea.sed and
relative humidity increased, a condition that,
along with an increase in absolute humidity,
occurred most frequently in the predawn
hours. Dipodomys ordii followed the same ac-
tivity pattern reported for D. merriami, a be-
havior reported to maximize the conservation
of water (Schmidt-Nielson 1964) or accom-
modate temporal competition with Dipo-
domys microps (Kenagy 1973), where their
distributions overlap. This, coupled with in-
creased activity as wind speeds decrease be-
low 3.2 km/hr and after periods of rain, sug-
gests that changes in activity may be related
to water conservation.
An additional 40 animals were marked and
their activity monitored for five days after 11
dominant animals had been removed from
the grid. All except 8 (most of which were
trapped near the grid border) of these 40 new
animals were juveniles and subadults. The
pooled activity pattern shifted to develop a
peak in the early evening hours and generally
declin ed thereafter (Fig. 1). Early evening ac-
tivity exposed the animals to less than optim-
al conditions for water conservation, but
seeds were more abundant on the soil surface
because of natural seed-drop and accumula-
tion during the day. Seeds of Oryzopsis
hymenoides were dropped to the ground in
rather large numbers while these data were
being gathered. The change in activity may
reflect the release from domination, early at-
tempts to reestablish social dominance, or
perhaps inexperience among the numerous
immature members of the population.
The optimum period of activity above-
ground during summer months was during
predawn hours when conditions for water
conservation were enhanced. Activity pat-
terns of the intact population of D. ordii were
highest during this predawn period of time.
When the .social order was disrupted in 1971
due to low rainfall and low vegetative pro-
duction and in 1973 when the dominant indi-
viduals were removed from the population,
activity was highest during less optimum
conditions for water conservation, but more
nearly optimal for seed availabilitv.
Terrell Johnson (1979) found that Per-
omyscus maniculatus is more efficient in har-
vesting seeds buried up to 0.25 in. in sand
when the moisture content of the sand is in-
creased. This ob.servation lends credence to
the harvesting strategies demonstrated by the
activity pattern of the intact D. ordii popu-
lation, since their peak activity occurred
when moisture was highest. Paradoxically, O.
hymenoides seeds were most abundant in the
early evening after thev had dropped from
the plants during the day. Rodents apparent-
ly selected harvesting times that either max-
imized seed availability or water con-
servation. From our data, it appears that
immature D. ordii were most active when
seeds were most abundant and adults tended
to optimize water conservation.
Since seed-drop is ephemeral and occurs
when voung animals are most abundant in
the population, strategies of D. ordii for sur-
vival until home ranges can be established
.seem enhanced by early temporal activity.
Annual replacement of older, often non-
reproductive adults that have established and
control home ranges within the population is
important to population survival, since dry
years with little or no reproduction are not
uncommon. Replacement of the breeding
population would facilitate survival of adults
during nonproductive years, until reproduc-
tion is again feasible. Replacement is likely
286
Great Basin Naturalist
Vol. 40, No. 3
only if the younger animals obtain a com-
petitive advantage from some source other
than size. This might be provided by the
"flush" of energy available to preadults that
feed during the evening hours when harvest
of high quality energy is optimized during
years with high seed production. This strate-
gy encourages the infusion of young animals
into the population and allows genetic fix-
ation of the activity patterns that optimize
the likelihood of survival for D. ordii until
the next breeding opportunity.
Literature Cited
BiDER, J. B 1968. Animal activity in uncontrolled terres-
trial communities as determined by a sand tran-
sect technique. Ecol. Monogr. 38:269-308.
BuRNHAM, K. P., .AND W. S. OvERTON. 1969. A simulation
studv of live trapping and estimation of popu-
lation size. Tech. Kept. No. 14, Oregon State
University, Corvallis, 152 pp.
G.\RciA, J. R., H. D. Smith, and C. D. Jorgensen. 1974.
A capture-release method for determining small
mammal activity. Proc. Utah .\cad. Sci., Arts and
Letters 51:1-11.
Harling, J. 1971. A technique for precisely timing cap-
tures of Perornysctts manicuhitus. Canadian J.
Zool. 48:1275-1277.
Johnson, T. K. 1979. Ability of desert rodents to find
buried seeds in desert range communities. Un-
published thesis. Brigham Young University, Pro-
vo, Utah. 23 pp.
Jorgensen, C. D., and C. L. Hayward. 1965. Mammals
of the Nevada Test Site. Brigham Young Univer-
sity Sci. Bull., Biol. Ser. 6(3): 1-81.
Jorgensen, C. D., D. T. Scott, and H. D. Smith. 1972.
Small mammal trapping simulator. Proc. 1972
Summer Computer Simulation Conf.: 1154-1168.
Ken AG Y, G. J. 1973. Daily and seasonal patterns of activ-
ity and energetics in a heteromvid rodent com-
munity. Ecology 54:1201-1219.
Manly, B. F. J. 1970. A simulation study of animal pop-
ulation estimation, using the capture-recapture
method. Jour. Appl. Ecol. 71:13-39.
Marten, G. G. 1973. Time patterns of Perornysctts activ-
ity and their correlation with weather. J. Mam-
mal. 54:169-188.
Platt, R. B., and J. R. Griffiths. 1964. Environmental
measurement and interpretation. Reinhold Publ.
Co., New York. 235 pp.
ScHMiDT-NiELSON, K. 1964. Descrt animals: phys-
iological problems of heat and water. Clarendon
Press, Oxford, England. 277 pp.
Scott, D. T., C. D. Jorgensen, and H. D. Smith. 1978.
Comparison of live and removal methods to esti-
mate small mammal densities. Acta Theriol.
23:173-193.
NEW RECORDS OF WESTERN TRICHOPTERA WITH NOTES ON THEIR BIOLOGY'
Bernard G. Swegniau" and Leonard C. Ferrington, Jr.-
Abstract.- Western records for 27 species of Trichoptera are given; a majority of the records are from the Bear-
tooth Mountains of northwestern Wyoming. In addition, examples of variation in the male genitiiia of Limnephilus
coloradensis (Banks) are figured and the female is described. Some comments regarding the larvae of Allomyia
(Inuinia) are pre,sented.
Twenty-two species of Trichoptera are re-
ported from Wyoming and Montana, includ-
ing 20 from the immediate vicinity of the
Beartooth Mountains, where the University
of Pittsburgh's Pymatuning Laboratory of
Ecology has offered a summer field course in
Alpine Ecology. In addition, 7 species were
collected near a pond at the Winding River
Campground near Rocky Mountain National
Park, Grand County, Colorado. Within the
Beartooth Mountains, immatures of many
common limnephilid genera were collected,
but the species remain unknown to us. Some
of the more common genera taken include:
Discoinoecus, Hesperophylax, Homophijlax,
Lenarchus, Limnephilus, and Psychoghjpho.
Considerable variability was observed in
the male genitalia of Limnephilus colorad-
ensis (Banks). Examples of this variation are
illustrated. Further, in copula specimens of
this species were collected, thereby making
possible the identification of the female that
is described and figured.
Collecting Sites
The following sites, with the exception of
the Grow Ventre and Winding River loca-
tions, are within the Beartooth Mountains of
the Absaroka range of northwestern (Park
Co.) Wyoming. The Star Lake site lies in
Park Co., Montana.
Inlet Run.— Elevation approximated 3140
m, 109° 29' W, 44° 58' N. The sample site is
on the more easterly of two first order, snow
melt streams that flow into Frozen Lake. A
majority of the specimens were collected
during a series of diel drift studies; however,
'Support for this paper has been provided by the Pymatuning Laboratory of Ecology.
■University of Pittsburgh and Pymatuning Laboratory of Ecology. Pittsburgh. Pennsylvania 15260.
occasional specimens were collected by
sweeping low vegetation or were picked
from rocks.
Frozen Lake.— Elevation 3070 m; same lo-
cation as above. Specimens were collected by
sweeping vegetation or were collected from
rocks in 1 m or less of water.
Chain Lakes.- Elevation 2880 m, 109° 31'
W, 44° 55' N. The sample .site was at the
southeast shore of lower Chain Lake near the
point where the stream draining Fantan Lake
enters. An extensive alpine meadow sur-
rounds the lake.
Sawtooth Lake.- Elevation 2835 m, 109°
28' W, 44° 54' N. The sampling site was on
an unnamed second order stream that drains
the western slope of Sawtooth Mountain.
This stream drains open alpine meadow and
flows into the ea.stern edge of Sawtooth Lake.
The specimens were collected from exposed
and submerged rocks.
Beartooth Butte.— Elevation approximatelv
2910 m, 109° 37' W, 45° 57' \. The sample
site is a large spring that originates on the
south face of Beartooth Butte at the base of
an open talus slope. The stream flows
through open meadow and enters Beartooth
Lake on its western shore at approximately
2710 m elevation.
Moose Bog.- Elevation 2740 m, 109° 37'
W, 44° 56' N. This bog is approximately 30
m south of U.S. Route 212, just ea.st of the
point where the gravel road to Clay Butte
lookout station begins. The specimens were
collected by sweeping the bog vegetation on
the Sphagnum mat near the largest area of
open water.
287
288
Great Basin Naturalist
Vol. 40, No. 3
Ghost Creek.- Elevation 2650 m, 109° 37'
W, 44° 56' N. The sample site was near the
point where the gravel road to the University
of Pittsburgh's research trailer crosses the
stream. The stream is second order and drains
intermittent patches of alpine meadows and
coniferous forest. Most specimens were col-
lected during diel drift studies; however,
some specimens were taken by sweeping veg-
etation.
Clark's Fork of Yellowstone River — Eleva-
tion 2060 m, 209° 41' W, 44° 51' N. The riv-
er at this point is fifth order and flows south-
east, draining mixed coniferous forest and
pasture lands.
Star Lafce.- Elevation 2940 m, 109° 55'
W, 45° 6' N. The sample site was located in
the stream just east of the outlet from Star
Lake. The stream width is approximately 2 m
at this point. This site is located in Montana.
Gros Ventre.- Elevation 1980 m, 110° 40'
W, 43° 36' N. The sample site was located in
Gros Ventre Campground approximately 15
miles northeast of Jackson Hole, Wyoming.
The specimens were picked from the win-
dows of the campground restrooms.
Winding River.- Elevation 2590 m, 105°
53'W, 40° 16' N. This sample site was lo-
cated just west of the entrance to Rocky
Mountain National Park, Colorado, in the
Winding River Campground. Specimens
were collected by sweeping the emergent
vegetation of a small pond and were picked
from the windows of the campground rest-
rooms.
Species Collected
Rhyacophilidae
Since less than 10 percent of the western
Rhyacophilia immatures are known (Flint, in
litt.), the following identifications are only
tentative. All identifications are based on our
use of Smith (1968).
Rhyacophila acropedes Banks.— Wyoming
(Park Co.): Ghost Creek, 14 August 1974, 10
larvae, and 15 August 1974, 7 larvae, collect-
ed L. Ferrington. 15 August 1974, 1 larva,
collected G. Goetz.
Rhyacophila hyalinata Banks.— Wyoming
(Park" Co.): Ghost Creek, 19 August 1977, 3
larvae, collected D. Ferrington. The ventral
surface of the head in these specimens is
darkened, but on the basis of distribution are
assigned to hyalinata rather than vocala
(Smith 1968).
Rhyacophila tucula Ross.— Wyoming
(Park Co.): Ghost Creek, 14 August 1974, 1
larva, 26 August 1978, 11 larvae, 19 August
1977, 3 prepupae. Sawtooth Lake vicinity, 23
July 1975, 1 larva; Inlet Run, 21 July 1978, 1
larva. Montana (Park Co.): Outlet of Star
Lake, 25 July 1975, 4 larvae. All by D. and L.
Ferrington.
Rhyacophila vacciia Milne.— Wyoming
(Park Co.): Sawtooth Lake vicinity, 23 July
1975, 2 larvae, D. Ferrington.
Rhyacophila vagrita Milne.— Wyoming
(Park Co.): Ghost Creek, 14 August 1974, 1
larva, L. Ferrington.
Rhyacophila verulla Milne.— Wyoming
(Park Co.): Ghost Creek, 26 August 1978, 2
larvae, 19 August 1977, 1 larva; Sawtooth
Lake vicinity, 23 July 1975, 4 larvae; Inlet
Run, 21 July 1978, 1 larva. Montana (Park
Co.): Outlet of Star Lake, 25 July 1975, 1 lar-
va. All by D. and L. Ferrington.
Glossosomatidae
Glossoma vebna Ross.— Colorado (Grand
Co.): Winding River Campground, 2 August
1978, 1 male, collected D. Ferrington.
Hydropsychidae
Arctopsyche grandis Banks.— Wyoming
(Park Co.): Clark's Fork Yellowstone River,
23 July 1978, 22 larvae, 6 pupae (1 pharate
male), 6 males, D. Ferrington.
Hydropsyche oslari Banks.— Wyoming (Te-
ton Co.): Gros Ventre Campground, 29 July
1978, 4 males, 26 females, D. Ferrington.
Hydroptilidae
Stactobiella delira Ross.— Colorado (Grand
Co.): Winding River Campground, 2 August
1978, 2 males, 1 female, D. Ferrington.
Limnephlidae
Anabolia bimaculata Walker.— Colorado
(Grand Co.): Winding River Campground, 2
August 1978, 1 male, D. Ferrington.
September 1980
SwEGMAN, Ferbington: Western Trichoptera
289
Chyranda centralis Banks.— Wyoming
(Park Co.): Ghost Creek, 27 July 1978, 1 fe-
male, collected L. Ferrington, identified A.
Nimmo.
Dicosmoecus gilvipes Hagen.— Wyoming
(Park Co.): Spring at base of Beartooth Butte,
15 August 1974, 1 female, G. Goetz. Uniden-
tified larvae of Dicosmoecus have also been
collected in large numbers from Inlet Run.
Ecclisomijia conspersa Banks.— Wyoming
(Park Co.): Inlet Run, 22 July 1978, 1 male,
L. Ferrington.
Ecclisomijia maculosa Banks.— Wyoming
(Park Co.): Spring at base of Beartooth Butte,
15 August 1974, 4 pupae (2 pharate males, 2
females), G. Goetz; Ghost Creek, 26 July
1978, 3 males, 27 July 1978, 3 males, 4 fe-
males, L. Ferrington. Frozen Lake vicinitv,
22 July 1978, 43 larvae, D. Ferrington. The
larval records are a tentative assignment
based on proximity of sites where adults were
taken. However, the single record of E. con-
spersa indicates two species occur in this
area.
Allomyia bifosa (Ross.)— Wyoming (Park
Co.): Inlet Run, 21-22 July 1978, 13 males, 2
females, 69 pupae (51 pharate males, 18 pha-
rate females). In addition to these adult-pupal
records, larval records of Allomyia include;
Ghost Creek, 14 August 1974, 1 larva, G.
Goetz, 14 August 1974, 8 larvae, and 15
August 1974, 1 larva, L. Ferrington, 18 Au-
gust 1977, 2 larvae, R. Seward, 19 August
1977, 5 larvae, and 26 August 1978, 1 larva,
L. Ferrington; Inlet Run, 10-11 August 1977,
89 larvae, 17-18 August 1977, 31 larvae, 21
July 1978, 4 larvae, all by L. Ferrington. Two
types of larvae were present in these collec-
tions. One type is without the flattened head
and pronounced carina, as in Allomyia
(Imania) scotti Wiggins (Wiggins 1977, Fig.
10.28). The second type has this character
and also possesses single gills dorsallv and
ventrally on segments 2, 3, 4, and 5. Wiggins
(pers. comm.) suggests that two species are
involved. He has associated gilled larvae with
Allomyia tripunctata Banks, which is known
from Wyoming. Adults of A. tripunctata,
however, have yet to be collected in the
Beartooth Mountains. These gilled larvae
were compared to gilled larvae of Allomyia
collected 17 July 1975 at St. Vrain Creek
above Peaceful Valley, Boulder Countv, Col-
orado by Dr. J. V. Ward. The specimens from
the two localities appear to be distinct, sug-
gesting that at least two species of Allomyia
in this area possess gills.
Limnephilus coloradensis (Banks).— Wyom-
ing (Park Co.): Ghost Creek, 19 August 1977,
1 male, 1 female, and 1 pair in copula, D.
Ferrington, Moose Bog, 9 August 1979, 10
males, 1 female, L. Brooks, 26
Fig, 1. Variation in the parameres (Lateral appendages sensu Ross 1938) of Limnephilus coloradensis (Banks). A
typical symmetrical stnicture, B-E. Variation in parameres showing auxiliary spines.
290
Great Basin Naturalist
Vol. 40, No. 3
males, 39 females, C. Sirianni. Considerable
variation in the male genitalia occm-s in this
species (Figs. lA-E). The simplest type and
probably the most common form (Fig. lA) is
somewhat different from the lectotype fig-
ured by Ross (Ross 1938, Fig. 76). Many spec-
imens also possess auxiliary spines (Fig.
IB-E) not unlike Limnephilus kennicotti
Banks. However, all types will key easily to
L. coloradensis in Ross and Merkley (1952).
In copula specimens have made possible the
correct association of the female. The origi-
nal description of the male is given by Banks
(1899).
Description of female: Forewing length
7-8 mm. Hindwing 6-7 mm. Hindwings
clear. Forewings with brown irregular patch-
es at stigma, distally between R4 and Mg and
in costal area between Mg and Cui^. Smaller
patches in discoidal and thyridial areas, and
along A,. Female genitalia as in Figures
2A-B. Tergite 9 small. Sternum 9 ventrally
divided into two distinct somewhat circular
lobes, roundly triangular when viewed later-
ally. Segment 10 reduced to a ventral flap.
Appendages of segment 10 greatly enlarged,
produced to points. Supragenital plate rec-
tangular. Median lobe of vulval scale
rounded apically, lateral lobes quadrate, pro-
duced laterad.
In general, the female bears some resem-
blance to Limnephilus kennicotti Banks. Both
species were placed in the fenestratus group
by Schmid (1955). In L. kennicotti the dorsal
body of segment 9 is indistinguishable from
segment 10 (see Nimmo 1971, Figs. 419, 420).
In L. coloradensis the dorsal lobe of segment
9 is distinct. In both species the appendages
of segment 10 are greatly enlarged and pro-
duced to points distally. These appendages
show considerable variation in L. colorad-
ensis. In both species the supragenital plate is
rectangular. The lateral lobes of the vulval
scale is more quadrate in L. coloradensis.
Limnephilus hageni Banks.— Colorado
(Grand Co.): Winding River Campground, 2
August 1978, 2 males, D. Ferrington. Wyom-
ing (Park Co.): Moose Bog, 18 August 1977, 4
females, collected L. Ferrington, females
identified A. Nimmo.
Limnephilus indivisus Walker.— Wyoming
(Park Co.): Moose Bog, 9 August 1979, 1 fe-
male, C. Sirianni.
Limnephilus janus Ross.— Colorado (Grand
Co.): Winding River Campground, 2 August
1978, 10 males, 8 females, D. and L. Ferring-
ton.
Limnephilus picturatus McLachlan.—
Wyoming (Park Co.): Moose Bog, 6 August
1977, 25 males, 22 females, 18 August 1977,
4 females, 1 female, L. and D. Ferrington, 9
August 1979, 33 males, 24 females, L. Brooks,
53 males, 66 females, C. Sirianni. Chain
Lakes, 12 August 1977, 2 males, 6 females, D.
app
Fig. 2. Female genitalia: A, ventral view; B, lateral view, segments 8 to 10. app. = appendages of segment 10.
September 1980
SwEGMAN, Ferrington: Western Trichoptera
291
Ferrington, 6 Augvist 1979, 57 males, 24 fe-
males, L. Brooks.
Limnephilus secludens Banks.— Colorado
(Grand Co.): Winding River Campground, 2
August 1978, 1 female, collected D. Ferring-
ton, identified A. Nimmo.
Neothremma alicia Banks.— Wyoming
(Park Co.): Ghost Creek, 15 August 1974, 33
larvae, 6 pupae (1 pharate male, 5 pharate
females), L. Ferrington; Sawtooth Lake, 23
July 1978, 4 larvae, 1 pharate male pupa, D.
Ferrington.
Oligophlebodes zelti Nimmo.— Wyoming
(Park Co.): Ghost Creek, 27 July 1978, 3
males, 3 females, collected L. Ferrington.
Some variation exists between these and Al-
berta specimens (Nimmo, pers. Comm.). This
species was previously known only from Al-
berta (Nimmo 1971).
Psychoglypha aff. subborealis Banks.—
Wyoming (Park Co.): Beartooth Lake, 17
August 1977, 5 larvae, R. Seward. These lar-
vae possess banded legs, as in P. subborealis
(Wiggins 1977).
Lepidostomatidae
Lepidostoma phwiale Milne.— Wyoming
(Teton Co.): Gross Ventre Campground, 29
July 1978, 8 males, 30 females, D. and L.
Ferrington.
Acknowledgments
We thank Deborah Ferrington and Lisa
Brooks for help in the field. Dr. Glenn Wig-
gins for his helpful comments regarding the
AUomyia larvae, and Dr. J. V. Ward for sup-
plying Alhmyia larvae from Colorado. Spe-
cial thanks are also extended to Dr. A. Nim-
mo for his identifications and helpful
comments. Dr. Oliver S. Flint, Jr. for obtain-
ing permission for us to examine material at
the U.S. National Museum of Natural History
and for checking the identifications of Hydro-
psyche oslari and Lepidostoma pluviale, and
to Dr. F. Schmid for his examination of L.
coloradensis. Drs. Flint and Schmid also pro-
vided critical comments of this manuscript.
Dr. Richard T. Hartman provided consid-
erable material support through the Pyma-
tuning Laboratory of Ecology.
Literature Cited
Banks, N. 1899. Descriptions of new North .American
neuropteroid insecis. Trans. .\m. Ent. See.
25:199-218.
Nimmo, A. P. 1971. The adult Rhyacophihdae and Lim-
nephilidae (Trichoptera) of .\lberta and eastern
British Columbia and their post-glacial origin.
Quest. Entomologicae. 7:234 pp.
Ross, H. H. 1938. Lectotypes of North American caddis
flies in the Museum of Comparative Zoology.
Psyche 45:1-61.
Ross, H. H., AND D. R. Merkley. 1952. An annotated
key to the nearctic males of Limnephilus (Tri-
choptera:Limnephilidae). .Am. Midi. .Nat.
47:435-455.
Schmid, F. 1955. Contributions a i'etude des Limnophi-
lidae (River drainage of Idaho with special refer-
ence to the larvae. .Ann. Ent. Soc. .Am.
61:655-674.
Wiggins, G. 1977. Larvae of the North .American cad-
disfly genera (Trichoptera). University of Toronto
Press, Toronto.
OBSERVATIONS OX SEASONAL VARIATION
LN DESERT ARTHROPODS IN CENTR-\L NEVADA
Robert D. Pietruszka-
AByrRACT- " " -i Malaise trap collections from terrestrial arthropod populations in central Nevada were
aQal^•zed for .; periods during the I97S growing season. Mites Acarina and ants Formicidae were the
taxa representee p% tne largest numbers of indi%iduals in pitfall trap collections throughout the season: Malaise col-
lections were composed mainlv of aerial taxa largeh Diptera and H%Tnenoptera . Peak arthropod abundance was
recorded during mid-June. Collection diversities for both trapping methods were generally low due to the abun-
dance of a few ta-xa. E\idence for spatial heterogeneity in arthropod populations was meager: temporal hetero-
geneitv in these populations, however, was more apparent.
Within shrub desert communities in-
vertebrate animals constitute a major part of
the biotic matrix \Fautin 19-t6\ forming an
important food resource for many consimier
species. Spatial and temporal variation in
such a resource base can affect the foraging
patterns of individual consumers iGill and
Wolf 1977. Mac-\rthiu- and Pianka 1966 > as
well as their intra- and interspecific ecologi-
cal relationships . Wiens 1976'.
Commonly, investigations of arthropod
populations are limited to faunistic invento-
ries e.g.. Allred et al. 1965. Beck and Allred
1968'. or to broad scale comparisons e.s;.. Al-
hed 1973. .\lh-ed and Gertsch 1976, Gertsch
and .\llred 1965 1. The present report repre-
sents a preliminar\" analysis of one part of an
ongoing investigation of lizard ecology" in the
Great Basin Desert. Here I will deal with the
arthropod food base at a single location, fo-
cusing upon the relative success of two sam-
pling schemes, seasonal changes in arthropod
abundance and diversity", and the apparent
degree of local spatial and temporal hetero-
geneity' in arthropod populations.
Study Site and Methods
The research site is in Fairsiew \'allev, Ne-
vada, a relatively flat basin ranging in eleva-
tion from 1370 to 1500 m. Vegetation within
the valley is dominated by Atriplex confer-
tifolia, Sarcobatus baileyi, and Oryzopsis
hymenoides and generally t\pifies the shad-
sc-ale zone common to much of interior Ne-
vada (Billings 1949 1. Average monthlv tem-
peratures for the site varv" from about 0 C to
just over 23 C; average monthly precipitation
varies from about .5 cm to just over 1.5 cm:
mean growing season is 142 days.
Data were analyzed from an insect sam-
pling plot estabhshed appro.ximately 9.6 km
N of Frenchman, Churchill County, where
49 pitfall traps were arrayed along cardinal
compass directions at intervals of 5 m Fig.
1l Traps were randomly located with respect
to the vegetation; the total linear distance of
each line was 120 m. Each pitfall trap mea-
sures 98 mm in diameter by 144 mm in
depth: a funnel insert prevents escape of ar-
thropods once captured. A Malaise trap
Townes 1972) was placed at appro.ximately
the center of the two lines iFig. L.
Malaise and pitfall traps were opened for a
period of 48 hours, followed by a closed peri-
od of generally equal duration. This schedule
was maintained from 13 May to 24 August
1978 and yielded a new sample at roughly
four-day intervals. Each trap contained a
standardized amount of 5 percent formalde-
hyde solution. The captures of each "arm" of
the pitfall traplines for each trapping period
were combined. Thus, each of the N-, S-, and
W-arm samples contained the contents of 12
traps, and the E-arm sample contained those
of 13 traps. Contents of NIalaise trap samples
were maintained separately.
To examine the major seasonal patterns in
the prospective arthropod food base, samples
were analvzed for four trapping periods des-
ignated 13 May. 12 June. 14 July, and 12
292
September 19S0 Pietruszka: Season.\i, V.\riation of Arthropods
293
Fig. 1. Pitfall and Malaise trap placements. Dots ref>-
resent pitfall traps; M represents Malaise trap.
Au2;iist. These spanned the major part of the
surface-active season for lizards in 1978.
Arachnids caught were identified to the or-
dinal level: insects caught were identified to
family or superfamily where practicable, us-
ing Borror. DeLong, and Triplehom J976\
Borror and \Miite (1970), and Chu (1949*.
Samples from each arm of the pitfall trap-
ping grid were analyzed separately, as were
Malaise samples. Total counts of identified
groups were then determined for each
sample. Arthropod diversity within a sample
was assessed using B = 1 — pf. where p,
equals the proportion of individuals in cate-
gory i ^MacArthur 1972). The relative de-
grees of spatial and temporal heterogeneity
were assessed using a similarity inde.x, S,
where
S = l-4(Z|p^-p,.,|) ,
and p^^, and p,, are the proportions of sam-
ples X and y in category i ^Schoener 1970).
Results
Trapping success.— For the four trapping
periods analyzed, the two methods employed
amassed a total catch of 7176 arthropods.
The vast majoritv of these. 6117. were col-
lected along pitfall traplines: 1059 arthro-
pods were collected bv Malaise trapping. On
a per-trapping period basis. Malaise trapping
yielded an average of 265 captures. This is a
substantially lower capture rate (up to 50
percent lower than when these traps are
used in forested habitats Matthews and Mat-
thews 1971). Pitfall traps also \ielded a high-
er number of captiu-es per trapping period,
averaging 382 per trapline arm, or just over
1500 captures per trapping grid. However,
capture rates for the two methods are not di-
rectly comparable due to the greater "at
risk" area for pitfall traps.
The composition of collections obtained bv
the t^vo methods also differed substantially.
Pitfall collections were dominated bv mites
(Acarina>, which comprised almost half the
total collection. These were followed, in or-
der of numerical importance, by H\Tnenoi>-
tera (the vast majority- of which were ants),
Coleoptera. and Diptera. In all, largely or
completely terrestrial forms comprised ap-
proximately 88 percent of the arthropods col-
lected. Malaise trap collections, by c-ontrast
were dominated by Diptera. which formed
over 70 percent of the total collection.
H\"menoptera, Homoptera. Lepidoptera. and
Coleoptera combined to form just over one-
quarter of the total collection Table 1).
These data are consistent with those from
Malaise traps used in forested areas in which
Diptera, H\"menoptera. Hemiptera including
Homoptera'. and Lepidoptera constitute at
least 90 percent of each collection (Matthews
and Matthews 1971).
Seusorwl clianges in abundance and diver-
sity.— Arthropod abundance appeared to
peak during mid-June, approximately 1.5
months after the last of the spring rains.
Numbers of arthropods declined rapidly
thereafter to moderate levels. This seasonal
trend is closely reflected in collections from
pitfall traps but not from Malaise trapping
(Fio. 2.\\ The low number o* captures on 12
June mav reflect an actual decrease in aerial
insects, but it is more Ukely that this is a re-
flection of local changes in wind conditions,
to which this technique is highly susceptible
^Matthews and Matthews 1971). This inter-
pretation is strengthened by the observation
that aerial insects occurred in approximately
equal numbers in the 12 June and 14 July
Malaise samples.
294
Great Basin Naturalist
Vol. 40, No. 3
Table 1. Siiminary of arthropods collected in pitfall and Malaise trap samples.
Pitfall trap
Malaise trap
Acarina
Araneida
Scorpion ida
Solpiigida
Coleoptera
Carabidae
Collembola
Diptera
Hemiptera
Homoptera
Taxa
Number of
specimens
Relative
abundance
Number of
specimens
Relative
abundance
3051
0.4988
Anthribidae
Buprestidae
Curculionidae
Dascillidae
Histeridae
Leiodidae
Melyridae
Nitidulidae
Pedilidae
Staphylinidae
Tenebrionidae
Sminthuridae
Poduridae
Anthomyiidae
Bibionidae
Bombyliidae
Calliphoridae
Cecidomyiidae
Chironomidae
Conopidae
Dolichopodidae
Empididae
Lauxaniidae
Muscidae
Mycetophilidae
Pipunculidae
Psychodidae
Ptychopteridae
Sarcophagidae
Sciaridae
Simuliidae
Syrphidae
Tachinidae
Therevidae
Tipulidae
Xylophagidae
Acalypterate muscoids
Diptera larvae
Lygaeidae
Miridae
Nabidae
Pentatomidae
Tingidae
Aphididae
Cercopidae
Cicadellidae
Coccoidea
Nymph /larvae
104
0.0170
16
0.0026
2
0.0003
30
0.0049
17
0.0028
5
0.0008
2
0.0003
5
0.0008
2
0.0003
244
0.0399
1
0.0002
1
0.0002
3
0.0005
243
0.0397
38
0.0062
7
0.0011
24
0.0039
1
0.0002
3
0.0005
1
0.0002
37
0.0060
18
0.0029
1
0.0002
8
0.0013
4
0.0007
0.0002
1
0.0002
3
0.0005
45
0.0074
1
0.0002
18
0.0029
1
0.0002
42
0.0069
2
0.0003
84
0.0137
18
0.0029
2
0.0003
2
0.0003
1
0.0002
10
0.0016
5
0.0008
102
0.0167
2
0.0003
1
0.0002
4
29
3
5
197
5
1
4
11
3
9
41
203
21
1
1
3
248
1
60
11
0.0009
0.0057
0.0019
0.0009
0.0019
0.0009
0.0038
0.0274
0.0028
0.0047
0.1860
0.0047
0.0009
0.0038
0.0104
0.0028
0.0085
0.0009
0.0387
0.1917
0.0198
0.0009
0.0009
0.0028
0.2342
0.0028
0.0038
0.0009
0.0567
0.0104
September 1980
Pietruszka: Seasonal Variat
ION OF
Arthropods
295
Table 1 continued.
Pitfall trap
Malaise
trap
Hymenoptera
Andrenidae
6
0.0010
Apidae
21
0.00.34
1
0.0009
Braconidae
13
0.0123
Chalcidoidea
51
0.0083
87
0.0822
Chrysididae
6
0.0010
Dryinidae
2
0.0003
Fonnicidae
1577
0.2578
13
0.012.3
Halictidae
66
0.0108
5
0.0047
Ichneumon idae
1
0.0002
1
0.0009
Mutillidae
5
0.0008
Pom pil idae
4
0.0007
2
0.(K)19
Sphecidae
46
0.0075
3
0.0028
Lepidoptera
Cosmopterygidae
19
0.0031
38
0.0359
Lycaenidae
1
0.0002
Pyralidae
5
0.0008
4
0.00,38
Unidentified larvae
12
0.0020
Neuroptera
Chrysopidae
1
0.0009
Coniopterygidae
3
0.0005
1
0.0009
Henierobiidae
18
0.0029
1
0.0009
Myrmeleontidae
17
0.0028
Unidentified larvae
3
0.0005
Orthoptera
Acrididae
2
0.0003
Blattidae
2
0.0003
Gryllacrididae
18
0.0029
1
0.0009
Mantidae
1
0.(XK)2
Thysanoptera
Heterothripidae
4
0.(XK)7
6
0.0057
Phlaeothripidae
7
0.0011
Thripidae
9
0.0015
Unidentified larva
(campodeiform)
1
0.(K)02
Isopoda
1
0.0002
Among the major arthropod groups occur-
ring in seasonal samples, mites most closely
follow the general trend. It is quite likely, in
fact, that mite populations are the major fac-
tor underlying the observed seasonal pattern.
The much greater abundance of mites tends
to mask other groups, such as the Hymenop-
tera, Diptera, and Coleoptera, which tend to
remain at low to moderate levels of abun-
dance throughout the season (Fig. 2B).
Diversity values based on pitfall trapping
and Malaise trapping, respectively, were sub-
stantially different from one another during
all trapping periods of the 1978 season (Fig.
3). Among pitfall samples arthropod diversity
is generally low due to the high abundance of
both mites and ants (see Table 1). The trend
toward increa.sing diversity reflects the rela-
tive decrease in mite abundance in late .sea-
son samples. Among Malaise samples diver-
sity values reflect, in part, the lower total
catch afforded by this method. Trapping
dates with the highest diversity values, 12
June and 12 August, had catches that were
approximately 28 and 8 percent, respective-
ly, of the catches for the remaining trapping
periods. These samples contained fewer taxa
more equitably represented, yielding greater
apparent diversity. It is likely that the values
for 12 June and 12 August Malaise samples
are inordinately high due to the sensitivities
of the technique mentioned earlier. Never-
theless, that there should be relatively greater
diversity of aerial insects seems reasonable,
if for no other reason than their greater
mobility.
Spatial and temporal heterogeneity of ar-
thropods— As mentioned above, spatial and
temporal variation in arthropod abundance
may affect not only characteristics of individ-
ual consumer behavior, but also the ecologi-
cal relationships within and between species.
As an approach to spatial variation on a rela-
tivelv small scale (minimum area effect of the
296
Great Basin Naturalist
Vol. 40, No. 3
8-1
Hymenoptera
13
May
12
June
14
July
12
August
Fig. 2. Seasonal changes in arthropod abundance: a,
as reflected by pitfall and Malaise trap samples; b, sea-
sonal changes in the major taxa of these samples.
trapping grid is probably on the order of 1.4
ha), I calculated similarity values for all pos-
sible combinations of grid arms for each
trapping period. An average similarity value
(S) was then obtained as a measure of the
overall spatial heterogeneity over the trap-
ping grid. Immediately apparent from this
analysis is the high degree of similarity (low
heterogeneity) between grid arms at all
trapping periods (range = .730-.893, Fig. 4).
Yet, there does appear to be a trend toward
increasing arthropod patchiness with decreas-
ing abundance levels. The trend is not statis-
tically significant, however, based upon these
data.
13
May
12
June
14
July
12
August
Fig. 3. Seasonal changes in arthropod diversity as re-
flected in pitfall and Malaise trap samples. Pitfall diver-
sities represent the average diversity for the four grid
arms at each sampling date.
Though evidence for spatial variation in
arthropod numbers during 1978 is meager,
temporal variation is much more apparent.
Average similarity values, based upon all pos-
sible comparisons of each grid arm over all
trapping dates, were substantially lower than
for spatial variation: N-arm = .621; E-arm
= .648; W-arm = .672; S-arm = .692.
These data indicate a substantial change in
the arthropod fauna throughout the active
season over a relatively small area. Com-
parisons of Malaise trap collections support
this interpretation (S = .384), but, as men-
tioned above, the collections for two of the
dates may be suspect.
Discussion
Desert habitats are characterized by both
cyclic and unpredictable climatic changes on
micro- as well as macrogeographic scales
(Cloudsley-Thompson 1968, Logan 1968). As
a result, these habitats are typified by periods
of pulsed production. It is particularly note-
worthy that in the first four months of 1978
Fairview Valley received 255 percent greater
than normal rainfall; average temperatures
between April and August were below nor-
mal (U.S. Weather Bureau data). Such a com-
bination of climatic events may have provid-
ed for a longer than normal production pulse,
resulting in a marked increase in arthropod
abundance throughout the season. The nu-
merical dominance by mites and ants of col-
September 1980 Pietruszka: Seasonal Variation of Arthropods
297
12 Aug
14 July
13 May
0.5 15 25
Arthropods/ Trapping Period ( x 10^)
Fig. 4. Spatial heterogeneity of arthropods as reflect-
ed in the average similarity, S, between pitfall trap grid
arms at each sample date.
lections spanning the entire season seems to
argue for this possibiHty. Moreover, it has
been suggested that at extremely high popu-
lation levels habitat patchiness will be re-
duced, and localized areas may even become
unifonn in their species distributions (Wiens
1976). Indeed, this seems to be what occurred
in 1978. If this hypothesis is correct, then ar-
thropod patchiness would be predicted to be
more apparent during dry years when abun-
dance levels are low. The data to test this
prediction have been gathered (for 1979, a
substantially drier year) but have not yet
been analyzed.
Finally, it is clear from these analyses that
to effectively monitor arthropod populations
no single methodology is sufficient. The com-
binati ai of pitfall and Malaise trapping pro-
vides a reasonable balance of terrestrial and
aerial forms from desert habitats. Never-
theless, specific situations and goals will ulti-
mately determine the techniques to be used.
Literature Cited
Allred, D. M. 1973. Scorpions of the National Testing
Station, Idaho. Great Ba.sin Nat. 33:251-254.
Allred, D. M., and VV. J. Gertsch. 1976. Spiders and
scorpions from northern Arizona and southern
Utah. J. Arachnol. 3:87-99.
Allred, D. M., D. E. Johnson, and D. E. Beck. 1965. A
list of some beeflies of the Nevada Test Site.
Great Ba.sin Nat. 25:5-11.
Beck, D. E., and D. M. Allred. 1968. Faunistic inven-
tory-BYU ecological studies at the Nevada Test
Site. Great Basin Nat. 28:132-141.
Billings, W. D. 1949. The shadscale vegetation zone of
Nevada and eastern Galifornia in relation to cli-
mate and soils. .\mer. .Midi. Nat. 42:87-109.
BoRROR, D. J., D. .M. DeLonc, and C. A. Triplehgrn.
1976. .\n introduction to the study of insects. 4th
ed. Holt, Rinehart and Winston, New York. 852
pp.
BoRROR, D. J., and R. E. White. 1970. A field guide to
the insects of .\merica north of Mexico. Hough-
ton Mifflin Go., Bo.ston. 404 pp.
Ghu, H. F. 1949. How to know the immature insects.
Wm. G. Brown Go. Publishers, Dubuque, Iowa.
234 pp.
Gloudsley-Thompson, J. L. 1968. The Merkhivat Je-
bels: a desert community. Pages 1-20 in G. VV.
Brown, ed.. Desert biology. .Academic Press, .New
York.
Fautin, R. W. 1946. Biotic communities of the Northern
Desert Shrub Biome in western Utah. Ecol. Mon-
ogr. 16:251-310.
Gertsch, W. J., and D. M. .-Vllred. 1965. Scorpions of
the Nevada Test Site. Brigham Young Universitv
Sci. Bull., Biol. Ser. 6(4): 1-15.
Gill, F. B., a.nd L. L. Wolf. 1977. Nonrandom foraging
by sunbirds in a patchy environment. Ecolog\'
58:1284-1296.
Logan, R. F. 1968. Gauses, climates, and distribution of
deserts. Pages 21-50 in G. W. Brown, ed.. Desert
biology. .Academic Press, New York.
Mac.\rthur, R. H. 1972. Geographical ecolog)'. Harper
and Row, New York.
Mac.\rthur, R. H., and E. R. Pianka. 1966. Optimal
use of a patchy environment, .^mer. Nat.
100:60.3-609.
Matthews, R. W., and J. R. Matthews. 1971. The .Ma-
laise trap: its utility and potential for sampling
insect populations. Michigan Entomologist.
4:117-122.
Schoener, T. W. 1970. Nonsynchronous spatial overlap
of lizards in patchy habitats. Ecology 51:408-418.
Townes, H. 1972. A light-weight Malaise trap. Ent.
News 83:2.39-247.
Wiens, J. A. 1976. Population responses to patchy envi-
ronments. .\nn. Rev. Ecol. Svst. 7:81-120.
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TABLE OF CONTENTS
Spatiotemporal variation in phenolog)' and abundance of floral resources on short-
CTrass prairie. V. J. Tepedino and N. L. Stanton 197
Dog owners and hvdatid disease in Sanpete County, Utah. Peter M. Schantz and
Ferron L. Andersen 216
New grass distribution records for Arizona, New Mexico, and Texas. Stephan L.
Hatch 221
A comparison of epiphvtic diatom assemblages on living and dead stems of the com-
mon grass Phragmites atistralis. Judith A. Grimes, Larry L. St. Clair, and Sam-
uel R. Rushforth 22.3
Poisonous plants of Utah. Jack D. Brotherson, Lee A. Szyska, and William E. Even-
son 229
The successional status oi Cupressus arizvnica. Albert J. Parker 254
A self-pollination experiment in Pintis edulis. Ronald M. Lanner 265
Comparative floral biology of Penstemon eatonii and Penstemon cyananthus in cen-
tral Utah: a preliminar\^ study. Lucinda Bateman 268
Differential habitat utilization by the sexes of mule deer. Michael M. King and H.
Duane Smith 2/ -3
Temporal activitv patterns of a Dipodomys ordii population. Clive D. Jorgensen, H.
Duane Smith, and James R. Garcia 282
New records of western Trichoptera with notes on their biolog). Bernard G. Sweg-
man and Leonard C. Ferrington, Jr 287
Observations on seasonal variation in desert arthropods in central Nevada. Robert D.
Pietniszka 292
HE GREAT BASIN NATURALIST
^ lume 40 No. 4
December 31, 1980
Brigham Young University
MUS. COMP. ZOO'
LIBRARY
^MM
f^
GREAT BASIN NATURALIST
Editor. Stephen L. Wood, Department of Zoology, Brigham Young University, Provo, Utah
84602.
Editorial Board. Kimball T. Harper, Botany; Wilmer W. Tanner, Life Science Museum;
Stanley L. Welsh, Botany; Clayton M. White, Zoology.
Ex Officio Editorial Board Members. A. Lester Allen, Dean, College of Biological and Agricul-
tural Sciences; Ernest L. Olson, Director, Brigham Young University Press, University
Editor.
The Great Basin Naturalist was founded in 1939 by Vasco M. Tanner. It has been published
from one to four times a year since then by Brigham Young University, Provo, Utah. In gener-
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taining to the biological and natural history of western North America are accepted. The
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5-81 650 50104
ISSN 0017-3614
The Great Basin Naturalist
Published at Provo, Utah, by
Brigham Young University
ISSN 0017-3614
Volume 40
December 31, 1980
IMPACT OF THE 1975 WALLSBURG FIRE ON
ANTELOPE BITTERBRUSH {PURSHIA TRIDENTATA)
Fred J. W'agstaff
.\bstract.— Antelope bitterbrush {Pur.sliia tiidentcita) is a preferred browse species that is susceptible to decreases
ill population density due to fire. The reduction in density of this species due to fire was determined bv sampling
areas within and adjacent to the burn. The 1975 burn caused a significant reduction in the population density of
bitterbmsh. It \yas also determined that rate of growth was lower for plants within the burn.
In the summer of 1975 a fire burned sever-
al himdred acres of mule deer winter range
in Wasatch County, Utah. The burned area
includes the area from the junction of the
Wallsburg road southeast to the crest of the
west Daniels Canyon ridge and to the north-
east along Highway 40 to near the Midway
Junction. Deer Creek Reservoir is just across
the highway to the northwest of the burned
area.
The study plots are near the northeast cor-
ner of the burn in an area where a population
density of bitterbrush was great enough to
permit quantitative analysis of the response
of this species to fire. Burned and imburned
areas were studied along with some islands
that escaped burning. Bitterbrush {Purshia
tridentata) was selected as an indicator spe-
cies because of its status as a preferred
brow.se plant on mule deer winter ranges.
Any factor that causes significant changes
in the structure of the plant communities on
winter ranges is of concern. One of the most
significant agents known is fire. Fire has oc-
curred naturally since time began and is a
major factor in determining the .structure of
many plant communities. In other commu-
nities, man-caused fire has introduced an
agent of change that has modified vegetation
over large areas.
In a situation where prefire structure is so
important, the impact of fire on structure of
the plant community should be known. Will
plants be killed? How long will the impacts
last? Will value of the area as mule deer win-
ter range be completely lost? Can deer move
to another winter range? These and other
questions occur and need to be answered if
the impacts of fire are to be understood.
The area chosen for study has been burned
several times in the last one hundred years.
Fire has occurred at irregular times and over
different portions of the area. This has led to
a mosaic of vegetation types and age struc-
tures. The 1975 fire was much larger than
most of the past fires and affected a signifi-
cant portion of mule deer winter range in the
area. With passage of four years, the in-
ception of the postfire plant succession
should be identifiable.
Three major hypotheses were fornuilated
to determine some of the relative impacts of
the 1975 fire. First, the density of bitterbmsh
had been significantly reduced by the fire.
Secondly, use by mule deer is less in the
burned area. Third, the shift in deer u.se has
'U.S. Forest Service Interniountain Forest and Range Experiment Station Sl.nib Sciences Laboratory, 735 North 500 East. Provo. Utah 84601.
299
3()0
Great Basin Naturalist
Vol. 40, No. 4
had a detrimental effect on surviving bitter-
bnish plants.
Literature Review
Literature pertaining to bitterbrush is ex-
tensive, with over 200 references identified.
The following references support the results
of the study.
FiirsJiid tridcntata (antelope bitterbrush) is
highly desirable as browse on deer winter
range (Bissell et al. 1955, Giunta et al. 1978,
Hoskins and Dalke 1955, Julander 1952,
Leach 1956, Longhurst et al. 1952, Mace
1957, Reynolds 1960, Smith et al. 1954,
Smith 1952). Since bitterbmsh is so highly
preferred, it can be used as an indicator spe-
cies for use on an area bv game animals and
game winter range conditions.
There have been numerous articles written
about the impacts of fire on bitterbrush
(Blaisdell 1950, 1953, Blaisdell and Mueggler
1956, Countryman and Cornelius 1957, Fer-
guson and Basile 1966, Komarek 1965, Miller
1963, and Pechanec et al. 1954). They have
determined that browse production of bitter-
brush plants that have been burned has
lagged behind miburned control plants for
several years. Blaisdell (1950) also showed
that relative densities of bitterbrush in
burned and imbiuned areas differed signifi-
cantly. Nord (1965) developed data that dem-
onstrates the existence of fire-resistant eco-
types where most plants in the population
resprout after fire. Blaisdell (1953) and others
have shown that variables of fire intensitv,
fuel loading, and soil moisture affect re-
sprouting. Even nonsprouting types will have
some survivors, particularly where the fire
does not burn intensely.
Results
The following data were collected from six
100 ft- quadrats in the burned and adjacent
unburned area. Two of the quadrats were at
the lower edge of the burn in the sagebrush-
bitterbrush tvpe and four near the middle of
the burn in the oak-sagebrush tvpe. The rela-
tive density of live bitterbrush plants is
shown in Table 1.
There is a striking difference between the
nimiber of living plants in the burned and un-
burned areas. None of the burned plots had
anv surviving old plants because the fire was
intense enough to kill the tops of all bitter-
brush plants. It appears the fire mav have
been hotter at the lower part of the burn be-
cause there were no relic bitterbnish plants.
In the midslope plots, relics were found for
most shrubs of various species.
All the young bitterbrush plants in the
burned area were from resprouting crowns.
There were voung plants in all the unburned
plots, indicating the species is successfully re-
producing in the study area.
In Table 2, the data collected on the
current-year twig growth are presented.
Twenty twigs per plant on 20 plants (200
twigs in burned areas and 200 in unbiu'iied)
were measured to determine if there was anv
difference in current-year growth. The differ-
ence in twig growth was found to be statistic-
ally significant at the 90 percent level. In
other words, the burned plants were growing
at a slower rate.
For each of the six plots, mule deer fecal
pellet groups were counted. All pellet groups
were counted without regard to pellet age.
Clearly, the unl)urned areas have received
heavier use than the adjacent burned areas, as
shown in Table 3.
Table L Relative density of bitterbrush plants on six plots.
Burned
Slope
location
Bitte
rbrush
plants
Quadrat
New
Old
Dead
Total
1
Yes
Mid
2-
0
22'
24
2
Yes
Mid
f-i-
0
22'
28
3
No
Mid
(i
15
1
22
4
No
Mid
9
15
1
24
5
Yes
Lower
2-
0
0
2
6
No
Lower
3
10
0
13
Includes relics probably killed by fire.
Includes resprouting after fire.
December 1980
Wagstaff: Impact of Fire ox Bitterbrush
301
Takle 2. 1979 lcn<4tli (if tuitrs on 20 liitterhrush
plants.
Table 3. Number of mule deer fecal pellet groups for
each of six plots.
Unhiirned area plants
Aveniije
Plant twig length
mmiber (inches)
Burned area plants
Average
Plant twig length
number (inches)
9.5
1
9.3
2
9.4
3
7.05
4
10.15
5
10.05
6
9.8
7
6.55
8
8.45
9
9.9
10
9.02
y
Discu:
SSION
9.2
7.75
9.55
7.6
6.95
7.45
8.45
9.1
6.8
7.15
8.0
There are several obvious differences be-
tween the burned and unburned plant popu-
lations. It is evident the fire had a detrimen-
tal impact on the density of bitterbrush
plants and production of this component of
the plant community. A relatively low per-
centage of the bitterbrush plants were able to
resprout and, therefore, most burned area
plants were eliminated from the community.
Sagebnish was also largely absent in the
burned areas, but in adjacent unburned areas
it was a significant part of the community.
The marked change in the community spe-
cies composition caused by the fire is still
very much in evidence. Also the size of the
browse plants has been altered to the point
that much of the burned area vegetation
would be totally covered by several inches of
snow. This physical barrier would limit use of
the burned area even though considerable
forage may occur there.
Another obvious factor is the difference in
occurrence of fecal pellet groups between
burned and adjacent unburned areas. Since
the measurements were taken in adjacent
areas, the reason for the significant difference
seems easiest to explain on the basis of physi-
cal availability of browse diuing the winter.
If there are no significant differences in nu-
tritive value or availability, one would expect
essentially equal use near the boundary of the
two areas. The difference in use must occur
because of an absolute difference in the
amount of browse available, which was prob-
ably compounded by snow coverage.
Plot
Slope
position
Number of
pellet
groups
Mid
Burned
8
Mid
Burned
5
Mid
I'nburned
28
Mid
Unburned
.36
Low
Burned
4
Low
Unburned
16
All the bitterbrush plants that were exam-
ined during this study exhibited substantial
twig growth and appeared to be vigorous and
healthy. Growth form of the old plants in the
unburned areas showed a fairlx' open fcjriu
that indicates little use by sheep and/or deer.
These plants did not exhibit the clubhead
form indicative of sustained heavy use; nei-
ther was there evidence of browsing on large-
diameter twigs.
Growth of the Ijitterbrush plants in un-
burned areas was greater, as has been docu-
mented in other areas by Blaisdell and
Mueggler (1956). This difference in the
growth rate is expected to continue for sever-
al years. In terms of total production of hit-
terbnish forage, the burned area has lapsed
considerably since the fire and will most
likely continue to do .so for a long time. The
density of plants has been reduced, as well as
the size of plants. There are fewer plants of
smaller size in the burned areas than were
there prior to the fire. Since no evidence of
new seedlings could be foinid, it is reasonable
to conclude the burned area production will
lag for nianv years.
Deer herd populations are often directly
tied to winter range availability and condi-
tion because it is the element most often in
shortest supplv. It is not known whether this
is the case with the mule deer using the study
area. There is need for concern because deer
numbers are increasing and winter range in
the area is decreasing due to changing land
ase. Both of these trends cannot continue in-
definitelv without deer niunhers reaching the
limit of available winter range.
CONCLL SIO.NS
It is easv to conclude that the 197.5 fire
was detrimental to bitterbnish, but it is not
302
Great Basin Naturalist
Vol. 40, No. 4
easy to conclude that the numbers of deer
have been reduced because of it. This is due
to populations of deer being within the ca-
pacity of the remaining winter range. The
area is neither producing the winter forage
for deer that it was prior to the burn nor are
deer numbers anywhere near historic high
levels.
If this area is indeed a critical winter range
when population numbers are larger and
other factors holding populations down are
temporary, additional concerns arise.
Thought should be given to introducing a
fire-tolerant ecotype of bitterbnish with the
hope of hybridizing this trait into the com-
munity, and effective means of reducing fire
occurrence and spread should be developed.
Literature Cited
BissELL, H. D., B. Harris, H. Strong, and F. James.
1955. The digestibility of certain natural and arti-
ficial foods eaten by deer in California. Calif.
Fish and Game 41(1)1.57-78.
Blaisdell, J. p. 1950. Effects of controlled burning on
bitterbni.sh on the upper Snake River plains. U.S.
For. Serv., Int. For. and Range Exp. Sta. Res.
Pap. 20, .3 pp.
19.53. Ecological effects of planned burning of
sagebnish-grass range on the upper Snake River
plains. U.S.D.A. Tech. Bull. 1075, .39 pp., illus.
Blaisdell, J. P., and W. F. Mueggler. 1956. Sprouting
of bitterbnish (Purshia tridentata) following burn-
ing or top removal. Ecology 37:.365-370, illus.
Countryman, C. M., and D. R. Cornelius. 1957. Some
effects of fire on a perennial range type. J. Range
Manage. 19:39-41, illus.
Ferguson, R. B., and J. V. Basile. 1966. Topping stiniu
lates bitterbrush twig growth. J. Wildl. .Manage.
.30(4): 8.39-841.
Giunta, B. C, R. Steve.ns, K. R. Jorge.nsen, and A. P.
Plummer. 1978. Antelope bitterbrush— an impor-
tant wildland shrub. Utah State Div. Wildl. Re-
sources Publ. 78-12.
HosKiNS, L. W., AND p. D. Dalke. 1955. Winter browse
on the Pocatello big game range in southeastern
Idaho. J. Wildl. Manage. 19:21.5-225.
JuLA.NDER, O. 1952. Forage habits of mule deer during
the late fall as measured by stomach content
analyses. U.S. For. Serv. Int. For. and Range Exp.
Sta. Res. Note-2. 5 pp.
KoMAREK, R. 1965. Fire and changing wildlife habitat.
Proc. Tall Timbers Fire Ecol. Conf. 2:35-43.
Leach, H. R. 1956. Food habits of the Great Basin deer
herds of California. Calif. Fish and Game
43(4):243-308, illus.
LoNGHURST, W. M., A. S. Leopold, and R. F. Das.ma.nn.
1952. A survey of California deer herds, their
ranges and management problems. Calif. Dept.
Fish and Game Bull. 6. 1.36 pp., illus.
Mace, R. LI. 1957. Oregon's mule deer. Oregon State
Game Comm. Wildlife Bull. No. 3. 25 pp.
Miller, H. A. 1963. Use of fire in wildlife management.
Proc. Tall Timbers Fire Ecol. Conf. 2:19-30.
NoRD, E. C. 1965. Autecology of bitterbnish in Califor-
nia. Ecol. Monogr. .35:307-.3.34.
Pechanec, J. F., G. Stewart, and J. P. Blaisdell. 1954.
Sagebrush burning— good and bad. U.S.D..\.
Farmer's Bull. 1948. .34 pp.
Reynolds, T. A., Jr. 1960. The mule deer— its history,
life history, and management in Utah. Utah State
Dept. Fish and Game Inform. Bull. 60-4. .32 pp.
Smith, \. D., and R. Hubbard. 1954. Preference ratings
for winter deer forages from northern Utah
ranges based on browsing time and forage con-
sumed. J. Range Jv-lanage. 7(6):262-265.
SxuTii, J. G. 1952. Food habits of mule deer in Utah. J.
Wildl. Manage. 16(2): 148-155, illus.
TERRESTRIAL VERTEBRATE FAUNA OF THE KAIPAROWITS BASIN
N. Diiane Atwood', Clyde L. Pritchctt', Richard D. Porter', and Benjamin W. Wood'
.\bstr^ct.- This report inehides data collected during an investigation by Brighani Young University personnel
from 1971 to 1976, as well as a literature review. The fauna of the Kaiparowits Basin is represented by 7 species of
mnphihians (1 salamander, 5 toads, and 1 tree frog), 29 species of reptiles (1 turtle, 16 lizards, and 12 snakes), 183
species of birds (plus 2 hypothetical), and 74 species of mammals. Geographic distribution of the various species
within the basin are discussed. Birds are categorized according to their population and seasonal status. .Avian habitat
relationships are discussed, and extensions of range are reported for 5 species of birds. Three threatened or endan-
gered avian species occur in the basin. Four avian species seem to have declined significantly in numbers in recent
\ears.
The early activities and exploration of
trappers, missionaries, and government sur-
vey workers provided little information to
om- knowledge of fauna in the Kaiparowits
Basin. Most of these early expeditions skirted
around the basin on all sides or were con-
fined to the depths of the Colorado River
Canyon. The Domingues-Velez de Escalante
party traveled along the north side of the
Colorado River between Lee's Ferry and the
well-known Crossing-of-the-Fathers. During
this portion of their journey, it became neces-
sary for the party to eat their horses (Auer-
bach 1943). No mention is made in Father
Escalante's journal of any fauna observed.
Records of Powell's expeditions of 1869-1870
and 1871-1872 indicate observations of wild-
life were limited to the larger game animals,
probably those that could be used for food.
In 1892 the American Museum of Natural
History sent an expedition into the San Juan
region just east of the Kaiparowits Basin (Al-
len 1893). Wetherill, Flattum, and Sterns
(1961) made a trip by boat up the Colorado
River from Lee's Ferry to Rainbow Bridge.
Both expeditions recorded the animals ob-
served.
The number of scientific investigations af-
ter those of Powell into the Glen Canyon
were few indeed (Crampton 1959). The Na-
tional Park Service recognized this fact and,
in measure, filled some of the gaps by send-
ing out an expedition that descended by boat
the San Juan and Colorado Rivers to Rock
Creek. From this point, they spent about two
weeks on the southwest end of the Kaiparo-
wits Plateau. The results of this work were
reported by Hall (1934). Others such as
Gregory (1917, 1938, 1945, 1947, 1948) and
Gregory and Moore (1931) have made sub-
stantial contributions to our knowledge of the
Colorado River drainage, particularly with
respect to the geology. Observations and
comments were also recorded regarding the
biota.
Members of the Department of Zoology at
Brigham Young University' (BYU) initiated a
.series of biological investigations of the up-
per Colorado River Basin; of these, the fol-
lowing were within the Kaiparowits Basin: In
July 1927, a group visited Lee's Ferry and
Bryce Canyon, and in 1936. a party of four
biologists spent 810 man hours in the Esca-
lante River drainage. In the fall of 1937, V.
M. Tanner and C. L. Hayward studied in the
Paria Valley. A party headed by D. E. Beck
entered the region in the fall of 1938 and
1939, exploring the course of the Escalante
River. Beck also .spent the last part of May
and earlv June of 1940 in the area east of
Willow Tank Spring down to the Escalante
River. In 1946, 1952, and 1953, other BYU
expeditions entered the area in both Kane
and Garfield Counties. The Navajo Mountain
and Wahweap Drainage were visited in 1955
and 1958 (Hayward et al. 1958).
'U.S. Forest Service, Provo, Utah 84601.
■Department of Zoology, Brigham Young I'niversity. Prove, Utah 84602.
U.S. Fish and Wildlife'Service, Provo, Utah 8460r(.325 North .300 West. Mapleton, Utah 84663).
'Department of Botany and Range Science. Brigham Young University, Provo, Utah 84602.
303
304
Great Basin Naturalist
Vol. 40, No. 4
11 ,; .A 30/ ^^J' V 7 Aa;^'^ ;|smok* NfV^j -i ,• , Ky'x \ ^^'vA
1/ S ,v ^-x^;^ ;,
NIPPIE BENCH "^^^
0'
...^
Fig. 1. Map of Brigham Young University Xavajo-Kaiparowits study sites, Kane and San Juan Counties, Utah, and
Coconino Count\. Arizona.
Benson (1935) published an important pa-
per on the fauna and flora collected during
an expedition led by Alexander, Kellogg, and
Benson in the Navajo Mountain region in
1933. They had six stations as follows: one
five miles .south of the mountain, 10-12 June;
one at War God Spring on top of the moun-
tain at 8400 feet, 13-20 June; one at Bridge
Canyon, 21-24 June, two on the mesa south
of the mountain; and the other near Navajo
Mountain Trading Post, 25-26 June. In addi-
tion, naturalists assigned to Bryce Canvon
National Park have, since 1932, made contri-
butions toward our knowledge of the fauna
and flora of the region.
Woodbury and Russell (1945), in their
comprehensive report on the birds of the
Navajo countrv, presented data on specimens
collected and observations made in Glen
Canvon, on Navajo Mountain, and on Kai-
parowits Plateau. Behle and three associates
made a boat trip down the Colorado River
13-17 April 1947. They made observations at
various places between Hite and Lee's Ferry,
December 1980
Atwood et al.: Kaiparowits Vertebrates
305
including the mouth of the Escalante River,
river mile 88; Hidden Passage, river mile 76;
Aztec Creek, river mile 68.5; Crossing-of-the-
Fathers; near Creek, river mile 40.5; and
Lee's Ferry. Behle and associates also collect-
ed birds at or near the confluence of Calf
Creek and the Escalante River, 11-14 June
1953, and 7-9 May 1954 (Behle et al. 1958;
Behle 1960). In 1958, Behle and Higgins
(1959) made some observations at Hole-in-
the-Rock (20 October), and the confluences
of Kane Creek and the Escalante River with
the Colorado River (19 October); birds were
observed by Harold Higgins and Gerald
Smith on a river trip extending from 1 Julv to
9 August 1958 from Hite to Lee's Ferry
(Behle and Higgins 1959). We have not given
the precise dates in the species accounts giv-
en below of birds collected or seen bv Behle
and his associates on June 1953 and Mav
1954 trips, or for Benson's (1935) trips in
1933, becau.se of the short duration of their
investigations. If desired, such can be obtain-
ed from the literature. The dates of observa-
tions from our investigations, however, are
reported herein.
In 1957, the National Park Service in-
itiated a salvage program for the Glen Can-
\"on area preparatory to the constniction of
Glen Canyon Dam. These data were pub-
lished by the University of Utah and the Mu-
seum of Northern Arizona in their respective
journals. Earlier (as a result of a reconnais-
sance trip, 8-14 August 1957, and as a part of
tlie same project), Woodbury et al. (1959)
had prepared an annotated checklist of the
birds of the proposed Glen Canvon Reservoir
area from various sources, including pub-
li.shed and unpublished manuscript records.
In addition, Behle and Higgins (1959) pub-
lished a number of previouslv impubli.shed
bird observations from Woodburv and Rus-
sell's field notes from the Rainbow Bridge-
Monument Valley expedition.
In June 1971, Brigham Young University
and Northern Arizona University initiated
the Environmental Impact Studies for the
Navajo and later in 1972 for the proposed
Kaiparowits Generating Stations. Extensive
collections and observations of the terrestrial
vertebrate fauna and flora have been made
by these groups.
This report on the terrestrial vertebrate
fauna is based on species reported in the lit-
erature and collections or field observations
made by BYU personnel from 1971 to the fall
of 1976. The various taxa in the annotated
lists are arranged phylogenetically following
the order given in Schmidt (1953) for the am-
phibians and reptiles; the .\merican
Ornithologists Union (AOU) Check List, Fifth
Edition (1957) and subsequent supplements
for the bird.s; and Hall and Kelson (1959) for
the mammals.
Most of the studies dealing with the distri-
bution and ecology of amphibians and rep-
tiles within the Kaiparowits Basin have l^een
restricted to areas along the Colorado River.
The stretch of river generally known as Glen
Canyon Gorge has been the area most com-
monly studied. The most comprehensive list
(28 taxa) of amphibians and reptiles for this
area was by W. W. Tanner (1958a). Wood-
bury et al. (1959) published a similar list, but
it contained only 19 ta.xa. Both of these stud-
ies were designed to obtain data prior to the
constrution of Glen Canyon Dam and sub-
sequent development of Lake Powell. .-Xs part
of the Navajo-Kaiparowits environmental
baseline studies. Toft (1972) prepared a field
key ba.sed on the above literature and field
observations and collections made during the
summer of 1971 and 1972. .\dditional data
have been added from subsecjuent studies
made during the period of 1973-1975. These
studies represent to date the most extensive
field studies into the interior of Kaiparowits
Basin.
Amphibians
Ambystomidae (Salamanders)
Ambystoma trigrimim nehulosum Hallow-
ell. Utah Tiger Salamander. V. NL Tanner
(1930) and Weight ^1932) Bryce Canvon. W.
W. Tanner (1975).
Pelobatidae (Spadefoot Toads)
Scaphiopus hammondi Baird. Hammonds
Spadefoot Toad. V. M. Tanner (1930) and
Weight (19.321 Bryce Canvon.
Scaphiopus intermontanus Cope. Great
Basin Spadefoot Toad. V. M. Tanner (1940a)
306
Great Basin Naturalist
Vol. 40, No. 4
Tablf. 1. Vegetation types and locations of perniaiient sites for Biighaiii Young University Navajo-Kaiparowits
study.
Site
No.
Vegetation type
Oryzopsis-Stipa-Ephedra
Vanclevea
Location
At the base of Cedar Mt. on loo.se,
deep sand, 3.2 mi W of GCC
through the highway fence. T 4.3 S
R 2E S\\' Part of the NW quarter
of Sec. 21.
Junipenis-Bouteloua
On Cedar Mt. 4.1 mi on from Site
1. .\pprox. 20 vds N of fence on left
of road. Exclosure 100 yd N of
fence and W 30 yd R 44 S T 2E
NE corner of Sec. 3.
Hilaria-Ephedra
Cedar Mt. .\pprox. 1.3 mi on from
Site 2. Road is headed W and ca 50
vd before Pole Line on left. R 44
S T 2E Center of Sec. 10, ca 100
vd from state line. Kane Co., UT.
Coleogyne
Cedar Mt. 0.8 mi on from Site 3
after turning left under Pole Line.
R 6E T 42.N SW part of the NE
quarter of Sec. 5.
.•\triplex corrugata- A triplex
confertifolia
.\pprox. 7.0 mi East GCC near
U.S. W. B. and Dames and Moore
weather station on Dakota
formation on right, ca 75 yd from
W. B. station.
Pinyon-Juniper- Artemisia
Smokij Mt. 14.2 mi from Last
Chance Jet. Turn right .4 mi Site
on right of road. On coal mine road
turn right at top of Smoky (survey
marker) 0.1 mi then left .4 mi T
41S R 4E NE part of the SE quarter
of Sec. 8.
10.
Gravia-Coleogvne-Hilaria
Gravia-Hilaria
Atriplcx-Kochia-
Artemisia spinescens
Orvzopsis
Smoky Mt. 12.3 mi from Last
Chance Jet. on top. E of road near
dead juniper. T 41S R 8E NE part
of the SE quarter of Sec. 21.
Smokif Mt. 9.1 mi from Last
Chance Jet. (on left of road) or 5.2
mi from coal mine road jet. on
right side of road. T 41S R 9E SW
part of the SE quarter of Sec. .34.
Base of Sntokii Mt. 2 mi on from
Ahlstrom Point Jet. on left. Turn off
road in bottom of small wash to
dead end at ca .2 mi Site 20 yd N.
T 42S R 5E NW part of the NW
quarter of Sec. .30.
Alil.strotn Point Road 4.7 mi from
Ahlstrom Point Jet. T 43S R 5E SE
part of the NW quarter of Sec. 8.
December 1980
Tal)Ie 1 tontiiuied.
Atwood et al.: Kaiparowits V'ertebrates
307
Vegetation type
Coleogvne
Coleotrvne
Grayia-Ephedra-Orvzopsis
Boiiteloua-Hilaria
Grayia-Coleogyne-Bouteloiia
Popiilus-Tainaiix-L\ ciimi
Tainari.\-Chr\'sothainn\is
Spoiol)
(Joleosivne
Location
Ahl.strom Point Road 5.0 mi or .3
mi from Site 10. T 4;3,S R 5E .\E
part of the SE (|iiartL-r of Sec. 8.
\il)))lc Bench In first well-
developed Goleogyne ca .3.5 mi
from jet. at top after leaving Tibbet
Spring. On left of road, there i,s a
.small ridge on right. T 42 S R .3E
NE part of the S\\' qnarter of Sec.
9.
\ipple Bench .\pprox. .6 mi on
fiom Site 12. Tnrn np wash to the
right ca 50 ft into Sec. 17 from
marker, which is .1 mi off road up
the woods. T 42 S R 3E \E part
of the NE qnarter of Sec. 17.
\ip})le Bench .Appro.x. 1.1 mi S of
SE corner 17 on W of road. 2.1 mi
S Site 13 T 42 S R 3E SE part of
the SW ([iiarter of Sec. 21.
Lmt ('liance Creek 31.8 mi from
C.CXJ tnrn \ip creek bottom ca 1 mi
T 415 R 5E SE part of the SE
quarter of Sec. 4.
.Salt Wash at Middle Branch Creek
Across the creek and ca 40 vd
downstream from washed out stock
watering pond. T 41S R 4E SE part
of the SE quarter of Sec. .3.5.
Staleline 7.6 mi from (iC^C on Utah
side of border turn left toward
Lone Rock .Marina from U.S. 89.
.\pprox. .2 mi on right of road.
Statcline Same as above except on
left of road 75 \cl and on a slight
rise.
Coleogyne
Winihnill 9.3 mi S of Faue on 89 on
left of highwav on pullout bv
reflector post. T .39.\ R 8E SE part
of the XW (luarter of Sec. 10.
Bouteloua-Hilaria-
Muhlenber<;ia
Pinvon-Juniper
Coleog)aie
M mi post .5.35 ca 12 mi S of Page.
50 vd \ of post and 20 \d E of
road on pullout. T .39N'R 8E SW
part of the .\E Cjuarter of Sec. 20.
Echo Cliffs 19.3 mi S of Page turn
left and on ridge ca '4 mi.
Xavajo Plant Site (undecided)
308
Great Basin Naturalist
Vol. 40, No. 4
Table 1 continued.
Site
No.
Vegetation type
Location
23.
24.
25.
26.
27.
28.
29.
30.
31.
32.
33.
3.3a.
33h.
33c.
Coleogyne-Ephedra-Grayia
Hilaria-Ephedra-Grayia
Atriplex confertifolia
Atriplex corrugata
Pinyon-Juniper
Artemisia tiidentata
Pinyon-Jiiniper
Grass-Ephedra
Coleogvne
Orvzopsis
Moist hanging gardens
with thin-leaved genera of
trees and shrubs various
genera of herbaceous plants.
Cercis, Cladium, Rhamnus
Ostrva, Cirsiium. Rubus
Quercus. Gcltis. Populus,
Baccharis, grass/forlj
1.2 mi N Tibbet Spring to jet. and
thence 1.6 mi E on Cathy's Flat
Road 50 yd N of road. T 41S R 3E
NE part of the SE quarter of Sec.
33.
2.1 mi E Site 2.3. .\ppro.\. .2 mi
beyond end of road. T 42S R 3E SE
part of the NE quarter of Sec. 3.
To be selected near Cath\"s Flat.
1(X) yd W of small twin flat-tops at
Last Chance Summit on S of road
ca 13 mi E of Warm Creek Jet. T
42S R .5E east central part of the
SE quarter of Sec. 25.
The Pine ,\pprox. 3 mi from cow
camp at head of Wesses Canyon. T
40 S R 2E part of the SE quarter of
Sec. 2.
1-1 mi W Drip Jet. 40 yd SW from
Dead Juniper on S of road. T 40S R
3E NE part of the SW quarter of
Sec. 7.
Farthest distance out on Drip
Point. T 40S R 3E SE part of the
NE quarter of Sec. 21.
T 41S R 1 W NE part of the SW
quarter of Sec. 27. Brigham Plains
Bench, ca 5 mi N of U.S. Highway
89, E of Paria River.
Grand Bench ca 50 mi E of Glen
Canyon City. T 42S R 6E.
Grand Bench ca 54 mi E of Glen
Canvon Citv. T 42S R 6E.
In Driftwood Canvon on the .N side
of Lake Powell ca'l mi NW of
Rainbow Bridge Canyon. T 43S R
8E.
Ribbon Canyon, between San Juan
Drainage and Hole-in-the-Rock on
the E side of Canyon. T 41S R lOE.
Reflection Canyon (Cottonwood
Gulch), between San Juan drainage
and Hole-in-the-Rock on the W
side of canvon. T 42S R 9E.
December 1980 Atwood et al.: Kaiparowits Vertebrates
309
Table 1 continiifd.
Site
No. Vegetation type Location
.33d. Quercus, Cercis, Cirsiuin 1 pii N of confluence of
Forh/grass Colorado/San Jnan Rivers on E
side of canyon. T 42S R 9E ,5.5 mi E
of Glen Canyon Citv in Last
Chance Creek. T 41S R 6E.
.34.
Taniarix-dcsert siiiuh
K 42S H9E.55ini Euftilen
Canyon C^itv in Last C^hance
Creek. T 4IS R 6E.
Willow Tank Spring and confluence Calf
Creek/Escalante River; Hayvvard et al.
(1958) Navajo Mountain; Russell and Thomp-
son (1964) Bryce Canyon; BYU (1972)
Grosvenor Arch.
Bufonidae (Toads)
Bufo cognatus Sav. Great Plains Toad.
BYU (1971) Site 15.
Biifo punctattis Baird & Girard. Red-
spotted Toad. V. M. Tanner (1940a) Willow
Tank Spring; Woodbury et al. (1959) Bridge
Canyon, Rock Creek, Hidden Pa.ssage, Aztec
Canvon, Rainbow Bridge, and Warm Creek;
BYU (1971-1973) Warm Creek, Escalante
drainage, Wahweap Creek, Driftwood Can-
yon, Reflection Canyon, and Three Garden.
Bufo woodhoiisei Girard. Woodhouses
Toad. V. M. Tanner (1940a) Tropic, Esca-
lante, and Escalante River; Hay ward et al.
(1958) Navajo Mountain; Woodbury et al.
(1959) Rock Creek and Kane Creek; Russell
and Thompson (1964) Bryce Canvon; BYU
(1972-1973) Cottonwood 'Wash Spring, Re-
flection Canyon, and Tibbet Spring.
Hylidae (Treefrogs)
Hyla arenicolor Cope. Canyon Tree Frog.
V. M. Tanner (1940a) confluence of the Esca-
lante/Colorado River; Hayward et al. (1958)
Paria Valley; Woodbury et al. (1959) Bridge
Canyon and Rock Creek; BYU (1972-1973)
Driftwood Canyon and Three Garden.
Ranidae (True Frogs)
Rana pipens brachycephala Cope. West-
em Leopard Frog. V. M. Tanner (1940a) con-
fluence of the Escalante/Colorado Rivers;
Woodbury et al. (1959) Bridge Canyon, Hole-
in-the-Rock, Hidden Pas.sage, Rock Creek,
West Canyon, Padre Creek, and Warm
Creek; Russell and Thompson (1964) Br\ce
Canyon; BYU (1972) Escalante River, Reflec-
tion Canyon, and Ribbon Canvon.
Reptiles
Emydidae (Water and Bo.x Turtles)
Chrysemys picta belli Gray. Western
Painted Turtle. Woodbury et al. (1959) Rock
Creek, Labvrinth Canvon, and three miles
above Face Canyon; Miller (1966) Hole-in-
the-Rock.
Iguanidae (Iguanid Lizards)
Crotaphytus collaris subssp. Western C^ol-
lard Lizard. Records of C. c. haileyi Stejnegar
are: V. M. Tanner (1940a) Paria River and
Henrieville; Woodbury et al. (1959) Rainbow
Bridge and Rock Creek; W. W. Tanner
(1958a) collected specimens between Lee's
Ferry and Tuba City and indicated, "inter-
gradations of .subspecies C. c. haileyi and C.
c. auriceps occur in the region south of the
San Juan River and west of its confluence
with Colorado River." BYU (1971-1974) Sites
4 and 11, .south ba.se of Navajo Mountain.
Glen Canyon, and Nipple Bench. C. r. bicin-
tores Smith & W. W. Tanner was named
from specimens collected at Crossing-of-lhe-
Fathers. An additional collection of this ani-
mal was made by BYU (1971) east of Smoky
Mountain.
Crotaphytus wislizenii punctatus Baird &
Girard. Long-nosed Leopard Lizard. \'. .M.
Tanner (1940a) Willow Tank Spring; Hay-
ward et al. (1958) Navajo Mountain; Wood-
310
Great Basin Naturalist
Vol. 40, No. 4
bury et al. (1959) Last Chance and Lee's Fer-
ry; W. W. Tanner and Banta (1963) Hole-in-
the-Rock, Lone Rock, Willow Tank Spring,
Catstairs Canyon, Crossing-of-the-Fathers,
Navajo Mountain Trading Post, and Lower
Wahweap Creek; BYU (1971-1973) Sites 1,
14, and 27, and Cottonwood Wash. Tanner
and Banta (1977). W. W. Tanner (1980, pers.
comm.).
Sauromalus obesus multiforaminatus Tan-
ner & Avery. Upper Colorado River Chuck-
walla. V. M. Tanner (1940a) Warm Creek;
Hayward et al. (1958) Paria Valley; Wood-
bury et al. (1959) Rainbow Bridge, Rock
Creek, and Last Chance drainage; W. W.
Tanner and Avery (1964) Crossing-of-the-Fa-
thers, Hole-in-the-Rock, Warm Creek, and
one mile upstream from Glen Canyon Dam;
BYU (1971-1972) Navajo Creek, Warm
Creek Bay, Glen Canyon Dam, Grand Bench,
and Navajo Creek.
Holbrookia maculata approximans Baird.
Lesser Earless Lizard. BYU (1972) Site 20,
Tietso Spring, and 13 miles south of Page
along U.S. Highway 89.
Sceloporus magister cephaloflacus Tanner.
Orange-headed Desert Spiny-Lizard. V. M.
Tanner (1940a) Willow Tank Spring and
Wahweap Creek; W. W. Tanner (1954a) Kai-
parowits Plateau, Lone Rock, Catstairs Can-
yon, Escalante River, and 15 miles northwest
of Hole-in-the-Rock; Hayward et al. (1958)
Navajo Mountain; Woodbury et al. (1959)
Bridge Canvon, Rock Creek, and Kane
Creek; BYU (1971-1974) Sites 1, 2, 8, 12, 14,
16, 17, and 23, 5 miles south of Page, Tibbet
Canyon, Tibbet Spring, Wahweap Bay area,
Cottonwood Wash/U.S. 89, and Three Gar-
den.
Sceloporus undulatus elongatus Stejnegar.
Northern Plateau Lizard. V. M. Tanner
(1940a) Cannonville, Escalante, and Calf
Creek/Escalante River; W. W. Tanner
(1954d) Paria River drainage north of U.S.
Highway 89; Hayward et al. (1958) Navajo
Mountain; Russell and Thompson (1964)
Bryce Canyon; BYU (1973) Tibbet Canyon.
Sceloporus graciosus graciosus Baird & Gi-
rard. Great Basin Sagebrush Lizard. V. M.
Tanner (1930), Weight (1932), and Presnall
(1935) Bryce Canyon; V. M. Tanner (1940a)
Calf Creek/Escalante River, Tropic, and Es-
calante/Colorado Rivers; Havward et al.
(1958) Navajo Mountain; BYU (1971-1974)
south base of Navajo Mountain and at Sites 1,
10, 13, 14, 21,23, and 27.
Ufa stansburiana uniformis Pack & Tan-
ner. Upper Colorado Basin Side-blotched
Lizard. V. M. Tanner (1940) Escalante, Wil-
low Tank Spring, and Calf Creek/Escalante
River; W. W. Tanner (1954d) Paria River
drainage north of U.S. Highway 89; Hayward
et al. (1958) Navajo Mountain; Woodbury et
al. (1959) Rainbow Bridge Trail and lower
Bridge Canyon; Russell and Thompson (1964)
Brvce Canvon; BYU (1971-1974) Sites 1, 2, 4,
6, 7, 8, 9, 10,1 3, 14, 17, 18, 19, 20, 21, 22,
23, 27, 28, and 30, Glen Canyon City, Four
Mile Bench, Three Garden, and Tibbet Can-
yon.
Urosaurus ornata wrighti Schmidt. Colo-
rado Tree Lizard. W. W. Tanner (1954d)
Paria River drainage north of U.S. Highway
89; Hayward et al. (1958) Navajo Mountain;
Woodbury et al. (1959) Bridge Canyon, Rock
Creek, and Last Chance; BYU (1971) three
miles west of Site 19.
Phrynosoma douglasii hernandesi Girard.
Short-horned Lizard. V. M. Tanner (1930),
Weight (1932), and Presnall (1935) Bryce
Canyon; BYU (1972) Sites 14 and 23, and
Warm Creek Bay. W. W. Tanner (1975).
Phrynosotna platyrhinos calidairum Cope.
Sonoran Desert Horned Lizard. W. W. Tan-
ner (1954d) Paria River drainage north of
U.S. 89; Woodbury et al. (1959) Rock Creek
and Lee's Ferry; BYU (1971) Utah-Arizona
state line north of Page along U.S. Highway
89.
Xantusidae (Night Lizards)
Xantusia vigilis utahensis Tanner. Utah
Night Lizard. W. W. Tanner (1957) records
topotvpes taken from Trachyte Creek, Gar-
field Co., Utah; W. W. Tanner (1958b) in-
dicates this species possibly is found within
the area defined as the Kaiparowits Basin.
Teidae (Whiptail Lizards)
Cnemidophorus tigris Baird & Girard.
Western Whiptail. Two subspecies occur in
the Kaiparowits Basin, viz. C. t. tigris (Baird
& Girard) reported by V. M. Tanner (1930),
Weight (1932), and' Presnall (1935) Bryce
December 1980
Atwood et al.: Kaiparowits Vertebrates
311
Canyon and V. M. Tanner (1940a) Calf
Creek /Escalante and Paria River drainage;
and C. t. septentrionalis Burger reported by
Woodbury et al. (1959) from Rainbow Bridge
Trail, Beaver Creek, and lower Bridge Can-
yon, BYU (1971-1973) Sites 1, 2, 6,^^13, 16,
17, 19, 20, 22, 23, 27, and 30, and the south
base of Navajo Mountain.
Cnemidophorus velox Springer. Plateau
Whiptail. Woodbury et al. (1959) Rock
Creek; Schmidt (1953) lists C. sacki innotatus
from Kanab, Kane Countv.
Cnemidophorus sexlineatus perplexus
Baird & Girard, Six Lined Racerunner. V. M.
Tanner (1940a) Canonville and Escalante.
Scincidae (Skinks)
Eumeces skiltonianus utahensis Tanner.
Great Basin Skink. Russell and Thompson
(1964) Bryce Canyon beneath logs and stones.
Colubridae
Thamnophis cyrtopsis cyrtopsis Kennicott.
Western Black-necked Garter Snake.
Schmidt (1953) lists this species for Utah, Ari-
zona, southern Colorado, New Mexico, and
southward.
Thamnophis elegans vagrans Baird & Gi-
rard. Wandering Garter Snake. V. M. Tanner
(1930), Weight (1932), and Presnall (1935)
Bryce Canyon; V. M. Tanner (1940a) Tropic
and confluence Calf Creek /Escalante River;
Woodbury et al. (1959) Rock Creek.
Masticophis taeniatus taeniatus Hallow-
ell. Desert Striped Whipsnake. W. W'. Tan-
ner (1954d) Paria drainage north of U.S.
Highway 89; Woodbury et al. (1959) Beaver
Creek, Kaiparowits Plateau, and Hole-in-the-
Rock; Russell and Thompson (1964) Bryce
Canyon.
Salvadora hexalepis mojavensis Bogert.
Mojave Patch-nosed Snake. W. W. Tanner
(1953) near the old town site of Adairville,
Kane County, Utah; W. W. Tanner (1954c)
Wahweap Creek, one mile southeast of Lone
Rock; Hayward et al. (1958) Paria Valley;
Woodbury et al. (1959) Rock Creek; BYU
(1971) Paria River three miles south of U.S.
Highway 89 and at Glen Canyon City.
Arizona elegans philipi Klauber. Painted
Desert Glossy Snake. W. W. Tanner (1964)
14 miles south of Page, 9 miles west of Page,
and 2 and 5 miles west of the Paria River all
along U.S. Highway 89; BYU (1972) Sites 1
and 3, Arizona-Utah border between Page
and Glen Canyon City, and 5 miles northwest
of Page.
Pituophis meUinoleucus deserticola Stejne-
gar. Great Basin Gopher Snake. Presnall
(1935) Bryce Canyon; V. M. Tanner (1940a)
Tropic, Escalante, Wahweap Creek, and
W'illow Tank Spring; W. W. Tanner (1954d)
Paria River drainage north of U.S. Highway
89; Hayward et al. (1958) Navajo Moimtain;
BYU (1971-1973) Sites 1, 2, 14, 17, and 20.
Lampropeltis getulus californiac Blain-
ville. California King Snake. \'. M. Tanner
(1940a) 30 miles south of Escalante; W. W.
Tanner (1958b) undoubtedly extending well
into the upper Colorado Basin; BYU
(1971-1973) one observed dead on U.S. High-
way 89 by Glen Canyon City and another
collected west of Cockscomb Ridge near U.S.
Highway 89.
Rhinocheilus lecontei lecontei Baird & Gi-
rard. W'estern Long-nosed Snake. W. W.
Tanner (1964) Wahweap road at junction
with U.S. 89 and Buck Tank Draw.
Sonora semiannulata isozona Cope. West-
ern Ground Snake. BYU (1972) seven miles
southeast of Glen Canyon City on U.S. High-
way 89. The specimens collected by BYU ex-
tend the range of this subspecies into Kane
County and east almost to the Colorado
River.
Hypsiglena torquata deserticola Tanner.
Desert Night Snake. W. W. Tanner (1954b)
northeastern Kane County, Utah, in the area
south and east of the Vermillion Cliffs; Rus-
sell and Thompson (1964) Bryce Canyon.
Hypsiglena torquata loreala Tanner.
Plateau Spotted Night Snake. Woodbury et
al. (1959) Labyrinth Canyon; BYU (1973) Re-
flection Canyon.
Tantilla planiceps utahensis Blanchard.
Utah Black-headed Snake. W. W. Tanner
(1954c) Paria River drainage 38 miles east of
Kanab in Catstairs Canvon.
Crotalidae (Rattlesnakes)
Crotalus viridis lutosus Klauber. Great
Basin Rattlesnake. V. .\1. Tanner (1930),
Weight (1932), and Presnall (1935) Bryce
312
Great Basin Naturalist
Vol. 40, No. 4
Canyon; W. W. Tanner (1958a) indicates that
this subspecies ranges east at least to the
Paria River. Pritchett (1962) extended the
range to the plateau east of the Paria.
Crotalus viridis nuntius Klauber. Hopi
Rattlesnake. W. W. Tanner (1958a) southeast
portion of Navajo Mountain near the Utah-
Arizona border and at the confluence of Es-
calante River, Willow Tank Spring, and
Wahweap Creek; W. W. Tanner (1958a)
north Escalante/ Colorado River junction;
BYU (1972-1974) Sites 2, 3, 10, 14, 22, and
28, Tibbet Canyon, two miles southeast of
Glen Canyon, Tibbet Spring, and Paria,
Utah.
Crotalus viridis concolor Woodbury.
Midget-faded Rattlesnake. V. M. Tanner
(1940a) Tropic, Escalante River, Willow
Tank Spring, and Wahweap Creek; W. W.
Tanner (1958a) north of Escalante/Colorado
River junction; BYU (1972-1974) Sites 2, 3,
10, 14, 22, and 28, Tibbet Canyon, two miles
southeast of Glen Canyon City, Tibbet
Spring, and Paria, Utah.
Birds
Present avian classification systems have
divided Fringillidae into two families and
have separated them in their positions in the
phylogenetic order and have removed other
species such as the Bushtits {Psaltripanis)
from tlie families where they have been well
established. Additionally, new positions in
the phylogenetic order of a number of
Passerine families have been proposed. Hay-
ward et al. (1976) have discussed these di-
verse taxonomic changes in considerable de-
tail and have deviated from the Fifth Edition
of the AOU Check-list (1957) to conform
with some of the proposed changes. In view
of the present diversity of opinion regarding
avian classification and because the AOU
Check-list committee has not yet made a de-
cision on the proposed changes, we have fol-
lowed the Fifth Edition of the AOU Check-
list and its supplements.
Our ornithological data were collected
from July 1971 through Febniary 1974; the
number of individual birds seen on a monthly
or seasonal basis and the number seen in vari-
ous vegetational associations on a seasonal
basis are biased, since only two years data
were collected for the months of March
through June. Also no data were collected in
December 1972. The data are biased further
by the fact that not every observer deter-
mined numbers, nor did they always report
the vegetational associations in which they
saw the species. Furthermore, the vegeta-
tional associations were not sampled equally
for bird species composition and numbers;
some were sampled more intensively than
others. Therefore, our data pertaining to
avian seasonal population trends and their
preferences for certain vegetational associ-
ation are only suggestive. Additionally, since
birds usually show decided preferences for
the architectural structure of the vegetation
rather than the plant species composition of
specific associations, many of the plant asso-
ciations have been lumped into Pi7ius-Juni-
perus or Juniper us associations, often referred
to as woodland, riparian {Populus fremontii,
Salix, Tamarix) associations, and desert
shrubs. Additional data are available on habi-
tat relationships of the vertebrate animals
discussed here by comparing the sites (Fig. 1,
Table 1) where the animal was seen or col-
lected with the vegetational associations
found at that site. All river miles listed for
the Colorado River are reckoned from Lee's
Ferry.
In the following species accounts, numbers
followed by localities and dates refer to the
number of specimens taken.
Podicipedidae (Grebes)
Podiceps nigricollis C. L. Brehm. Eared
Grebe. BYU (1971) 50 seen at Warm Creek
Bay, 8 Nov.; BYU (1972) 12 seen at river mile
56, 7 on Lake Powell near mouth of San Juan
River, and 17 at Three Garden on Lake Pow-
ell, 2 Nov. Behle et al. (1958) regarded the
species as a regular spring migrant through
the Kanab area. Late fall resident on Lake
Powell.
Aechmophorus occidentalis (Lawrence).
Western Grebe. BYU (1971) Lake Powell 6
seen on Warm Creek Bay, 8 Nov. and one
seen on Wahweap Bay, 1 Dec. A total of 80
Western Grebes were counted at the follow-
ing localities of Lake Powell 2 Nov. 1972:
river miles 12, 14, 17, 18 and 20, Dangling
Rope Canyon, Driftwood Canyon, and
December 1980
Atwood et al.: Kaiparowits Vertebrates
313
Navajo Creek. A 29 June 1973 observation on
Lake Powell suggests either a straggler or
possible nesting. Late fall transient south-
eastern Utah (Behle 1960).
Podilymbus podiceps (Linnaeus). Pied-
billed Grebe. BYU (1971) on Lake Powell,
one seen at Warm Creek Bay, 8 Nov. and an-
other at Wahweap Bay, 1 Dec; BYU (1972)
36 were recorded at the following localities
on Lake Powell: Gregory Butte near Rock
Creek Bay, Dangling Rope Bay, and river
miles 12 and 15, 2 Nov. Late fall resident on
Lake Powell.
Pelecanidae (Pelicans)
Pelecanus erythrorhynchos Gmelin, White
Pelican. Presnall (1937) Bryce Canyon;
Woodbury and Russell (1945) three miles be-
low Rock Creek on Colorado River, 28 July
1937. Rare transient.
Ardeidae (Herons and Bitterns)
Ardea herodias Linnaeus. Great Blue He-
ron. Behle (1948) Aztec Creek, Rock Creek,
Last Chance, and river miles 3, 6, and 21;
Woodbury and Russell (1945) one at Lee's
Ferry, Aug. 1909 as recorded by the Nelsen
and Birdseye trip; Behle and Higgins (1959)
from previously unpublished observations of
Woodbury and Russell on the Rainbow
Bridge-Monument Valley expedition, 23
herons seen from 1 to 11 Aug. 1938; they
were seen from 2V2 miles below Lee's Ferry
to river mile 69, 3100-3200 ft; Behle (1960)
common from Hite to Lee's Ferry, May
through Oct.; BYU one at Warm Creek/Lake
Powell, 8 Nov. 1971 and 3 May 1972; BYU
(1973) one at Three Garden alcove, 21 Mar.
The presence of a nesting colony at river
mile 117 (Woodbury et al. 1959) indicates
that the species also nests in the area. Not un-
common resident between 21 Mar. and 8
Nov.
Egretta thiila (Molina). Snowy Egret.
Woodbury and Russell recorded in their un-
published field notes observing this species at
Forbidding Canyon along the Colorado River
18 July 1937 and seeing tracks on shore of
Colorado River between river miles 63-69
(Behle and Higgins 1959); Behle (1960) Hig-
gins saw 17 at Wahweap Creek and river
mile 17 and one at the dam site, 7 .\ug. 1958;
BYU (1972) one seen in Populus at Navajo
Creek, 27 Apr.; BYU (1973) one seen by Rob-
ert Whitmore at junction of Paria Riv-
er/Colorado River, 6 June (riparian vegeta-
tion), and 2 more were seen bv Whitmore
feeding from a sandbar in the Colorado River
one mile north of Lee's Ferrv, 15 June.
Spring and summer resident, possiblv breeds.
Nycticorax nycticorax (Linnaeus). Black-
crowned Night Heron. \\'oodbur\- and Rus-
sell (1945) specimen: river mile 64, 4 Aug.
1938 and six were seen in 65 miles, i.e., be-
tween river mile 63 and 2'/2 miles below
Lee's Ferry, between 4 and 11 .\ug. 1938;
Behle (1948) two were seen 16 .\pr. 1947 at
Crossing-of-the-Fathers near Kane Creek and
another pair seen at Wahweap Creek. None
were seen during the present investigation.
Summer and spring records suggest breeding.
Threskiornithidae (Ibises and Spoonbills)
Plegadis chihi (Vieillot). White-faced Ibis.
Behle and Higgins (1959) reported that ood-
bury saw two at river mile 13, 13 Sept. 1957.
Uncommon transient along the Colorado Riv-
er.
Anatidae (Swans, Geese, and Ducks)
Branta canadensis (Linnaeus). Canada
Goose. Grater (1947) Bryce Canyon; Behle
and Higgins (1959) noted that Woodbury saw
two at river mile 50, 12 Sept. 1957; Behle
and Higgins (1959) observed a pair of adults
with six young that were unable to fly at riv-
er mile 106, 16 July 1958; they considered
the species a common sinnmer resident and
transient in Glen Canyon. Sununer resident,
possiblv nesting near Lake Powell.
Anas platyrhynchos Linnaeus. .Mallard.
Behle (1948) six were seen near Hole-in-the-
Rock; BYU (1971) three were seen on a small
islet at Warm Creek/Lake Powell, 8 Nov.
Uncommon transient along the Colorado Riv-
er and Lake Powell.
Anas strepera Linnaeus. Gadwall. Behle
(1948) saw a pair at Hole-in-the-Rock, five at
the mouth of San Juan River, and two at
Wahweap Creek; BYU (1972) five were seen
on a pond at Grosvenor's Arch, 2 Mar. and
two at Lake Powell /Cottonwood Wash. 22
314
Great Basin Naturalist
Vol. 40, No. 4
July (Ephedra-gr ass); BYU (1973) three on a
pond near Coyote Creek, 5 June. Uncommon
spring and summer resident.
Anas acuta (Linnaeus). Pintail. Behle
(1948) six near Hole-in-the-Rock; BYU (1973)
two on a pond at Wiregrass Spring, 5 Nov.
Spring and fall transient.
Anas crecca (Linnaeus). Green-winged
Teal. Woodbury and Russell (1945) one, Col-
orado River between Rock Creek and Lee's
Ferry, 17 Aug. 1937; Behle and Higgins
(1959) Woodbury saw seven at river mile 82,
11 Sept. 1957; BYU (1972) three at Wahweap
Creek Bay on a rain pond, 30 Oct. Transient
Lake Powell and the Colorado River.
Anas discors Linnaeus. Blue-winged Teal.
Behle (1948) near Klondike Bar; Behle and
Higgins (1959) Higgins saw six near mouth of
Rock Creek at river mile 58, 28 July 1958.
Uncommon spring and summer transient Col-
orado River and Lake Powell.
Anas cyanoptera Vieillot. Cinnamon Teal.
Woodbury and Russell (1945) one,
Paria /Colorado Rivers, 11 Aug. 1938; Behle
(1948) mouth of Wahweap Creek; Behle et
al. (1958) found one dead on highway south
of Escalante, 8 May 1954; Behle and Higgins
(1959) reported that in 1958 Woodbury saw
one between river mile 63 and 50, 5 Aug. and
one at V2 mile below Lee's Ferry, 11 Aug.
Spring and summer transient.
Anas americana Gmelin. American Wid-
geon. BYU (1972) two at Rainbow Landing
on Lake Powell, 2 Nov. Uncommon transient
on Lake Powell.
Aythyinae (Diving Ducks)
Aythya valisineria (Wilson). Canvasback.
Behle (1948) near mouth of Last Chance
Creek. Uncommon transient.
Bucephala clangula (Linnaeus). Common
Goldeneye. Behle (1948) saw 12 at Crossing-
of-the-Fathers, 12 at river mile 36, and 16 at
river mile 31; BYU (1972) one at Coyote
Creek Pond, 8 Dec. Spring and winter tran-
sient.
Bucephala albeola (Linnaeus). Bufflehead.
BYU (1972) five on Lake Powell at Warm
Creek, 7 Feb.; three on gravel pit pond at
Wahweap Bay, 27 Mar.; and a female on
Colorado River at Last Chance Bay, 5 July.
Uncommon spring and summer transient.
Cathartidae (American Vultures)
Cathartes aura teter Friedmann. Turkey
Vulture. Presnall (1934) Bryce Canyon; Behle
and Higgins (1959) reported that Woodbury
saw two along the Colorado River between
miles 13 and zero, 10 Aug. 1938; Woodbury
and Russell (1945) one, Navajo Mountain
Trading Post on 26 July 1936 and another on
8 Aug. 1936 near Lee's Ferry, and observed
on Kaiparowits Plateau; BYU (1971) two seen
12 miles south Page, Ariz., 24 July; BYU
(1972) Site 2 (flying), 23 July; Site 12, 16 Apr.
1972 (Coleogyne); Site 20; Driftwood Can-
yon, 24 June; three seen at Dry Rock Creek,
3 May; and one the confluence of the San
Juan River/Lake Powell, 17 July. Woodbury
and Russell (1945) reported the Turkey Vul-
ture as a common summer resident almost
everywhere in the Navajo country area in the
1930s. They observed it nearly every day
from 1 June through 15 Sept. over a period
of years. Yet, present observations covering a
three-year span yielded fewer than a dozen
sightings, suggesting a drastic decline in num-
bers. This is particularly significant when one
considers, as noted by Woodbury and Russell
(1945), "that this conspicuous bird is seldom
missed in observations in both time and
space," a factor that tends to overemphasize
its occurrence by comparison with less con-
spicuous birds (Woodbury and Russell 1945).
Uncommon summer resident from 28 Mar. to
1 Oct. (in the general area).
Accipitridae (Eagles, Hawks, and Harriers)
Accipiter gentilis atricapillus (Wilson).
Goshawk. Woodbury and Russell (1945) Nav-
ajo Mountain, i.e.. War God Spring, 23 July
1936; Russell and Thompson (1964) Bryce
Canyon. Uncommon summer resident.
Accipiter striatus velox (Wilson). Sharp-
shinned Hawk. Presnall (1934) Bryce Point;
Woodbury and Russell (1945) Navajo Moun-
tain, 13 July 1933, and Kaiparowits Plateau,
30 July 1937, and young birds out of nest
being fed by parents on Kaiparowits Plateau,
5-6 Aug. 1937; Benson (1935) base of Navajo
Mountain, mid- June 1933; BYU (1971) Site 8,
1 Aug. and one Navajo Mountain, 13 Oct.
Fairly common summer resident and spring
and fall transient.
December 1980
Atwood et al.: Kaiparowits Vertebrates
315
Accipiter cooperii (Bonaparte). Cooper's
Hawk. Presnall (1934) and Grater (1947)
Bryce Canyon; Woodbury and Russell (1945)
specimens: Navajo Mountain, 9 Aug. 1935
and 25 July 1936 and two seen nesting on
Kaiparowits Plateau, 5 Aug. 1937; Behle and
Higgins (1959) Rock Creek river mile 58, 28
July 1958; BYU (1971) two near Site 1 in
Jiinipenis, 14 and 18 July; BYU (1973) one at
Wahweap Creek, 30 Apr. {Tamarix in Salt-
wash). Summer resident, spring and fall tran-
sient.
Buteo jamaicensis calurus (Cassin). Red-
tailed Hawk. Presnall (1934) Bryce Canyon;
Woodbury and Russell (1945) two at Navajo
Mountain, 7 July 1936 at 3250 ft and seen at
Beaver Creek, War God Spring, Soldier Seep
(Navajo Mountain), and on Kaiparowits
Plateau; Behle and Higgins (1959) reported
that Woodbury found them common along
Colorado River, 4-23 July 1936, and one nest
with young hawks was also noted; Behle and
Higgins (1959) abundant in Glen Canyon, 1
July to 9 Aug. 1958. BYU (1971-1973") Sites
1, 2, 6, 12, 13, 15, 18, 20, 22, 23, 27, and 30,
and Cottonwood Wash, Tibbet Canyon, Ce-
dar Mountain, Grosvenor Arch, Wahweap
Creek, and specimen: Cockscomb /Highway
89, 14 Jan. 1972. Recorded every month;
April-July (21 seen) and November-Januarv
(17 seen). The months January-June are un-
fairly represented since no records were ob-
tained for these months in 1971. During the
winter months most observations were in
Juniperus and associated species (17), and the
remainder were in grass (2), grass-shrub (2),
perched on a ledge (2), and in Tamarix (1).
During the spring and summer nesting season
they were seen flying over desert shrubs (9)
consisting of Atriplex, Vanclevea, Chryso-
thamnus, Artemisia, and shrub-grass. Four of
the nine were in Coleogijne; three others
were in Juniperus and/or woodland, and one
each in grass and washbottom situations. Per-
manent resident.
Buteo swainsoni Bonaparte. Swainson's
Hawk. Presnall (1934) reported this species
to be common at Bryce Canyon during the
summer.
Buteo lagopus (Pontoppidan). Rough-
legged Hawk. In 1935 Long (1937) observed
one in Bryce Canyon. Sparse winter resident.
Buteo regalis (Gray). Ferruginous Hawk.
Long (1937) and Russell and Thompson
(1964) Bryce; BYU (1973) one perched on
power lines south of Glen Canyon near
Warm Creek, 27 Apr. Uncommon transient.
Aquila chrysaetos canadimsis (Linnaeus).
Golden Eagle. Presnall (1934) Rainbow
Point; Grater (1947) Bryce Canyon; Wood-
bury and Russell (1945) Navajo Mountain and
Glen Canyon; Behle and Higgins (1959)
sighted adults and immatures near mouth of
Aztec Creek, 26 July 1958; BYU (1971-1974)
Sites 1, 6, 12, and 23, Glen Canyon City.
Church Wells, Cockscomb, Grosvenor Arch,
Last Chance Creek, Tibbet Canyon, Wah-
weap Creek, and Warm Creek. Recorded 42
times at 26 locations during every month of
the year except October. Fewest birds were
seen October to November and .March to
April, and most May- August followed bv De-
cember-Febniary. Only one eagle was seen
during our study, in January. The vegeta-
tional type over which they were flving or in
which they were perched during Janu-
ary-June consisted of desert shrubs (6), cliffs
or rock ledges (4), grasslands (2), woodland
(1), and washbottom (1). In August, Septem-
ber, and December they were recorded in
mixed shrubs and cliffs (4). Common per-
manent resident.
Haliaeetus leucocephalus (Linnaeus). Bald
Eagle. Behle and Higgins (1959) report that
Woodbury observed one flying on the east
side of the Kaiparowits Plateau below the
cliffs north of Glen Canyon, 4 Aug. 1938;
BYU (1972) one seen near Rock Creek Bay
and another at confluence of San Juan Riv-
er/Lake Powell, 2 Nov.; BYU (1975) Last
Chance Wash, Dec. Sparse fall transient; the
4 Aug. bird seen by Woodbury may haNC
been a postnesting eagle from the small nest-
ing populations in central Arizona.
Circus cyaneus hudsonius (Linnaeus).
Marsh Hawk. Woodbury and Russell il945^
one, Kaiparowits Plateau, 9 Aug. 1937; BYU
(1971) two at Site 17, 27 Aug. (gras.slands);
two at Site 20, 6 Nov. (grassland): BYU (1972)
specimen (male): Wahweap Greek. 17 Jan.;
three seen at Wahweap Creek near Glen
Canyon City, 7 Feb., 18 Apr., and 3 July
(washbottom). Uncommon permanent resi-
dent; more common spring and fall transient.
316
Great Basin Naturalist
Vol. 40, No. 4
Falconidae (Falcons)
Falco mexicanus Schlegel. Prairie Falcon.
Presnall (1934) and Russell and Thompson
(1964) Bryce Point; Woodbury and Russell
(1945) Navajo Mountain; one seen in Glen
Canyon in 1938 between river miles 41 and
25, 8 Aug.; and three on cliffs between miles
13 to zero, 10 Aug.; Behle and Higgins (1959)
report that in 1938 the species was seen by
Woodbury in two places between river miles
78 and zero, 4 July; Behle (1948) pairs (prob-
ably nesting) near mouth of Escalante River
and junction of Bridge and Aztec Canyons;
BYU (1971) five sightings; Site 3, 16 Aug.
(grasslands); Site 15, no date; Glen Canyon
City/Wahweap Creek, 1 June 1972 (grass-
lands); Church Wells, 14 Nov. 1971 (grass-
lands); and Nipple Creek (chasing dove), 23
May 1973 (washbottom). Common summer
resident, less common in spring and fall;
probably a permanent resident.
Falco peregrinus anatum (Bonaparte).
Peregrine Falcon. Russell and Thompson
(1964) Bryce Canyon; Woodbury and Russell
(1945) saw peregrines at Navajo Mountain at
three different sites, 4-10 July 1936, and at
Beaver Creek, Navajo Mountain, 2-16 Aug.
1936; Behle (1960) Glen Canyon near Wah-
weap Creek, river mile 17, 6 Aug. 1958; BYU
(1971) two observations believed to be per-
egrines (both uncertain) at Site 15, 8 Oct. Un-
doubtedly the species formerly nested in
Glen Canyon, along the Colorado River, and
its side canyons as well as on Navajo Moun-
tain. With the construction of Lake Powell,
the area may now be more suited for pere-
grines than formerly (Porter and White
1973). Since early investigators did not exam-
ine the area specifically for peregrines, they
probably underestimated the size of the
population. The entire area now needs to be
thoroughly surveyed, especially Lake Powell.
Falco columbarius bendirei Swann. Mer-
lin. Grater (1947) Bryce Canyon.
Falco sparverius sparverius Linnaeus.
American Kestrel. Presnall (1934) Bryce Can-
yon; Benson (1935) Navajo Mountain; Wood-
bury and Russell (1945) two 4 Aug. 1935 and
11 July 1936 (9,500-10,000 ft, Picea-Ahies
and Pinus flexilis), and seen on Kaiparowits
Plateau and at Lee's Ferry; Behle et al.
(1958) one seen 10 miles south of Escalante in
stand of Juniperus, 6 May 1954; Hayward et
al. (1958) Escalante drainage along stream-
sides; BYU (1971-1974) Sites 1, 2, 3, 7, 10,
12, 14, 16, 19, and 26; one mile south of Glen
Canyon City, two specimens: 26 Aug. and 13
Sept. 1971; Brigham Plains, Cottonwood
Wash, Grosvenor Arch, Smoky Mountain,
Driftwood Canyon, Tibbet Canyon, Church
Wells, Crosby Canyon (nesting on cliff face,
28 Apr. 1973), Lee's Ferry, and Wahweap
Creek. Very common (over 75 birds seen at
31 sites); earliest spring sightings, 20 March
1972 (1 Apr. 1973) and latest fall sightings, 7
Nov. 1971 (3 Aug. 1972 and 9 Aug. 1973).
The greatest numbers were seen in April (16),
June (17), and July (22). Most observations
were in desert shrubs in March-July (29).
They were also seen in Tamarix (2), Juniperus
(1), saltwash (1), Juniperus in March and
April (4), grassland (4), and Tamarix in
May- July. The shrubs represented among the
desert shrubs were Vanclevea, Atriplex, Graij-
ia, Chrifsothamnus, Artemisia tridentata, Co-
leogyne, and Epiiedra. Summer resident, a
few may winter.
Tetraonidae (Grouse)
Dendragapus obscurus (Say). Blue Grouse.
Presnall (1934), Grater (1947), and Russell
and Thompson (1964) Bryce Canyon. Per-
manent resident.
Centrocercus urophasianus urophasianus
(Bonaparte). Sage Grouse. Russell and
Thompson (1964) Bryce Canyon. Permanent
resident.
Phasianidae (Chukars, Pheasants, and Quail)
Lophortyx garnbelii gambelii Gambel.
Gambel's Quail. Hayward et al. (1958) Calf
Creek area; BYU (1971) one at Site 13, 15
Oct. {Coleogyne) and five birds in Cotton-
wood Wash; also specimen, 16 Dec. Uncom-
mon permanent resident.
Phasianus colchicus Linnaeus. Ring-
necked Pheasant. Behle et al. (1958) observed
and heard in the fields around Escalante, 7
and 8 May 1954. Uncommon permanent resi-
dent near areas of cultivation.
Alectoris chukar (Gray). Chukar. BYU
(1972) Cottonwood Wash. Sparse permanent
resident.
December 1980
Atwood et al.: Kaiparowits Vertebrates
317
Rallidae (Rails, Gallinules, and Coots)
Fulica americana Gmelin. American Coot.
Woodbury and Russell (1945) below mouth
of Bridge Canyon, 15 July 1936; Behle and
Higgins (1959) river mile 47, flushed from S«-
hx, 30 July 1958; BYU (1971) on Wahweap
Creek, three miles east of Glen Canyon Citv,
6 Nov. {Tamarix around pond); BYU (1973)
Site 14, 9 May. Uncommon summer, spring,
and fall resident.
Charadriidae (Plovers)
Charadrius vociferus vociferus Linnaeus.
Killdeer. Woodbury and Russell (1945) one,
Kaiparowits Plateau, 11 Aug. 1937 (pond,
7000 ft); Behle et al. (1958) confluence Calf
Creek/Escalante River; BYU (1971) two,
north of Church Wells, 28 Sept.; BYU (1972)
two near Glen Canyon City, 14 July and two
at Wahweap Creek, 13 June (Tamarix); BYU
(1973) one at Coyote Creek, 5 June. Uncom-
mon summer resident and spring and fall
transient.
Scolopacidae (Sandpipers and Willet)
Actitis macularia (Linnaeus). Spotted
Sandpiper. Woodbury and Russell (1945) one.
Rock Creek /Colorado River, 20 July 1937
(3300 ft), and as reported from their impub-
lished field notes (Behle and Higgins 1959)
they observed 17 individuals from river mile
44 to Lee's Ferry, 7-10 Aug. 1938; Behle and
Higgins (1959) common along Colorado
throughout Glen Canyon during summer of
1958 and nesting along many of the smaller
streams; Behle et al. (1958) confluence Calf
Creek/Escalante River; BYU (1973) one at
Coyote Creek Pond, 5 June and 2 at junction
Paria/ Colorado Rivers, 6 June. Formerly a
common summer resident, this species seem-
ingly has declined in numbers since the con-
struction of the Glen Canyon Dam and the
filling of the reservoir.
Catoptrophorus semipalmatus inornatus
(Brewster). Willet. Presnall (1937) Bryce
Canyon; BYU (1972-1973) Four Mile Bench,
no date. Sparse transient.
Calidris mauri (Cabanis). Western Sand-
piper. Woodbury and Russell (1945) near
Lee's Ferry, 1 1 Aug. 1938. Sparse transient.
Phalaropodidae (Phalaropes)
Steganopus tricolor \'ieilIot. Wilson's
Phalarope. Woodbury and lUissell (1945)
specimen from flock on Kaiparowits Plateau.
2-3 Aug. 1938. Sparse fall transient.
Laridae (Gulls and Terns)
Larus californicus Lawrence. California
Gull. BYU (1971) Warm Ocek Bav. 8 .Nov.;
BYU (1973) Lone Rock Ba>, 8 Aug. Sparse
fall transient.
Lams delawaremis Ord. Ring-bilkd (iiili.
Behle (1948) three at Last Chance Creek and
again at the mouth of Kane Creek, on the
Colorado River. Sparse spring transient.
Columbidae (Doves and Pigeons)
Columba fasciata fasciata Say. Band-
tailed Pigeon. Benson (1935) Navajo Moun-
tain; Behle et al. (1958) Bryce Canvon. Sum-
mer resident.
Zenaida macroura (Linnaeus), .Mourning
Dove. Presnall (1934) Br\ce Canvon; Benson
(1935) Navajo Mountain; Tanner (1940a) Es-
calante River drainage; Woodburv and Rus-
sell (1945) Rock Creek, 24-26 July; Kaiparo-
wits Plateau, 3 Aug. (1937); and War (iod
Springs at Navajo Mountain, 16 .Vug. 1935.
common breeder up to 9000 ft elevation on
Navajo Mountain; Behle et al. (1958) con-
fluence of Calf Oeek/Escalante River; Behle
(1960) abundant Coyote Gulch (Whitney)
north side of Kaiparowits Plateau. Davis
Gulch, and along Escalante River 55 miles
southeast of Escalante, Aug. 1957; Behle and
Higgins (1959) abundant in Glen (.'anyon.
summer 1958; BYU (1971-1973) specinK-n;
one mile west Site 2 on 23 July 1971, Sites 1.
6, 10, 13, 14, 19, 28, 30. and 34. Glen Canyon
City, Cannonville, Cottonwood Wash.
Grosvenor Arch, Tibbet Sprinu, Lee's Ferry.
Nipple Creek, Wahweap Creek. Warm
Creek, Driftwood Canyon, and Hiree (harden
one mile above confluence of .San Juan with
Lake Powell. During the .\pril and .May mi-
gration period most Mourning Doves were
encountered in washbottoins 102'. 5 were
seen in grasslands, 14 in desert shrubs, and 4
were distributed more evenly among varioas
vegetational types; in contrast, during fall
318
Great Basin Naturalist
Vol. 40, No. 4
migration fewest were seen in washbottoms.
For example, 24 were seen in grasslands, 22
on desert shrubs, 21 in Popidus, 12 in Tama-
rix, 15 in woodland, 6 in hanging gardens, 3
in open fields, 2 in rocks, and 2 in washbot-
toms. In AugiLst-November grasslands were
most used where 39 individuals were seen,
followed by desert shrubs (27) and woodland
(2). This species was sighted 155 times with
1335 individuals recorded. The earliest spring
record was 15 Apr. (1972) and the latest fall
record was 1 Nov. (1973). The peak of spring
and fall, passage through the area was in May
and in August, respectively. The monthly to-
tals for 1972 and 1973 combined are as fol-
lows: Apr. (29), May (121), June (72), July
(68), Aug. (463), Sept. (109), Oct. (197), and
Nov. (2). Abundant summer resident.
Strigidae (Typical Owls)
Otus asio (Linnaeus). Screech Owl. Russell
and Thompson (1964) Bryce Canyon. Per-
manent resident.
Otus flammeolus (Kaup). Flam mula ted
Owl. Woodbury (1939) indicates that during
July 1936 H. N. Russell, Jr., collected a speci-
men in Salix near War God Spring on Navajo
Mountain, 6 July 1936. Others were seen or
heard calling from 6, 12, 20, and 21 July;
they made additional observations at Beaver
Creek, eight miles north of Navajo Mountain,
8 and 9 Aug. Resident, breeding in Piniis
ponderosa and pygmy conifers.
Bubo virginianus pallescens Stone. Great
Homed Owl. Presnall (1934) and Behle et al.
(1958) Bryce Canyon; Woodbury and Russell
(1945) Navajo Mountain at 9000 ft; BYU
(1971-1973) specimen: Site 1, near Glen
Canyon City, 29 Aug. 1971; Site 1, 16 Feb.
1972 (grasslands); Tibbet Spring, 15 Apr.
1972 {Populus fremontii, washbottom); BYU
(1973) Four Mile Bench, 13 May. Permanent
resident.
Glaucidium gnoma californicum Sclater.
Pygmy Owl. Russell and Thompson (1964)
Bryce Canyon. Permanent resident.
Athene cunicularia hypugaea (Bonaparte).
Burrowing Owl. Phillips et al. (1964) south-
east portion of the Kaiparowits Basin; BYU
(1972) Cottonwood Wash, 9 Aug. (grassland);
five miles west of Glen Canyon City, 4 June
and 21 July; four miles west of Glen Canyon
City {Artemisia fUifoIio) sitting in burrow on
side of road; U.S. 89/Cottonwood Wash
Road, 22 and 23 July; BYU (1973) three to
four miles west of Glen Canyon City, 5 June
(open grassland) nest under construction
nearby; BYU (1974) Utah/Arizona state line
north of Page on U.S. 89. Uncommon sum-
mer resident.
Strix occidentalis lucida (Nelson). Spotted
Owl. Woodbury (1939) reported that this
species was collected 13 Aug. 1936 in Navajo
Canyon by H. N. Russell, Jr.; Behle (1960)
Davis Gulch, a tributary of the Escalante
River, 55 miles southeast of Escalante, Aug.
1957; two were seen several times in a small
side canyon of Glen Canyon near river mile
101, 17 July, and another at the mouth of the
Escalante River, 19 July 1958. Uncommon
permanent resident.
Asio otus (Linnaeus). Long-eared Owl.
BYU (1973) Site 30, 9 June (grassland). Per-
manent resident.
Caprimulgidae (Goatsuckers)
Phalaenoptilus nuttallii nuttallii (Audu-
bon). Poorwill. Woodbury and Russell (1945)
Navajo Mountain Trading Post, 22 July 1936,
and Kaiparowits Plateau, 29 July and 3 Aug.
1937 (all in pygmy conifers); common in
Aug. On Navajo Mountain, nests were found
containing two downy young 23 July 1936 in
pygmy conifers, a family of three poorly
feathered juveniles in pygmy forest, Kaiparo-
wits Plateau, 7000 ft, 29 July 1937, and sim-
ilar family on 3 Aug. not far away in Arte-
uiisio-Quercus; Russell and Thompson (1964)
Bryce Canyon; BYU (1971) Paria Plateau, 29
Sept.; BYU (no dates) Site 3 and Church
Wells. A recent decline in numbers is evi-
denced by the paucity of sightings in the
present study as compared to the relative
abundance of the species in earlier studies.
Uncommon summer resident.
Chordeiles minor henryi Cassin. Common
Nighthawk. Presnall (1934) Bryce Point; Ben-
son (1935) Navajo Mountain; Tanner (1940a)
Escalante drainage; Woodbury and Russell
(1945) top Navajo Mountain, 14 July 1936;
BYU (1971-1973) Sites 2, 3, 4, and 28; Glen
Canyon Dam; one mile west, six miles east,
and 2.5 miles southwest of Grosvenor Arch;
and Four Mile Bench. Nighthawks were seen
December 1980
Atwood et al.: Kaiparowits Vertebrates
319
IS early as 19 May (1972) and as late as 5
3ct. (1972). Monthly distribution of observa-
:ions during our study is one in May, 8 in
[une, 26 in July, 37 in Aug., none in Sept.,
ind one in Oct. During June and Julv night-
lawks were recorded in the following vege-
:ation types and in the following niuubers:
3oleogyne (2), mixed shrub (1), Graijia (1),
'iiniperus (6), and grassland (7); from August
^n they were distributed as follows: Coleog-
/tie (1), grass (35), Juniperus (9), and Arte-
nisia (1). Common summer resident.
Apodidae (Swifts)
Aeronatites saxatalis saxatalis (Wood-
louse). White-throated Swift. Presnall (1934)
3ryce Canyon; Benson (1935) Navajo Moun-
:ain; Tanner (1940a) Kaiparowits Basin;
^oodburv and Russell (1945) one, Kaiparo-
wits Plateau, 27 Julv 1937 (7000 ft); Behle et
il. (1958) Escalante River/Calf Creek; Hay-
A'ard et al. (1958) Escalante drainage; Behle
1960) Coyote Gulch and Kaiparowits
Plateau, Aug. 1957; Behle and Higgins (1959)
^ive numerous observations for lower Glen
Hanyon from the field notes of Woodbury
ind Russell. In these notes a colony of swifts
was observed in a crevice of a cliff at the
iiouth of Rock Creek, 20-26 July 1937; six at
iiile 2 on the San Juan River, 1 Aug. 1938,
md others were seen between miles 41 and
15, 8 Aug., and at mile 25, 9 Aug. BYU
1971-1973) Sites 1, 2, 3, 4, 10, and 23, Cot-
:onwood Wash, Hackberry Canyon, Lee's
Ferry, Kelly Grade, Tibbet Spring, Wiregrass
spring, Reflection Canyon, Ribbon Canyon,
md Three Garden one mile above San Ju^n
•onfluence with Lake Powell. Although most
;wifts were seen near cliffs, those migrating
were seen away from cliffs and over various
<inds of vegetation, including Coleogyne,
^rass, Grayia, woodland, and Tamarix.
Where cliffs were near rivers or streams they
kvere seen above riparian streamside. In 1972
:he earliest spring sighting was 28 Apr.; in
1973 it was 1 May. The latest fall sighting
was 8 Aug. in 1972, when 150 birds were
»een. Sightings in April and August contain
large numbers of swifts (50-150). In May and
[une they were usually seen singly or in twos
3r threes; 9 were seen in one flock. In July
they were seen mostly in groups containing
manv birds. In earlv julv some sightings con-
tained 1, 2, 5, and 10 birds. From 1971-1973
the species was sighted 37 times and 324 in-
dividuals were recorded. Common summer
resident.
Trochilidae (Hummingbirds)
Archilochns alcxandri (Bourcier & Mul-
sant). Black-chinned Hununingbird. Presnall
(1934) Bryce Canyon; Benson (1935) one,
Navajo Mountain; Woodbury and Russell
(1945) one, Navajo Mountain, 9 July 1936
{Piniis ponderosa) and seen on the Kaiparo-
wits Plateau; Behle et al. (1958) nest with
eggs confluence of Calf Creek /Escalante Riv-
er; Behle (1960) Kaiparowits Plateau, Aug.
1957; BYU (1972) Tliree Garden, 25 June.
Summer resident.
Selasphorus platycercus platycercus
(Swainson). Broad-tailed Hununingbird. Pres-
nall (1934) and Russell and Thompson (1964)
Bryce Canyon; Benson (1935) Navajo Moun-
tain; Woodbury and Russell (1945) three,
Navajo Mountain, 17 July 1936 {Populiis
treinulokles), 9 Aug. 1935 {Finns ponderosa,
8,500-10,500 ft), and Kaiparowits Plateau, 1
Aug. 1937 (deciduous shrub, 7000 ft); Behle
(1948) Aztec Creek; Behle et al. (1958) con-
fluence of Calf Creek /Escalante River; Hay-
ward et al. (1958) Calf Creek area; Behle and
Higgins (1959) Aztec Creek, 26 July 1958;
BYU (1972) Reflection Canyon and Three
Garden. Summer resident; the species is
likely more common than these few sightings
suggest because a number of hummingbirds
were not identified and others were mis-
identified.
Selasphorus rtifus (Gmelin). Rufous Hum-
mingbird. Woodbury and Russell (1945) the
Nelson and Birdseye trip noted the species
from Fort Defiance to Lee's Ferry; Wood-
bury and Russell (1945) four, Navajo Moun-
tain, 9 Aug. 1935, and 11-17 July 1936, pon-
derosa zone, 7000-10,400 ft); Russell and
Thompson (1964) Bryce Canyon; Woodbury
and Russell (1945) found it mostly at altitudes
above 5300 feet. Summer resident.
Stellida calliope (Gould). C:alliope Hum-
ingbird. Behle et al. (1958) five miles north-
west of E.scalante; Behle (1960) one mile up
Aztec Creek from river mile 68.5. 26 July
320
Great Basin Naturalist
Vol. 40, No. 4
1958 (believed to be this species). Sparse
migrant.
Alcedinidae (Kingfishers)
Megaceryle alcyon (Linnaeus). Belted
Kingfisher. Behle (1960) one seen at the
Crossing-of-the-Fathers (mouth of Kane
Creek); river mile 41, by Clayton White, 2
Apr. 1954. Permanent resident, more com-
mon in summer.
Russell (1945) three, Navajo Mountain, 18
Aug. 1935; six, 16 July 1936 {Picea-Abies and
Pinus ponderosa between 8500-10,000 ft);
this expedition had nearly 50 records, chiefly
from Navajo Mountain, distributed through
June, July, and August. Summer resident.
Picoides pubescens leucurus (Hartlaub).
Downy Woodpecker. Long (1937), Grater
(1947), and Russell and Thompson (1964)
Bryce Canyon. Summer resident, descending
to river bottoms in winter (Behle 1960).
Picidae (Woodpeckers)
Colaptes auratus cafer (Linnaeus). Red-
shafted Flicker. Presnall (1934) Bryce Can-
yon; Benson (1935) Navajo Mountain; Wood-
bury and Russell (1945) Navajo Mountain, 5
Aug. 1934, 10 Aug. 1935, and 22 July 1936
(6700-9500 ft including pygmy forest, Pinus
ponderosa, and Picea-Abies zones; most com-
monly observed from 1 Mar. to 31 Oct.);
Behle et al. (1958) eight miles south of Esca-
lante, 20 Sept. 1935; BYU (1971) two speci-
mens: Site 1, 9 and 15 Oct.: lower Cedar
Moimtain, 7 Oct. (woodland); BYU (1972)
Site 2, one seen 11 Apr. {Jtinipems); one mile
east of Glen Canyon City, 28 Jan. (wood-
land); Cottonwood Wash, 29 Sept. (wash-
bottom); BYU (1973) Site 28, 13 June {Jiini-
penis); Four Mile Bench; Lee's Ferry;
Wiregrass Spring; and Tibbet Spring, 8 Feb.
(washbottom). Permanent resident, uncom-
mon.
Melanerpes lewis (Gray). Lewis' Wood-
pecker. Presnall (1934) Bryce Canyon. Sum-
mer resident.
Sphyrapicus variiis (Linnaeus). Yellow-
bellied Sapsucker. Presnall (1934) and Russell
and Thompson (1964) Bryce Canyon; Wood-
bury and Russell (1945) one, Navajo Moun-
tain, 23 July 1936, with other observations,
11-23 July; BYU (1971) Nipple Spring, 15
Oct. (riparian woodland); Site 12; specimen:
Site 15, 9 Oct. Summer reident and migrant.
Sphyrapicus thyroideus (Cassin). William-
son's Sapsucker. Presnall (1934) and Behle et
al. (1958) Bryce Canyon. Summer resident.
Picoides villosus leucothorectis (Oberhol-
ser). Hairy Woodpecker. Presnall (1934)
Bryce Canyon; Benson (1935) three, Navajo
Mountain; Behle et al. (1958) confluence of
Calf Creek /Escalante River; Woodburv and
Tyrannidae (Tyrant Flycatchers)
Tyrannus verticalis Say. Western King-
bird. Woodbury and Russell (1945) the Nel-
son and Birdseye trip recorded a few at Lee's
Ferry, 23-24 Aug. 1909; Woodbury and Rus-
sell (1945) two. Rock Creek /Colorado River,
22 July 1937 (streamside trees, 3300 ft); Kai-
parowits Plateau, 13 Aug. 1937 {Salix, 8000
ft); Hay ward et al. (1958) Escalante drainage;
Behle and Higgins (1959) several sightings in
lower Glen Canyon, early Aug. 1938; Behle
and Higgins (1959) field notes of Woodbury
and Russell for 1938, give the observation of
eight kingbirds on Colorado River between
miles 63 and 2V2 miles below Lee's Ferry,
5-11 Aug.; BYU (1971) Site 3, 13 July; BYU
(1972) Site 1, 3 Aug. {Jiiniperus); between
Sites 1 and 2, 3 June (Jtinipems); Site 3, 3
June (grass). Site 2, two seen 4 June {Jiini-
perus); Site 21, 24 June (woodland); BYU
(1973) Site 12, 9 May; Nipple Bench, 1 Aug.;
and Tibbet Spring, 16 June (Popuhis-Tama-
rix); Glen Canyon City, 28 Apr.; Cottonwood
Wash, 5 June (Taniarix); and Paria River/U.S.
89, three seen 6 June (stand of Populus).
Earliest spring date, 28 Apr. (1973); latest fall
date, 3 Aug. (1972). Common summer resi-
dent.
Tyrannus vociferous vociferous Swainson.
Cassin's Kingbird. Presnall (1934) and Russell
and Thompson (1964) Bryce Canyon; Wood-
bury and Russell (1945) two, Kaiparowits
Plateau, 13 Aug. 1937 (chaparral at 7000 ft)
and Rock Creek/Colorado River, 26 July
1937 (Querciis at 3300 ft); Behle et al. (1958)
one, 10 miles south of Escalante, 9 May 1937;
Behle (1960) frequently seen in side canyons
of Glen Canyon, 1 July to 9 Aug. 1958; BYU
(1973) Site 3, three believed to be this species
seen 6 Aug.; Cottonwood Wash, two mated
December 1980
Atwood et al.: Kaiparowits Vertebrates
321
pairs seen each day, 5 and 6 June (Popuhis)-,
Paria River/ U.S. Highway 89 and Lee's Fer-
ry, two seen each day, 6 and 7 June {PopuUis-
Salix-Tamarix). Summer resident.
Myiarchus tyrannulus (Muller). W'ied's
Crested Flycatcher. BYU (1973) this rare spe-
cies was seen in semiarid grassland several
times at Site 3 by Clyde Pritchett betwen 1
and 9 Aug.: two were seen 1 Aug., three on 3
Aug., one on 4 Aug., three on 6 Aug., and
two on 9 Aug. Behle and Perry (1975) report
this species as a rare summer resident only in
extreme southwestern Utah. Hayward et al.
(1976) cite only those records given bv Behle
and Perry. Thus, Pritchett's observations
seemingly extend the range of this species to
southeastern Utah. Sparse migrant.
Myiarchus cinerascens cinerascens (Law-
rence). Ash-throated Flycatcher. Presnall
(1934) Bryce Canyon; Benson (1935) Navajo
Mountain; Woodbury and Russell (1945) five.
Rock Creek/Colorado River, 21-26 July
1937 (canyon shrubs, 3300 ft) and three miles
north Navajo Mountain Trading Post, 12 July
1934 and 27 July 1936 (pygmy forest, 6000
ft); Behle et al. (1958) 10 miles south of Esca-
lante, 9 May 1937, and confluence of Calf
Creek/Escalante River; Behle and Higgins
(1959) give at least 13 observations from
Woodbury and Russell's field notes for the
Colorado River between river mile 75 and Vz
mile below Lee's Ferry, 3-10 Aug. 1938;
Higgins (Behle and Higgins 1959) found this
flycatcher to be very abundant along the
length of Glen Canyon and side canyons, 1
July to 9 Aug. 1958; Behle (1960) junction
Colorado and Escalante Rivers, river mile 88,
19 July 1958; BYU (1972) Site 1, 27 June and
31 July (Atriplex); Reflection Canyon, 4 July
(washbottom); Three Garden, 3 July (Hang-
ing Garden); confluence of San
Juan/Colorado Rivers, 24 June; BYU (1973) 8
miles north of U.S. 89 on Cottonwood Wash
road, 5 June (possibly nesting in Tdiiuirix)- 13
miles north U.S. Highway 89 on Cottonwood
Wash road, 8 June (Populu.s); Brigham Plains
Road, 8 June {Populus); Cow Camp on Four
Mile Bench, 13 June; Hackberry Canyon, 8
June (scattered bnish); 2.5 miles north Lee's
Ferry, 14 June (scattered brush); Paria River
(1 mile south of Paria), 7 June (dense Salix);
Tibbet Spring, three seen 5 June (open shrubs
and Populus).. and Wiregrass Spring, two seen
5 June (Taniarix). Common summer resident.
Sayornis nigricans (Swainson). Black
Phoebe. Woodbury and Russell (1945) Lee's
Ferry, observed bv E. W. Nelson, 23-26 Aug.
1909; Phillips et al. (1964) south central por-
tion of the Kaiparowits Basin. Occasional mi-
grant or summer resident.
Sayornis saya saya (Bonaparte). Say's
Phoebe. Benson (1935) Navajo Mountain,
21-24 June 1933; Tanner (1940a) Escalante
drainage; Woodbury and Russell (1945) sev-
en, Navajo Mountain, 13 Aug. 1935; Warm
Creek, 16 July 1936; Rock Creek, 21 July
1937; Kaiparowits Plateau, 4 Aug. 1937;
Navajo Mountain Trading Post, 27 July 1936
(pygmy forest, Salix, desert bnish, Populus
tremuioides, and Salix, 3190-9000 ft); Behle
et al. (1958) Escalante drainage; Behle and
Higgins (1959) three, Colorado/Escalante
River, 19 July 1958; common all along Glen
Canyon, summer 1958; they believed this tax-
on nested in cliffs and larger trees; one in
Warm Creek Canyon, 18 Oct. 1958 {Salix,
Tamarix, Baccharis, and Pltichea); in addition,
Behle and Higgins (1959) reported from the
Rainbow Bridge-Monument Valley expedi-
tion field notes, common along Colorado Riv-
er and tributaries, 4-22 July 1938, and over
29 seen between mile 75 and V2 mile below
Lee's Ferry, 4-11 Aug. 1938; Russell and
Thompson (1964) Bryce Canyon; BYU (1961)
specimen (male): Paria, 20 May; BYU
(1971-1973) one mile .south Site 2, 28 Aug.
1971; Sites 3, 7, 12, 13, and 23, Church
Wells, Cottonwood Wash at miles 5, 8, and
11 north of U.S. Highway 89, Paria River,
and Nipple Spring; one, four miles north of
Church Wells. Phoebes were sighted 13 times
(17 individuals) at 10 localities. Habitat in
which phoebes were seen, based on only six
of 13 sightings, consisted of a rocky cliff,
rockv areas, scattered bru.sh, Chrysothamnus,
and Populus. The earliest spring date was 5
June (1973). The latest fall observation was
30 Sept. (1971). Common summer resident.
Empidonax traillii (Audubon). Willow
Flycatcher. Woodbury and Russell (1945)
five, Navajo Mountain, 26 July 1936; Kai-
parowits Plateau, 3 Aug. 1937; Lee's Ferry
(nest), 11 Aug. 1938, and 25 Aug. 1909; and
two miles below San Juan/Colorado Rivers, 3
Aug. 1938 (Salix, Tamarix, and Quercus,
322
Great Basin Naturalist
Vol. 40, No. 4
3200-7000 ft); Phillips et al. (1964) south
central portion of the Kaiparowits Basin;
Higgins (Behle 1960) considered this species
common to stands of Salix, Tamarix, Bac-
charis, and Pluchea along Colorado River in
Glen Canyon, summer 1958; Woodbury and
Russell (1945) considered it primarily as a
bird of .streamside thickets; BYU (1973)
Hackberry Canyon, 8 June, male displaying
(Tamarix); Lee's Ferry, 7, 14, 15 June, singing
(Tamarix with Salix); and Paria /Colorado
Rivers, 7 and 14 June (dense Salix and Tama-
rix). Summer resident.
Empidonax oberholseri Phillips. Dusky
Flycatcher. Woodbury and Russell (1945)
several, Kaiparowits Plateau, 31 July; 6 Aug.
1937 (also female seen with three half-grown
nestlings); and 13 Aug. 1936 (nest in
Quercus); BYU (1973) one (female), Tibbet
Spring, 2 May. Summer resident.
Empidonax wrightii Baird. Gray Fly-
catcher. Woodbury and Russell (1945) one,
Navajo Mountain Trading Post, 26 July 1936
(pygmy conifers, 6500 ft); BYU (1973) two,
female, two miles east Glen Canyon City, 29
Apr. and male, Tibbet Spring, May. Summer
resident.
Empidonax difficilis hellmayri Brodkorb.
Western Flycatcher. Woodbury and Russell
(1945) Beaver Creek/Navajo Mountain area,
7 Aug. 1936 (Quercus, 6000 ft); Behle and
Higgins (1959) one, Kane Creek /Colorado
River, 2 Aug. 1958. Summer resident.
Contopus sordidulus veliei Coues. West-
em Wood Pewee. Presnall (1934) Bryce Can-
yon; Woodbury and Russell (1945) two, Nav-
ajo Mountain, 12 Aug. 1935 and 13 Aug. 1936
(Pinus ponderosa, 8500-9000 ft); Hayward et
al. (1958) Escalante drainage; BYU (1971)
Site 4 (Paria Narrows), 30 Sept. Uncommon
summer resident and migrant.
Nuttallornis borealis (Swainson). Olive-
sided Flycatcher. Presnall (1934) Bryce Can-
yon; Woodbury and Russell (1945) several,
Beaver Creek /Navajo Mountain area, 6 and 9
Aug. 1936, and War God Spring on Navajo
Mountain, 13 Aug. 1935; seen near Lee's Fer-
ry, 10 Aug. 1938; six birds observed at War
God Spring, 13 Aug. 1937 (Pinus ponderosa,
6500-9000 ft); Behle and Higgins (1959) seen
at river mile 3.5 and V2 mile below Lee's Fer-
ry; and BYU (1961) one, Paria, Utah, 20 May.
Summer resident.
Alaudidae (Larks)
Eremophila alpestris leucolaema Coues.
Horned Lark. Behle et al. (1958) 10 miles
southeast of Escalante, 8 May 1954; Russell
and Thompson (1964) Bryce Canyon; Behle
and Higgins (1959) Higgins saw flock near
mouth Kane Creek, river mile 40, 2 Aug.
1958; one, between Warm and Wahweap
Creeks, 25 March 1958 (3800 ft); BYU
(1971-1974) 4, (males) T 43S R2W S24, near
tank, 7 Dec. 1971; Sites 1, 3, 10, 12, 13, 15,
and 23; Warm Creek; Church Wells; Cotton-
wood Wash; Smoky Mountain; and Tibbet
Canyon; Summit Navajo Mountain; Esca-
lante Valley/Little Valley Road; three miles
north U.S. Highway 89 on Cottonwood Wash
Road. A total of 1639 larks or sightings were
seen during the study, at over 20 locations
and from 97 sightings. A number of sightings
were recorded as many or common. The spe-
cies is decidedly more frequent in the late
fall/winter months, i.e., November through
February, when at least 1220 were recorded.
In March through June only 87 individuals
were recorded, with only 3 in April and one
in May. The numbers increased to 32 in June.
In July and August they greatly increased
(165 individuals plus five flocks containing
many individuals). Only 11 were seen in Sep-
tember and 156 in October. A January flock
contained 600 birds. The species shows a
marked affinity for grasslands. During the
nonbreeding season of September-April, 482
individuals were recorded in grassland vege-
tation, compared to 75 in desert shrubs and
15 in saltwash. In May and June, 4 were seen
in grass and 18 in desert shrubs. In July and
August, 45 (one flock) were seen in Juni-
penis/ grass, and 12 in grass.
In April through June, Honied Larks were
seen either singly (11 of 16 sightings), in pairs
(2), or in small flocks containing 6 (2) or 8 (1)
birds. They congregated into larger groups or
flocks in July, half of which contained 10-15
birds; one, about 45 birds; one, 5 birds, and
three, one bird each. Additionally, four flocks
containing many birds were also encountered
for which numbers were not obtained. As the
season progressed the Honied Larks were
seen in increasingly larger flocks. For ex-
ample, from July through September the av-
erage number of birds per sighting was 9.6
December 1980
Atwood et al.: Kaiparowits Vertebrates
323
range 1-45); for October and November it
A'as 18.1 (range 10-30); and for December
;hrough March it was 53.2 (range 1-600).
During the non-nesting season, July through
Vlarch, only 7 larks were seen singly, and
)nly one sighting each contained 2 and 3
)irds; five contained 4 birds; three, 5 birds;
wo, 6 birds; one, 7 birds; two, 9 birds, seven,
10 birds; eight, 20 birds; one, 23 birds; one,
15 birds; four, 30 bird; three, 50 birds; two,
LOO birds; and one, 600 birds. Additionallv,
here were five flocks for which no coimt was
iiade. Common vear-roimd resident of the
Hinrndinidae (Swallows)
Tachycineta thalassina lepida Mearns.
/iolet-green Swallow. Presnall (1934) Bryce
Ilanyon; Benson (1935) Navajo Mountain;
^Voodbury and Russell (1945) one, four miles
lorth of Navajo Mountain Trading Post, 27
uly 1936 (pygmy conifers, 6000 ft) and nest
bund in old woodpecker hole, Navajo Moun-
ain; Behle (1948) Aztec Creek; Behle et al.
1958) confluence Calf Creek /Escalante Riv-
M-, 1954; Behle and Higgins (1959) Rainbow
3ridge-Monument Valley expedition field
lotes, seen frequently along Colorado River,
1-22 July; over 28 seen from river mile 69 to
iver mile 41, 4-8 Aug. 1938; seen frequently
hroughout Glen Canyon; Phillips et al.
1964) eastern part of the Kaiparowits Basin;
3YU (1972) Site 30, 23 Julv; Cedar Mountain,
U July {Ephedra); Site 20', 21 July; Site 16, 2
lime {Tcunarix); Cottonwood Wash, 26 June
Populus); Nipple Bench, 1 Aug.; San
uan/Colorado Rivers, 24 June; Tibbet
spring, 17 June (Tamarix); Four Mile Bench,
18 June {Artemisia); BYU (1973) Lee's Ferry,
15 June; 7 at Paria /Colorado Rivers, 14 June
Tamarix); Site 7, 4 July; Site 14, 10 May;
lear Site 28, 13 June {Juniperus/grsLSs); and
site 34, 5 June (open bnish). Summer resi-
dent.
Iridoprocne bicolor (Vieillot). Tree Swal-
ow. BYU (1973) Lee's Ferry, one seen 6
fune. Sparse migrant.
Riparia riparia riparia (Linnaeus). Bank
5wallow. Russell and Tliompson (1964) Bryce
Canyon. Summer resident.
Stelgidopteryx ruficollis (Vieillot). Rough-
winged Swallow. Woodbury and Russell
(1945) specimens at river mile 63, 4 Aug.
1938 and Lee's Ferry, 23-26 Aug. 1909;
Behle and Higgins (1959) Woodbury and
Russell's field notes from 1938, six observed
between San Juan /Colorado Rivers and five
miles up the San Juan, 2 Aug.; nine seen
along the Colorado River, 3-4 Aug.; BYU
(1971) Wahweap Creek, 10 Aug.; BYU (1973)
Paria/Colorado Rivers, two seen in Tamarix,
14 June. Summer resident.
Petrochelidon pyrrhonota pyrrhonota
(Vieillot). Cliff Swallow. Presnall (1934)
Bryce Canyon; Tanner (1940a) Kaiparowits
Basin; Behle et al. (1958) confluence Calf
Creek/Escalante River; Hayward et al.
(1958) Escalante drainage; Behle and Higgins
(1959) Higgins found it common in Glen
Canyon, nests seen attached high canyon
walls, July 1958; BYU (1973) Site 13, 11 June;
Site .30 nine seen, 8 June (nesting in rocks and
Juniperus); Nipple Spring, 12 June; and Tib-
bet Spring, 10 June. Fairly common summer
resident.
Corvidae (Jays, Magpies, Ravens,
and Nutcrackers)
Perisoreus canadensis capitalis (Ridgway.
Gray Jay. Presnall (1934) and Russell and
Thompson (1964) Bryce Canyon. Permanent
resident.
Cyanocitta stelleri macrolopha Baird. Stel-
ler's Jay. Presnall (1934) Bryce Canyon; Ben-
son (1935) four, Navajo Mountain, 13-20 July
1933; Woodbury and Russell (1945) several,
Navajo Mountain, 13 July 1933, 12 Aug.
1935, 10 July and 10 Aug. 1936, (numerous in
Pintis ponderosa, 9000 ft); Grater (1947) and
Russell and Thompson (1964) Bryce Canyon.
Permanent resident.
Aphelocoma coerulescens woodhouseii
(Baird). Scrub Jay. Benson (1935) Navajo
Mountain area; Long (1937) and Russell and
Thompson (1964) Bryce Canyon; Woodbury
and Russell (1945) four, Navajo Mountain, 2
and 25 Julv 1936 and Kaiparowits Plateau, 21
and 31 July 1937 {Pinus ponderosa, Populus
tremuloides, pygmy conifers, and Amelan-
chier, 6500-8500 ft); Tanner (1940a) Kaiparo-
wits Basin; Behle et al. (1958) Calf
Creek/Escalante River; Hayward et al.
(1958) Escalante drainage; BYU (1971) ^k
324
Great Basin Naturalist
Vol. 40, No. 4
mile south Site 1; BYU (1972) one mile south
Glen Canyon City. Permanent resident.
Pica pica hudsonia (Sabine), Black-billed
Magpie. Russell and Thompson (1964) Bryce
Canyon; BYU (1971) near Nipple Spring, 16
Oct. (riparian) and Wahweap Creek, four
seen 18 Dec. (Tatnanx); BYU (1972) Wah-
weap Creek, four seen 17 Jan. (salt wash) and
Glen Canyon City, 13 Feb. (city dump). The
magpie seemingly is absent during the spring
and summer, with a few moving into the area
during fall and winter.
Corvus corax sinuatus Wagler. Common
Raven. Presnall (1934) Bryce Canyon; Benson
(1935) one, Navajo Mountain; Woodbury and
Russell (1945) up to 10,000 feet elevation on
Navajo Mountain; Behle (1948) river mile 31;
Behle et al. (1958) 10 miles south of Esca-
lante, 8 May 1954; BYU (1971-1973) Sites 1,
2, 3, 4, 5, 10, 11, 19, 20, 21, and 22, Page,
Cottonwood Wash, Buckskin Gulch, Warm
Creek, Church Wells, Grand Bench, Grosve-
nor Arch, Smoky Mountain, Tibbet Canyon,
Wahweap Creek, Driftwood Canyon, San
Juan/Colorado Rivers, Reflection Canyon,
Lee's Ferry, Three Garden, one mile above
confluence of San Juan River and Lake Pow-
ell, and east of Lone Rock/Lake Powell.
Sight records are distributed throughout
every month of the year. The greatest num-
ber of individuals was seen during December
(85) and February (97) and the least during
January (6), April (7), and May (10). A total
of 407 individuals were counted during the
study. Of 164 sightings of ravens, 82 (50 per-
cent) of them were of single birds, 43 (26 per-
cent) were of pairs, 11 (7 percent) of triples,
and 15 (9 percent) were quadruples. Four
sightings contained 5 birds and four, 6 birds,
and one each consisted of flocks containing 7,
10, 25, and 60. The flocks containing 25 and
60 birds were at city dumps. Ravens were
seen in all vegetational types, but the obser-
vations were not distributed evenly among
them; of 178 individuals 57 percent were
seen in grasslands and 25 percent were seen
in desert shnibs. The remaining 32 birds were
seen in Juniperus (5 percent), washbottoms (4
percent), Populus (1 percent) and mis-
cellaneous situations (8 percent). Ninety-
eight additional ravens were seen at city
dumps. Permanent resident.
Gymnorhinus cyanocephaliis Wied. Pin-
yon Jay. Presnall (1934) Bryce Canyon;
Woodbury and Russell (1945) three, Kaiparo-
wits Plateau, 2 and 3 Aug. 1937, and Navajo
Mountain Trading Post, 31 July 1936 (pygmy
forest, 6500-7000 ft); Woodbury encountered
a flock of 100 birds on Kaiparowits Plateau, 4
Aug. 1937; Behle et al. (1958) eight miles
south of Escalante, 19 Sept. 1935; Hayward
et al. (1958) Escalante drainage; BYU
(1971-1973) Sites 1, 2, 3, 4, 14, 21, 27, 28, 29,
and 30; Cannonville; Tibbet Spring; and Cot-
tonwood Wash; Paria Plateau, 29 Sept.;
lower Cedar Mountain; five miles west Site
27. Pinyon Jays were recorded every month
of the year with the largest numbers in Aug.
(310). Numbers observed monthly were as
follows: 5 were seen in Sept., 14 in Oct., 6 in
Nov., 16 in Dec, 3 in Jan., 35 in Feb., 31 in
Mar., 22 in Apr., 8 plus one flock in May, 14
plus one flock in June, and 39 in July. A total
of 503 individuals (plus two flocks un-
counted) were seen from 1971 to February
1974. Most sightings (25) and individuals
(254) where vegetational type was recorded
were in pygmy conifers. Six sightings and 16
individuals were in desert shrubs consisting of
Coleogyne, mixed grass-shrubs, Grayia-grass,
and Artemisia. A sighting of 30 individuals
was in reseeded grass. Fourteen of the 52
sightings were of single birds; 4 were of
doubles; 6 contained 3 to 5 birds; 8, 6 to 10
birds; 3, 15 birds; 2 each, 20 and 30 birds;
and one each consisted of 16, 17, 35, 37, and
100 birds. Permanent resident.
Nucifraga columbiana (Wilson). Clark's
Nutcracker. Presnall (1934) Bryce Canyon;
Benson (1935) specimen: Navajo Moun-
tain/Bridge Canyon, 13 July 1933 (seen
everyday in mid-June 1933); Woodbury and
Russell (1945) six specimens: Navajo Moun-
tain, July 1936-37; Colorado River side can-
yon next below Bridge Canyon, 13 July 1936
{Popiihis, Pinus ponderosa, and Picea-Ahies
forest from 3200 to 10,000 ft); BYU (1971) in
Picea-Ahies at summit Navajo Mountain, 13
Oct. Permanent resident higher mountains,
moves lower after nesting.
Paridae (Bushtits, Chickadees, and Titmice)
Parus atricapiUus garrintis Behle. Black-
capped Chickadee. Russell and Thompson
December 1980
Atwood et al.: Kaiparovvits Vertebrates
325
(1964) Bryce Canyon; Presnall (1934) also re-
ported P. a. septentrionalis in Bryce Canyon.
However, Behle et al. (1958) and Behle and
Perry (1975) place the area of Utah jnst west
of the Kaiparowits Plateau in a zone of in-
tegradation between P. a. nevadensis of west-
ern Utah and P. a. gairinus of extreme east-
em Utah. Permanent resident.
Pariis gamheli Ridgway. Mountain Chick-
adee. Presnall (1934) Bryce Canyon; Wood-
bury and Russell (1945) three at Kaiparowits
Plateau, 5-6 and 12 Aug. 1937, and one at
Navajo Mountain, 10 July 1936 {Populus
tremuhides, Pinus ponderosa, and pvgmv
conifers, 7000-9500 ft); BYU (1971) Lake
Powell /Wann Creek inlet, 8 Nov. and sum-
mit of Navajo Mountain, 12 Oct. Behle
(1960) described the race wasatchensis,
which included the population of the Kai-
parowits Basin. Hayward et al. (1976) follow
Snow (1967a), who considers wasatchensis to
be a synonym of inyoensis. Permanent resi-
dent of mountains, moves lower in winter.
Parus iyiornatus ridgwayi Richmond.
Plain Titmouse. Behle et al. (1958) con-
fluence of Calf Creek /Escalante River; Rus-
sell and Thompson (1964) Bryce Canyon;
BYU (1971) specimen: Paria Plateau! 29
Sept.; BYU (1972) Sites 3 and 21, 25 June
(pygmy conifers) and Navajo Mountain. P. i.
griseus which is now a synonym of P. i. ridg-
wayi was listed by Presnall (1934) for Bryce
Canvon and collected bv Woodburv and Rus-
sell (1945) four, Kaiparowits Plateau, 31 July
1937 and the Navajo Mountain area, 26 Julv
1936 (pygmy conifers, 6500-7000 ft). This
species should be more common in the pyg-
my conifers than our few observations sug-
gest. Woodbury and Russell (1945) reported
it in pygmy conifers every month of the year;
they considered it a permanent resident.
Psaltriparus minimum (Townsend). Com-
mon Bushtit. Long (1937) and Russell and
Thompson (1964) Bryce Canyon; Woodbury
and Russell (1945) six, Kaiparowits Plateau, 1
Aug. 1937, and Navajo Mountain, 8, 24, and
31 July 1936 and 14 Aug. 1937; Hayward et
al. (1958) benches of the Escalante drainage;
BYU (1971) one, T43S, R2W Sec 24, 7 Dec;
Site 2, 30 .seen 7 Dec. (woodland); also seen
in Tibbet Canyon, 11 Dec. (ChrysotJuimniis).
Permanent resident. This taxon is placed in
Aegithalidae hy Snow (1967b).
Sittidae (Nuthatches)
Sitta carolinensis nelsoni Mearns. White-
breasted Nuthatch. Presnall (1933, 1934, and
1936) and Russell and Thompson (1964)
Bryce Canyon; Benson (1935) three, Navajo
Mountain, 13 Aug. 1935 (mixed coniferous
forest, 9()()() ft). Woodbury and Russell (1945)
found this species a regular, but not abun-
dant, inhabitant of the pygmy forest, Pinus
ponderosa zone, and more sparingly in the
Picea-Ahics forests. Permanent resident.
Sitta canadensis Linnaeus. Red-breasted
Nuthatch. Presnall (1934 and 1936) Bryce
Canyon and Woodbury and Rus.sell (1945)
three, Navajo Mountain, 14 July 1936, and
Kaiparowits Plateau, 11 Aug. 1937 {Picea-
Ahies-Populus zone at 7000-10,400 ft); saw
one to six birds each day in Picea-Abies forest
on Navajo Mountain, 6' and 13-18 July 1936.
Woodbury and Russell (1945) believed that
the species probably nests in Picea-Abies for-
est of mountain tops and canyon heads and
spreads to other habitats following nesting
season. Permanent resident.
Sitta pygmaea melanotis van Rossem. Pyg-
my Nuthatch. Presnall (1934) Bryce Canyon;
Benson (1935) two, Navajo Mountain and ob-
served daily; Woodbury and Russell (1945)
three. Navajo Mountain, 11 Aug. 1935 and 10
and 21 July {Pinus ponderosa, 8000-8500 ft);
very common and abundant in Pinus ponde-
rosa forest, but not in Picea-Abies forest on
Navajo Mountain, 1 July- 13 Aug. 1936;
adults feeding well-grown voung, 4 Jidv;
Woodburv and Russell foimd it very abun-
dant and primarily in the Pinus ponderosa
zone, restricted to elevations between
7500-9820 ft. Permanent resident.
Certhiidae (Creepers)
Certhia familiaris Linnaeus. Brown
Creeper. Presnall (1934) Bryce Rim; Wood-
burv and Russell (1945) one, Navajo Moun-
tain, 12 Aug. 1935 (Pinus ponderosa, 9000 ft);
seen at Navajo Mountain, 18 Julv 1933 (dense
forest) and 6-13 July to 13 Aug'. 1936 (Picea-
Abies forest). Woodbury and Ru,sse!l found it
there during June, July, and Augu.st in the
Picea-Abies forest with winter records at
lower elevations. Permanent resident.
326
Great Basin Naturalist
Vol. 40, No. 4
Cinclidae (Dippers)
Cinclus mexicanus unicolor Bonaparte.
Dipper. Benson (1935) Bridge Canyon at
northwest base of Navajo Mountain; Tanner
(194()a) Kaiparowits Basin. Woodbury and
Russell (1945) believed that the dipper did
not occur on Navajo Mountain and that it
had a definitely limited distribution in Nav-
ajo country; they gave sight records for
Bridge Canyon, 6 July 1933; several records
in 1934 and one record 7 Aug. 1935. Per-
manent resident.
Troglodytidae (Wrens)
Troglodytes aedon parkmanii Audubon.
House Wren. Presnall (1934) Navajo Trail;
Benson (1935) one, Navajo Mountain, 14 July,
and seen daily in fallen Pinus ponderosa,
July; Woodbury and Russell (1945) three,
Navajo Mountain, 14 July 1933, 16 Aug.
1935, and 9 July 1936 (common in thickets of
Arctostaphylos, Ceanothiis, and Roso under
Finns ponderosa, 8500-9000 ft); Tanner
(1940a) Kaiparowits Basin. Summer resident.
Thryomanes bewickii eremophilus Ober-
holser. Bewick's Wren. Woodbury and Rus-
sell (1945) one, two miles north of Navajo
Mountain Trading post, 27 July 1936 (pygmy
conifers, 6500 ft); they reported its regular,
though not abundant, occurrence in pygmy
conifers on the lower slopes of Navajo Moun-
tain from 22 June to 9 Aug., 1934-1938.
Catherpes mexicanus conspersus Ridgway.
Canyon Wren. Presnall (1934) Bryce Canyon;
Woodbury and Russell (1945) six, Lee's Fer-
ry, 26 Aug. 1909 (Nelson and Birdseye trip)
and 18 July 1936; Rock Creek /Colorado Riv-
er, 21 July 1936; two miles north of Navajo
Moimtain Trading Post, 2 Aug. 1936; Navajo
Mountain, 6 Aug. 1935 and 2 Aug. 1936 (in
cliffs of some size, bearing cracks for nesting,
3100 ft to 10,000 ft); Behle (1948) Hidden
Passage Canyon, Lee's Ferry, and mouth of
Aztec Canyon; Behle et al. (1958) one, con-
fluence Calf Creek/ Escalante River, 1954;
Hayward et al. (1958) Escalante drainage;
Behle and Higgins (1959) recorded a consid-
erable number of individuals in the Glen
Canyon and Navajo Mountain areas, 21-23
June and July 1933, 4-22 July 1936, and 3-10
Aug. 1938. In summer of 1958 they found the
species abundant along the full length of
Glen Canyon; BYU (1971) Cockscomb Ridge;
29 Sept.: BYU (1972) mouth of Escalante Riv-
er, 25 June (hanging garden); BYU (1973)
Hackberry Canyon, 8 June (open cliffs); Paria
River, 14 June (cliffs); Lees Ferry eight seen
7, 14, and 15 June (rocks); Driftwood Can-
yon; Three Garden; and Paria/Colorado Riv-
ers, 15 June (rocks). Permanent resident.
Salpinctes ohsoletus obsoletus (Say). Rock
Wren. Benson (1935) several seen, Navajo
Mountain; Woodbury and Russell (1945)
four, Navajo Mountain, 11 Aug. 1935 and 13
July 1936; above Lee's Ferry, 19 July 1936
{Artemisia, pygmy forest, Artemisia-Arctosta-
phijlos under Pinus ponderosa, 3100-10,000
ft); seen at Kaiparowits Plateau and Beaver
Creek /Navajo Mountain area, 8 Aug. 1936;
Behle (1948) Lee's Ferry; Behle et al. (1958)
Escalante River, 7 May 1954 and 10 miles
south of Escalante, 8 May 1954; Hayward et
al. (1958) Escalante drainage; Woodbury (in
Behle and Higgins 1959) one on 23 July 1937,
and 20 in August 1938 at Rock Creek, Last
Chance Creek, Lee's Ferrv, and various dis-
tances along the Colorado River from river
mile 50 to river mile 18. Russell and Thomp-
son (1964) Bryce Canyon; BYU (1958) one
(male), Paria Basin, 9 June; BYU (1961) one
(male), Paria, 20 May; BYU (1971) Sites 2 and
15; Church Wells area, 28 Sept.; Grand
Bench, 9-10 Oct.; Cottonwood Wash; Hack-
berry Canyon; Lee's Ferry; Nipple Spring;
Smoky Mountain; Tibbet Spring; Ribbon
Canyon; and Cockscomb Ridge, 29 Sept.;
BYU (1973) 82 wrens of this species were
seen at 15 localities in June and July. The
earliest spring record was 5 June 1973 with
nesting on 8 June. The latest fall record was
10 Oct. 1971. The Rock Wren is nearly obli-
gate to cliffs and rocks. Seventeen of the indi-
viduals observed were in rocks or cliffs and
one in Popuhis. Summer resident, mav winter
sparingly (Woodbury and Russell 1945) in
Kaiparowits Basin.
Mimidae (Mockingbirds and Thrashers)
Mimus polyglottos leucopterus (Vigors).
Mockingbird. Benson (1935) Navajo Moun-
tain area; Tanner (1940a) base Kaiparowits
Plateau, June 1936; Woodbury and Russell
(1945) one, five miles south Navajo Mountain
December 1980
Atwood et al.: Kaiparowits Vertebrates
327
Trading Post, 30 July 1965 {Artemisio, 6500
ft); few seen at Lee's Ferry and flats south of
Navajo Mountain; Behle et al. (1958) one, 10
miles south of Escalante, 8 May 1954 (fairly
common); Higgins saw one in Atriplex confer-
tifolia at mouth of Escalante Canvon, 19 julv
1958; Russell and Thompson (1964) Bryce
Canyon; B\TJ (1971) Last Chance Wash, 25
Sept. {Clmjsothammis); BYU (1972) Site 27, 1
Jime (woodland); Cedar Mountain, 28 July
{Epliedm-grdss); Site 24, 28 June (shrubs);
BYU (1973) near Site 1, 9 and 21 May {Juni-
perus); Site 2, 2 seen 30 April (shrubs) and 3
July {Juniper us); near Site 3, 28 May (wood-
land) and 2 July (grass); Cottonwood Wash, 5
June (riparian); Nipple Spring, 12 June {Pop-
ulus); Tibbet Spring, 5 and 10 June (open
brush); Site 34, 4 seen, 10 July; and between
Sites 6 and 8 on Smoky Mountain, 20 were
seen 24 July {Grayia-Coleogync). Summer
resident between 30 April and 25 September.
Dumetella carolinensis (Linnaeus). Gray
Catbird. Woodbury and Russell (1945) Lee's
Ferry, as recorded by the Nelson and Birds-
eye trip of 1909; Phillips et al. (1964) Lee's
Ferry. Summer resident.
Toxostoma bendirei (Coues). Bendire's
Thrasher. Woodbury (1939) near Escalante, 9
May 1937; Behle et al. (1958) give the local-
ity of the aforementioned specimen as five
miles south of Escalante; however, they did
not observe this species at this site, 8 May
1954; Hayward (1967) Wahweap Creek, 20
May 1956; BYU (1971) Site 15, 25 Sept.
{Chnjsothamnus). Summer resident.
Eoroscoptes montanus (Townsend). Sage
Tlirasher. Woodbury and Russell (1945) one,
Lee's Ferry, 26 Aug. 1909, as recorded by
Nelson and Birdseye's trip of 1909; Behle et
al. (1958) two miles south of Escalante, 9
May 1954, and common in Escalante Valley,
May 1954; Russell and Thompson (1964)
Bryce Canyon; BYU (1971) Site 12, 15 Oct.;
Site 14, 15' Oct.; Tibbet Spring, 13 Aug.; Site
15, 11 Oct.; and Grand Bench, 9 Oct. Un-
common migrant.
Turdidae (Thnishes, Solitaires, and Bluebirds)
Turdus migratorius propinquus Ridgway.
Robin. Presnall (1933 and 1934) Bryce Can-
yon; Benson (1935) Navajo Mountain; Wood-
bury and Russell (1945) three, Navajo Moun-
tain, 21 July and 16 Aug. 1936, and
Kaiparowits Plateau, 11 Aug. 1937 {Pinus
ponderosa, 7000-9000 ft); Hayward et al.
(1958) Escalante drainage; Behle and Higgins
(1959) seen at many places in Glen Canvon;
most abundant in large vegetation at mouths
of side canyons; BYU (1972) Tibbet Canyon,
1 Aug. (Coleogyne); BYU (1973) two, Wah-
weap Lodge, 31 Jan. (grass); Woodbury and
Russell (1945) consider Navajo Mountain and
the Kaiparowits Plateau to be chief breeding
area of this species in the Navajo country of
Utah. They list a juvenile from Navajo Moun-
tain taken 16 Aug. 1936. Summer resident, a
few apparently winter.
Catharus guttatus auduboni (Baird). Her-
mit Thrush. Presnall (1934) and Grater (1947)
Bryce Canyon; Benson (1935) two specimens:
Navajo Mountain (10,000 ft, species com-
mon); Woodbury and Russell (1945) speci-
men: Navajo Mountain, 14 July 1936 (Picea-
Abies forest 10,000 ft); common (estimates
100 pairs) in Picea-Abies forest 6 and 11-18
July 1936 (nesting 16 July; absent 13 Aug.);
BYU (1971) summit of Navajo Mountain, 13
Oct. Summer resident and transient in higher
mountains and transient in lowland.
Sialia mexicana bairdi Ridgway. Western
Bluebird. Presnall (1933 and 1934) and Rus-
sell and Thompson (1964) Bryce Canvon;
Benson (1935) and Woodbury and Russell
(1945) specimens at Navajo Mountain, 16
Aug. 1935 and 4 July 1936 (Pinus ponderosa);
Behle et al. (1958) one, confluence of Calf
Creek/Escalante River, 1954; BYU (1971)
one (male), Cottonwood Wash, 7 Nov.; BYU
(1972) Site 23, 25 .seen 3 Mar. (mixed shrubs);
and 11 miles east Grosvenor Arch, one seen
(not identified to species), 17 June (wood-
land). Permanent resident.
Sialia currucoides (Bechstein). Mountain
Bluebird. Presnall (1933 and 1934) and Grat-
er (1947) Bryce Canyon (also late summer to
late winter at Cedar Breaks); Tanner (1940a)
Escalante drainage; Behle et al. (1958) Paria,
6 March 1946; Hayward et al. (1958) Esca-
lante drainage; BYU (1971) one, Paria
Plateau, 29 Sept.; BYU (1972) one, one mile
south Glen Canyon City, 7 Feb.; Sites 3 and
6; Site 23, three seen on 24 Feb. (Coleogyne);
Four Mile Bench, 25 seen 26 Oct. (wood-
land); BYU (1973) one (ad. male), eight miles
east Glen Canyon Citv, 2 May; Warm Creek
328
Great Basin Naturalist
Vol. 40, No. 4
Bay, two seen 10 June; and Four Mile Bench,
13 June (near a spring). Simnmer resident, a
few may winter.
Myadestes townsendi townsendi (Audu-
bon). Townsend's Solitaire. Presnall (1933
and 1934) and Grater (1947) Bryce Canyon;
Behle et al. (1958) five miles west of Esca-
lante, 20 Sept. 1935. Transient.
Sylviidae (Old World Warblers,
Gnatcatchers, and Kinglets)
Polioptila caerulea amoenissinia Grinnell.
Blue-gray Gnatcatcher. Presnall (1934) and
Russell and Thompson (1964) Bryce Canyon;
Benson (1935) Navajo Mountain; Tanner
(1940a) Kaiparowits Basin; Woodbury and
Russell (1945) five, Navajo Mountain Trading
Post, 25 July 1936; Rock Creek/Colorado
River, 26 July 1937; Kaiparowits Plateau,
30-31 July and 10 Aug. {Artemisia, wood-
land, Quercus, Amelanchier, 3300-7000 ft);
Hayward et al. (1958) Escalante drainage;
BYU (1971) Last Chance, 25 Sept. (Tamarix);
BYU (1973) Sites 15 and 28 {Junipems-grass)-
Lee's Ferry, 14-15 June {Tamarix and Salix);
Cottonwood Wash/Paria River, 6 July; Rib-
bon Canyon; Paria /Colorado Rivers, 7 and
14 June (Tamarix); Tibbet Spring, two seen 5
June (Popuhis); and Wahweap Creek, 30 Apr.
and 7 Jime. Summer resident (30 April to 25
Sept.).
Regulus satrapa Lichtenstein. Golden-
crowned Kinglet. Russell and Thompson
(1964) Bryce Canyon; BYU (1971) north of
Church Wells; one, summit of Navajo Moun-
tain, 13 Oct. Permanent resident.
Regulus calendula (Linnaeus). Ruby-
crowned Kinglet. Presnall (1934) and Russell
and Thompson (1964) Bryce Canyon; Behle
(1948) Lee's Ferry; Woodbury and Russell
(1945) two, Navajo Mountain, 14 Aug. 1935
and 15 July 1936 {Picea-Ahies, Pinus ponde-
rosa forest, 9000-10,000 ft); breeds on top of
Navajo Mountain. Permanent resident.
Bombycillidae (Waxwing)
Bombycilki cedrorum Vieillot. Cedar Wax-
wing. Behle et al. (1958) Bryce Canyon. Un-
common transient, some may breed (Behle et
al. (1958).
Ptilogonatidae (Silky Flycatchers)
Phainopepla nitens lepida Van Tyne.
Phainopepla. BYU (1973) near Tibbet Spring,
5 June; seen by Robert Whitmore in open
brush. This species was previously known in
southern Utah only from lower Santa Clara
Valley, Washington County, to Kanab, Kane
County (Hayward et al. (1976). Summer resi-
dent.
Laniidae (Shrikes)
Lanius excubitor Linnaeus. Northern
Shrike. BYU (1973) Site 3, 3 Aug. Rare tran-
sient.
Lanius ludovicianus Linnaeus. Log-
gerhead Shrike. Benson (1935) Navajo Moun-
tain area; Tanner (1940a) Escalante drainage;
Woodbury and Russell (1945) one, Kaiparo-
wits Plateau, 12 Aug. 1937; Rock
Creek/Colorado River, 26 July 1937; Navajo
Mountain Trading Post, 31 July 1936 {Popii-
his, Artemisia, and Amelanchier, 4000-7000
ft); also in Artemisia flats, Sarcobatus, Xan-
thocephalum, Atriplex confertifolia. Yucca,
and Ephedra types of cover (Woodburv and
Russell 1945); Behle et al. (1958) 10 'miles
southeast of Escalante, 8 May 1954; Russell
and Thompson (1964) Bryce Canyon; BYU
(1971) Sites 1 and 2; specimens at two miles
southwest of Site 3, 28 Aug. and two miles
west of Site 3, 20 Aug.; Sites 6 and 10; Cot-
tonwood Wash, 7 Nov.; Four Mile Bench;
Glen Canyon City; Pump Canyon Spring;
Tibbet Spring; one. Warm Creek, two miles
below road to Escalante, 6 Dec; BYU (1973)
one (ad. female), two mile southwest Glen
Canyon City, 29 Apr. It is a permanent resi-
dent of the area according to Woodburv and
Russell (1945). We saw 155 individuals be-
tween 1971 and 1973. Sightings (73) were
distributed throughout the year, except for
Feb. and Mar. when none were seen. Wood-
bury and Russell (1945) believed the species
to be more plentiful in late summer after
emergence of young. This agrees with our
findings. We found the species very sparse in
Nov. (2), Dec. (1), and Jan. (1); absent in Feb.
and Mar.; and somewhat more frequent in
Apr. (10), May (3), and June (3). It was com-
mon in July (28), reaching a peak in Aug.
(56), and declining in numbers in Sept. (27)
i
December 1980
Atwood et al.: Kaiparowits Vertebrates
329
and Oct. (16). It was most frequently encoun-
tered in woodlands, Vanclevea-grdss, Coleog-
ijne, Ephedra-grsiss, and Atriplex-grass in this
order of abundance. Common permanent
resident.
Stumidae (Starlings)
Sturnus vulgaris vulgaris Linnaeus. Star-
ling. Russell and Thompson (1964) Bryce
Canyon; BYU (1971) two (males), Glen Can-
yon City, 2 Dec; two seen 8 Dec; BYU
(1972) Glen Canyon City, two seen 5 Mar.;
BYU (1973) Paria River /U.S. 89, 6 June. The
starling is a newcomer to the area. Behle et
al. (1958) reported sightings of this species in
Jan. 1941 and Sept. 1948 for Kane County,
but the observations by Russell and Thomp-
son (1964) appear to be the first published
observations of this species in the area of the
current report. Earlier observers (Behle 1960,
Woodbury and Russell 1945, Hayward 1976)
did not report the species. The species is
known in Utah to first occur as a migrant or
winter visitant, followed later by permanent
residency (Behle and Perry 1975). A similar
pattern of occurrence in the Kaiparowits
area suggests that the species is establishing
itself here as a breeder also. Permanent resi-
dent.
Vireonidae (Vireos)
Vireo bellii Audubon. Bell's Vireo. BYU
(1973) Lee's Ferry, 7 June. Until this vireo
was seen at Lee's Ferry by Robert Whitmore
it was known in Utah only from southwestern
Utah (Hayward et al. (1976), where it is an
imcommon summer resident (Behle and Perry
1975). Sparse summer resident.
Vireo vicinior Coues. Gray Vireo. Behle et
al. (1958) specimen: confluence (Calf
Creek /Escalante River, 1954; BYU (1973)
Site 28, 13 June by Lloyd Pack in Juniperiis-
grass association. Behle et al. (1958) report
this species as an uncommon resident of
woodlands of the Kanab area and adjacent
high plateaus. Although Behle and Perry
(1975) consider it a common summer resident
of southern Utah, in the area of the Kaiparo-
wits Plateau it seems to be rather sparse, as
suggested by only one sighting in 2V2 years.
Sparse summer resident.
Vireo solitarius (Wilson). Solitary Vireo.
Presnall (1934) and Russell and Thompson
(1964) Bryce Canyon; Behle et al. (1958) con-
sider V. s. plumbeus to be a fairly common
summer resident in the lower reaches of
Bryce Canyon. They observed this taxon in
canyons vegetated with Quercus gainheUii,
Acer negundo, and Populus frernontii; Wood-
bury and Russell (1945) three (all V. s. cas-
sinii), Navajo Mountain, 7 Aug. 1935, 11
Aug. 1936, and Lee's Ferry, 25 Aug. 1909, as
recorded by the Nelson and Birdseye trip
{Populus tre7nuloides and riparian forest,
Quercus and Salix, 3100-9500 ft); it is prob-
able that V. s. plwnheus breeds in the Kai-
parowits Plateau and Navajo Moimtain areas,
although none have been seen. Vireo s. cas-
sinii, on the other hand, is a migrant. Wood-
bury and Russell (1945) considered V. s.
plumbeus as a siunmer resident of Pinus pon-
derosa, Quercus, and woodlands of the mesa
tops and slopes of the Navajo country. Sum-
mer resident and transient.
Vireo gilvus (Vieillot). Warbling Vireo.
Benson (1935) and Woodbury and Russell
(1945) Navajo Mountain area, 17 Aug. 1935
and 4 Aug. 1936, and Kaiparowits Plateau, 2
Aug. 1937; Tanner (1940a) Kaiparowits Ba-
sin; Behle et al. (1958) Bryce Canyon. Al-
though specimens from Garfield County,
Utah, have been assigned to V. g. swainsonii,
Worthen (1968) has questioned the presence
of the race swainsonii in Utah (also see Hay-
ward et al. 1976); it is probablv a migrant
(Behle and Perry 1975). The breeding popu-
lation represents the race leucopolius (Behle
and Perry 1975). Woodbury and Russell
(1945) found this latter race in stands of
Qiwrcus, streamside Populus fremontii and
Salix, and Populus tremuloides. On Navajo
Mountain they encountered it in Populus
tremuloides at 9000-10,500 ft. They also
found breeding adults and two young, 16 to
18 July 1936. We did not encounter it. Sum-
mer resident.
Parulidae (Wood Warblers)
Vermivora celata (Say). Orange-crowned
Warbler. Behle and Higgins (1959) Hole-in-
the-Rock/Colorado River, 20 Oct. 1958 (ri-
parian, 3266 ft); BYU (1971) Site 15, 9 Oct.
Uncommon migrant.
330
Great Basin Naturalist
Vol. 40, No. 4
Vermivora ruficapilla ridgwayi van Ros-
sem. Nashville Warbler. Woodbury and Rus-
sel (1945) one, Navajo Mountain, 11 Aug.
1935 {Pinus pundewsa, 8500 ft). Transient.
Vermivora virginiae (Baird). Virginia's
Warbler. Presnall (1934) Bryce Canyon;
Woodbury and Russell (1945) Navajo Moun-
tain, 17 July 1936 {Populus tremuloides) and
Kaiparowits Plateau, 9 Aug. 1937 (pygmy
forest, 7000-10,000 ft); Behle et al. (1958)
confluence Calf Creek/Escalante River,
1954; Russell and Thompson (1964) Bryce
Canyon; BYU (1971) Site 4; Vi mile south
Paria River Bridge, 3 Sept. According to
Woodbury and Russell (1945), this warbler is
a breeder of the deciduous bnish and tree
zones between 6000-10,000 ft. Summer resi-
dent.
Vermivora luciae (Cooper). Lucy's War-
bler. Woodbury (1939) and Woodbury and
Russell (1945) two, river mile 41,8 Aug. 1938
{Salix and Rhus, 3180 ft); Colorado River side
canyon, 13 July 1936 {Salix thickets, 3200 ft);
seen at mouth of Paria River, 1938; V2 mile
below Lee's Ferry, 30 July to 11 Aug. 1938;
yoimg out of nest, banks Colorado River, two
miles below San Juan River, 17 July 1931;
and single birds on the Colorado at river mile
41, 8 Aug. and one mile above Lee's Ferry,
10 Aug.; Woodbury and Russell (1945) be-
lieve this species to be a sparse inhabitant of
Salix and brush thickets along the Colorado
and San Juan Rivers; Behle (1948) Lee's Fer-
ry; Behle et al. (1958) one, junction Calf
Creek/Escalante River, 4 July 1938; BYU
(1973) Lee's Ferry and one mile south of
Paria, Utah, 7 June (Salix and Tamarix). Sum-
mer resident.
Dendroica petechia morcomi Coale. Yel-
low Warbler. Woodbury and Russell (1945)
four, three miles below confluence San
Juan/Colorado Rivers, 3 Aug. 1938; six at
Rock Creek/Colorado River, 20-23 July
1937; one from Kaiparowits Plateau, 4 Aug.
1937; one from Navajo Mountain, 13 Aug.
1935; one from Lee's Ferry,25 Aug. 1909;
two along Colorado River, 11 July 1936; one
and nest at river mile 63, 5 Aug. 1938; and
two at river mile 50, 6 Aug.; (Quercus, Salix,
riparian thickets, Artemisia, Populus tremu-
loides, 3120-7000 ft; nested below
6500-7000 ft in riparian Salix, brush or Popu-
lus); Behle and Higgins (1959) give the addi-
tional observational records from Wood-
bury's notes as follows: "many were seen at
several points, six at two, and two at another,
plus three family groups at one and six at an-
other, from river mile 75 to the mouth of
Paria Creek; common breeder 1 July to 9
Aug. 1958 in riparian vegetation along Glen
Canyon"; Hayward et al. (1958) Escalante
drainage; Russell and Thompson (1964) Bryce
Canyon; BYU (1972) Reflection Canyon, 4
July and confluence San Juan/Colorado Riv-
ers, 24 June (Fop«/MS-riparian); BYU (1973)
Lee's Ferry, three seen 7 June (Tamarix) and
one seen 15 June (Tamarix); one mile south of
Paria near Paria River, 7 June (dense Salix);
and Paria River/Colorado River, 7 June
(Tamarix-Salix). Summer resident.
Dendroica coronata auduboni (Town-
send). Yellow-rumped (Audubon's) Warbler.
Presnall (1933) Bryce Canyon; Benson (1935)
Navajo Mountain; Woodbury and Russell
(1945) three, Navajo Mountain, 11 Aug.
1935; nests in Pinus ponderosa, 2 July 1936
(8500 ft; breeds 8000-10,000 ft, spring migra-
tion dates in general area of Navajo Moun-
tain are 13 Apr. -25 Mav; and fall, 20
Aug.-14 Oct.); Behle (1948) Lee's Ferry;
Behle et al. (1958) 10 miles west of Escalante,
7 May 1941; BYU (1971) one, summit Navajo
Mountain, 13 Oct.; and 25 at Site 15, 9-10
Oct.; BYU (1973) Crosby Canyon Bay, two
seen courting, 28 Apr. (Tamarix); and two
seen 5V2 miles south of Tibbet Springs, 1
May. The intermountain race D. auduboni
memorabilis is now in synonymy with D. a.
auduboni (Townsend) and the species has
been placed in Dendroica coronata. Summer
resident and fall and spring transient.
Dendroica nigrescens (Townsend). Black-
throated Gray Warbler. Presnall (1934) Bryce
Canyon; Benson (1935) Navajo Mountain;
Woodbury and Russell (1945) six specimens:
Kaiparowits Plateau, 31 Julv and 4, 5, and 9
Aug. 1937; Navajo Mountain, 14 Aug. 1935
and 2 Aug. 1936 (pygmy forest, 6000-9000
ft; a nearly obligate breeder in pygmy forest,
migrating through most other vegetational
types); BYU (1971) Site 12 and specimen
(male): V2 mile north Nipple Spring, 16 Oct.
Summer resident and transient spring and
fall.
Dendroica townsendi (Townsend). Towns-
end's Warbler. Presnall (1933 and 1934)
December 1980
Atwood et al.: Kaiparowits Vertebrates
331
Bryce Canyon; Woodbury (1939) two, Nav-
ajo Mountain, 10 Aug. 1936 (also seen 13
Aug. 1935); Kaiparowits Plateau, 12 Aug.
1927 (pvgmv forest, Piniis ponderosa,
6000-700'(Ht).' Transient.
Dendroica occidentalis (Townsend). Her-
mit Warbler. Woodbury and Russell (1945)
two, Navajo Mountain, 11 Aug. 1935 and 13
Aug. 1936 [Pinits ponderom, 8500-10,000 ft).
In 1935 this normally rare species was a com-
mon member of the band of warblers on
Navajo Mountain during Aug. (Woodbury
and Russell 1945). Transient.
Dendroica graciae graciae Baird. Grace's
Warbler. Presnall (1934) Bryce Canyon;
Woodbury and Russell (1945) seen at Navajo
Moimtain, 15-16 June 1938, and one collect-
ed but discarded, 15 June; Benson (1935) saw
and heard them in Pinus ponderosa, but not
numerous (Woodbury and Russell 1945).
Summer resident.
Oporornis tolmiei (Townsend). MacGil-
livray's Warbler. Woodbury and Russell
(1945) one, Navajo Mountain, O. t. tolmiei;
Behle (1960) 5 Aug. 1936 {Salix in canyon
bottoms); BYU (1971) Glen Canyon City;
Kaibab Wash, 29 Sept. Summer resident.
Geothlypis trichas occidentalis Brewster.
Common Yellowthroat. Woodbury and Rus-
sell (1945) two at Rock Creek/Colorado Riv-
er, 20-26 July 1927 (large family flocks also
seen in streamside thickets); one near Bridge
Canyon, 12 July 1936; one from river mile
60, 6 Aug. 1938; and one from river mile 25,
9 Aug. 1938 (including nest ); nest observed
at river mile 63, 5 Aug. 1938 (Salix); family
flocks mouth of Rock Creek, 20-26 July
1937; usually found in Salix reeds, canes,
brush, or Ti/phits in canyons or valleys below
5500-6000 ft; Behle et al. (1958) confluence
Calf Creek /Escalante River, 2 May 1954;
Behle and Higgins (1959) 165 individuals
seen, plus many others including 12 families
(out of nest) at various points along the Colo-
rado River by Woodbury and others during
the Rainbow Bridge-Monument Valley expe-
dition. Higgins (Behle and Higgins 1959) saw
both adults and immatures in the Salix fringe
at Lee's Ferry, 8 Aug. 1958, and found the
species nesting abundantly during the sum-
mer of 1958 (Behle 1960). Woodbury and
Russell (1945) noted that the habitats of the
Yellow Warbler and Yellowthroat overlap.
but the warbler extended landward into Pop-
ulus fremontii, whereas the yellowthroat ex-
tended toward the moist areas containing
Typhus and Jiincus. It is interesting that none
were seen during this study. It is not known if
this is because of the creation of Lake Powell
or due to insufficient observations.
Icteria virens (Linnaeus). Yellow-breasted
Chat. Woodbury and Russell (1945) three,
Colorado River, 11 July 1936 and river miles
41 and 50, 7 and 8 Aug. 1938 (stream bank
thickets, Salix, and Rhii.s); Behle et al. (1958)
confluence of Calf Creek /Escalante. 1954,
i.e., /. f. auricollis in dense vegetation. Behle
and Higgins (1959) give the following addi-
tional records from the field notes of Wood-
bury and Russell for 1938 for the Colorado
River from the Rainbow Bridge-Monument
Valley expedition: 80 were counted from riv-
er mile 50 to V2 mile below Lee's Ferry near
the mouth of Paria Creek, between 7 and 1 1
Aug. Higgins found them to be abundant
breeders along the length of Glen Canyon.
He saw them every day of the trip in the
densest streamside vegetation. Russell and
Thompson (1964) Bryce Canyon; BYU (1973)
Lee's Ferry, a total of 12 were seen 6, 7, and
14-15 June; and along the Paria River near
Paria, Utah, three were seen 7 and 14 June.
We saw them in the Tamarix and Salix along
the rivers. Summer resident.
Wilsonia pusilla pilcolata (Pallas). Wil-
son's Warbler. \Voodbur\ and Russell (1945)
Navajo Mountain, one or two seen daily in
Populus tremuloides, 9-16 Aug. 1935, and
one, 1 Sept. 1934; Russell and Thompson
(1964) Bryce Canyon; BYU (1973) lower
Wahweap Creek, 30 Apr. Woodbury and
Russell give the migration date through the
area as 4-26 May in the spring and 9
Aug. -26 Oct. in the fall. Transient.
Ploceidae (Weaver Finches)
Passer domesticus (Linnaeus). House Spar-
row. Woodbury and Russell (1945) Lee's Fer-
ry/Paria River, several birds seen 10 Aug.
1938; Behle et al. (1958) Escalante; BYU
(1971) Glen Canyon City, .30 seen 2 Dec. and
5 seen 8 Dec. Permanent resident.
332
Great Basin Naturalist
Vol. 40, No. 4
Icteridae (Meadowlarks, Blackbirds,
and Orioles)
Sturnella neglecta neglect a Audubon.
Western Meadowlark. Presnall (1934) Bryce
Canyon; Behle et al. (1958) near Escalante in
cultivated fields; Hay ward et al. (1958) Esca-
lante drainage; BYU (1971) Site 3 (no date,
semiarid grasslands); and Paria Plateau, 29
Sept. Uncommon permanent resident.
Xanthocephalus xanthocephalus (Bona-
parte). Yellow-headed Blackbird. Woodbury
and Russell (1945) Lee's Ferry as recorded by
Nelson and Birdseye's trip, 23-26 Aug. 1909;
Russell and Thompson (1964) Bryce Canyon;
Behle and Higgins (1959) river mile 25, three
females seen, 6 Aug. 1958; BYU (1972) Warm
Creek Bay, one pair seen on shore of Lake
Powell, 3 May. Migration through the area is
given by Woodbury and Russell (1945) as
April-May passing north and 12 July-11 Oct.
passing south. Transient.
Agelaius phoenicetis (Linnaeus). Red-
winged Blackbird. Behle (1948) Aztec Creek,
two seen; Russell and Thompson (1964)
Bryce Canyon; Hay ward et al. (1958) Esca-
lante drainage; BYU (1973) one mile above
Lee's Ferry (flying), 15 June. Transient.
Icterus parisorum Bonaparte. Scott s Ori-
ole. Benson (1935) south of Navajo Mountain;
BYU (1972) Site 4, 25 June {Coleogijne)- BYU
(1973) three miles northwest of Site 3, 29
Apr. (Pintis edulis); two miles north Site 3, 28
May (woodland); Site 6, 3 May {Junipenis);
one mile east of Site 7, 4 July; and Site 28, 13
June {Juniperus-grsLSs). All 1973 observations
except the one July, were in Juniperus-grass
or pygmy forest. No vegetation was given for
the July sighting. Summer resident.
Icterus galhula bullockii (Swainson).
Northern Oriole. Benson (1935) Beaver
Creek/Navajo Mountain; Woodbury and
Russell (1945) two, north foot and southwest
foot of Navajo Mountain, 6 Aug. 1935 and 1 1
Aug. 1936 {Populus fremontii near woodland
and Quercus community, 5500-6000 ft);
Behle et al. (1958) confluence Calf Creek/
Escalante River, 1954; Hay ward et al. (1958)
Escalante drainage; Behle and Higgins (1959)
field notes of Rainbow Bridge-Monument
Valley expedition of 1938: seven were seen
along the Colorado River from river mile 75
to '/2 mile below Lee's Ferry, 3-11 Aug.; Rus-
sell and Thompson (1964) Bryce Canyon;
BYU (1961) one male and three females,
Paria, 20 May; BYU (1973) Paria River/U.S.
89, two seen 6 June; and Cottonwood Can-
yon, 8 June; Hackberry Canyon, 8 June (male
displaying); and Lee's Ferry, adult and fledg-
ling seen in Tamarix and Salix 15 June. Ex-
cepting the 15 June sighting, all others were
in stands of Populus fremontii. Summer resi-
dent.
Euphagus cyanocephalus (Wagler).
Brewer's Blackbird. Presnall (1934) Bryce
Canyon; Tanner (1940a) Escalante drainage;
Behle et al. (1958) confluence of Calf
Creek/Escalante River; Hayward et al.
(1958) Escalante drainage; Behle and Higgins
(1959) 5 were seen at mouth of Kane Creek,
river mile 41, 1 Aug. 1958; BYU (1973) Lee's
Ferry, three sightings totaled 53 birds, 7 June
(Tamarix); Glen Canyon City, 50 seen 28
Apr. (grasslands); and Cottonwood
Wash/Brigham Plains road, 2 seen 8 June (ri-
parian vegetation). Transient, a few may nest
and a few may winter.
Molothrus ater obscurus (Gmelin). Brown-
headed Cowbird. Woodbury and Russell
(1945) specimen: Colorado River at Rock
Creek, 22 July 1937 (riparian thicket of
Tamarix and Salix, 3100-3300 ft) and one egg
found in the nest of Willow Flycatcher near
Lee's Ferry, 11 Aug. 1938; Behle et al. (1958)
Bryce Canyon; Behle and Higgins (1959) Col-
orado River/Rock Creek, 28 July 1958; BYU
(1972) Site 3 on Cedar Mountain, 25 July
(Ephedra-grsiss); BYU (1973) Cottonwood
Wash, 5 June (Popuhis); and Lee's Ferry, one
seen 7 June (Tamarix) and two seen 15 June
(brush). Summer resident.
Thraupidae (Tanagers)
Piranga ludoviciana (Wilson). Western
Tanager. Benson (1935) one, War God Spring
on Navajo Mountain, 14 June 1933; Wood-
bury and Russell (1945) six, two at Navajo
Mountain (War God Spring and Beaver
Creek Canyon), 13 July, 7 Aug. 1936; one at
Navajo Mountain Trading Post, 26 July [no
vear]; one at Colorado River/Rock Creek, 27
July 1937; one at Rock Creek, 28 July 1937;
and one at Kaiparowits Plateau, 6 Aug. 1937;
usually found in Piniis pondcrosa, Populus
fremontii and other riparian trees, chaparral.
December 1980
At WOOD ET AL.: Kaiparowits Vertebrates
333
and Quercus. 3300-8500 ft; Behle and Hig-
gins (1959) Aztec Canyon at river mile 68.5,
7 July 1958; mouth of Kane Creek at river
mile 41, 1 Aug.; and one mile upstream from
mouth of Escalante River, several seen of
both sexes, 19 July 1958. Woodburv and Rus-
sell (1945) saw this species during the breed-
ing season in stands of Pinus ponderosa and
Picea-Abies on Navajo Mountain; Russell and
Thompson (1964) Brvce Can von; BYU (1971)
Site 16 in Tamarix, \H Aug.;' BYU (1972) one
mile south of Site 1, 8 June. Summer resident.
Fringillidae (Grosbeaks, Finches,
Sparrows, and Buntings)
Pheucticus melanocephalns melanoce-
phalus (Swainson), Black-headed Grosbeak.
Benson (1935) Navajo Mountain; Woodbury
and Russell (1945) two, Navajo Mountain, 15
Aug. 1935 and 4 Aug. 1936 {Salix and Picea-
Abies forest, 6500-10,300 ft); they give post-
breeding dates 23 July to 15 Aug. 1934-38 at
Navajo Mountain Trading Post, Beaver
Creek Canyon, and Lee's Ferry. They in-
dicated that it breeds at canyon heads and
moimtains mostly below the coniferous for-
est, among Querciis, but mostly above the
pygmy forest; it may descend into the
streamside fringes of lower elevations. Wood-
bury and Russell (1945) record extreme oc-
currence dates as 8 May and 22 September;
Hayward et al. (1958) Calf Creek area; Behle
and Higgins (1959) mouth of Aztec Creek at
river mile 68.5 in stands of Quercus, 26 July
1958; river mile 46 in Salix, 30 July 1958;
and from Woodbury and Russell's field notes
(Behle and Higgins 1959) between river miles
63 and 50, 5 Aug. 1938; Russell and Thomp-
son (1964) Bryce Canyon; BYU (1971) Site 4
and V2 mile south of Paria Bridge, 30 Sept.
Summer resident.
Guiraca caerulea interfusa (Dwight &
Griscom). Blue Grosbeak. Presnall (1934)
Bryce Canyon; Tanner (1940a) Kaiparowits
Basin; Woodbury and Russell (1945) 2 at riv-
er mile 13, 9-10 Aug. 1938; 3 near Lee's Fer-
r\', 17 July 1936 and 9-11 Aug. 19.38 (Tama-
rix and Salix 3100 ft); in addition 14 birds
seen including a nest and two nestlings,
10-11 Aug. 1938; Higgins (Behle and Higgins
1959) foimd it an abundant breeder in dense
streamside vegetation along the length of
Glen Canyon; BYU (1971) Site 16, 8 Aug.
(Tamarix); BYU (1972) 3 .seen in Cottonwood
Wash, 22 July (Tamarix); BYU (1973) Lee's
Ferry, a total of 5 birds seen in Tamarix,
6-15 June. Common summer resident in
riparian vegetation.
Passenna cyanea (Linnaeus). Indigo Bun-
ting. BYU (1973) Cottonwood Wash, 6 June.
One was seen by Robert Whitmore in Tama-
rix-Populiis fremontii association about six
miles up Cottonwood Wash from U.S. High-
way 89. Lazuli Buntings were also present.
Whitmore (1975) noted apparent com-
petition between males of the two species in
the Paria River area. Other sightings include
Lee's Ferry where Whitmore (1975) observed
the species in Tamarix. As noted bv Whit-
more, the species apparently has moved into
the Paria River drainage since the in-
vestigations of Woodbury and Russell (1945)
and Behle et al. (1958). VVhitmore (1975) also
summarized the expansion of the species into
Utah. Summer resident.
Passerina amoena (Say). Lazuli Bunting.
Tanner (1940a) Kaiparowits Basin; Wood-
bury and Russell (1945) two, Navajo Moun-
tain, 13 Aug. 1935, and Colorado River/ Rock
Creek, 20 July 1937 (riparian thicket, under
shrubs, Populus tremuloides, and Pinus pon-
derosa, 3300-9000 ft); Russell fovuid it fairly
conniion on lower Navajo Mountain, 10-16
Aug. 1936; Behle and Higgins (1959) Kane
Creek and river mile 41, 1 Aug. 1958; BYU
(197.3) in Cottonwood Wash at six (five seen
5-6 June), eight (two seen 6 June), and nine
(six seen 6 June) miles north of U.S. Highway
89, Kane County, Utah. Two sightings were
in Atriplcx canescens, one was in Juniperus-
Tainarix. and the remainder were in Tamarix.
Whitmore (1975) noted the Lazuli Bunting
nesting in the Paria River area. Extreme
dates of occurrence are 15 May and 28 Aug.
(Woodbury and Russell 1945). Summer resi-
dent.
Hesperiphona vespertina (Cooper). Eve-
ning Grosbeak. Presnall (1934) Bryce Point;
BYU (1972) Cottonwood Wash, four seen in
stands of Piniis-Jiiniperus, 26 Oct. Fall and
winter visitant.
Carpodacus cassinii Baird. Cassin's Finch.
Woodbury and Russell (1945) two specimens:
Navajo Mountain 9 and 14 Aug. 1935 (9000
ft); it is a sparse breeder in stands of Pinus
334
Great Basin Naturalist
Vol. 40, No. 4
ponderosa; Russell and Thompson (1964)
Brvce Canyon. Summer resident.
Carpodacus mexicanus frontalis (Say).
House Finch. Woodbury and Russell (1945)
two, Navajo Mountain Trading Post, 26 July
1936 and mouth of Rock Creek, 23 and 26
July 1937 {Populus frernontii and riparian
vegetation, 33()()-65()0 ft); Behle (1948) Az-
tec Creek and Lee's Ferry; Behle et al. (1958)
confluence Calf Creek /Escalante River,
1954, and Paria River, 6 Mar. 1946; Hayward
et al. (1958) Escalante drainage; Russell and
Thompson (1964) Bryce Canyon; Behle and
Higgins (1959) Higgins found them abundant
in Glen Canyon during summer 1958; they
were seen nearly every day in streamside
vegetation, on the terraces, and on the hill-
sides; Behle and Higgins (1959) give the fol-
lowing additional unpublished data from
Woodbury and Russell's field notes: "species
common and conspicuous between river
miles 78 and zero of the Colorado River,
4-22 July 1936; along the Colorado River, 22
were comited and many others seen between
river mile 75 and Paria Creek, 3-11 Aug.
1938"; BYU (1971) one, Warm Creek, 8
Nov.; Wiregrass Spring; Lee's Ferry; three
(ad. males), Tibbet Spring, 16 Oct.; Wah-
weap Creek; Escalante River Bay; Driftwood
Canyon; T43S R2W Sec 19, 7 Dec; Three
Garden; 2 miles south Church Wells, 10
Nov.; T43S R2W Sec 24, 7 Dec; along the
Escalante River; Paria Plateau, 29 Sept., and
mouth of Long Canyon 18 Dec; BYU (1972)
Driftwood Canyon; Three Garden; mouth of
Escalante Canyon, 24-25 June; and Wire-
grass Spring, 22 July; BYU (1973) Highway
273/Wahweap Creek, 20 Apr.; five miles
north of Site 34, 1 May; left fork Tibbet
Spring, 5 June; Sites 6, 23, 28, and 30, 6-13
June; Sites 2 and 3, 12 July-9 Aug.; Cotton-
wood Wash, 9 miles north Highway 89; Lee's
Ferry, Paria/Colorado Rivers, Paria River,
Tibbet Spring, and Wahweap Creek, 30
Apr.-15 June; Glen Canyon City, 17 Aug.
The greatest numbers were seen in the win-
ter. Three large flocks were seen 8 Nov.
1971, and two flocks each containing about
300 birds and another containing 50 were
seen 7-18 Dec. 1971. From 1971-1973, 2
finches were seen in Apr., 46 in June, 8 in
July, and 12 in Aug. In April-June 1973,
when House Finches are expected to nest, 8
were seen in Tamarix, 2 in Pinus edulis, and
one each in Coleogijne, Juniperus-gra.ss, and
Salix; 2 were seen during July in semiarid
grasslands and Tamarix. The 4 seen in June
1972 were in hanging gardens. The Novem-
ber and December 1971 flocks were in semi-
arid grasslands and Juniper us. The August
1973 sightings were in semiarid grassland
(10), Artemisia (5), woodland (2), Vanclevea-
grass (1), and Glen Canyon City (1). Per-
manent resident; abundant in winter.
Pinicola enucleator (Linnaeus). Pine Gros-
beak. Presnall (1934) Bryce Canyon Rim.
Summer resident.
Leticosticte tephrocotis tephrocotis (Swain-
son). Gray-crowned Rosy Finch. Behle et al.
(1958) Paria River (see Black Rosy Finch).
Winter resident or transient.
Leticosticte atrata Ridgway. Black Rosy
Finch. Behle et al. (1958) Paria River, about
300 mixed with Gray-crowned Rosy Finch, 6
Mar. 1946; BYU (1971) Warm Creek; seven
miles above Escalante; Warm Creek drainage
near Tibbet Spring, 200-400 seen 6 Dec; 3
females, 2 males, Tibbet Canyon, 7 Dec;
Nipple Bench above Tibbet Canyon, 1000
seen 9 Dec; and Vi mile up long Canyon, 200
seen 16 Dec; BYU (1972) three miles south
of Tibbet Spring in Tibbet Canyon, 200 seen
14 Feb. and Grosvenor Arch pond, 100 seen
26 Oct.; BYU (1973) Tibbet Canyon, 25 seen
25 Jan. and Tibbet Spring, 100 seen 1 Mar.
The earliest fall sighting was 26 October; the
latest spring sighting was 6 March. They
were encountered in Chnjsothamnus (1000),
washes (300), woodlands (200), Artemisia
(100), and salt wash (25). Locally abundant,
winter resident.
Cardiielis pinus pintis (Wilson). Pine Sis-
kin. Presnall (1934) Bryce Canyon; Benson
(1935) Navajo Mountain; Woodbury and Rus-
sell (1945) Russell saw from one to four birds
almost daily on Navajo Mountain, 13-21 July
{Picea-Abies forest); BYU (1971) Grand
Bench, 5 Oct. Summer resident and transient.
Carduelis tristis pallida (Mearns). Ameri-
can Goldfinch. Behle et al. (1958) confluence
of Calf Creek /Escalante River 1954; Long
(1937) Bryce Canyon, 23 Nov. 1935; Russell
and Thompson (1964) Bryce Canyon; BYU
(1971) Grand Bench, 5 Oct. and three (males)
Nipple Spring, 6 Dec; BYU (1972) Cedar
Mountain. Permanent resident.
December 1980
Atwood et al.: Kaiparowits Vertebrates
335
Carduelis psaltria hesperophila (Oberhol-
ser). Lesser Goldfinch. Presnall (1934) and
Behle et al. (1958) Bryce Canyon; Woodburv
and Russell (1945) Navajo Mountain, two
adults and three young seen on 10, 11, and 13
July 1936 and Navajo Mountain Trading
post, two or three in Populus fremontii,
27-29 July 1936; believed to nest in larger
stands of Populus and Salix (Monson Expedi-
tion in Navajo Country as recorded in Wood-
burv and Russell 1945). Permanent resident.
Loxia curvirostra Linnaeus. Red Crossbill.
Presnall (1934) and Russell and Thompson
(1964) Bryce Canyon; Woodbury (1939) 2,
Navajo Mountain obtained from a flock of
12, i.e., one L. c. bendierei, 13 Aug. 1935, the
other L. c. stricklundi, 12 Aug. 1935; Wood-
bury and Russell (1945) flock of 30-40 cross-
bills were seen many times by Russell in
stands of Pinus ponderosa and Picea-Ahies on
Navajo Moimtain, 2-23 July 1936. Transient.
Pipilo chlorurus (Audubon). Green-tailed
Towhee. Presnall (1934) Bryce Canyon;
Woodbury and Russell (1945) two, Kaiparo-
wits Plateau at 7000 ft in bushes of Amekm-
chier, 3 and 12 Aug. 1937; seen several times
in Amelanchier and Querctts in the heads of
canyons on Kaiparowits Plateau, late July
and early Aug. 1913 [1937], believed to be
nesting. Extreme dates for the general area
by Woodbury and Russell (1945) were 22
Aug.- 13 Oct. in the fall and 29 April- 11 May
in the spring. Summer resident.
Pipilo erythrophthalmus montanus
Swarth. Rufous-sided Towhee. Benson (1935)
three, Navajo Mountain in thickets beneath
Pinus ponderosa; Woodbury and Russell
(1945) three, Navajo Mountain, 3 July and 7
Aug. 1936, and 11 Aug. 1935, and five from
Kaiparowits Plateau, 30 July- 10 Aug. 1937
{Archtostaphylos under Pinus ponderosa, Sa-
lix, Populus treniuloides, chaparral, i.e., very
common in bushy thickets, 6000-8500 ft);
Behle et al. (1958) Escalante drainage; Behle
and Higgins (1959) one of two seen was col-
lected at mouth of Aztec Creek, 26 July
1958; eight seen Navajo Creek Canyon, 9
July 1936 (Rainbow Bridge-Monument Val-
ley expedition); Russell and Thompson (1964)
BWce Canyon; BYU (1972) Three Garden lo-
cated one mile above the confluence of San
Juan River and Lake Powell, 4 July (hanging
gardens). Summer resident, a few mav win-
ter.
Passerciilus sandwichensis nevadensis
Grinnell. Savannah Sparrow. Presnall (1934)
Bryce Canyon; Tanner (1940a) Kaiparowits
Basin; Behie et al. (1958) 10 miles south of
Escalante, 1954; Hay ward et al. (1958) Esca-
lante drainage. Transient.
Ammospiza leconteii (Audubon). Le-
Conte's Sparrow. [Hypothetical] BYU (1973)
Site 2, one seen by Clyde L. Pritchett, 1 Aug.
This species is considered bv Behle and Perrv
(1975) and Hay ward et al. (1976) to be of ac-
cidental, rare occurrence in Utah. They note
a specimen from Provo, Utah, 24 Dec. 1927,
and three observations there on 10 Mar.
1928; one was also seen at Moab, 19 Apr.
1966. This adds one additional sight record
for the state. Transient.
Pooecetes gramineus confinis Baird. Ves-
per Sparrow. Presnall (1934) Bryce Canyon;
Tanner (1940a) Kaiparowits Basin; Behle et
al. (1958) 10 miles south of Escalante, 1954;
BYU (1972) one mile above confluence of the
San Juan /Colorado Rivers, 25 June (hanging
garden pool); BYU (1973) one (ad. female),
two miles southwest Glen Canyon City, 30
Apr.; Site 1, two seen 30 .\pr.; Site 8, 3 May;
and one mile north of Site 10, several flocks
seen, 1 May; 1973 sightings were in Van-
clevea-grsLss, Grayia-Coleogyne, and Ephedra-
grass. Siunmer resident, more common in mi-
gration.
Chondestes grammacus strigatus Swain-
son. Lark Sparrow. Presnall (1934) Bryce
Canyon; Tanner (1940a) Kaiparowits Basin;
Woodbury and Russell (1945) two, Kaiparo-
wits Plateau, 4 Aug. 1937; Navajo Mountain
Trading Post {Artemisia iridcniata,
6000-7000 ft); Behle and Higgins (1959) Col-
orado River/San Juan River and river mile
75, 2-3 .\ug. 1938 (Woodbury's field notes of
Rainbow Bridge-Monument Valley Expedi-
tion, 1938); Hayward et al. (1958)' Escalante
drainage; Behle et al. (1958) Bryce Canyon,
confluence of Calf Creek /Escalante River,
and Hayward et al. (1958) Escalante drain-
age; Behle et al. (1958) Bryce Canyon, con-
fluence Calf Creek /Escalante River, and 10
miles south of Escalante, 7 May 1954. Ex-
treme dates of arrival and departure for the
general area are 28 Apr. and 14 Sept., re-
spectivelv (Woodbury and Russell 1945).
336
Great Basin Naturalist
Vol. 40, No. 4
Nesting takes place on open flats in medium
dense brush and often in savannah at the
woodland edges or openings (Woodbury and
Russell 1945). Summer resident.
Aimophila cassinii (Woodhouse). Cassin's
Sparrow. [Hypothetical] BYU (1973) Sites 1,
2, and 3, small flocks common, 30 Apr.;
Wahweap Creek, 30 Apr.; and Tibbet Can-
yon, small flocks common, 1 May. The Cas-
sin's Sparrow was placed on the hypothetical
list of the birds of Utah by Behle and Perry
(1975). The present observations made by
Lloyd Pack are questionable. He possibly ob-
served Brewer's and Chipping Sparrows in-
stead of Cassin's Sparrows. Transient.
Amphispiza bilineata deserticola Ridg-
way. Black-throated Sparrow. Woodbury and
Russell (1945) 3, Rock Creek /Colorado Riv-
er, 21 July 1937; Navajo Mountain Trading
Post, 27 July 1936 and 6 Aug. 1935 (desert
brush at 3200-6500 ft and Artemisia triden-
tata at 5800-6500 ft); Woodbury and Russell
(1945) give extreme dates of occurrence in
the Navajo County as 12 Apr. and 1 Sept.
with exception of 20 birds seen 29-30 No-
vember; Behle and Higgins (1959) report that
during the Rainbow Bridge-Monument Val-
ley expedition Woodbury observed this spe-
cies at river mile 63, 5 Aug. 1938, river mile
50, a few seen 6 Aug. 1938, four miles up
Last Chance Creek, a few seen 7 Aug. 1938,
river mile 41, 2 seen 8 Aug. 1938, and mouth
of Paria Creek, 3 seen 11 Aug. 1938; Behle et
al. (1958) 10 miles south of Esca-
lante/Kaiparowits Plateau, Aug. 1957; Behle
and Higgins (1959) mouth of Aztec Creek
and Woodbury saw more than 6 along the
Colorado River from mile 63 to the Paria
Creek, 5-11 Aug.; BYU (1971-1973) Sites 1,
2, and 30; Glen Canyon City, Cottonwood
Wash/ Paria River; Tibbet Spring; and one
mile above confluence of San Juan/Colorado
Rivers. Our earliest spring observation of this
species is 1 May (1973). The latest in the fall
is 8 Aug. (1973). Numbers seen were greatest
in Aug. (35) and least in May (6); in addition,
20 were seen in June and 12 in July. Fifteen
of the birds were seen in scattered bnish con-
.sisting of Artemisia tridentata-Ephedra and
other desert species, 4 in Juniperus or Juni-
penis- Artemisia, and 2 in Tamanx. Wood-
bury and Russell (1945) believe this species is
closely associated in its distribution with Co-
leogyne, from the lower part of the pygmy
forest downward. Common summer resident.
Amphispiza belli nevadetisis (Ridgway).
Sage Sparrow. Benson (1935) south of Navajo
Mountain; Woodbury and Russell (1945) 2,
Warm Creek, 15 July 1936, and five miles
south Navajo Mountain Trading Post, .30 July
1936 {Salix, Artemisia, 3100-6500 ft); Wood-
bury and Russell (1945), commonest bird of
open Artemisia flats south of Navajo Moun-
tain, with at least 100 seen in a single day;
Russell and Thompson (1964) Bryce Canyon;
BYU (1971) one mile south Glen Canyon
City, 10 Nov.; Grand Bench, 10 Oct.; Warm
Creek Inlet, 8 Nov. and 6 Dec; Tibbet Can-
yon, near spring, 4 Dec; and two miles east
of Nipple Butte, 6 Dec; BYU (1972) Site 1,
25 Jan. and 7 Feb.; Tibbet Spring, 25 Jan.;
BYU (1971-1973) Sites 2, 3, 6, 10, 12, 13, 14,
15, 23, and 28 and Cottonwood Wash. Sage
Sparrows winter in the area; the largest num-
bers were seen in Dec. (67) and Jan. (34).
None were seen in Mar., 17 in Apr., 12 in
June, one in July, 18 in Aug., none in Sept.,
and 3 in Nov. Although this species is ex-
pected to occur in Artemisia during the nest-
ing season (May-June), it was encountered in
Graijia (2), Coleogijne-Vanclevea (19), Arte-
7nisia tndentata (1), Artemisia-Juniperus (1),
and Atriplex (3). During the remaining period
they were recorded in Mahonia (40), Arte-
misia tridentata (25), Cercoearpiis (10),
Chnjsothammis (6), Juniperus (2), Juniperus-
Artemisia (1), Ephedra (1), grass (1), Atriplex
(1), and unidentified shrubs (13). The Dec.
and Jan. birds were recorded in Cercocarpus
(10), Ephedra-Atriplex-Chrysothamnus (4),
Mahonia (40), Artemisia tridentata (25), and
mixed shrub-grass (5). Common permanent
resident.
Jtinco hyemalis (Linnaeus). Dark-eyed
Junco. In the thirty-second supplement to the
American Ornithologists Union Check-list of
North American Birds (Auk 90:411-419,
April 1973), the committee on classification
and nomenclature considered most of the
Juneo species conspecific with /. hyemalis.
Following this pattern we are combining all
the juncos in the Kaiparowits Basin, except
one, into the hyemalis complex. The former
specific names now become the subspecific
names. Junco caniceps is currently under
December 1980
Atwood et al.: Kaiparowits Vertebrates
337
study and therefore is presently maintained
as a separate species.
Junco hyemalis oreganus (Townsend) (=/.
oreganus moiitanus and /. o. shiifeldti). Pres-
nall (1934) /. o. oreganus {=}. shufcldti) at
Bryce Canyon; Grater (1947) Bryce Canyon;
Behle et' al. (1958) confluence Calf
Creek/Escalante River, 1954; and BYU
(1971) Site 12; Site 15, 10 Oct.; Cottonwood
Wash, 13 Nov.; Tibbet Spring, 4 Dec; and
Wahweap Creek, 16 Dec; BYU (1972) Wah-
weap Creek, 28 Jan.; Tibbet Spring, 25 Jan.;
and Cockscomb, 28 Jan. The earliest fall ob-
servations of juncos are 10 Oct. 1971 and 26
Oct. 1972; the latest spring sighting is 28 Jan.
1972. The greatest numbers were seen Dec.
(48 + a large flock). Juncos were observed in
Tamarix (35), Cercocai-pus (10), mixed shrubs
(10), Mahonia (3), Chnjsothamnus (3), Popu-
his fremontii-Tamarix (1). Common transient
and winter visitant.
Junco caniceps caniceps (Woodhouse).
Gray-headed Junco. Presnall (1934) Bryce
Canyon; Benson (1935) Navajo Mountain,
breeding bird, common 1-23 July 1933;
Woodbury and Russell (1945) seven, Navajo
Mountain, 14 Aug., 1935 and 2-18 July 1936
{Populus tremuloides, Finns, Picea-Ahies,
8500-10,000 ft); Behle et al. (1958) con-
fluence Calf Creek/Escalante River, 1954;
BYU (1971) xNavajo Mountain summit, 12
Oct.; Cottonwood Wash, 7 Nov., and a large
flock was seen, 7 Nov. (Populus fremontii)
and one was seen in Dec [Juniperus). Found
on Navajo Mountain as a breeding bird be-
tween 1 to 23 July 1936 (Woodbury and Rus-
sell 1945). Common summer resident of Nav-
ajo Mountain, winters in lowlands.
Spizella arborea (Wilson). Tree Sparrow.
BYU (1971) one (male). Warm Creek Inlet, 8
Nov. Rare transient or winter visitant.
Spizella passerina arizonae Coues. Chip-
ping Sparrow. Presnall (1934) Bryce Canyon;
Benson (1935) Navajo Mountain area; Tanner
(1940a) Kaiparowits Basin; Woodbury and
Russell (1945) river mile 69, 3 Aug. 1938; 3
specimens: Navajo Mountain Trading Post,
23 July 1936; Navajo Mountain, 16 Aug.
1935, and Navajo Mountain Trading Post, 23
July 1936 {Pinus ponderosa, pygmy forest,
65()0-90(X) ft); also observed 69 miles above
Lee's Ferry, 3 Aug. 1938; Behle et al. (1958)
confluence Calf Creek/Escalante River, 13
Jime (stand of Junipcrus-grass); Russell and
Thompson (1964) Bryce Canyon; BYU (1973)
Site 28 {Artemisia tridentata); Woodbury and
Russell (1945) reported seeing 1074 individ-
uals of this species (15 sightings) between 12
Apr. and 13 Oct. 1936-38. Our paucity of re-
cords may be due to the difficulty of in-
experienced observers in distinguishing
among the various species of small sparrows.
It is probable that many were misidentified,
e.g., Cassin's Sparrow as previously men-
tioned as well as Brewer's Sparrow. Summer
resident, spring, and fall transient.
Spizella hreweri breweri Cassin. Brewers
Sparrow. Tanner (1940a) Kaiparowits Basin;
Woodbury and Russell (1945) three, Kaiparo-
wits Plateau, 31 July 1937; Navajo Mountain,
8 Aug. 1936; and Navajo Mountain Trading
Post, 30 July 1936 (Artemisia tridentata, pyg-
my forest, 6500-7000 ft); observed at Lee's
Ferry, 14-26 Aug. 1909, bv E. W^ Nelson;
Behle et al. (1958) confluence Calf
Creek/Escalante River, 1954, and 10 miles
south of Escalante, 8 May 1954; BYU (1971)
one. Site 2, 28 Aug.; one. Church wells, 28
Sept.; and one Paria Plateau, 29 Sept.; BYU
(1973) Site 4, 6 June; Site 6, 10 seen 4 July;
Site 23, 5 June; Site 28, 13 June; Cottonwood
Wash road, U/2 miles north of U.S. Highway
89, 6 June and 13 miles north of U.S. High-
way 89, 8 June; Hackberry Can-
yon/Cottonwood Wash, two seen 8 June; and
Kelly Grade/Smoky Mountain, 3 July. Al-
though the species is primarily a bird of Arte-
misia tridentata, most sightings were in open
or scattered desert shrub and one was in a
Juniperus-grass association. Common summer
resident and migrant.
Zonotrichia quenila (Nuttall), Harris
Sparrow. BYU (1973) one specimen (ad.
male), three miles east of Glen Canyon City,
30 Apr. Winter visitor of lower and warmer
vallevs.
Zonotrichia leucophrys (Forster). White-
crowned Sparrow. Tanner (1940a) Kaiparo-
wits Basin; Behle (1948) Lee's Ferry; Behle et
al. (1958) specimens at confluence Calf
Creek/Escalante River (Z. /. gambelii and
oriantha) and 10 miles south of Escalante,
7-8 May 1954; Russell and Thompson (1964)
Bryce Canyon; Presnall (1934) recorded Z. /.
gambelii from Bryce Canyon; it is a common
migrant in Bryce Canyon fall and spring.
338
Great Basin Naturalist
Vol. 40, No. 4
occurring in mixed flocks of Z. /. oriantha;
BYU (1961) two (females), old Paria townsite,
20 May; BYU (1971) Site 15, 9 Oct. (salt wash
vegetation); Cottonwood Wash, 7 Nov.; BYU
(1972) Tibbet Spring, 30 seen 15 Apr.; BYU
(1973) Wahweap Creek, 30 Apr. Transient
and winter visitant.
Passerella iliaca (Merrem). Fox Sparrow.
BYU (1971) one (not kept), Wahweap Creek,
4 Nov. Transient.
Melospiza lincolnii alticola (Miller &
McCabe). Lincoln's Sparrow. Behle et al.
(1958) Bryce Canyon. Transient.
Melospiza melodia montana Henshaw.
Song Sparrow. Russell and Thompson (1964)
Bryce Canyon; Behle and Higgins (1959)
Kane Creek at river mile 41, 2 seen 18 Oct.
1958; BYU (1971) Site 12 (Coleogyne); Glen
Canyon City; and one (not kept). Nipple
Spring, 15 Oct. Transient.
Summary of Birds of Kaiparowits Region
(183)
Permanent Residents (.36)
Biiteo janiaicensis calurtis
Aqtiila chnjsaetos canadensis
Dendragapus ohscii nis
Centrocerciis uropliasianus mopluisianiis
Ottis ash
Otus flammeolus
Bubo virainiantis pallescens
Glaucidium gnoma caUfornicinn
Asio ottis
Perisoreus canadensis capitalis
Ctjanocitta stelleri macrolopha
Aphelocoma coerulescens woodhuuscii
Corvus corax sinuattis
Ci/ninorliinus cijanocephalus
Nncifraga colwnhiana
Pants atricapilltis garrinus
Panis gainbcli
Partis itiornattis ridgivai/i
Psaltripants ntinim us
Sitta carolinensis nelsoni
Sitta canadensis
Sitta pijgniuea nielanotis
Certhia familiaris
Cinchts inexicanus unicolor
Troglodijtes aedon parknianii
Catlierpes niexicanus consperstis
Si(tli(t niexicatia hairdi
Reguhts satrapa
Regultis calenditia
Lanitts ludovicianus
Sturntts vulgaris ntlgaris
Passer domestic us
Car}wdacits niexicanus frontalis
Carduelis tristis pallida
Carduelis psaltria hesperophila
Aniphispiza belli nevadensis
Uncommon or Sparse Permanent Residents (7)
Ualiaeettts leucocephalus
Lophortyx gambelii gandjelii
Phasianus colchicus
Alectoris ehukar
Strix occidentalis lucida
Colaptes auratits cafer
Sturnella neglecta neglecta
Spring-Siinnner Residents-May Be I'ncommon (3)
Egretta tint la
Nycticorax n ycficorax
Anas strepera
Summer Residents— Common or Abundant (57)
Ardca hcrodias
Accipiter striatus velox
Buteo sivainsoni
Faico niexicanus
Actitus macularia
Cohimba fasciata fasciata
7,enaida macroura
Chordeiles minor lienryi
Aeronautcs saxatalis saxatalis
ArcliUochits alexandri
Selasphorus platycents platycerus
Selasphorus rufus
Spli yrapicus tli yroideus
Picoides villosus leucothorectus
Picoides pubescens leucttrus
Tyrannus verticalis
Tyrannus vociferans vociferans
Myiarchus cinerascens cinerascens
Sa yorn is n igrica n s
Sayornis say a say a
Enipidonax traillii
Enipidonax oberliolseri
Enipidonax icrigli tii
Enipidonax difficilis lielhnaiiri
Sutallornis borcalis
Ereniophila alpcstris leucohienui
Tacliycineta thalassina lepida
Riparia riparia riparia
Stelgidopteryx ruficollis
Petrochclidon pyrrhonota pyrrhonota
Tliryomancs beivickii ereniophilus
Minius polyglottos leucopterus
Duniettella carolinensis
Toxostonia bendirei
Catharus guttatus auduhoni
Polioptila caerulea anioenissima
Phainopepla nitens lepida
Vireo gihiis
Vennivora virginiae
Virniivora luciae
Dcndroica petechia morconii
Dendroica graciae gruciae
Geothlypis triclias occidentalis
Icteria virens
Icterus parisorunt
Icterus iialbula bullockii
December 1980
Atwood et al.: Kaiparowits Vertebrates
339
Molotlinis ater ohscurus
Fimngii ludoiiciana
Fhciicticus inelanoccphalits ruchinoccpluilus
Gtiinicd caerulcd interfusa
Passerina cijiineu
Passerina amociui
Carpoddciis cassin n
Pinicohi cniiclcdtor
Pipilo rhlorurtis
Chondcstcs oranuiidcus strigattis
Amphifipizu hilincdld dcsciiiroUi
Summer Residents, Some Mav Winter (6)
Brdtita cdiuidcnsis
Falco spdrverius spdnciiiis
Salpinctes obsoletus oh.wlctiis
Tardus migmtorius propiiupitis
Skdui curmroides
Pipilo en/throphthdhmi.s uiontdniis
Summer Residents, Transient or Migrant during Spring
and/or Fall (10)
Accipitcr cooperii
Sphyrdpicu.s vdiiiis
Vireo solitariiis
Dendroica cororidta duduhoni
Dendroicd niorescens
Opowrnis tolmiei
Cardiu'lis piniis pinus
Pooecetcs grdmineiis con f inns
Spizella passerina arizonae
Spizella breweri hreweri
Uncommon Summer Residents, Mav Be Spring-Fall
Transients (12)
Carthartes aura tetcr
Accipiter gentilis atricapillus
Falco peregrinus anatum
Falco colnmbarius bcndirei
Fulica anwricana
Cbaradrius vocifcrus lociferus
Athene cuniculdria hijpugaea
Phalaenoptilus nnttallii nuttaUii
Melanerpes lewis ,
Contopus sordidulus veliei
Vireo bellii
Vireo vicinior
Late Fall/Fall-VVinter Residents (3)
Podiceps nigricollis
Podih/nduts podiceps
Pica pica hudsonid
Winter Residents (4)
Buteo logopus
Leucosticte dtratd
Junco hyenialis oreganus
Zonotrichia cpicrula
Transients, Some May Nest, Some Mav Winter (20)
Aruis creccd
Bucephdla chingula
Myadestes towiisendi toicnscndi
Vennivora ruficapilla ridgtcayi
Dendroica toicnscndi
Dendroica occidenlalis
Wdsoniu pusdla pileolata
Xdnthocephdlus xdnthoceplialus
Agelaius phocn iceus
Euphdgus cydnocephalus
Leucosticte tephrocotis tephrocotis
Loxia cunirostra
Passerculus sandwicliensis nevadensis
Animonspiza leconteii
Airnophila cassinii
Jttnco caniceps canicc})s
7.onotri(liia Icucuphrys
Passerelld ilidca
Melospiz^i lincolnii alticola
Melospiza melodia niontana
Uncommon Spring-Summer Transients (3)
Anas discors
Bucephdld albeola
Larus delawarensis
Spring-Fall Transients (2)
Anas dcutd
Anas cyanoptcra
Fall Transients (3)
Steganopus tricolor
Larus californicus
Hesperiphona vespcrtina vcspcrtina
Uncommon Transients (8)
Pie gad is chili i
Anas platyrhynchos
Aruis amcricana
Aydiya vdlisinerid
Buteo regdlis
Oreoscoptes i>i(intdnus
Bombycilld ccdrorum
Vennivord crlata
Sparse Transients (5)
Catoptrophorus scinipalinatus inorudtus
Calidris inauri
Stellula cdlliope
Myiarchus tyrannulus
Iridoprocne bicolor
Rare Transients or Winter Visitants (4)
Accluuophorus occidentalis
Pelecanus eryth rorh ynchos
Lanius excubilor
S})izclla arborcd
Formerly More Abundant Than Indicated h\ This
Study (4)
Cathartes dura teter
Actitis macularia
PJidldcnoptilus nuttallii nuttallii
Geothylypis trichas occidentalis
Range Extensions into Kaiparowits Basin (5)
Myiarchus tyrannulus
Phainopepla nitens lepida
Sturnus vulgaris vulgaris
Passerina cyanea
Anunospiza leconteii
340
Great Basin Naturalist
Vol. 40, No. 4
Mammals
In general we have followed Durrant's
(1952) classification with regard to the no-
menclature and taxonomy of mammals.
Soricidae (Shrews)
Sorex obscurus obscurus Merriam. Dusky
Shew. This species has been collected on
Boulder Mountain just north of Escalante
(Durrant 1952); it probably occurs on the
portion of Boulder Mountain in the Kaiparo-
wits Basin. This is within the range of the
species as indicated by Durrant.
Sorex palustris naingator (Baird). Water
Shrew. Reported by Tanner (1940a) from the
Kaiparowits Basin.
Sorex merriami leucogenys Osgood. Mer-
rian Shrew. Benson (1935) reported this spe-
cies as S. leucogenys, but in 1939 combined it
with S. merriami as a subspecies. This was
based on five specimens from War God
Spring on Navajo Mountain.
Vespertilionidae (Verpertilionid Bats)
Myotis yumanensis yumanensis H. Allen.
Yuma Bat. Durrant and Dean (1959) collect-
ed 10 specimens from along Glen Canyon
and 8 from a large colony at the mouth of the
Escalante River.
Myotis volans interior Miller. Hairy-
winged Myotis. Reported by Benson (1935)
from Rainbow Bridge.
Myotis leibii melanorhinus (Merriam).
Small-footed Bat. Durrant and Dean (1959)
made a collection at the mouth of Kane
Creek on the Colorado River.
Lasionycteris noctivagans (LeConte). Sil-
ver-haired Bat. Tanner (1940a) reported this
species from the Kaiparowits Plateau.
Pipistrellus Hesperus Hesperus H. Allen.
Western Pipistrelle. Benson (1935) reported a
collection from Rainbow Bridge and Tanner
(1940a) reported it from the Kaiparowits
Plateau. Durrant and Dean (1959) indicate
that this is the most common bat in Glen
Canyon. They collected two individuals at
Lee's Ferry and one from river mile 91.
Cockrum (1960) also collected this species at
Lee's Ferry. Pritchett (1962) reports this spe-
cies occurring on both sides of the Cocks-
comb Ridge; in 1961-1962 it was the most
common bat flying in the early evening.
Eptesicus fuscus pallidus Young. Big
Brown Bat. Benson (1935) reported this spe-
cies from Navajo Mountain Trading Post.
Antrozous pallidus pallidus (LeConte).
Pallid Bat. The occurrence of the pallid bat
in the Kaiparowits region has not been re-
ported in the literature. Pritchett (1962) re-
ports it as the most common late-flying bat
(after 2300 hours) near the Old Paria town-
site. He collected specimens there in 1961
and 1972. BYU (1973) observed them at Sites
2 and 3.
Molossidae (Free-tailed Bats)
Tadarida brasiliensis mexicana (Saussure).
Brazilian Free-tailed Bat. Durrant (1952) in-
dicates the Brazilian free-tailed bat occurs
throughout southern Utah, but did not list a
reference for any being collected in the envi-
rons of the Kaiparowitz Plateau. Hardy
(1941) records them as occurring in Zion Na-
tional Park. BYU did not collect this species
during this study, or did Pritchett (1962).
However, they are one of the most common
bats in Carlsbad Caverns in New Mexico and
we are including them as part of the fauna of
the Kaiparowits Plateau.
Leporidae (Hares and Rabbits)
Sylvilagus nuttallii (Bachman). Nuttall
Cottontail. This species is restricted to the
higher elevation with S. n. grongeri (Allen)
occurring in Bryce Canyon and vicinity as re-
ported by Presnall (1934). The subspecies S.
n. pinetis (Allen) was reported for Navajo
Mountain by Benson (1935) and is apparently
restricted to the mountain ranges east of the
Colorado River. One was observed by BYU
near Site 33, which is one mile north of the
confluence of the Colorado and San Juan
Rivers.
Sylvilagus audubonii (Baird). Desert Cot-
tontail. Durrant (1952) reported S. a. ari-
zonae (Allen) to be found seven miles south-
west of Tropic, Utah. The range of the
subspecies S. a. warreni Nelson in the Kai-
parowits is uncertain. Benson (1935) reported
tlie latter subspecies from the mesa top south
of Navajo Mountain. The Desert Cottontail
December 1980
Atwood et al.: Kaiparovvits Vertebrates
341
or their signs have been observed throughout
most of the Kaiparowits Basin, but no speci-
mens have been retained (BYU 1971-1974).
Lepus americanus bairdi Hayden. Snow-
shoe Kabliit. Russell and Thompson (1964)
list this species as occurring in Brvce Canyon.
Durrant (1952) includes the northwest por-
tion of the Kaiparowits Basin within its
range.
Lepus townsendii Bachman. White-tailed
Jack Rabbit. Durrant (1952) includes the
northwest portion of the Kaiparowits Basin in
the range of this species. Russell and Thomp-
son (1964) list it for Bryce Canyon. We did
not find this species on anv of our collection
sites.
Lepus californicus Gray. Black-tailed Jack
Rabbit. It is common throughout Utah west
of the Colorado River. This animal has been
observed by BYU (1971-1974) crews at
nearly every site studied. They have been
common in Butler Valley, Four Mile Bench,
Smoky Moimtain, and Cedar Mountain. The
subspecies L. c. texianus Waterhouse was ob-
served by our crews east of the Colorado Riv-
er at Sites 19, 20, 21, and 22 near Navajo
Mountain (BYU 1971-1974). Benson (1935)
reported it for the mesa top south of Navajo
Mountain.
Sciuridae (Squirrels, Prairie Dogs)
Eutamias minimus consobrinus (Allen).
Least Chipmimk. Presnall (1934 and 1938)
reported this species for Bryce Canyon. We
have a single collection from Brigham Plains
and have observed it at Site 2 (BYU 1972 and
1973).
Eutamias dorsalis utahensis Merriam.
Cliff Chipmunk. Durrant (1952) cites a speci-
men which was collected eight miles south of
Escalante. Pritchett (1962) collected it on the
Cockscomb Ridge. BYU records (1972-1973)
are for Sites 2, 27, and 28.
Eutamias quadrivittatus (Say). Say Chip-
mimk. Presnall (1934) reported that chip-
niimks were common at Bryce Canyon. Dur-
rant (1952) examined a specimen of E. q.
adsitus (Allen) from Bryce Canyon. Tanner
(1940a) collected a specimen from the north-
em part of the Kaiparowits Basin. We have a
single record from Site 27 (BYU 1974). The
subspecies E. q. hopiensis Merriam was
reported by Benson (1935) from Rainbow
Bridge and Navajo Mountain. Durrant and
Dean (1959) reported two collections from
the mouth of Kane Creek. BYU records
(1971-1974) are from sites 2 and 15, near
Page, along the Colorado River in Driftwood
and Reflection Canyons, and in Three Gar-
den one mile above the San Juan confluence
with Lake Powell.
Marmota flaviventris engelharti Allen.
Yellow-bellied Marmot. The only records of
this species for the Kaiparowits Basin are
those of Patraw and Gray (1932) and Presnall
(1934) for Bryce Canyon.
Ammospermophilus leucurus (Merriam).
Antelope Ground Squirrel. According to
Hansen (1955), two subspecies occur in the
Kaiparowits Basin, viz. A. I. escalante (Han-
sen) on the west .side of the Colorado and A.
/. cinnamomeus (Merriam) on the east side.
BYU crews (1971-1974) have recorded the
former subspecies for Sites 1, 2, 3, 4, 8, 10,
14, 16, 18, 23, 27, 28, and 30, and in all the
drainages west of the Colorado. The latter
subspecies was reported bv Benson (1935)
from Rainbow Lodge. (>ockrum (1960) col-
lected it at Lee's Ferry, and BYU
(1971-1972) collected it at Sites 19, 20, 21,
and 22, and at Three Garden.
SpermophiUis spilosoma crtjptospilotus
Merriam. Spotted Ground Squirrel. This spe-
cies was observed by Benson's party on 13
June 1934, five miles .south of the summit of
Navajo Mountain (Benson 1935). BYU (1972)
records are from Sites 20 and 21.
Spermophilus variegatus (Erxleben). Rock
Squirrel. Presnall (1934) reported that S. v.
utali was frequently seen in Bryce Canyon
and Tanner (1940a) reported it for the Kai-
parowits Basin. BYU (1972-1973) records are
from Cottonwood Wa.sh, Last Chance, Pet
Hollow, and Grosvenor Arch. Durrant and
Dean (1959) observed S. r. utali sporadically
throughout Glen Canyon and obtained one
.specimen of S. v. <[,raimnurus (Say) at .\ztec
Creek. BYU personnel also observed speci-
mens on both sides of Lake Powell. Benson
(1935) reported that this species has been ob-
.served at Rainbow Lodge and Navajo Trad-
ing Post.
Spermophilus lateralis lateralis (Say).
Golden-mantled Ground Squirrel. Presnall
(1934) reported it as common on the rim of
342
Great Basin Naturalist
Vol. 40, No. 4
Bryce Canyon. Tanner (1940a) reported it for
the Kaiparowits Basin.
Cynomys parvidens Allen, Prairie Dog.
Presnall (1934) reported two small prairie
dog towns in Bryce Canyon, "one near Fairy-
land (on the rim) and another about two
miles farther north. A total of about two doz-
en occupied burrows." Tanner (1940a) re-
ported this species for the Kaiparowits Basin.
Tamiasciurus hudsonicus dixiensis Hardy.
Red squirrel. Presnall (1934, 1937) indicates
that this species was quite common in Bryce
Canyon during the 1920s. During 1933 he
observed only two families and six in 1934; in
1935, however, they were quite numerous.
Glaucomys sobrinus lucifugus Hall.
Northern Flying Squirrel. This species was
collected 10 miles southwest of Bryce Can-
yon by Lowell Hansen and reported by Tan-
ner (1940b). Russell and Thompson (1964)
listed it as a rare species in Bryce Canyon.
Geomyidae (Pocket Gophers)
Thomomys bottae (Eydoux & Gervais).
Botta Pocket Gopher. Benson (1935) referred
the material described by Goldman (1937) to
T. h. alexandrc Goldman based on material
collected on Navajo Mountain. According to
Durrant (1952) and Cockrum (1960) this sub-
species is known only from Navajo Mountain
and tlie vicinity south of the San Juan and
east of the Colorado River. Thomomys h. ah-
sonus Goldman, according to Durrant (1952),
occurs only on the west side of the Colorado
and lists a specimen from Escalante. Our
(BYU 1971) only collection is from Site 3 on
Cedar Mountain. However, their tunnels are
scattered throughout the region. Presnall
(1934) reported that pocket gophers were
abundant in Bryce Canyon and are probably
referable to the latter subspecies.
Heteromyidae (Kangaroo Rats, Pocket Mice)
Perognathiis flavus hopiensis Goldman.
Silky Pocket Mouse. Benson (1935) indicates
that four specimens were collected five miles
southeast of Navajo Mountain Trading Post
and tliree from the environs of the trading
post. Durrant (1952) says they are confined to
the region east of the Colorado River in
loose, sandy, sparsely vegetated areas.
Perognathus apache Merriam. Apache
Pocket Mouse. Benson (1935) reports three
collections of P. a. apache Merriam from the
vicinity of Navajo Mountain. According to
Durrant and Dean (1959) and Cockrum
(1960), this subspecies is restricted to the area
south of the San Juan River on the east side
of the Colorado. P. a. caryi Goldman is re-
stricted to the east side of the Colorado River
above the San Juan River (Durrant and Dean
1959). The only specimens of this subspecies
from the Kaiparowits Basin are those collect-
ed by BYU (1972) at Three Garden above the
confluence of the San Juan/Colorado Rivers.
Perognathus longimembris (Coues). Little
Pocket Mouse. This species is represented in
the Kaiparowits Basin by two subspecies, one
occurring on each side of the Colorado River.
PerognatJius I. arizonensis Goldman is re-
stricted to the west side of the river and was
reported by Durrant and Dean (1959) from
Kane Creek and at river mile 34. Our records
(BYU 1971-1974) are from Sites 1, 2, 3, 4, 5,
6, 7, 8, 9, 10, 11, 12, 13, 14, 15, 17, 18, 23,
27, 28, and 30. Benson (1935) described P. I.
acrus (Benson) from material collected at
Rainbow Bridge (type #58624). Durrant and
Dean (1959) took one specimen from river
mile 43. Our records (BYU 1971-1972) are
from Sites 19, 20, 21, and 22; all are east of
the Colorado River.
Perognathus ampins ammodytes Benson.
Arizona Pocket Mouse. Cockrum (1960) in-
dicates that this species has been taken along
the Echo Cliffs which is the southern bound-
ary line for the Kaiparowits Basin.
Perognathus formosus domisaxensis Cock-
rum. Long-tailed Pocket Mouse. Durrant and
Dean (1959) collected four specimens from
Kane Creek, i.e., one from river mile 34 and
three at Lee's Ferry, and indicated, "Ecologi-
cally, these pocket mice differ somewhat
from those kinds farther west. They appear
to favor more rocky areas and occur on hill-
sides and at the junction of hillsides and stabi-
lized terraces." Our records (BYU
1971-1974) are from Sites 4, 9, 10, 15, 27, 28,
and 30. In addition, material from Three
Garden located one mile north of the con-
fluence of the San Juan on the east side of the
Colorado River has been tentatively referred
to this species. When additional material is
December 1980
Atwood et al.: Kaiparowits Vertebrates
343
obtained and analyzed, it may prove, how-
ever, to be P. intennedius crinitus.
Perognathus intermedius criyiittis Benson.
Rock Pocket Mouse. .According to Durrant
and Dean (1959), this subspecies is confined
to the east side of the Colorado River south
of the San Juan River. No specimens were
obtained by them until they reached Aztec
Creek at river mile 68.5. Between this point
and river mile 28 they collected 15 speci-
mens. Benson (1934b) reported a collection
from Navajo Mountain Trading Post and two
specimens from Rainbow Bridge. Cockrum
(1960) obtained specimens from Lee's Ferry.
BYU (1973) obtained specimens from Three
Garden near the confluence of the San Juan
and Colorado River Canyons. These speci-
mens document the range of this species
north of the San Juan River.
Dipodotnys ordii Woodhouse. Kangaroo
Rat. Two subspecies occur in the Kaiparowits
Basin and, of the two, D. o. cupidineus Gold-
man is restricted to the west side of the Colo-
rado south of the Escalante River. Specimens
have been collected by Durrant and Dean
(1959) at Lee's Ferry and Kane Creek; Pres-
nall (1934) for Bryce; Tanner (1940a) for the
Kaiparowits Basin; and BYU (1971-1974) for
Sites 1, 2, 3, 4, 6, 7, 8, 10, 13, 14, 15, 17, 27,
28, 30, and 33.
Dipodcnnys o. longipes (Merriam) is con-
fined to the area south of the San Juan River
and east of the Colorado (Durrant and Dean
1959). Benson (1935) reports nine collections
from south of Navajo Mountain. Our records
(BYU 1971-1972) are from Sites 19 and 33.
Castoridae (Beaver)
Castor canadensis repetitintis Goldman.
Beaver. Prior to the coming of the white
trappers, beavers were widespread through-
out the length of the Colorado River and its
tributaries. Tliis abundance of beaver enticed
the famous French trapper and fur trader
Denis Julien to navigate the Colorado in
1836 as far south as Cataract Canyon. The
best-known trapper in the Canyon was Nath-
an Galloway; in 1895, 1896, and on other oc-
casions he traveled from (ireen River to
Lee's Ferry with William Richmond (Cramp-
ton 1959). Durrant and Dean (1959) reported
that the populations in Glen Canyon were
among the largest in the state. During the
course of their expedition, 30 June to 9 .Au-
gust 1958, they observed beaver or their signs
daily. They indicated that these bank
dwellers fed almost entirely on willows and
only on one occasion was Tamarix utilized.
The creation of Lake Powell has nearly elim-
inated their main source of food and greatly
reduced their numbers. Observations by BYU
field crews (1971-1974) indicate that beavers
utilize representatives of nearlv every woody
species in the drainages of the Colorado.
They have, of necessity, been pushed into the
side canyons, particularlv where permanent
springs or seeps exist. It appears that they
have attempted to build dams in some of
these regions to create a more suitable habi-
tat.
Cricetidae (Voles, Rats, Mice)
Reithrodontomys megalotis (Baird). West-
ern Harvest Mouse. The subspecies R. m.
megalotis (Baird) is restricted to the west side
of the Colorado River in the Glen Canyon
area. Cockrum (1960) collected it at Lee's
Ferry; Durrant and Dean (1959) collected
specimens at river mile 43 and 93; and BYU
(1971-1974) collected it at Sites 2, 3, and 10.
Durrant and Dean (1959) collected R. m. az-
tecus J. A. Allen on the east side of the Colo-
rado at river miles 43 and 83.
Peromyscus eremicus eremicus (Baird).
Cactus Mouse. Cockrum (1960) reported it
from the south side of the Colorado at Lee's
Ferry.
Peromyscus tnaniculatus (Wagner). Deer
Mouse. Durrant and Dean (1959) have estab-
lished that P. m. sonoriensis (LeConte) is the
onlv subspecies of Deer Mouse in Glen Can-
von and that it was the most common mam-
mal in Glen Canyon during their study. They
collected specimens from Lee's Ferry to river
mile 78 on the Colorado. Presnall (1934) re-
ported this subspecies from Bryce Canyon;
Benson (1935) indicated that it was the most
abundant mammal in the region of Navajo
Mountain; and BYU (1971-1973) collected it
at Sites 1, 2, 3, 6, 7, 8, 15, 19, 22, 27, 28, and
30, and Navajo Mountain.
Peromyscus crinitus Merriam. Canyon
Mous. Two subspecies of the Canyon Mouse
occur in the Kaiparowits Basin, viz. P. c.
344
Great Basin Naturalist
Vol. 40, No. 4
auripectus (Osgood) and P. c. stephensi
(Meam.s). Durrant and Dean (1959) noted
that subspecies auripectus occured only on
the east side of Glen Canyon and collected
specimens at river miles 21, 28, 69, 78, and
83. Benson (1935) reported one from Rain-
bow Bridge and Navajo Mountain Trading
Post. Additional records are: BYU (1972) Site
19, BYU (1971-1974) Three Garden above
confluence of the San Juan and Lake Powell,
and Cockrum (1960) at Lee's Ferry and Nav-
ajo Mountain.
Peromyscus c. stephensi, as reported by
Durrant and Dean (1959), occurs throughout
the Kaiparowits Basin west of the Colorado
River. They made collections at river miles
23, 41, 56, and 88, and at Lee's Ferry; BYU
(1971) collected this subspecies at Sites 5, 6,
8, and 12; BYU (1972) also collected it at
Sites 23, 27, and 30.
Peromyscus boylii (Baird). Brush Mouse.
Two subspecies occur in tlie Kaiparowits Ba-
sin, viz. P. b. roioleyi (Allen) and P. b. uta-
hensis Durrant. The former species is re-
stricted to the east side of the Colorado and
has been collected at river miles 28, 43, 69,
78, and 83 by Durrant and Dean (1959).
Cockrum (1960) cites a specimen from Rain-
bow Lodge at the southwest base of Navajo
Mountain and BYU (1972-1974) cites speci-
mens from Three Garden near the San Juan
confluence and at Pool Garden in Reflection
Canyon.
The latter subspecies, Peromyscus b. uta-
Jiensis, occurs only on the west side of the
Colorado and has been collected by Durrant
and Dean (1959) from river miles 34, 41, and
56 and by BYU (1971) from Site 8 and from
Reflection Canyon. Durrant and Dean in-
dicate that this subspecies was nearly as com-
mon as the deer mice. They are reported to
be excellent climbers, preferring brushy habi-
tats among cliffs and rockv areas.
Peromyscus truei (Shufeldt). Pinyon
Mouse. Benson (1935) reported a collection
from the mesa top near Navajo Mountain and
one from the Trading Post. Presnall (1934)
reported this species from Bryce Canyon.
During the course of our study (BYU
1971-1974), we have records for this species
from Sites 2, 3, 6, 1.3, 14, 19, 20 21, 23, 27,
28, and 30, and Navajo Mountain.
Peromyscus difficilis nasutus (Allen) ( =
Peromyscus nasutus nasutus Allen). Long-
nosed Deer Mouse. Benson (1935) reported
this species from Rainbow Bridge in rocky
places where stands oi Juniperiis occur.
Onychomys leucogaster (Wied-Neuwied).
Northern Grasshopper Mouse. Durrant
(1952) indicates that O. I. melanophrys Mer-
riam is confined to the west side of the Colo-
rado River. Presnall (1934) reported it from
Bryce Canyon and BYU (1971-1974) from
Sites 1, 2, 3, 4, 6, 8, 14, 17, 18, and .30; O. /.
pallescens Merriam is confined to the east
side of the Colorado River (Durrant 1952).
Available records are: Benson (1935) several
specimens from 5 and 7 miles south of Nav-
ajo Mountain and BYU (1971-1972) from
Sites 19, 20, and 22. One specimen (No. 181
from Nipple Bench) collected by BYU (1971)
appears to be an intergrade between these
two subspecies.
Neotoma albigula laplataensis Miller,
White-throated Wood Rat. Durrant and
Dean (1959) indicate that this species is con-
fined to the east side of the Colorado River in
Glen Canyon. Collections were made at river
miles 69 and 78. Benson (1935) reported a
collection from Rainbow Bridge and the
mesa top south of Navajo Mountain. Studies
by BYU (1971-1974) in Glen Canyon and vi-
cinity confirm the observations of Durrant
and Dean. In addition, collections by BYU
(1971-1972) were made at Sites 19 and 21.
Neotoma lepida monstabilis Goldman.
Desert Wood Rat. Durrant and Dean (1959)
state that, "members of this species are the
counter parts of N. albigula and A', mexicana
of western side of the Colorado River." Col-
lections of N. /. monstrabilis were made at
river miles 41, 56, 88, and 91. Additional re-
cords for the Kaiparowits Basin are: Presnall
(1934) Bryce; Tanner (1940a) Kaiparowits
Basin; Pritchett (1962) Cockscomb; and BYU
(1971) Sites 1, 2, 3, 4, 6, 8, 12, and 13, Step
Garden, and Reflection Canyon.
Neotoma stephensi relicta Goldman.
Stephen's Wood Rat. Benson (1935) reported
one collection from Rainbow Bridge, four
specimens from south of Navajo Mountain on
tlie mesa top, and four from Navajo Moun-
tain Trading Post. We have not collected this
species at any of our sites. ^
December 1980
Atwood et al.: Kaiparowits Vertebrates
345
Neotoma mexicana inopinata Goldman.
Mexican Wood Rat. This species is restricted
to the east side of Glen Canyon (Durrant and
Dean 1959). Specimens were obtained bv
them from river miles 69, 78, and 83. Benson
(1935) reported it from War (iod Spring on
Navajo Moimtain. BYU (1972-1974) record
are from Three Garden just above the con-
fluence of the San Juan on the east side of the
Colorado River and from Ribbon Garden in
Ribbon Can von.
Neotoma ciyierea arizonae Merriam. Bush-
tailed Wood Rat. According to Durrant and
Dean (1959), N. cinerea is the onlv species of
wood rat that occiu-s on both sides of Glen
Canyon. Neotoma c. arizonae occurs only on
the east side and N. c. acraia on the west side.
However, no specimens of the latter sub-
species were reported bv Durrant and Dean.
They did collect specimens of A\ c. arizonae
from river miles 21, 43, and 83. Additional
records for the Kaiparowits Basin are: BYU
(1971-1972) Navajo Mountain, Site 27, and
Site 33.
Microtus mexicanus navajo Benson. Mexi-
can Vole. Benson (1934a and 1935) described
this species from specimens obtained from
Soldier and War God Springs on the east
slope of Navajo Mountain. Thev were col-
lected in a Ceanothus, Symplioricarpos,
Arctostaphylos, and Rosa vegetation type.
This taxon is known only from the type col-
lections.
Lagurus curtatiis intermedins (Taylor),
Sagebrush Vole. Presnall (1934) reported a
specimen from Bryce Canyon, but was
unable to assign a species name. Durrant
(1952) cites a specimen of this species from
David Hollow in Bryce Canyon that may be
the one cited by Presnall. We did not en-
counter this species at any of our trap sites.
Erethizontidae (Porcupine)
Erethizon dorsatum (Linnaeus). Porcu-
pine. According to Durrant (1952) two sub-
species occur in the Kaiparowits Basin, viz.
£. d. epixanthnm Brandt on the west side of
the Colorado River and E. d. consei Mearns
on the east side. Durrant and Dean (1959) re-
ported that expedition members observed
one animal in Navajo Canyon. Weight (1932)
and Presnall (1934) reported the former sub-
species from Bryce Canyon and BYU
(1971-1972) reported it from Sites 2, 3, and
4, and Reflection Canyon. Their signs are not
uncommon in stands of Finns ednlis.
Canidae (Coyotes, Foxes, and Wolves)
Canis latrans Say. Coyote. Two subspecies
occur in the Kaiparowits Basin, i.e., C. /. estor
Merriam and C. /. lestes Merriam. The for-
mer is by far the most conunon reported bv
Tanner (1940a) for the Kaiparowits Plateau;
bv BYU (1971) for Sites 3, 10, 11, 12, 15, and
Wahweap Creek; by BYU (1972) for Site 14,
Butler Valley, Cottonwood Wash, Covote
Creek, near Glen Canvon, and Warm Creek;
and by BYU (1973) for Sites 3, 6, 8, 23, 27,
and 28, and Smoky Mountain. The first re-
cord of this species in Glen Canyon was
made by members of Powell's first Colorado
River expedition in 1869. While in the vicin-
ity of the Escalante River, Sumner (Powell
1875) records as follows, "Dunn killed a half-
stayed coyote near camp, the only sign of an-
imal life we have seen for three davs." Dur
rant and Dean (1959) observed numerous
signs of coyotes in Glen Canyon proper and
in several side canvons. The senior author, in
company with S. L. Welsh and J. R. Mur-
dock, witnessed in 1971 the fall and death of
a coyote from a 200-foot cliff in Last Chance
Creek.
Canis I. lestes is known only from the
northern portion of the Kaiparowits Basin as
reported bv Presnall (1934) from Bryce Can-
yon. BYU (1973) observed coyotes north of
Canaan Mountain along High\\a\ 12 and
these may be C. /. lestes.
Canis lupus youngi Goldman. Wolf. .\c-
cording to Young and Goldman (1944) and
Cockrum (1960), the wolf formerly occurred
throughout this region, but no specimens
were cited. It is doubtful that it occurs in the
Basin at the present time.
Vulpes vulpes maeroura Baird ( = Vidpes
fulva maeroura Baird). Red Fox. .\t present
the red fox is rare in Utah due to hunting,
trapping, and use of poison bait stations. Sev-
eral records were reported prior to 1940.
Presnall (1934) reported that this animal was
occasionally seen at Bryce Canyon; Benson
(1935) reported one from the north side of
Navajo Mountain; Durrant and Dean (1959)
346
Great Basin Naturalist
Vol. 40, No. 4
indicate that one was seen two miles east of
Rainbow Bridge by a member of their expe-
dition. Durrant (1952) cites one from Wah-
weap Creek and states, "possibly the red fox
of the Colorado River is an undescribed sub-
species . . . and I tentatively refer them to V.
/. macroiira."
Urocyon cinereoargenteus scottii Mearns.
Gray Fox. The gray fox occurs throughout
the Basin and has been reported by Presnall
(1934) for Bryce Canyon and Benson (1935)
for Navajo Mountain; and Hayward et al.
(1958) reported their signs from nearly all
areas studied from the Paria to the Escalante
River. BYU (1971-1974) observed this animal
in Cottonwood Wash, Cockscomb, and on
Brigham Plains. Tliis species is much more
common than the Red Fox.
Ursidae (Bears)
Ursus americanus cinnamomum Audubon
& Bachman. Black Bear. Presnall (1934) in-
dicated that in 1922 a black bear was killed
in Bryce Canyon; and "Mr. Ruby Syrett and
Ranger Cope both say that black and grizzly
bears used to cross the Bryce Canyon region
at irregular intervals, apparently traveling
between the Parowan Moimtains on the west
and the Escalante Mountains on the east.
Bears are still found in both ranges, although
rarely." One of us (Atwood) observed in 1965
tracks of a bear on the south slope of the
Boulder Mountains. On 11 June 1975, tracks
of a solitary bear were observed by S. L.
Welsh at the head of Paradise Canyon about
11 miles north of Horse Mountain junction.
The left hind foot track measured 10.2 X
22.9 cm. In late June 1975, a small bear was
sighted at Canaan Peak by BYU personnel.
Ursus horribilis Merriam. Grizzly Bear.
Presnall (1934) indicates that they have
crossed through Bryce. For additional infor-
mation the reader is referred to the discussion
of the preceding species.
Procyonidae (Ring-tailed Cat, Raccoon)
Bassarisciis astutus (Lichtenstein). Ring-
tailed Cat. According to Durrant (1952) the
Colorado River serves as a barrier for two
subspecies. A specimen of subspecies B. a.
arizonensis Goldman was obtained from W.
Wilson, which he had shot in a chicken coop
at Rainbow Lodge in November 1932 (Ben-
son 1935). Wetherill, Flattum, and Stearns
(1961) recorded in their journal on 15 Janu-
ary 1931 that a ring-tailed cat came into
camp. They were in or near Bridge Canyon.
The entry by John Wetherill on 17 January
states, "Lots of skunks around, some ring-
tailed cats, fox. . . ." Bassariscus a. nevadensis
Miller has been reported by trappers and lo-
cal residents of Kaiparowits Basin to be fairly
common. Durrant and Dean (1959) observed
small five-toed tracks in the talus dust and
along the base of ledges throughout Glen
Canyon. During the course of our study we
have not met directly with this animal, but
have observed their tracks in the major drain-
ages.
Procyon lotor pallidus Merriam. Raccoon.
Hall and Kelson (1959) list this subspecies as
occurring throughout the Colorado and its
tributaries. Durrant and Dean (1959), how-
ever, were unable to find any evidence to
verify their presence in the Colorado drain-
age. Observations by BYU (1971-1974) con-
firmed the conclusions of Durrant and Dean.
Mustelidae (Weasels, Skunks, etc.)
Mustela erminea muricus (Bangs). Ermine.
Durrant (1952) reports one specimen from
Boulder Mountain north of Escalante. This
species probably occurred throughout the
Boulder Mountains and probably in the re-
gion of Bryce Canyon
Mustela frenata nevadensis Hall, Long-
tailed Weasel. Presnall (1934) reported this
species was common at Bryce Canyon. Ben-
son (1935) indicated that one was observed at
Rainbow Lodge in 1932. The Long-tailed
Weasel is probably the most widely dis-
tributed carnivore in Utah (Durrant 1952).
Taxidea taxus berlandieri Baird. Badger.
Presnall (1934) indicated that they were com-
mon on the rim of Brvce Canvon. Benson
(1935) observed fragments of badger taken by
trappers in the Navajo Mountain region. Tan-
ner (1940a) reported them from the Kaiparo-
wits Plateau. Durrant and Dean (1959) in-
dicated that this species was one of the most
common and abundant carnivores in Glen
Canyon. They observed their tracks and bur-
rows everywhere. Our records (BYU
December 1980
Atwood et al.: Kaiparowits Vertebrates
347
1971-1974) indicate their presence at Sites 1,
3, 13, 19, 27, and 34, and in every drainage in
the basin.
Spilogale gracilis Merriam. Spotted Skunk.
The Colorado River is a barrier for kiteral
movement of this animal with S. g. gracilis
Merriam confined to the east side. It was re-
ported by Benson (1935) for the Navajo
Mountain area. S. g. saxatilis (Merriam) is re-
ported by Presnall (1935) from Bryce Can-
yon. BYU (1972) noted their occurrence on
Cedar Mountain south of Glen Canyon City
and in Little Valley Creek at the south base
of the Kaiparowits Plateau. Durrant and
Dean (1959) observed signs of this species on
both sides of the Colorado River, and one
was observed by investigators from BYU at
Three Garden on the east side of Glen Can-
yon.
Mephitis mephitis estor Merriam. Striped
Skunk. Presnall (19.34) indicates that this spe-
cies was rare at Bryce Canyon. Durrant
(1952) reports that the subspecies M. m. estor
is limited to the Colorado River drainage in
.southern and eastern Utah. Specimens are
cited for both San Juan and Washington
Counties, but are not cited for the Kaiparo-
wits Basin.
Lontra canadensis nexa Goldman. River
Otter. Gregory (1938) reports otter from
Glen Canyon. Durrant and Dean (1959) ques-
tion his recording of the species in Glen Can-
yon, but do not discard the possibility entire-
iv.
Felidae (Cats)
Felis concolor kaihabensis Nelson & Gold-
man. Moimtain Lion. Benson (1935) indicates
that an animal was killed near Inscription
House by Navajos. Presnall (1934) reported
that an occasional lion wanders through
Bryce Canyon. Observations of this species
were made by BYU (1971 and 1973) in
Nipple Creek, Tibbet Canyon, and Willow
Creek. The observation in Willow Creek was
a female with two kittens. In addition, a
solid-black-colored lion was observed Octo-
ber 1976 on Cannon Peak.
Lynx rufus baileyi Merriam. Bobcat. Ap-
parently the only specimens available from
the Kaiparowits Basin are those collected bv
BYU (1973) in Tibbet Canyon, Buckskin
Gulch, and Cottonwood Canyon. Tracks and
scat have been observed throughout the re-
gion.
Cervidae (Deer and Elk)
Cervus elaphus nehoni V. Bailey. Wapiti,
Elk. Cope (1932) reported that A.'w. Ivins,
an early resident of this region, informed him
that, according to the Indians, elk once re-
sided in Bryce Canyon and the Paria Valley.
Mr. Amnion Davis, of Cannon ville, informed
him that he had found part of the head and
horns of a bull elk on Willis Creek just south-
east of Bryce Canyon Rim. This material is
now in the museum collection of Brvce. Pre-
snall (1934) reports further on the elk of
Bryce-Paria Valley as follows: "now extinct
in this region, but one was reported killed in
Willis Creek many years ago bv a Mr. John-
ston of Cannonville."
There is some question as to which is the
correct name, i.e., Cervus canadensis nelsoni
y. Bailey or Cervus elaphus L. Since the
question has yet to be resolved, this treat-
ment follows Durrant (1952).
Odocoileus hemionus hemionus (Rafi-
nesque). Mule Deer. No records of deer in
the Kaiparowits were made by early explor-
ers of the region. Pre.snall (1934) indicated
that they were common in Bryce Canyon.
During the course of our study (BYU
1971-1974) frequent sightings have been
made, especially along tributary canyons
along Lake Powell, .\nimals have been ob-
served onlv where water and forage are ade-
quate throughout the year and they appear
to be residents. The main areas of concentra-
tion (sens, lat.) are on Four Mile Bench,
Cockscomb. Canaan Mountain, Bryce Can-
yon, and the Kaiparow its Plateau.
Antilocapridae (Pronghom)
Antilocapra americana americana (Ord).
Pronghorn. Tanner (1940a) indicates tliat
pronghorn antelope were common in the
Kaiparowits Basin in pioneer days. Presnall
(1938) also reported their occurrence at
Brvce Canyon. Apparently the early herds
were himted too severely because records
since the beginning of this century are lack-
ing.
348
Great Basin Naturalist
Vol. 40, No. 4
In November 1971 some 104 antelope
were transplanted from northeastern Utah to
East Clark Bench, 10 miles west of Glen Can-
yon City, Utah. Since then, 40 separate sight-
ings of these animals have been recorded by
personnel of the U.S. Bureau of Land Man-
agement and BYU (1971-1974). These sight-
ings, for the most part, have been in Cotton-
wood Wash, East and West Clark Benches,
Coyote Creek, and as far north as Butler Val-
ley and on Nipple Bench. Sightings were in-
frequent in 1974 and even fewer in 1975; in
1976 a pair was known to range in the vicin-
ity of Nipple Spring.
Bovidae (Bovids)
Ovis canadensis canadensis Shaw, Moun-
tain Sheep. Powell's party made several ref-
erences to these animals and occasionally
used them for food. On 27 July 1869, Sumner
recorded (Powell 1875), "Killed two moun-
tain sheep today— a Godsend to us, as our
bread and rotten bacon is a poor diet for as
hard work as we have to do." Then on 3 Au-
gust below Music Temple, Sumner recorded,
"pulled out early, made a good run. Saw two
mountain sheep in a little valley on the south
side. How they got there I will leave others
to judge, as there is no outlet to the valley
that a man can climb. Killed one and chased
the other through the natiual pasture for an
hour and pulled out again." Gregory and
Moore (1931) reported sheep from Dark Can-
yon at the southeast base of the Kaiparowits
Plateau.
Durrant (1952) indicates that the species is
disappearing rapidly, and if adequate speci-
mens from extreme south central Utah are
available for study, they would probably
prove to be O. c. mexicanus.
Literature Cited
Allen, J. A. 1893. List of inamiiials collected by Mr.
Charles P. Rowley in the San Juan Region of Col-
orado, New Mexico, and Utah, with descriptions
of new species. Bull, .\nier. Miis. Natur. Hist. 5:
69-84.
American Ornithologists Union. 1957. Checklist of
North American birds. Lord Baltimore Press. Bal-
timore, Maryland. 691 pp.
.'\uERBACK, H. S. 194.3. Father Escalante's journal with
related documents and maps. Utah Hist. Quar-
terly 11:1-10.
Behle, W. H. 1935. Biological reconnaissance of Navajo
Mountain. University of California Piibl. in Zool.
40(14):4,39-456.
1948. Birds observed in April along the Colorado
River from Hite to Lee's Ferry. Auk 65:.303-306.
1960. The birds of southeastern Utah. University
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Behle, W. H., and H. G. Higgins. 1959. The birds of
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No. 7).
Behle, W. H., and M. L. Perry. 1975. Utah birds:
guide, check-list and occurrence charts. Utah
Mus. Nat. Hist., University of Utah, Salt Lake
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Benson, S. B. 1934a. Description of a race of the Mexi-
can vole, Microtiis mexicanus, from southeastern
Utah. Proc. Biol. Soc. Washington 47:49-50.
19.34b. Description of two races of Pewgnathus
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1935. A biological reconnaissance of Navajo
Mountain, Utah. Univ. Calif. Publ. Zool.
40:439-455.
BYU. 1971-1974. Navajo-Kaiparowits reference collec-
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CocKRUM, E. B. 1960. The recent mammals of Arizona:
their taxonomy and distribution. University of
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Cope, M. 1932. Elk formerly in the Paria Vallev. Zion
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Crampton, G. C. 1959. Outline history of the Glen Can-
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Durrant, S. D. 1952. Mammals of Utah. Universitv of
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Grater, R. V. 1947. Birds of Zion, Brvce, and Cedar
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1938. The San Juan country: a geographic and
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December 1980
Atwood et al.: Kaiparowits Vertebrates
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Hall, A. F. 1934. General report on the Rainbow
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Hall, E. R., a.nd K. R. Kelso.n. 1959. The mammals of
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350
Great Basin Naturalist
Vol. 40, No. 4
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pp.
A NEW SPECIES OF FOSSIL CHRYSOTHAMNUS (ASTERACEAE)
FROM NEW MEXICO
Loran C. Anderson'
.\bstr.'\ct.— The new. presmnahlx extinct species, C'lt
trated. The plant materials came from Holoeene packrat
species to extant taxa are discussed.
Study of Holoeene environments through
packrat {Neotoma sp.) middens in Chaco
Canyon, New Mexico, by J. L. Betancourt
and T. R. Van Devender (1980) provided
many samples of Chrysothamnus plant re-
mains. Most materials proved to be C. nau-
seosus ssp. bigelovii, a frequent constituent of
the present-day flora. Several involucres from
a midden (referenced as Mockingbird Canyon
No. 2) represent an undescribed, extinct spe-
cies of Clirysothamnus.
The plant materials of the new species
were in excellent condition; some were sub-
jected to anatomical study. One might ques-
tion iLsing the term fossil for such material,
but precedent has been set by Spilman (1976)
in his description of a new .species of beetle
from packrat middens. Spilman defined fossil
as "a specimen, a replacement of a specimen,
or the work or evidence of a specimen that
lived in the past and was naturally preserved
rather than biuied by man." Since this new
Chrysothamnus is apparently extinct, I
choose to describe it as a fossil.
Chrysothamnus pulchelloides L. C. Ander-
son, sp. nov.
Vegetatively unknown; involucres cylindri-
cal, 7-8 mm long, phyllaries .strongly gradu-
ated in five series in distinct vertical rows,
chartaceous-coriaceous, carinate with en-
larged -subapical co.sta, hyaline margined be-
low apex, acimiinate-cuspidate; disk flowers
4-6, corollas presumably yellow (tawny, as in
dried flowers of extant yellow-flowered spe-
cies), 4.3-4.6 mm long, corolla lobes lanceo-
late, erect, ca 0.5 mm long; stigmatic lines
shorter than appendages (ca 40 percent of to-
tal style branch length); achenes cylindric.
rysothiinnitts pulchelloides, is tormally described and illiis-
middens. .\natomical and phyletic relation.ships of the fossil
1-2 mm long (probably inunature), glabrous,
pappus of capillary bristles, nearlv as long as
corolla.
TYPE: New Mexico, San Juan Co.. (>haco
Canyon National Monument, shallow lenticu-
lar rock shelter in sandstone of .small alcove
at head of minor tributarv of Mockingbird
Canyon, 36° 3' 15" N, 107° 55' W, elev.
1927 m, /. L. Betancourt 6 T. R. Van Deven-
der s. n. in 1979 (Fig. 1; involucres at FSU!).
Midden material from Mockingbird Can-
yon No. 2 was dated at 1910 ±90 B.P. on
Juniperus nionospeiina twigs (/\-2111); other
fossils in the midden included Pinus eduHs,
Rhus aromatica, Cowania mexicana, and Ar-
temisia cf. tridentata. Present-day plants
growing on the talus immediatelv below the
midden include Artemisia htdoviciana, Cir-
sium pulchellus, and Stanleya pinnata; only a
few spindly Juniperus numosperma are found
in the vicinity.
Clirysothamnus puhheUoides is particu-
larly distinctive in its phyllaries that are
acuminate-cuspidate with subapical thick-
ened spots (Fig. 1). It is related to members
of .section Pulchelli (.see Anderson and Fisher,
1970, for sectional composition of the genus)
with its strongly ranked phyllaries and
glabrous achenes with long pappus; it resem-
bles C. pulcheUus in its short corolla lobes
and C. molestus in its hyaline margined phyl-
laries.
Two intact heads of C. pulchelloides were
revived and sectioned as in Anderson (1970).
The phvllaries have prominent secretory
canals and sclercnchyma distribution as in
section Pulchelli. .Vpical portions of the ph> 1-
laries are covered adaxially with glandniar
trichomes. Ovarian vasculature is abundant
'Department of Biological Science. Florida State University'. Tallahassee. Florida 32306.
351
352
Great Basin Naturalist
Vol. 40, No. 4
lO
Fig. 1. Camera lucida drawings of involucre and flow-
er of C. ])tilchcUoidcs. Note style branches are largely in-
cluded in corolla (stamens withered). Pappus bristles av-
erage 97 percent of corolla length, shortened here to
.show corolla lobes more clearly.
with 8-10(12) bundles. The vascular pattern
of the achene-coroUa transition follows pat-
terns "a" and "c" (Anderson, 1970) wherein
the additional ovarian bundles end blindly
distally and the style bundles are derived
from the ventral and dorsal corolla bundles.
Secretory canals a.s.sociated with the bundles
are abundant in the corolla, less frequent in
the achene, and ab.sent in the .style.
Anatomically, the fo.ssil species relates well
to section Pulchelli, but it is less specialized
in vasculature and secretory canal abundance
and also has less pronounced stigniatic lines.
A phylogenetic index of specialization was
developed for Chrysothamnus taxa from flor-
al data in Anderson and Fisher (1970). Extant
members of section Pulchelli have indices of
4.2-7.1; C. piilchelloides would have an index
of specialization of 3.5-lower than all extant
Chrysothamni except the least specialized
subspecies of C. parnji and C. nauseosus of
section Nau-seosi (which is considered the bas-
al section of the genus). The fossil species
may well be ancestral to some of the present
members of section Pulchelli.
To my knowledge, this report represents
the only record of a plant species extinction
documented for the Holocene in this region.
Drastic reduction of the tree species, Pinus
edulis and Juniperus monosperma, in the
Chaco Canyon area occurred during the
Holocene in relation to fuel demands of the
Anasazi culture (Van Devender and
Betancourt, pers. comm.). Desertification
probably continued with Navajo grazing ac-
tivities. These phenomena may have contrib-
uted toward the extinction of C. piilchel-
loides.
Acknowledgments
Special thanks are due Julio Betancourt
and Tom Van Devender for giving me these
and other fossil materials of Chrysothamnus.
They also kindly supplied radiocarbon dates
and descriptive data for the collection site.
Literature Cited
.\nderson, L. C. 1970. Floral anatomy of Chryso-
thamnus (Astereae, Conipositae). Sida 3:466-503.
Anderson, L. C, and P. S. Fisher. 1970. Phylogenetic
indicators from floral anatomx' in Chri/sothanmus
(Astereae, Conipositae). Phytomorphology
20:112-118.
Betancourt, J. L., and T. R. Van Devender. 1980.
Holocene environments in Chaco Canyon, New
Mexico: the packrat midden record. Natl. Park
Serv. Report, Albuquerque, New Mexico.
.SpiL.viAN, T. J. 1976. A new species of fossil Ptiniis from
fossil wood rat nests in California and Arizona
(Coleoptera, Ptinidae), with a postscript on the
definition of a fossil. Coleopt. Bull. 30:239-244.
NEW AMERICAN BARK BEETLES (COLEOPTEKA: SCOLYTIDAE),
WITH TWO RECENTLY INTRODUCED SPECIES
Stoplicii I.. Wood
.\hstrac;t.- Species luiiiied as new to science include: Coiioplitlioriis iiiirlioacciiKic. ('. Icocultiiit, //i//c.si;u/.s (iztccii\
(Mexico). Fhloeocleptus punctdtits (Closta Rica), I'sctidotliijsanocs (ilonins iN'enezuela). P. lecchi (California). Pitiio'^-
cm-s mcxiaiiuis. Aivpttis spcriosiis, Amplticnmits spcrtiis (Mexico), and Xijlchonis pnwstans (Panama). Also reported
,ire the first records of the notorions Xijlosiinilnis rompactus (Eichhoff) from .South America (Brazil), and the first
American records oi Xijlebomn fomicatus Eichhoff (Panama.) and .V. laliclus Eichhoff (New York and Pennsylvania).
As indicated in the above abstract, the fol-
lowing pages report the first .\nierican re-
cords of two species of Xylclwrus, the exten-
sion of the range of Xylosandrus compactus
(Eichhoff) into South America, and the de-
scription of 10 species of American Scolv-
tidae new to science. The species new to sci-
ence represent the genera Conoplithorus (2),
Hyh sinus (1), Fhloeocleptus (1), Pseudo-
thysanoes (2), Pityogenes (1), Xylehorus (1),
Araptus (1), and Amphicranus (1), and were
taken in California (1), Mexico (6), Costa
Rica (1), Panama (1), and Venezuela (1).
New Introductions
Xt/lehorus fomicatus Eichhoff
Xylchonis fornittittis Eichhoff, 1868. ISeriiner Ent.
Zeitschr. 12:151 (.Syntypes: Ceylon; not located)
This species has caused extensive economic
damage in southern Asia, Sri Lanka, In-
donesia, Micronesia, Africa, and Hawaii for a
half century or more in a verv large luunber
of host species.
A living specimen of this species was taken
in July 1979 from a small branch of a tree in
the Canal Zone, Panama. This constitutes the
first record of this species in any American
country. Since it is moderatelv aggressive, it
is expected to be of some economic impor-
tance in the tropical and subtropical areas
into which it spreads.
Xylehorus itilidus Eichhoff
Xillclxniis vdlidiis Eichhoff. 1875. Ann. .Soc. Ent. Bel-
Liicjue 18:202 (S\ntyi)es. female; Japan; apparent-
ly at Brussels Mus.
A breeding population of this Japanese
species was first found on Long Island near
New York City (Nassau Co.) in 1976. Several
additional collections were taken at Du
Pont's Farm near Newtown Square, Dela-
ware Co., Pennsvlvania, on 2-VII-1980. from
Quercus velutina, bv Ceorge Stevens.
This species breeds in the stumps and logs
or in the boles of a broad spectrum of trees.
Host genera in japan include: Abies, Acan-
thopanax, Acer, Betida, Curpinus, Castanea,
Cletha, Cryptomeria, Fagus. Julians, Mal-
lotus, Phellodcndron, Piuus, Prunus, Quercus,
and y.elkowd. It is apparenth' more aggresive
than native ambrosia beetles and should be of
economic concern.
Xylosandrus compactus (Eichhoff)
Xi/lchonis coiiipdcliis l'"-ichhoti. 1875, .\mi. Ent. Soc. Bel-
<4i(l\ie 18:201 iS\ntvpes. female. Japan: one syn-
t\pe in Schedl C'oll. at Vienna)
This destructive .species apparently origi-
nated in southern Asia and .spread to .Africa.
Micronesia, and adjacent areas more than a
half centurv ago. More recently it reached
Hawaii, Cuba, Florida, Georgia, and Loui-
siana.
'Life .Science Maseum and Department of Ztwloi^v. Bn^hain Young University. Provo. Utah 84602. Scolytidae contrihiition No. 70.
353
354
Great Basin Naturalist
Vol. 40, No. 4
On 7 December 1979 numerous collections
of this species were made in the vicinity of
Manaus, Amazonas, Brazil, from a variety of
hosts. This is the first documented record of
this species in South America. Its occurrence
at this remote locality in virgin forest sug-
gests that it is widespread in South America
and that it has been there for quite some
time. Those concerned with plant protection
should be advised of the range expansion of
this notorious pest species.
New Taxa
Conopthonis midioacanae, n. sp.
This species is distinguished from apach-
ecae Hopkins by the more slender body, by
the less densely punctured, smoother basal
half of the elytral disc, by the broader, more
gradual elytral declivity, and by other char-
acters cited below.
Male.— Length 3.9 mm (paratypes 3.0-4.2
mm), 2.4 times as long as wide; color very
dark reddish brown.
Frons as in apachecae except never with a
median crest or tubercle, a weak, transverse
impression usually present on upper half of
median half of area below upper level of
eyes.
Pronotum essentially as in apachecae ex-
cept slightly more slender.
Elytra resembling apachecae except 1.48
times as long as wide; discal striae with punc-
tures not as close, mostly in rows, interstriae
sparsely punctured, punctures only slightly
confused on basal fifth, surface smooth, not
wrinkled; declivity not as steep, less strongly
arched, sulcus deeper and much wider, tu-
bercles on interstriae 3 very small (less
strongly arched and more broadly sulcate
than in ponderosae); vestiture less abundant,
slightly coarser.
Female.— Similar to male in all respects.
Type locality.— Uniapan, Michoacan,
Mexico.
Type material.— The male holotype, fe-
male allotype, and 24 paratypes were taken
at the type locality in Febniary 1980, from
Pinus michoacana cones by Adolfo A. del Rio
Mora.
The holotype, allotype, and paratypes are
in my collection.
Conophthorus teocotum, n. sp.
This species is distinguished from ponde-
rosae Hopkins by the subacutely elevated
median carina on the lower half of the frons
in both sexes, by the totally obsolete punc-
tures on declivital striae 2 except near base,
and by other characters cited below.
Male.— Length 3.4 mm (paratypes 3.1-3.7
mm), 2.3 times as long as wide; color very
dark brown.
Front weakly, transversely impressed as in
ponderosae; median line on more than lower
half with a conspicuous, subacutely elevated
carina, end of carina somewhat tuberculate
at epistomal margin.
Pronotum as in ponderosae except asper-
ities averaging smaller, serrations on anterior
margin usually reduced, impressed points
rather numerous and sharply, distinctly im-
pressed.
Elytra as in ponderosae except punctures
on declivital striae 2 obsolete except on less
than basal fourth, declivity more broadly,
slightly less strongly impressed, tubercles on
declivital interstriae 3 slightly larger.
Female.— Similar to male except trans-
verse frontal impression more extensive,
slightly more conspicuous, carina slightly
shorter.
Type locality.— LIrupan Michoacan,
Mexico.
Type material.— The male holotype, fe-
male allotype, and two female paratypes
were taken at the type locality in March
1980, from Pinus teocote cones, by Adolfo A.
del Rio Mora.
The holotype, allotype, and two paratypes
are in my collection.
Hylesinus azteciis, n. sp.
This species is distinguished from califor-
nicus (Swaine) by the larger size, by the pres-
ence of a fine, low, median, frontal carina in
both sexes, by the less strongly concave male
frons and less strongly convex female frons,
and by differences in the declivital interstrial
setae described below.
Male.— Length 3.8 mm (paratypes 3.8-4.2
mm), 1.8 times as long as wide; vestiture of
dark brown and tan scales in a pattern similar
to calif ornicus.
December 1980
Wood: American Bark Beetles
355
Frons similar to californicus except very
shallowly concave from epistoma to upper
level of eyes, a low, median carina on lower
half, and granules on upper and lateral areas
of head conspicuously larger.
Pronotum similar to californicus except as-
perities smaller, piuictures smaller, less defi-
nite, and scales averaging much more slen-
der.
Elytra similar to californicus except inter-
strial crenulations more numerous, smaller,
confused (a median row not predominating);
declivital interstriae 1 less strongly elevated,
2 wider; groimd setae on declivital interstriae
2 in two indefinite ranks (never uniseriate);
erect setae always absent on 2, present on 1
and 3, each four to eight times as long as
wide, spaced within a row by distances great-
er (one to four times) than length of a seta.
Female.— Similar to male except frons less
strongly, more broadly impressed (irregularly
flattened); declivital interstriae 1 less strongly
elevated (vestiture not clearly sexually dimor-
phic).
Type locality.— Chapingo, Mexico, Mex-
ico.
Type material.— The male holotype and
seven paratypes were taken at the type local-
ity on 12-XII-1979, from Fraxinus uhdei, by
T. H. Atkinson. The female allotype and five
paratypes bear similar data except they were
taken on 17-VIII-1979.
The holotype, allotype, and paratypes are
in mv collection.
Phloeocleptus punctatus, n. sp.
This species is distinguished from trcs-
mariae (Schedl) by the slightly smaller size,
by the fringe of long setae at the upper mar-
gin of the female frontal concavity, by the
coarser strial punctures, and by other charac-
ters cited below.
Male.— Length 1.6-1.7 mm (females both
1.8 mm), 2.4 (female 2.6) times as long as
wide; color very dark brown.
Frons convex, a slight transverse impres-
sion just above epistoma; surface rugose-reti-
culate on lower half, more irregularly rugose
above, pimctures moderately coarse, rather
close. Antennal scape slender, elongate, orna-
mented by less than a dozen long setae.
Pronotum about as in tresmariae except an-
terior margin finely serrate and sparse vesti-
ture on posterior half of mixture of fine, slen-
der hair and stout scales.
Elytral outline about as in tresmariae;
striae distinctly impressed on posterior third
of disc, punctures at base rather small, gradu-
ally increasing to twice as large and very
deep at base of declivity; interstriae slightlv
wider than striae at base, narrower than
striae at base of declivity, punctures unise-
riate, fine at base, becoming replaced by
rounded granules near base of declivity. De-
clivity broadly, strongly convex, steep; strial
punctures decrease in size from base, moder-
ately coarse at apex; interstriae as wide as
striae on lower half, all imiseriately granulate
to apex. Vestiture of erect, uniseriate, inter-
strial scales, each three to four times as long
as wide, almost as long as distance between
rows, spaced within a row by about two-
thirds length of a scale.
Female.— Similar to male except more
slender; frons moderately concave almost
from eye to eye from epistoma to vertex; its
surface minutely irregular, punctures fine,
obscure, its upper margin ornamented by a
dense fringe of long hair, these setae equal in
length to about one-third diameter of con-
cave area; scape with a larger tuft of long
setae; anterior margin of pronotum imarmed;
elytral punctures and granules distinctly
smaller, interstrial scales each four to five
times as long as wide.
Type locality.— Santa Rosa National
Park, Guanacaste Province, Costa Rica.
Type material.— The female holotype,
male allotype, and three paratypes were
taken at the tvpe locality between 15 De-
cember 1979 and 6 January 1980, from the
phloem of an unidentified tree, by George
Stevens.
The holotype, allotype, and paratypes are
in my collection.
Pseudothysanoes atoinus, n. sp.
This species is distinguished from coluni-
hianus (Blackman) and other representatives
of the genus by the very small size and b\ the
apparent replacement of most strial punc-
tures by granules.
356
Great Basin Naturalist
Vol. 40, No. 4
Male.— Length 0.8 mm (paratypes 0.7-0.8
mm), 2.3 times as long as wide; color yellow-
ish brown.
Frons convex; surface shining and almost
smooth in central area, becoming reticulate
toward margins, pimctures fine, sparse, some
replaced by fine granules; vestiture sparse,
inconspicuous. Antennal scape elongate, or-
namented by several hairs; club without su-
tures, small, rather slender.
Pronotum as long as wide; outline typical
of genus; anterior margin armed by four
coarse, closely set denticles; posterior areas
shining, subreticulate in some areas, sparse
punctures obscure, replaced by fine granules
behind summit. Vestiture of inconspicuous
fine hair.
Elytra 1.4 times as long as wide, 1.5 times
as long as pronotum; striae not impressed,
punctures fine, distinct, those on posterior
two-thirds with a tubercle between punc-
tures; interstriae as wide as striae, shining,
surface irregular, indistinct, fine pimctures
replaced by coarse tubercles before declivity.
Declivity steep, convex; strial punctures
scarcely evident, all striae and interstriae
with rows of rather large, rounded tubercles.
Vestiture mostly confined to declivity, of
erect rows of slender strial and stout inter-
strial setae, all of uniformly rather short
length, setae on interstriae 2 apparently ab-
sent except at base of declivity.
Female.— Similar to male except slightly
more slender; anterior margin of pronotum
unarmed; strial and interstrial tubercles much
smaller (but present); all declivital setae slen-
der.
Type locality.— Finca Monasterios, Cau-
cagua, Miranda, Venezuela.
Type material.— The male holotype, fe-
male allotype, and 11 paratypes were taken
at the type locality in 1971, from Theobroma
cacao branches.
The holotype, allotype, and paratypes are
in my collection.
Pseudothysanoes leechi, n. sp.
This species is distinguished from pJiora-
clendri Blackman of the southwestern USA by
the larger size, by the much shorter, stouter
male declivital scales, and by the much long-
er setae on the female vertex. It is much
more closely related to the Mexican verdicus
Wood but is distinguished by the stouter
scales on the male declivity, by the much less
strongly impressed female frons, with the
setae on the vertex distinctly shorter and less
abundant, and by other characters cited be-
low.
Male.— Length 1.6 mm (paratypes, males
1.4-1.7 mm, females 1.6-1.8 mm), 2.4 (fe-
male 2.5) times as long as wide; color very
dark brown, vestiture pale.
Frons as in verdictis except median third
on lower half of frons more distinctly, con-
cavely impressed.
Elytra 1.45 times as long as wide, 1.7 times
as long as pronotum; as in verdicus except
strial punctures on disc slightly larger, inter-
strial granules distinctly larger, extending to
base, interstrial scales closer, shorter, those
on declivity about twice as long as wide,
each half to two-thirds as long as distance be-
tween rows.
Female.— Similar to female verdicus ex-
cept frons shallowly concave on median two-
thirds of lower two-thirds, setae on vertex
shorter, less abundant, tips of longest attain-
ing middle of frons; posterior areas of pro-
notum without reticulation; strial punctures
slightly deeper, interstrial punctures and
scales closer, scales distinctly shorter. A very
small tuft of hair on scape.
Type locality.— North side of Howell
Mountain, 3 km NNE Angwin, Napa Co.,
California.
Type material.— The male holotype, fe-
male allotype, and 19 paratypes were taken
at the type locality (reared) on 27-VIII-1980,
from Phoradendron flavescens var. villosum
(taken from Quercus kelloggii), by H. B.
Leech. Other paratypes emerged or were cut
from the same sample on the following 1980
dates: (3) 18- VI, (2) 21-VII, (4) 25-VII, (1) 6-
VIII, (1) 13-VIII, (1) 10-VIII.
The holotype, allotype, and part of the
paratypes are in the California Academy of
Sciences; the remaining paratypes are in my
collection.
Pityogenes mexicanus, n. sp.
This species is distinguished from merid-
ianus Blackman by the much larger size, by
the shorter, more strongly hooked upper
December 1980
Wood: American Bark Beetles
357
spines on the male elytral declivity, bv the
more regularly punctured discal interstriae,
and by the much more strongly convex fe-
male elytral declivity. The female frons of
this species and nwridiamis differs from all
other American Pityogencs in lacking a deep-
ly excavated central area.
Male.— Length 3.2 mm (paratypes 3.2-3.4
mm), 2.6 times as long as wide; color verv
dark brown.
Frons broadly granulate, a few fine punc-
tures interspersed; vestiture of fine, long,
moderately abimdant hair.
Pronotum essetially as in mcridianus ex-
cept minute, impressed points nuich more
numerous.
Elytra essentially as in meridianus except
interstrial punctures regular, about equal in
size to those of striae; upper declivital spines
slightly shorter, more strongly hooked, series
of tubercles on lower fourth of lateral margin
much lower and romided except lowest one
larger and pointed (male meridianus not at
hand; comparison based on Blackman's draw-
ing).
Female.— Similar to male except median
line on upper half of frons shallowly concave;
epistomal area on median third slightly pro-
tuberant, granulate, and ornamented by mod-
erately abundant, fine, short hair; frontal tu-
bercles smaller; declivity shallowly, narrowly
sulcate (more shallowly impressed than any
other American Pihjogenes), declivity with
stouter, more abundant vestiture than in
meridianus.
Type locality.— Parque Nacional Zoquia-
pan, Mexico, Mexico.
Type material.— The female holotype,
male allotype, and six paratypes were taken
at the type locality in August 1979, from a
Pinus luirtwegii branch (shaded out?), by T.
H. Atkinson.
The holotype, allotype, and paratypes are
in mv collection.
Araptus speciosus, n. sp.
A specimen of this species in the U.S. Na-
tional Museum, which had been incorrectly
identified as Neodryocoetes exquisitus Black-
man, led me to apply Blackman's name in-
correctly. His exquisitus ( = F. inceptis Wood)
must be referred to PityophtJiorus and the
very similar, misidentified specimens, here
named speciosus, to Araptus. In all probabili-
ty, both species should be in Araptus, al-
though the paucity of material for studv
makes resolution of the problem difficult.
This species is distinguished from exquis-
itus by the more broadly flattened female
frons, with longer, more broadly distributed
frontal vestiture; by the less distinctlv reticu-
late, more finely punctured pronotum; and
by the slightly shorter elytral vestiture.
Female.— Length 1.7 mm (paratvpes
1.4-1.7 mm), 2.7 times as long as wide; color
dark reddish brown.
Frons broadly flattened from epistoma to
vertex, shining, finely, closely punctured,
sparsely pubescent at center, densely orna-
mented by long yellow hair at sides and
above, longest setae on vertex extend two-
thirds distance to epistoma.
Pronotum 1.1 times as long as wide; widest
slightly behind middle; sides on posterior half
weakly arcuate, feebly constricted on ante-
rior half, then rather narrowly rovmded in
front; anterior margin armed bv four to ten
serrations; summit poorly developed, slightly
in front of middle; asperities on anterior
slope moderately large, arranged into dis-
continuous, irregular, subconcentric rows,
posterior areas mostly smooth, shining, some
specimens with very obscure indications of
reticulation, punctures moderately coarse,
deep, rather close. Glabrous.
Elytra 1.7 times as long as wide, 1.5 times
as long as pronotum; sides straight and paral-
lel on basal two thirds, rather broadl)
rounded behind; striae not impressed, pimc-
tures rather small, moderately deep; inter-
striae about twice as wide as striae, almost
smooth, shining, impunctate, impressed
points not clearly visible. Declivity steep,
rather broadly convex; sutural interstriae
feebly elevated; strial punctures smaller than
on disc, a few verv small interstrial punctures
also present. Vestiture confined to declivity,
consisting of a few interstrial bristles, each al-
most as long as distance between rows.
Type locality.— Five miles or 8 km south
of La Huerta, Jalisco, Mexico.
Type material.— The female holotype
and eight female paratypes were taken at the
type locality on l-VII-1965, No. 168, from a
Ficus twig, by me.
358
Great Basin Naturalist
Vol. 40, No. 4
The holotype and paratypes are in my col-
lection.
Amphicranus spectiis, n. sp.
This species is distinguished from spec-
tabilis Wood by the larger size, by the more
elongate antennal club, with more strongly
arcuate sutures, by the more shallowly im-
pressed elytral punctures, and by the more
strongly, more acutely elevated lateral mar-
gin of the elytral declivity from base to apex.
Male.— Length 2.9 mm, 3.2 times as long
as wide; color reddish brown (fully mature?).
Frons about as in spectahilis except surface
more finely pimctvired, raised median gran-
ular area slightly larger (occupying almost
meridian third), much more sharply defined.
Antennal club more slender, 1.5 times as long
as wide, sutures more strongly arcuate than
in spectahilis.
Pronotiun 1.6 times as long as wide; about
as in spectahilis except punctures on posteri-
or areas slightly smaller.
Elytra 1.8 times as long as wide, 1.13 times
as long as pronotum; similar to spectahilis ex-
cept punctures on disc very shallow,
obscurely impressed, declivity more deeply
excavated, lateral margin more acutely, more
strongly elevated, more strongly explanate
below, basal area of spine 1 protruding
slightly.
Type locality.— Pichucalco, Chiapas,
Mexico.
Type material.— The male holotype was
taken at the type locality on 26-III-1980,
from Theobroma cacao.
The holotype is in my collection.
Xyleborus praestans, n. sp.
This species is distinguished from meritus
Wood by the larger size, by the different de-
clivity, and by other characters cited below.
Female.— Length 3.9 mm, 2.9 times as
long as wide; color dark brown.
Frons and pronotum as in meritus except
pronotal summit more subacutely elevated.
Elytra about as in meritus except declivity
slightly steeper, more nearly convex, more
broadly rounded behind; interstrial punctures
on disc more nearly obsolete, irregular inter-
strial lines on disc more conspicuous; strial
punctures on declivity more distinct, not
larger than those on disc, interstrial tubercles
on 1 and 2 not as close, very slightly larger.
Type locality.— Cerro Punta, Chiriqui,
Panama.
Type material.— The female holotype
was taken in the vicinity of the type locality
on 31-V-1972, 6000-8000 ft, by f . L. and L.
J. Erwin.
The holotype is in the Canadian National
Collection.
HELD OBSERVATIONS ON THE RESPONSE OF THE RAH.HOAD VALLEY
SPRINGFISH {CRENICHTHYS XEVADAE) TO TEMPERATURE
Thomas M. Baus^li' and Briicf G. Browir
Abstract.— The presence of Crcniclithi/s ncvadac Hiihbs is verified from 37.8 to 18.3 C in the Bi^ Sprint^s aquatic
system.
The Railroad Valley springfish {Crenichthijs of Railroad Valley, Nye County, Nevada.
nevadae Hubbs) is a small (ca 5 cm) cvprino- One location in this valley is Big Springs on
dont found naturally only within the confines Lockes Ranch closely adjacent to Highway 6.
N
Flu. 1. Map of Lockes Ranch spring-stream-pond complex. .\ to B. spring and segment of stream to first down-
stream culxert; B to C, stream on west side of highway between two culverts; C to F, stream between second down-
stream culvert and pond.
'1020 Custer Avenue, Ogden. Ltah H4-1(I4.
'142 Eccles. Ogden, Utah 84401.
359
360
Great Basin Naturalist
Vol. 40, No. 4
On 21 March 1980, the authors, while in-
volved in other work, had the opportunity to
make a cursory survey of water temperatures
at several locations from the spring to the
terminus of the stream in a pond on Lockes
Ranch and to relate these temperatures to
the presence of C. nevadae. (The length of
the total aquatic system from spring to pond
is estimated to be about 500 yards.)
Temperatures were taken and observations
were made from 1459 to 1606 hours Pacific
Standard Time. The air temperature was
about 7.3 C. There were scattered clouds and
wind gusts accompanied by snow flurries.
All temperatures were taken with a West-
on thermometer (Model 2265) and the pres-
ence of C. nevadae in waters of various tem-
peratures was verified by both authors.
The spring-stream-pond complex (Fig. 1)
can be divided into three unequal segments:
Segment 1 (A-B) encompasses the spring and
that segment of the stream to the first down-
stream culvert. Segment 2 (B-C) is composed
of that segment of the stream on the west
side of the highway between the two cul-
verts. Segment 3 (C-F) is composed of the
segment of stream between the second down-
stream culvert and the pond.
The temperature gradient from spring to
pond was 37.8 to 17.8 C. The temperatures
at various points along the system, under the
climatic conditions described above, were as
follows:
Point Temperature (C)
Fish
A
37.8
Yes
B
32.2
Yes
C
26.7
Yes
D
20.0
Yes
E
17.8
No
F
17.8
No
The last fish noted by the authors were be-
tween points D and E at a temperature of
18.3 C. No fish were found below that tem-
perature nor were any seen in the shallow
water along the perimeter of the pond,
where temperatures were a uniform 17.8 C.
The presence of fish was verified visually or,
where vegetation made viewing impossible,
by capture in a fine mesh net. Those fish
noted at 18.3 C were taken in a net.
This cursory survey establishes a tempera-
ture profile for the Big Springs system of 37.8
to 17.8 C and a temperature tolerance range
for C. nevadae of 37.8 to 18.3 C.
WOODRAT NEST FLEA AXOMIOrSYLIA'S AMPIIIBOLUS
IN SOUTHEASTERN ORE(X)N
llaroki |, K<j;()sciie'
Ahsthact.- The lira .\/i()»nii/)si///i/v (iiiipliilxihis is reported from soutlieastern Oieuon, a ian<4e extension of aliont
173 km from tlie nearest reported totalities in northw esteiii I'tali.
To m\ kno\vlcd<j;e fleas in the yenus Ano-
miopsyllus have not been previonslv reeord-
ed in Oregon (Hubbard 1947, Barnes, Tipton
and Wildie 1977), although one or more
kinds of woodrats {Neotoma) that normally
host these interesting nest fleas are found
throughout the state. This note reports the
presence oi Anomiopsi/Uus amphihohis Wag-
ner, 1936, in southeastern Oregon, a range
extension of about 475 km from the nearest
localities mapped by Barnes et al. (1977) in
northwestern Utah.
My first specimen, a female (H.J.E.
No.6641), was collected 25 November 1968
from a bushy-tailed woodrat, Neotoma cine-
tea alticola, captured 11.2 km south of
Crane, Harney County, elevation 1290 m. A
male A. amphibolus (H.J.E. No. 8026) was
collected 21 April 1980 from a deer-mouse,
Pewnnjscus manicukiius ssp. caught less than
4 m from the woodrat den where the 1968
host was trapped.
This locality is near the northeastern limits
of the Great Basin in Oregon, in arid waste-
land characterized bv low annual precipi-
tation and vegetation dominated by desert
shrubs. Traps were set along a steep, dry
talus- and boulder-strewn hillside that includ-
ed several prominent lava outcrops. Spiny
hopsage, Grai/ia spinosa, is the dominant
plant. The area is treeless, the nearest scat-
tered stands of juniper being some miles dis-
tant. Other small mammals trapped here in-
cluded desert woodrats, Xeotoma lepida
nevadensis, canyon mice, Peromyscus crinitus
crinitu.s, and Great Basin pocket mice, Per-
ognathiis parvus parvus. Other fleas in the
subfamily Anomiopsvllinae found here were
Stenistomera huhhardi, S. (dpina. and Callis-
topsyllus terinus terinus.
The sympatric occurrence at this place of
the bushy-tailed woodrat and desert woodrat
was unexpected. Zonally, the habitat seemed
ideal for the latter species but too low and
arid for .V. cinerea, although Finley (1958)
found that lack of suitable den sites more
than type of vegetation limited the distribu-
tion of bushy-tailed woodrats in Colorado.
Field work in 1980 was done under Scien-
tific Taking Permit No. 06.3 courtesv of the
Oregon Department of Fish and W ilillife.
LiTKHATL RK (hTKl)
Bahms, a. M.. \'. J. Tipton, and J. A. Wiinn:. 1977.
The suhtamilv Anomiopsvllinae (H\s-
tiieliopsyllidae: Siphonaptcra). I. A revision of
the s^enns Anoniiopstilliis Raker. Creat Basin Nat.
.37: 138-206.
FiM.i:-^. K. B.. Jr. 19.5S. The woodrats of Colorado: dis-
tribution and ecologv. Univ. Kansas Publ.. Mns.
Nat. Hist. l()(6):213-.552.
Ill lUiAHi). C .\. 1947. Siphonaptera of western North
.\nieriea. Iowa State Colle'^e Press. .533 pp.
'297 West Durfee Street, Granlsvi
361
POSTEMERGENCE DEVELOPMENT AND INTERYEAR RESIDENCE OF
JUVENILE COLUMBIAN GROUND SQUIRRELS IN THE IDAHO PRIMITIVE AREA
(lliarlcs L. Elliott' and |erran T. FIjikIlms'
Abstract.— A colonv of Clohinihiaii giouiid squirrels in the Idaho Primitive Area was observed from 1976 to 1978.
Seven hodv measurements were recorded tor juveniles obtained in 1978. There was a lack of sexual dimorphism
among developing juveniles. The hind foot was the fastest developing feature. Juveniles obtain adult size their sec-
ond vear. Juvenile males exhibited the lowest interyear residency of either sex or age group examined.
Pengelley (1966), in a comparison of devel-
opmental patterns of four species of ground
squrrels (Genus: SpennopJiihis), noted that
developmental rates appeared to have an
adaptive value for the particular habitats oc-
cupied by each species. Thus an under-
.standing of the developmental pattern of a
particular species may provide insight into
the basic ecology of that .species. Information
on various aspects of growth and devel-
opment for several species of Spewiophilus
has been reported (Svihla 1939, Blair 1942,
Maver and Roche 1954, Tomich 1962,
Mckeever 1964, Neal 1965, Clark 1970, Iver-
son and Turner 1972, Zimmerman 1972,
Michener 1974, Turner et al. 1976), but little
data have been compiled for the Columbian
groimd squirrel (Spermopliiliis columhianus).
Levenson (1979) presented a growth rate
constant from birth to 50 days for S. colum-
hianus and Shaw (1925) traced the devel-
opment of Columbian ground squirrels from
birth to emergence from the den, but infor-
mation concerning postemergence devel-
opment is lacking.
Methods
A description of the Cold Meadows study
area and trapping procedure used have been
described elsewhere (Elliott and Flinders
1980). In addition to ground squirrels live
trapped, specimens were collected from out-
side the trap grid using a small caliber rifle.
Postemergence development reported here is
based on data acquired during the 1978 field
season.
Results and Discussion
Seven body measurements were taken on
each sex of juvenile (juvenile = young of the
year) Columbian ground squirrel live trapped
or shot (Table 1). No significant difference
(unpaired t-test) was found between measure-
ments of males and females (excluding body
weights for July or August). This lack of sex-
ual dimorphism in developing ground squir-
rels has been observed for other spermophiles
(Kiell and Millar 1978).
Table 1.
.\U
•an
measurements
1 ±Sl^) of juvenile Co'
lumbiau gi
ound s([uirrels eollectec;
1 at Cold M
eadous. Idaiio
Primitixe Ai
ea.
19'
78.
Foot
Ear
Tail
Total
Bodv
Zvgomatic
Condvlobasal
Date
Sex
N
length
length
length
length
weight
breadth
length
Julv 17-24
M
4
47 ± 1 mm
17±1
82 ±.5
2.58 ±14
200±.34g
27±2
49 ±3
F
8
46 ±2
17±]
82 ±7
2.58 ±15
173 ±34
27±1
51 ±3
c:
omliiued
12
46 ± 1
17±1
82 ± 6
2,58 ±14
182 ±.35
27±1
50 ±3
August 14-2
,1
M
9
.50 ± 1
18± 1
82 ±4
280 ± 7
311 ±43
.30 ±2
.52 ±3
1-^
9
49 ± 1
17± 1
81 ±.3
276±11
264 ±31
29 ± 3
49 ±1
c
ombined
18
49 ±1
18±1
82 ±.3
278 ±9
288 ± 44
29 ±2
50±2
'Dc'partiiR'nl of Botany and Rani^e Science, Brighain Yonng University, Provo, lUali 84fi()2
362
December 1980 Elliott, Flinders: Columbian Ground Squirrels
363
T.\BLE 2. Size of juvenile Columbian ground scjuirrels expressed as a percent of the size of adults taken during the
same collection period in Cold Meadows, Idaho Primitive Area, 1978.
Foot
Ear
Tail
Total
Body
Zygomatic
Condvlobasal
Date
Sex
length
length
length
length
weight
breadth
length
Julv 17-24
M
92
85
87
78
33
80
84
F
93
86
92
81
34
83
9()
Combined
92
85
89
79
32
82
88
August 14-21
M
98
89
87
84
53
88
89
F
99
89
91
87
51
89
87
Combined
98
89
89
86
52
89
88
The body dimensions were compared
(Table 2) to corresponding measurements of
76 adult squirrels taken in the same collec-
tion periods. The hind foot was the fastest de-
veloping item measured, a feature also noted
in S. riduirdsonii (Clark 1970), S. lateralis
(Clark and Skrvja 1969), S. pamjii (Kiell and
Millar 1978), s' tereticaiidus (Neal 1965), and
S. Jiamsii (Neal 1965). All mea.surements ex-
cept tail length and condylobasal length were
significantly larger (unpaired t-test, P<0.01)
during August than July. Shaw (1925) noted
Columbian ground squirrels did not complete
their growth cycle until the second season.
Based on 12 juveniles captured in 1977 and
recaptured in 1978, ground squirrels at Cold
Meadows also obtain adult size their second
year of life. The combined proportions for
both sexes in August (Table 2) indicate ap-
proximately 90 percent of the adult dimen-
sions (excluding body weight) are obtained by
the end of the first season. This delaying of
maturity has been observed in other species
of Sperrnophilus (Bridgwater 1966, Morton
and Tung 1970). Morton et al. (1974) noted
that in S. heldingi fattening and overall
growth were concurrent at first but that ca-
loric intake was then diverted primarily to-
ward lipid synthesis and storage for catabo-
lism during hibernation. This caloric
diversion resulted in a late season slowing of
increase in linear dimensions. The Columbian
ground squirrels at Cold Meadows are active
four months out of the year, hibernating for
the remaining period. The necessity to "trade
off" calories for body growth to develop
greater body reserves for hibernation may ac-
count for the inability of juveniles to attain
adult size their first season.
Utilizing capture-recapture data acquired
during 1976-1978, the percent interyear resi-
dence for each sex and age group of S.
Table 3. Percent interyear residence of Columbian
ground squirrels at Cold .Meadows, Idaho Primitive
Area, 1976-1978.
1977
1978
recaptures/
recaptures/
1976
1977
captures
Percent
captures
Percent
Adult
males
10/16
62.5
5/13
38.4
Adult
females
16/22
72.7
7/20
35.0
Juvenile
males
3/14
21.4
2/12
16.6
Juvenile
females
4/9
44.4
3/9
33.3
columbianus was calculated (Table 3). Juve-
nile males exhibited the lowest rate of fide-
litv. Michener and Michener (1971) observed
the .same residency pattern for S. richard-
sonii. Reasons for the absence of juvenile
male ground scjuirrels has been postulated to
be the result of dispersal (Evans and Hold-
enried 1943, Fitch 1948, McCarley 1966.
Quanstrom 1971, Ycaton 1972), exceptional
vulnerabilitv to predation (Schmutz 1979),
and/or overwinter mortality (Michener and
Michener 1977). We did not determine
which specific factor(s) resulted in the ob-
served low juvenile male interyear residency
for the Cold Meadows colony.
We thank the I'niversity of Idaho for per-
mission to use the facilities at the Ta\lor
Ranch Field Station, Idaho Primitive Area.
Blai
LiTER-\TURE Cited
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364
Great Basin Naturalist
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Clark, T. M., and D. D. Skkyja. 1969. Postnatal devel-
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Elliott, C. L., and J. T. Fllnders. 1980. Seasonal activ-
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Idaho Primitive Area. Great Basin Nat.
40:175-177.
Evans, R. C, and R. Holdenried. 1943. .\ population
study of the Beechey ground squirrel in central
California. J. Mammal. 24:231-260.
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KiELL, D. J., AND J. S. Millar. 1978. Growth of juvenile
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56:1475-1478.
Levenson, H. 1979. Sciurid growth rates: some correc-
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McCarley, H. 1966. Annual cycle, population dynamics
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McKeever, S. 1964. The biology of the golden-mantled
ground squirrel. Ecol. Monogr. 34:383-401.
Michener, D. R., and C. R. Michener. 1971. Sex ratio
and interyear residence in a population of
Spermophilus richardsonii. J. Mammal. 52:8.53.
1974. .\nnual cycle of activity and weight change
in Richardson's ground squirrel Spermophilus
richardsonii. Can. Field-Nat. 88:409-413.
Michener, G. R., and D. R. Michener. 1977. Popu-
lation structure and dispersal in Richardson's
groimd squirrels. Ecology 58:3.59-368.
Morton, M. L., and H-C. L. Tung. 1971. Growth and
development in the Belding ground squirrel
{Spermophiltis hchlintj^i heldiniii). J. .Mammal.
.52:611-616.
Morton, M. L., C. S. M.vxwell, and C. E. Wade. 1974.
Body size, body composition and behavior of
juvenile Belding ground s(|uirrels. Great Basin
Nat. .34:121-1.34.
Neal, B. J. 1965. Growth and development of the
round-tailed and Harris antelope ground squir-
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Pe.n'Gelley, E. T. 1966. Differential developmental pat-
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ground squirrel Spermophilus richardsonii rich-
ard.wnii. .\nim. Behav. 19:646-6.52.
SCHMUTZ, S. M., D. A. BOAG, AND J. K. ScHMlTZ. 1979.
Causes of unequal sex ratio in populations of
adult Richardons ground stjuirrels. Can. J. Zool.
.57:1849-18.55.
Shaw, W. T. 1925. Breeding and development of the
Columbian ground squirrel. J. Mammal.
6:106-113.
Svihla, a. 1939. Breeding habits of Townsends ground
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Tomich, p. Q. 1962. The annual cycle of the California
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Pub. Zool. 65:21.3-282.
Turner, B. N., S. L. Iverson, and K. L. Severson. 1976.
Postnatal growth and development of captive
Franklin's ground squirrels {Spermophilus frank-
linii). Amer. Midi. Nat. 95:9.3-102.
Yeaton, R. I. 1972. Social behavior and social organiza-
tion in Richardson's ground squirrel (Spermo-
philus richardsonii) in Saskatchewan. J. Mammal.
53:1.39-147.
Zimmmerman, E. G. 1972. Growth and age determina-
tion in the thirteen-lined ground squirrel,
Spermophilus trideeemlineatus. .\mer. Midi. Nat.
87:314-325.
FLOOD FREQUENCY AND THE ASSEMBLAGE OF DISPERSAL TYPES LN
HANGLNG GARDENS OF THE NARROWS, ZION NATIONAL PARK, UTAH
George P. Malanson' and Jeaiiiie Kay-
.\bstr.\ct.— Hanging gardens of the Narrows. Zion National Park, Utah, are plant commnnities growing at per-
manent seeps on the canyon walls. The gardens are isolated from each other and from other plant connnunities by
\ertical expanses of sandstone. Gardens consist mostly of herbaceous species less than 1 m tall. Though not individ-
ually species-rich, the hanging gardens are diverse as a group, and verv di.ssimilar.
This study considers two explanations of the heterogeneous distribution of species in hanging gardens. The assem-
i)lages of dispersal types in gardens of different spatial attributes and disturbance frequencies are examined. The C.^^
statistic, a log likelihood ratio test, analyzes the incidence of dispersal types among classes of three spatial and two
disturbance variables.
Tlie disturbance variables of expected flood frequency and soil depth segregate dispersal t\pes; and the spatial
\ariables of area, distance to possible seed sources, and relative isolation do not. Ferns and mosses, dispersing
through spores, dominate a heterogeneous higitive guild in the flood-prone gardens. Infre(|ucntlv flooded gardens
support more large-seeded species.
Zion National Park of .southwestern Utah is
renowned for the sheer canyons of the North
Fork of the Virgin River, which dissect over
600 m of Navaho sandstone. Compared to
the arid and semiarid environments of the
Colorado Plateau, the climate of the narrow
canyons is cool and moist, and direct sunlight
may penetrate for only a few hours per day.
Large expanses of the canyon walls support
no vegetation. Vascular plants, including
shrubs and trees such as Finns ponderosa, in-
habit the occasional crevices. Perhaps the
most beautiful plant commmiities of the can-
von faces are the haniiino; gardens, which in-
elude ferns and wildflowers among their spe-
cies. Hanging gardens are defined here as
insular plant communities growing at per-
manent seeps on canyon walls (Fig. 1).
Seeps occur where precipitation has per-
colated downward through a porous forma-
tion imtil meeting less permeable strata, and
it then flows laterally until a canyon inter-
sects this plane. The volume of water dis-
charged from permanent seeps in Zion Can-
yon varies from barely perceptible trickles to
gushing springs. The seeps, and consequently
the hanging gardens, assume a variety of
shapes. Seeps usually create a less steep, rela-
tivelv narrow ledge on the canyon wall. At
other .sites, vertical jointing concentrates the
seepage, or travertine deposits from calcium
carbonate in the water create bulging forms.
Relict potholes containing permanent .seep-
age water supplies also support hanging gar-
dens.
Hanging gardens illustrate several bio-
geographic problems. The effects of the un-
u.sual environments of hanging gardens on
species composition have not been adequate-
ly explained. Also, the i.solation of gardens by
steep rock surfaces provides another testing
groimd for concepts of island biogeography.
Most studies of island biogeography treat
broad areas where pathways of dispersal be-
tween sites are not restricted (Simberloff
1974). The linear course of Zion Canyon,
however, is an additional constraint on the
movement of propagules between hanging
gardens.
Earlier studies of hanging gardens (Welsh
and Toft 1976, Nebeker et al. 1977), island
biogeography (MacArthur and Wilson 1967,
Diamond 1975), and plant distributions (Piatt
1975, Levin 1976a, 1976b) suggest several ex-
planations of the assemblages of hanging gar-
den species. Species distributions are in-
fluenced by (1) habitat requirements and
tolerances, (2) species' abilities to disperse to
Department of Geography. University of California, l.os Angeles, California 90024.
Department of Geography. University of Utah. Salt Lake City, Utah 841 12.
365
366
Great Basin Naturalist
Vol. 40, No. 4
Fig. 1. A hanging garden in the Narrows, Zion National Park, Utah.
sites of different sizes and degrees of isola-
tion, and (3) time elapsed since sites were
opened by disturbance.
Few authors have investigated the vegeta-
tion of hanging gardens, and their con-
clusions emphasize the importance of habitat
and isolation as controls of plant distribution.
Woodbury (1933) outlined primary succes-
sion at seeps from algae to mosses and vascu-
lar plants. Welsh and Toft (1976) disclosed
the geographical affinities of species they
found at seeps of different morphology, call-
ing hanging gardens "relictual refugia" of
species not native to the region. Welsh and
Wood (1976) studied structure, and Wood
and Welsh (1976) measured productivity of
hanging gardens, finding stability of both.
Nebeker et al. (1977) examined floristic sim-
ilarity, flower size, and dispersal type in
hanging gardens, and concluded that the as-
semblages were "random assortments of indi-
viduals from the species pool capable of ex-
ploiting the environments of individual sites."
Malanson (1980) recently explored the
relationships of .species and habitat in hang-
ing gardens of Zion Canyon. Although the
gardens were floristically dissimilar, species
assemblages did not clearly vary according to
perceptible differences in physical environ-
ment. Tests of species presence across a
range of habitat and spatial variables pro-
duced few significant relationships. A few
gardens had relatively high levels of solar
radiation that might exclude several species.
He also found a species-area relationship
characteristic of small islands (Whitehead
and Jones 1969).
Malanson (1980) concluded that other fac-
tors must influence assemblages of hanging
gardens. This study examines the ideas that
spatial characteristics and disturbance his-
tories of the gardens affect the incidence of
dispersal types, and thereby the plant assem-
blages.
Methods
We sampled 29 of the 60 hanging gardens
observed in an 8 km section of the Narrows
and in 0.75 km of a tributary (Orderville
December 1980
Malanson, Kay: Hanging Gardens
367
'^ >^>^. V
Fit^. 2. Tlie location ot sampled han^iiit; t^arcloiis and the t()pf)<4rai)li\ of the Narrows.
Canyon) between June and September 1977 of the i;arden.s, spaced at 2 m intervals and
(Fig. 2). Sites were selected for approach- perpendicular to the long axis. Species were
ability, though five were reached only identified in the field or at the Garrett Her-
through technical climbing, and for variety barium. University of Utah, and were sub-
of garden sizes. Species presence was record- sequently classified according to dispersal
ed along line transects spanning the lireadth tvpe, according to Dansereau and Lems
368
Great Basin Naturalist
Vol. 40, No. 4
(1957). Seed descriptions in floras of the
western United States (Arnow and Wyckoff
1977, Davis 1952, Flowers 1973, Hitchcock
and Cronquist 1973, Munz and Keck 1970)
facilitated this classification. Malanson (1980)
provides additional information on sample
design.
We measured three spatial variables to test
the applicability of island biogeographic hy-
potheses to the distribution of hanging gar-
den species. These variables are (1) area of
the gardens, (2) relative isolation from other
gardens, and (3) distance of gardens from the
terminus of the Gateway to the Narrows
Trail. Area was derived from vegetation sur-
vey transects. Isolation is defined as the sum
of the distances from each garden to its three
nearest neighbors, as determined from a top-
ographic map (ZNHA 1977). This arbitrary
measure was suggested by the stepping-stone
effect whereby species colonize one "island"
from another (MacArthur and Wilson 1967).
The terminus of the Gateway to the Narrows
Trail coincides with the entrance to the Nar-
rows (Fig. 2). The trail receives much pedes-
trian tourist traffic, but use declines markedly
beyond it. The distance provides a crude in-
dex of gardens' accessibility to animal-
dispersed or riparian species originating out-
side the Narrows environment.
Because flash floods are a common envi-
ronmental disturbance of Zion Canyon, we
evaluated the susceptibility of gardens to in-
undation. The discharge of the North fork of
the Virgin River is recorded at a USGS gauge
10 km downstream from the Narrows. Mad-
dox. Hart, and Hawkins (1977) calculated the
expected return periods for instantaneous
peak flows from magnitude and frequency
data recorded at this gauge. After measuring
the various canyon widths and elevations
where hanging gardens are located, we esti-
mated the probable frequencies of flooding
for each garden. Two classes of flood fre-
quency were used: less than 7.5 yr and great-
er than 15 yr. These classes should be distinct
and allow some margin of error without
overlap. We also measured soil depth, assum-
ing that high-velocity floods would scour soil
from affected gardens.
The Gh statistic for heterogeneity (Sokal
and Rohlf 1969) was used to disclose signifi-
cant differences in the incidence of the four
Table 1. Incidence of tested dispersal types in classes
of the variables and relationship of soil depth and flood
frequency in hanging gardens.
Class
Spore
D
Wind
ispersal Ty
Plume
pe
Fleshy
Area
0-10
21
10
4
3
(m2)
10-25
19
11
5
2
25-50
14
8
4
0
<50
19
28
12
12
Isolation
0-80
10
13
7
5
(ni)
81-160
26
24
12
19
161-240
14
5
3
1
241-320
11
9
1
0
<320
12
/
2
2
Distance
500-1630
11
25
9
5
(m)
1631-2760
8
6
1
0
2761-3890
23
11
5
0
3891-5020
21
12
9
8
5021-6150
10
4
1
4
Soil deptl
1 0-1
19
12
1
1
(cm)
1-2
29
18
10
10
2-4
8
1
0
2
4-8
1
1
0
3
<8
12
25
14
11
Flood
<7.5
51
23
12
14
(yr)
>15
20
25
11
12
Flood (yr
)
Soil depth (cm)
0-1 1-2 2-4
4-8
<8
<7.5
7
9
2
0
1
>15
1
1
0
1
4
most common dispersal types between classes
of the spatial variables and soil depth (Table
1). Onlv spores, wind-blown, plumed, and
fleshy types were abundant enough to pro-
vide meaningful tests. We used a probability
of p = .05 for significance. Because we made
multiple comparisons, the .05 chance of Type
I error applies to the individual tests where
df = 3, but the probability of "experiment
wide error" is much higher. To limit experi
ment wide error to .05 we judged individual
tests at p = .001 (Gabriel 1966).'^Single tests of
dispersal type and of soil depth between the
two flood frequency categories were made at
p = .05.
Results
Forty-eight species were counted in the 29
hanging gardens (Table 2). The frequency of
species occurrence ranges from 1 to 17. Only
13 species were found in more than four gar-
dens. The number of species per garden
ranges from 2 to 20. The average richness is
December 1980
Mal.\nson, Kay: Hanging Gardens
369
Table 2. Hanging garden plant species.
Species
Frequency
Diaspora type
Abies concolor
1
heavy
Acer ncfiimdo
3
winged
Adiantum capiUus-venehs
15
spore
Acluintuin pcdattini
6
spore
Aiiuirarilhiis (dhiis
1
heavy
Anopludis ituirficiritaccd
4
plumed
ApocynuDi ciinnubinuin
1
plumed
Acjuileoiu spp.
13
wind-blown
Aralia raccmosa
12
tlesh)
Artemisia litdovicidrui
1
u iiul-hlown
Aster eatonii
8
jiluined
Berheris repcns
3
flesln
Brickelia grandifhna
Bromus ciliatus
1
3
spins
plumed
Calainagrostis seopii lorn in
5
plumed
Cirsiutn ariz^niicinit
1
spin)
Ctjstopteris frd'^ili.s
Dodecdtheon pulcltcllum
17
9
spore
wind-blown
Drijopteris filix-inas
Ek'ocharis sp.
2
3
spore
wind-blown
Epipactis ^i^igantea
Equisetum hijemcde
3
1
heavy
heavy
Fraxinus velutinu
3
winged
Galium aparine
4
glandular
Hepaticae
Hcuchera versicolor
10
2
spore
wind-blow n
Jumiis sp.
2
windblown
Lobelia cardinalis
3
wind-blovMi
Mimulus cardinalis
13
wind-blown
Mimulus ^uttatus
1
heavy
Muhlenben^ia andina
1
wind-blown
Mtddeid>erf:^iu mexicana
2
wind-biouii
Sasttirtiitm officinale
2
witid-blowii
Poa nevadensis
2
wind-blown
Rhus radicans
2
fleshy
Ritbtis leucodermis
3
fleshy
Rumex sp.
Salix sp.
1
1
wind-blown
wind-blown
Smilacina steUata
~
flesh \
Sphagnum sp.
Sphagnaceae
Taraxacum officimdis
14
9
4
spore
spore
plumed
Thalictrum fendleri
2
wind-blown
Viola spp.
3
expulsi\e
unidentified # 1
1
unidentified #2
1
unidentified #3
2
unidentified #4
1
7.3 species, but in the seven gardens found to
be infrequently flooded the average is 12.
These seven gardens contain 80 percent of
the rare species. All but one species identi-
fied are perennial.
The sizes of the hanging gardens vary
greatly, from 2 to 100 m among samples.
Most values of isolation are low. Twenty-four
gardens are less than 300 m from the nearest
three neighbors. All but four distances from
sampled gardens to the Gateway to the Nar-
rows trail are clustered between 500-2000 m
and 300()-5()00 m.
Nineteen sampled hanging gardens are
within the range of flash floods with an ex-
pected recurrence interval of 7.5 yr. Only
seven gardens are high enough on the canyon
walls to escape flood crests with a 15 yr ex-
pected recurrence interval, and gardens 28
and 29 probably never have been inundated.
Three gardens could not be put unequivo-
callv in either cla.ss.
Individual tests at p = .05, df = 3 indicate a
higher incidence of spore dispersal types in
the smaller and more isolated gardens and a
larger proportion of the heavier, plumed, and
fleshy types in the larger and less isolated
gardens. However, when applving the
p = .001 level to limit the probability of Type
I error within the groups, the statistic re-
vealed no significant differences in the 26
spatial tests (Table 3).
Among the 10 soil depth tests, the Gh sta-
tistic indicated a significant difference in dis-
persal type between the shallowest (0-1 cm)
and the deepest (8 cm) classes. The incidence
of dispersal types and soil depths significantly
differed between the two classes of flood sus-
ceptibility. The frequently flooded gardens
usually have thin soils and a high incidence
of spore dispersal t\ pes, and the heavier,
plumed, and fleshy dispersal types and deep-
er soils are more common in the infrequently
flooded eardens.
Discussio.N AM) Conclusions
The length of time seeps are available for
colonization between disturbances seems an
important control of plant assemblages. The
mosses and ferns disperse by microscopic
wind-blown spores and can establish them-
selves rapidly in recently flooded gardens.
However, at least seven species disperse
through spores, and a few individuals of other
dispersal tvpes do establish themselves in fre
quentlv flooded gardens, so garden commu-
nities are not necessarily similar. The postu-
lated susceptibility of 65 percent of this
sample to frequent floods may explain wh\
spore-dispersed plants were the mo.st com-
mon tvpes. Nebeker et al. (1977) found bird-
dispersed types to be mo.st common in hang-
370
Great Basin Naturalist
Vol. 40, No. 4
Table 3. Significant differences disclosed by Gh statistic (p = .05 or less) for incidence of dispersal types in cate-
eories of spatial variables, soil depth, and flood susceptibility and incidence of soil depth classes in categories of flood
susceptibility.
Probability of a Type I
error among tests
.40
.80
.80
.80
.0,5
.01
Area (m^)
10-25
25-50
<50
0-10
-
.05
.05
10-25
-
—
-
25-50
—
—
—
Isolation (m)
81-160
161-240
241 -.320
<.320
0-80
—
-
.05
—
81-160
.05
.01
-
161-240
-
—
241-320
—
Distance (m)
1631-2760
2761-3890
3891-5020
5020-6150
500-1630
.01
.05
.05
1631-2760
—
—
—
2761-3890
—
—
3891-5020
—
Soil depth (cm)
1-2
2-4
4-8
<8
0-1
.05
-
.02
.001
1-2
—
—
.05
2-4
-
.01
4-8
—
Flood (yr)
<15
7.5
.05
Soil depth (cm)
Flood
15
Flood <7.5
.01
ing gardens of Arches and Canyonlands Na-
tional Parks. Where garden habitats are sel-
dom disturbed, large-seeded species are prob-
ably more successful competitors in plant
succession, because their larger propagules
provide more energy to their seedlings.
Tlie term hanging gardens is useful from
the standpoint of vegetation physiognomy,
but it has little relevance to floristic composi-
tion and ecology. The species compositions of
29 hanging gardens in the Narrows of Zion
Canyon were quite di.ssimilar, and variables
of the gardens' physical environment do not
afford a convincing explanation of the differ-
ences (Malanson 1980). Dispersal seems to be
more important in influencing plant assem-
blages than the early succcssional environ-
ment represented by .soil depth, because Ma-
lanson (1980) did not find many species
limited by that variable.
The spatial variables of area, distance to a
likely seed .source, and relative isolation com-
monly are used by island biogeographers to
predict species distributions. Hanging gar-
dens in the Narrows, however, are not de-
monstrably in equilibrium, and, without fur-
ther investigation, we cannot support a spa-
tial explanation of their plant assemblages.
Mo.st species of the Zion Narrows could be
considered "fugitive species" sensu Piatt
(1975), in being both perennial and com-
paratively vagile (annual colonizers are clas-
sified as "ruderals"). According to this meth-
od, following flash floods, the able dispersers
would quickly occupy open spaces. During
later stages of colonization, perennials would
be more successful than annuals. Dispersal of
annuals to a new site already occupied by
perennials would have a low probability of
success because of the scarcity of favorable
spots. There may also be a high ri.sk of seed
loss from any established annuals, given the
restricted habitat spaces available and the
vertical nature of the Narrows environment.
At the gardens situated above the flood
crests, the fugitive species would give way to
species with larger seeds.
December 1980
Malanson, Kay: Hanging Gardens
371
A minority of seven gardens support the
conclusion by Nebeker et al. (1977) that gar-
den assemblages are "random" collections of
plants. The majority of gardens in the Nar-
rows apparently maintain their dissimilar,
fugitive assemblages through response of dis-
persal types to frequent disturbance.
Acknowledgments
We appreciate the assistance of Kimball T.
Harper, Paul A. Kay, Kezia M. Nielsen, Gar-
ry F. Rogers, Lois A. Arnow, Robert W. Aus-
tin, Walter E. Westman and Jonathan D.
Sauer. This research was fimded in part by
grants from the University of Utah Student
Research Grants in Geography, the Zion Nat-
ural History Association, and Sigma Xi, the
Scientific Research Society of North Ameri-
ca.
Literature Cited
.\rnow, L. a., and a. M. Wyckoff. 1977. Flora of the
central Wasatch front. University of Utah Press,
Salt Lake City.
D.\NSEREAU, P., AND K. Lems. 1957. The grading of dis-
persal types in plant communities and their eco-
logical significance. Contr. Inst. Bot. Univ.
Montreal 71:1-52.
Davis, F. J. 1952. Flora of Idaho. Wm. D. Brown Co.,
Diibuque, Iowa.
Diamond, J. M. 1975. Assemblv of species communities.
Pages 342-344 in M. S. Cody and J. M. Diamond,
eds.. Ecology and evolution of communities. Bel-
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Flowers, S. 1973. Mo.sses of Utah and the West. Brig-
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Gabriel, K. R, 1966. Simultaneous test procedures for
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Hitchcock, C. L., and A. Cronquist. 1973. Flora of the
Pacific Northwest. University of Washington
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Levin, S. .\. 1976a. Population dynamic models in
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1976b. Spatial patterning and the structure of
ecological communities. Pages 1-35 in S. A.
Levin, ed., Some mathematical questions in biol-
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Mac.\rtiilr, R. ., and E. O. Wilson. 1967. The theo-
ry of island biogeography. Princeton University
Press, Princeton, New Jersey.
.Maddo.x, J. D., G. E. Hart, and R. H. Hawkins. 1977. A
water resource inventory of the .North Fork of
the Virgin River, Zion National Park. Depart-
ment of Forestry and Outdoor Recreation, Utah
State University, Logan.
Malanson, G. P. 1980. Habitat and plant distributions
in hanging gardens of the Narrows, Zion National
Park. Great Basin Nat. 40:178-182.
Mu.NZ, P. A., and D. D. Keck. 1970. A California flora.
University of California Press, Berkelev.
Nebeker, G. T., K. T. Harper, J. D. Brothehson, and S.
L. Welsh. 1977. Characteristics of plants of com-
mon occurrence in hanging gardens of the Colo-
rado Plateau, Utah. Unpublished manuscript. De-
partment of Botany and Range Science, Brigham
Young University. Provo, Utah.
Plait, W. J. 1975. The colonization and formation of
equilibrium plant species a.s.sociations on badger
disturbances in a tall grass prairie. Ecol. .Monogr.
45:28.5-.3()5.
Simberloff, D. S. 1974. Equilibrium theorv of island
biogeographv and ecologv. .\nnu. Rev. Ecol. &
Syst. 5:161-182.
SoKAL, R. R., AND F. J. RoHLF. 1969. Biometry. W. H.
Freeman, San Francisco.
Welsh, S. L., and C. .\. Toft. 1976. Biotic comnnmities
of hanging gardens in .southeastern Utah. Pages
468-498 in J. R. Murdock, W. L. Welsh, and B.
W. Wood, eds., Navaho-Kaiparowits environmen-
tal baseline studies 1971-1974. Center for Health
and Environmental Studies, Brigham Young Uni-
versity, Provo, Utah.
Welsh, S. L., and B. W. Wood. 1976. Structure of a se-
lected hanging garden. Pages 499-511 in J. R.
Murdock, W. L. Welsh, and B. W. Wood, eds.,
Navaho-Kaiparowits environmental baseline
studies 1971-1974. Center for Health and Envi-
romnental Studies, Brigham Young L'niversitv,
Provo, Utah.
Whitehead, D. R., and C. E. Jones. 1969. Small islands
and the equilibrium theorv of insular bio-
geography. Evolution 23:171-179.
Wood, B. W., and S. L. Welsh. 1976. Productivity of
hanging gardens. Pages 512-.521 in J. R. .Mur-
dock, W.' L. Welsh, and B. W. Wood, eds., Nav-
aho-Kaiparowits environmental baseline studies
1971-1974. Center for Health and Environmental
Studies, Brigham Young University, Provo, Utah.
Woodbury, A. M. 1933. Biotic relationships of Zion
Canyon, Utah, with special reference to succes-
sion. Ecol. Monogr. 3:147-245.
Zion Natural History .\ssociation. 1977. Topograph-
ic map of Zion National Park and vicinity.
Springdale, Utah.
ZONATION PATTERNS IN THE POTHOLES OF KALSOW PRAIRIE, IOWA
Jack D. Brotherson'
,\bstract.— Kalsow Prairie, a mesic prairie remnant in central Iowa, was acquired by the state in 1949 and later
established as a botanical reserve. This study concerns a complex of 14 potholes and adjacent drainage areas within
the prairie. The potholes varied in depth and degree of fill and are thus usefid in studies of plant succession. All 14
potholes exliibit zonation patterns. A total of 36 transects were taken in the various comnuuiity zones. Canopy cover
data were taken in every zone. The zones are ordinated into three-dimensional space as well as clustered. Inter-
specific association patterns are elucidated. A successional sequence is proposed.
There are prairies three, six, ten, and twenty leagues
in length, and three in width, surrounded by forests of
the .same extent; beyond these, the prairies begin again,
so that there is as much of one sort of land as of the
other. Sometimes we saw the grass very short, and, at
other times five or six feet high; hemp, which grows nat-
urally there, reaches a height of eight feet.
A settler would not there spend ten years in cutting
down and burning trees; on the day of his arrival, he
could put his plough into the ground. (Louis Jolliet)
These are the finest and most fertile countries in the
world. . . . From time to time there are vast prairies
where the gra.ss is ten or twelve feet high at all sea-
sons. . . . No settler arriving in the country will not find
at first enough to support plenteously a large family, or
will not, in less than two years" time be as comfortably
settled as in any place in Europe. (Douay)
Such are the early accounts from explorers
and settlers of presettlement lowan vegeta-
tion (Dondore 1926). A government survey
begun in March 1832, when Iowa was still a
territory, and completed in August 1859, first
documented the original extent of this vast
grassland area. That survey indicated that in
the 1850s about 85 percent of Iowa was cov-
ered by grassland (U.S. Government 1868,
Hayden 1945, Hewes 1950, Dick-Peddie
1955).
Accounts by the first explorers, surveyors,
and settlers in the area describe three major
types of landscape in Iowa: (1) woodland, (2)
well-drained prairie, and (3) poorly drained
prairie and marsh (Dondore 1926, Berry
1927, Hewes 1950). The woodlands were
confined to stream valleys and adjacent
slopes, and the prairie was .said to occupv the
remaining portions of the land (Berry 1927,
Hewes 1950). The well-drained prairie areas
were the most extensive except in portions of
the state subjected to late Wisconsinan gla-
ciation; there the poorly drained prairie and
marsh were more common (Hewes 1950,
Hewes 1951, Hewes and Frandson 1952). The
well-drained prairie was described as highly
fertile (Dondore 1926, Berry 1927), whereas
the poorly drained prairie was filled with wa-
ter most of the season and "capable of pro-
ducing nothing but wild rice, frogs, and mos-
quitoes" (Hewes and Frandson 1952).
The characteristics of Iowa prairie in terms
of vegetation types, structure, and general
ecology of the dominant species were the
subjects 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
grassland communities and generally con
eluded that water relations, as affected by
climate, soil, and topography, were respon-
sible for local variations in the structure and
distribution of prairie vegetation. "In varying
the water relations of soil and air they merely
bring about changes in the groupings of the
dominant grasses and accompanying segrega-
tions and rearrangements of the forbs"
(Weaver and Fitzpatrick, 1934).
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 ge-
rardi)—iound on the lower moist slopes
and well-aerated lowlands.
2. Slough Grass type [Sportino pectinata)—
found on poorly aerated and wet soils of
sloughs and natural drainage systems.
'Department of Botany and Range Science. Brighani Young University. Provo, Utah 84602.
372
December 1980
Brotherson: Kalsow Prairie, Iowa
373
3. Tall panic grass-wildrve type [Panicum
virgatum and Elymus canadensis)—
found on soils intermediate between
Slough Grass and Big Bluestein tvpes.
4. Little Bluestem type (Schizocherium
scoparius)—moiii important upland tvpe
on well-drained soils.
5. Needle Grass type {Stipa spartea)—
found on the uplands, often occurring
as a narrow zone following the shoul-
ders of ridges.
6. Prairie dropseed type (Spowbohis
hcterolepis)—ionn(\ locally on the driest
upland sites.
Mover (1953), Aikman and Thorne (1956),
Ehrenreich (1957), and Kennedy (1969) have
presented ecological and taxonomic descrip-
tions of four state-owned native prairie
tracts. The vegetation complexes studied
were basically limited to upland prairie.
Those studies include information on soils,
microclimate, topography, and management,
as well as extensive literature reviews.
These prairies are presentlv owned bv
state agencies and were purchased and set
aside as natural areas with the intent that the
various typical landscapes, wild flowers, and
wildlife of the native tall-grass prairie region
be preserved for posterity. It was also in-
tended that these areas serve as game and
wildlife sanctuaries; as examples of the native
prairie soil types, where comparisons could
be made with cultivated soils of the same soil
association; and as prairie reserves where sci-
entific investigations could be undertaken on
problems concerning the native flora and
fauna of the various topographic, climatic,
and prairie districts throughout Iowa. The
prairies were also meant to serve as reference
points by which future generations could
compare the postsettlement influences of
man on Iowa (Hayden 1946, Moyer 1953,
Aikman 1959, Landers 1966).
One such area is Kalsow Prairie, 160 acres
of unplowed grassland in Pocahontas Coimty,
Iowa. Criteria for its purchase dictated that
this area satisfy the requirements of a game
preserve, contain one or more soil types of an
association, and include several regional veg-
etation types (Hayden 1946). The prairie was
purchased in 1949 by the Iowa State Con-
servation Commi.ssion and since its purchase
has been the object of several studies dealing
with its vegetation, soils, management, in-
sects, response to fire, mammals, and nema-
todes (Mover 1953, Ehrenreich 1957, Esau
1968, Richards 1969. Brennan 1969, Norton
and Ponchillia 1968, Schmitt 1969).
The present investigation was undertaken
to provide information on the phvtosociolog\'
and ecological relationships of poorl\- drained
prairie and marsh areas of Kalsow Prairie. It
includes information on species composition
and distribution, zonation patterns, and inter-
relationships within and between these zones.
Methods
General
This study was begun in the spring of 1967
and continued through the following vear
(1968) and into the summer of 1969. Kalsow
Prairie is one of four state-owned Iowa
prairies. It is located five miles northwest of
Manson, Iowa, and comprises the \E '4 of
Section 36, Belleville Township, T 90N, R
32W, Pocahontas County. It occurs in a part
of north central Iowa that was glaciated dur-
ing the most recent advances of the W'is-
consinan glaciation and within the Clarion-
Nicollet-Web.ster soil association area (Ruhe
1969). The area was chosen for study on the
basis of its vegetational composition, i.e., flo-
ristic richness and the presence of several
pothole areas (poorly drained prairie and
marsh).
Taxonoiu)'
Voucher specimens were collected in du-
plicate throughout the growing seasons of
1967 and 1968. All specimens were identi-
fied, and identical sets have been deposited
in the herbaria of Iowa State University,
Ames, Iowa, and Brigham Young University,
Provo, Utah. Nomenclature follows Pohl
(1966) for the grasses, Gilly (1946) for the
sedges, and Glea.son (1952) for the forbs.
Community Types
Kalsow Prairie contains within its bound-
aries a complex of potholes and correspond-
ing drainage ways. The vegetation of these
communitv types was analyzed itsing two
374
Great Basin Naturalist
Vol. 40, No. 4
separate approaches. The first involved the
identification and listing of all species, and
the second utilized random sample plots to
determine percent cover, composition, and
interspecific relationships of species within
these subcommunities.
Quadrat Analysis
The vegetation of each area was sampled
by using a 20 X 50 cm (1000 cm^) quadrat.
Twenty quadrats were located along tran-
sects on a restricted basis to reduce bias and
GRAZED AREA
Map of the KALSOW PRAIRIE
A areas affected by soil drift from adjacent fields
yCi^ potholes and drainage
— location of 1000 foot bisect
Fis;. 1. Map of Kalsow Fraiiic showing locations ot potholes and drainagf ways.
December 1980
Brotherson: Kalsow Prairie, Iowa
375
to keep adjacent quadrats at fairlv e(jual dis-
tances apart. The number of samples varied
with the subcommunity or zone, but a total
of 720 quadrats were taken within the com-
munity. Sampling was done between 1 Au-
gust and 15 September each vear when most
species had reached their maximum growth.
Cover estimates were made for each quadrat
through use of Daubenmire'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 cano-
pies of different species were commonly in-
terlaced or superimposed over the same area;
therefore coverage percents often total great-
er than 100 percent.
Commiuiity Analysis
Analysis of these areas was accomplished
by dividing the sites into subimits or zones
(Fig. 1) based upon location and dominant
species. Each subvmit was then sampled by
randomly locating a starting point and then
placing a quadrat every 3 m along a transect.
Twenty quadrats were taken for each zone.
Data Analysis
General descriptive data.— Data collected
from quadrat, mapping, and topographic
studies were used to describe the vegetation
of each zone. Frequency values and average
cover values were determined for all species
in every stand. Cover values were deter-
mined by summing the midpoints of the
cover-class ranges and dividing by the num-
ber of sample quadrats in each zone.
Ordination analysis.— An ordination tech-
nique proposed by Orloci (1966) was em-
ployed to ordinate vegetation units within
the different subcommimities listed above.
Raw data were first summarized by hand cal-
culation and then transferred to punch cards.
The entities to be ordinated (i.e., plant spe-
cies or stands of vegetation) were projected
as points into n-dimensional space. Such
points were positioned by attribute scores
through the application of the R and Q-tech-
niques of factor analysis (Orloci 1967). Once
established, this multidimensional array of
points was then reduced to a three-dimen-
sional system. This was accomplished by se-
lecting the two most different stands or spe-
cies and placing one at zero and the other at
some distance along the abscissa. All other
stands or species under consideration were
then positioned linearly in relationship to
these two extremes. This action thus estab-
lished the X-axis. The above process was re-
peated until all points had been established in
three-dimensional space (i.e., Y and Z axes
had been added). Coordinate values for the
X, Y, and Z axes were given as output from
the computer.
Cluster analifsis.- Cluster analyses were
performed by calculating a similarit\' index
(SI) (Ruzicka 1958) in percent from the for-
mula:
SI =
2 min (Xi, Yi)
X max (Xi, Yi)
and then clustering the indices using un-
weighted pair-group clustering techniques
(UPGMA) (Sneath and Sokal 1973). The UP-
GMA method computes the average sim-
ilarity of each unit to the clu.ster, using arith-
metic averages. It is widelv used and has
been found to introduce less distortion than
other methods (Kaesler and Cairns 1972).
Interspecific association analysis.— Expres-
sions of interspecific association were at-
tempted using Cole's Index (1949). Step one
in the computation of the index involved the
accumulation of 2 X 2 contingency tables
(Fig. 7). Actual calculation of the index in-
volved the following three sets of formulas:
when ad < be:
C. ± Sdc =
ad-bc
(a + c)(c + d)
(a + b)(b + d) n(a + b)(l) + c)
when be > ad and d < a:
C, ± Sdc =
ad-bc
(b + d)(c + d)
(a + b)(a + c) n(a + b)(a + c)
when be > ad and a > d:
ad-bc (a + b)(a-Hc)
C, ± Sdc = ^^^^j^^^._^j^ ± n(b-l-d)(c + d)
where C; = Cole's Index of Interspecific
Association
Sdc = standard deviation of Cole's in-
dex
n = total number of samples
376
Great Basin Naturalist
Vol. 40, No. 4
and a, b, c, and d represented the four cells of
the 2 X 2 contingency table.
Tests of statistical significance were per-
formed by means of the Chi-square test. The
Chi-squares were computed by the formula:
(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 represented the different
cells of the 2x2 contingency table.
In all cases, a single degree of freedom was
used. Chi-quare values greater than 3.84
were significant at the 5 percent level, while
values greater than 6.63 were significant at
the 1 percent level.
Data representation.— The three-dimen-
sional graphic representation of data ob-
tained from ordination analysis was drawn by
the computer. Such representation was ac-
complished through the use of a plotting
technique developed and programmed by
Mr. Howard Jesperson, Agricultural Experi-
ment Station, Iowa State University.
Results and Discussion
Marean and Jones (1903) gave the follow-
ing description of the landscape in central
Iowa:
Low knolls are separated by saucerlike depressions in
which impounded water often stands the year around.
In many cases these low-lying areas have been reclaimed
by artificial drainage, but in the main rainwater which
falls upon the upland has to escape by seepage or evapo-
ration. Little ponds and marshes are foimd in almost in-
numerable places scattered all over the coimtry.
These saucerlike depressions have been es-
timated as covering more than 50 percent of
that part of Iowa subjected to late
Wisconsinan glaciation (Hewes 1950). They
were early recognized as supporting a dis-
tinct vegetation from that of the adjacent up-
land prairie (Yapp 1909, Sherff 1912, Shimek
1915, and Berry 1927). The grasses of these
areas were described as being "ten to twelve
feet tall all season" (Dondore 1926). These
and later descriptions indicate that the pot-
holes and drainage ways were often charac-
terized by very discrete zones of vegetation
(Sherff 1912, Shimek 1915, Schaffner 1926,
Fig. 2. Three-dimensional ordination of pothole and drainage zones found in Kalsow Prairie, with numbers corre-
sponding to pothole and drainage mimbers from Figure 1 except 3.3-36, which are prairie edge areas.
December 1980
Brotherson: Kalsow Prairie, Iowa
377
Hayden 1943, Trauger 1967). Three to four
zones were generally recognized, yet in all
cases little information was given on the rela-
tionships of these zones to one another either
floristicallv or spatially. Some authors (Sherff
1912, Schaffner 1926, Hayden 1943), how-
ever, indicated that succession was taking
place in these areas and proposed the follow-
ing successional scheme:
I Pond center
II Sedge zone
III Slough grass zone
IV Dry margin of slough grass
V Andropogon gerardi
VI Upland prairie
Within the boundaries of Kalsow Prairie
there exists a complex of 14 potholes and cor-
responding drainage ways (Fig. 1). These
areas are found scattered throughout the 160
acres at different elevations. They also vary
in depth and degree of fill. These character-
istics make them extremely useful in studies
of plant succession and zonation. All 14 pot-
holes studied exhibited strong zonation. Each
zone was subsampled 20 times for cover and
then averaged to obtain a characteristic veg-
etation for each zone. The 36 zones were or-
dinated into three-dimensional space using
Orloci's (1966) method (Figs. 2 and 3) and
then clustered according to Sneath and Sokal
(1973) (Fig. 4).
Following the ordination and cluster analy-
ses, the zones were then grouped into six
units as .shown in Figures 3 and 4. TTiis pro-
cedure seemed justified since each zone rep-
resented a rather discrete vegetational unit.
After grouping, the data from all zones in-
cluded in each new unit were averaged and
placed in Table 1. These six groups (with one
exception, Group 5) correpond in reality to
the suggested successional sequence shown in
Figure 5. Table 1 is so designed that columns
1 through 6 represent values from the center
of each pothole through a transition to up-
land prairie, respectively. The positioning of
each species within Table 1 was done by
50-
IV
,.\
y
\ -36
32/ -<ll./
0 02<
024 O 0 =
323 Q28
Oio
ill
/^\T
V&VI
\ I
Y
50
100
Fiu. 3. Two-dimensional ordination of pothole and drainage zones of Kalsow Praine. grouped as shown in Table 1
with minor exceptions, llie factors responsible for the ordination are nnknown.
378
Great Basin Naturalist
Vol. 40, No. 4
Table 1. Average percentage cover values in the six groups according to Orloci ordination for the pothole and
drainage communities.
Species
1
3
Polygonum coccineum*
Lysimachia hybrida^
Scirpus fluviatilis^
C^arex atherodes^
Spartina pectinata^
Carex aquatilis*
Carex retrorsa
Phalaris anmdinacea
Sagittaria latifolia
Eleocharis sp.
Calamagrostis canadensis^
Apocvnuni sibiricum
Lvcopus americanus
Convolvulus sepium
Teucrimn canadense
Carex meadii
Iris virginica
.\sclepias incarnata
Hordeum jubatum
Runiex crispus
Panicum capillare
Cirsium altissimum
Asclepias sidlivantii
Zizia aurea
Pycnanthemum virginanum
Elymus canadensis
Thalictrum dasycarpuni
Helenium autumnale
Helianthus laetiflorus
Anemone cylindrica
Solidago rigida
Gentiana andrewsii
Agrostis hiemalis
Heliopsis helianthoides
Ciciita maculata
Lythnuu alatum
Aster ericoides
Panicum virgatuni
Lathynmi palustris
Silphium laciniatum
Eryngiimi yuccifolium
Desmodium canadense
Liatris pycnostachya
Vernonia fasciculata
Rosa sufhdta
Fragaria virginiana
Senecio pauperculus
Solidago gymnospermoides
Audropogon gerardi
Poa pratensis
Solidago canadensis
Sporobolus heterolepis
.\ster simplex
Galium obtusum
Carex lasiocarpa*
Helianthus grosseserratus
66.15
19.50
5.55
2.45
.05
26.74
..39
22.94
10.56
1.89
1.44
.80
.39
.20
2.60
.40
.24
.41
31.15
1.55
.53.45
.80
.75
.20
.05
3.15
.40
5.27
.55
.11
.06
17.54
14.51
3.27
2.64
1.21
.73
26.73
.98
.18
.16
.10
.01
.01
.07
.01
.01
.01
.01
.01
.55
1.15
.01
.01
.02
.24
.32
.58
.01
.86
.35
5.55
.65
2.58
.36
.12
12.53
3.42
2.10
2.21
.01
.29
57.10
2.08
1..36
.14
.76
.26
.14
.01
.02
.21
.07
.06
.53
.10
.01
.17
.60
.01
2.01
1.56
7.43
1.98
.57
.07
2.20
6.05
.02
6.42
.75
1.07
.12
.60
.02
.02
.05
.07
.10
.15
.15
.20
.20
.20
.20
.20
.22
.22
.22
.22
.25
.25
.35
.37
.40
.40
.55
.55
.72
1.32
1.52
1.70
2.20
2.35
3.05
4.07
5.22
5.65
10.15
13.10
^Species picked by the three-dimensional ordination as indicator species.
December 1980
Brotherson: Kalsow Prairie, Iowa
379
assigning those species with the highest val-
ues for Group 1 at the top and those species
with the highest values for Group 6 at the
bottom of the list. It was then possible to de-
termine from tlie table the characteristic dis-
tributional patterns of man\' of the species as
well as their positions of importance within
each zone (i.e., Helianthus grosseserratus is
mainly restricted to Groups 5 and 6 and is
the dominant species of Group 6).
The species of these different zones were
also ordinated into three-dimensional space
(Fig. 6). The ordination did not yield groups
of stronglv associated taxa but rather picked
out eight species exhibiting distinct and dif-
ferent distributional patterns. It placed all
other species within the areas covered by the
circles A, B, and C. When the results of Fig-
ure 6 are compared with those of Table 1, it
can be seen that the species picked bv this
method as indicator species are those taxa
which represent the dominants or sub-
dominants c.f Groups 1 through 6.
A lOOO-foot bisect of the area noted in Fig-
ure 1 was taken in an effort to correlate the
distribution of the dominants of each zone
with elevation and topography. This informa-
tion has been summarized in Figure 7. The
data show that elevation changes of 6 to 12
inches altered the distribution patterns of the
zone dominants.
Attempts to pick groups of associated spe-
cies through the application of Cole's Index
are shown in Table 2 and in Figures 8 and 9.
Figure 8 represents a clearly definable cluster
and includes the dominant species of Groups
1 through 3 of Table 1. These species are
Carex atfwrodes, LijswiacJxia lu/hrida. Poh/-
PERCENT SIMILARITY
-c
^
Hi polygonum COCCINEUM LTSMACHIA HTBtdDA
'lai POLVCONUM COCCINEUM
14' POLYGONUW COCCINEUM -SCIRPUS FLUVIATIliS
1 15 I POLTCONUM COCCINEUM - SClW^US FLUVIATIttS
9, POLYGONUM COCCINEUM - SOflPus FLUVUTIks
i6i POLYGONUM COCCINEUM - SCIRPUS FLUVIAflLIS -CAREX ATHEBOOES
1121 POLYGONUM COCCINEUM -SCtRPUS FLUVIATILIS -CA«I ATMEROOCS
1171 POLYGONUM COCCINEUM - SCIflPUS FLUVIATILIS -CAREX ATHEROOES
171 POLYGONUM COCCINEUM -CAREX ATHERODES
(191 POLYGONUM COCCINEUM - CAREX ATMEROOtS
^13i POLYGONUM COCCINEUM -CAREX ATHEROOES
II POLYGONUM COCCINEUM CAREx AOUATiLIS
131 CAREX AOUATILIS
|29) POLYGONUM COCCINEUM -SPARTIna PECT
]"
LAMOCnOSTIS CANADENSIS
l32» SPARTI
PECT
-CALAMOGROSTIS CANADENSIS
' CALAMOGROSTIS CANADENSIS
-CALAMQOROSTIS CANADENSIS
- CALAMOGROSTIS CANADCNSIS
-CALAMOGROSTIS CANADENSIS
-CALAMOGROSTIS CANADENSIS
CALAMOGROSTIS CANADCNSIS
- CALAMOGROSTIS CANADENSIS
- CALAMOGROSTIS CANAOCNilS
-CALAMOGROSTIS
- CALAMOGROSTIS
SPARTINA PECTtMATA-CALAMOG«OSTl
CALAMOGROSTIS CANADENSIS
CALAMOGROSTIS CANADENSIS
CANADCNSIS
28 CALAMOGROSTIS CANADCNSIS
— 3Si PRAiW'l COGC
— 36 PRAWK EDUC
\3T' PR»>"it EDGB
i34l PRAIRIE I
Fig. 4. Phenogram of 36 potholo zones as developed from cluster analysis tSneath and Sokai 1973). Croups are a.s
shown in Table 1 with minor exceptions.
380
Great Basin Naturalist
Vol. 40, No. 4
gonum coccineum, and Scirpus fluviatilis.
Figures 8b and 9 show several definable clus-
ters and include taxa found in Groups 4
through 6 of Table 1. The cluster of species
POLYGONUM COCCINEUM-LYSIMACHIA HYBRlDA I
i
POLYGONUM COCCINEUM- SCIRPUS FLUVIATILIS II
I
POLYGONUM COCCINEUM -CARE X ATHERODES m
1/ ^
SPARTINA PECTINATA ^CAREX AQUATILIS IV&V
CALAMOGROSTIS CANADENSIS
CALAMOGROSIIS CANADENSIS VI
I
PRAIRIE EDGE VII
Fig. 5. Suggested successional sequence for potholes
of Kalsow Prairie.
designated by A in Figure 9 contains species
found entirely in Groups 4 and 5 of Table 1.
Those clusters identified by the letters B and
C of this same figure contain only plants
found in Group 6 of Table 1 and correspond
in reality to the prairie edge. Cluster A and
Clusters B and C are bridged by a single spe-
cies {Aster simplex) that is found growing
mainly along the border between Groups 5
and 6 of Table 1.
The vegetation of the potholes and drain-
age ways of Kalsow Prairie can best be de-
scribed as a series of five zones (Fig. 5), each
of which exhibits different spatial and floris-
tic properties. This characteristic zonation
can be expected to repeat itself from pothole
to pothole when controlling environmental
factors are similar. The zones themselves are
best described by starting at the center of the
potholes and moving toward the prairie edge.
Zone 1 (Group 1 of Table 1, etc.) is found at
the center of the deepest potholes and is
lOO—
SO—
X
o o
09 O'
-7f-
Y
50
100
220
Fig. 6. Two-dimensional ordination of species found in pothole and drainage areas of Kalsow Prairie; A, B, and C
= clusters of species not showing distinct distribution patterns, d = Polyg,onu)n coccincuui (usually in center of
pothole), e = Sci>/;(/.s fluviatilis. f = Q/re.v atherodes. g = Lijsiutachia hyhrida, h = Spartina pectinata. i = Carcx
(Ujuatilis, i = Carex la.siocurfm, k = Calamagrostis canadensis (usually in outer zone of pothole complex).
December 1980
Brothersox: Kalsow Prairie, Iowa
381
Table 2. Cole's Index values expressing positive interspecific association in pothole and drainage communities.
Species
Species
X2a
Ct"
sde,
Andropogon gerardi
Anemone cvlindrica
Apocvnuni sibiricum
.Aster simplex
Calamasrrostis canadensis
Carex atherodes
Carex meadii
Carex aquatilis
Carex lasiocarpa
Carex retrorsa
Convolvulus sepium
Desmodium canadense
Eleocharis sp.
Elvmus canadensis
Eryngium ynecifoli\im
Fragaria virginiana
Galium ohtusum
Helenium autunniale
Heliantlnis grosseserratus
Iris virginica
Eryngium \aiccifolium
Liatris pycnostaclna
Sporobolus heterolepis
Fragaria virginiana
Panicum virgatum
Thalictrum das\carpum
Calamagrostis canadensis
Carex lasiocarpa
Carex retrorsa
Spartina pectinata
Calamagrostis canadensis
Carex lasiocarpa
C.alium ohtusum
Heliantlius grosseserratus
Poa pratensis
Spartina pectinata
Carex acjuatilis
Carex lasiocarpa
Carex retrorsa
Spartina pectinata
Polygonum coccincum
Scirpus fluviatilis
Rosa suffulta
Carex lasiocarpa
Carex retrorsa
Spartina pectinata
Carex retrorsa
Spartina pectinata
Spartina pectinata
Galium ohtusum
.\ster ericoides
Liatris pycnostachya
Ratihida columnifera
Senecio pauperculus
Phalaris anmdinacea
.\ster ericoides
Galium ohtusum
Helianthus grosseserratus
Lathvrus palustris
Desmodimn canadense
Liatris pycnostachya
Senecio pauperculus
Thalictrum dasycarpum
Sporoholus heterolepis
Andropogon gerardi
Ciaiium ohtusum
Helianthus grosseserratus
Poa pratensis
Senecio pauperculus
Solidago canadensis
Sporoholus heterolepis
Helianthus grosseserratus
Poa pratensis
Spartina pectinata
Helianthus grosseserratus
Poa pratensis
Solidago canaclciisis
Poa pratensis
Rosa sufhilta
Vemonia fasiculata
119.80
.30
.02
130.35
.38
.03
65.90
.22
.02
89.71
.19
.02
84.61
.24
.02
91.73
..33
.03
41.22
.68
.10
30.42
.57
.10
9.44
.18
.05
19.63
..37
.08
.36.47
.56
.09
31.79
.51
.09
212.39
.51
.03
231.49
.51
.03
85.80
.18
.01
19.10
.31
.07
49..30
.17
.02
338.21
.68
.03
81. .33
.19
.02
200.44
.41
.02
153.47
.93
.07
120..34
.47
.04
28.47
..38
.07
73..33
.49
.05
53.85
.24
.03
19.61
.20
.04
111.72
.70
.06
18.3.90
.40
.02
43.76
.35
.05
8.76
..39
.13
.54.86
.28
.03
87.08
.42
.04
200.56
.28
.02
189.69
.71
.05
28.62
.25
.04
65.57
..33
.04
4..34
..39
.18
.30.83
1.00
.18
62.34
.49
.06
64.57
..33
.04
44.21
.32
.04
.33.25
..32
.05
91.73
.33
.03
.54.86
.28
.03
.37.61
.19
.03
46.46
.70
.10
104.86
1.00
.09
19.29
.2.5
.05
37.61
.19
.a3
.30.90
.27
.04
75.87
.19
.02
242.41
.60
.03
86.58
.20
.02
13.73
.29
.07
5..54
.60
.2.5
131.97
.2.5
.02
112.89
.20
.01
7..38
.29
.10
23. 1 1
..32
.06
8.39
.29
.10
382
Great Basin Naturalist
Vol. 40, No. 4
Table 2 continued.
Species
Species
Lathyrus paliistris
I .iatris pycnostachya
I.Nsiniachia hybrida
Lvthnnn alatuin
Panicuni virgatum
Poa pratensis
Polygonum coccineum
Rosa sufftilta
Solidago canadensis
Zizia aurea
Poa pratensis
Solidago gymnospermoides
Thalictruni dasycarpiun
Senecio pauperculus
Sporobolus heterolepis
Polvgoniun coccineum
Scirpus fluviatilis
Vernonia fasciculata
Poa pratensis
Teucrium canadense
Solidago canadensis
Scirpus fluviatilis
Andropogon gerardi
Solidago canadensis
Senecio paupercidus
Desm odium canadense
^hi-square
"Cole's Index
'^Standard dexiation Cole's Index
-rNNW
FEET
X^a
1000
Sd&,
9.53
.20
.06
62.18
.30
.03
75.89
.18
.02
44. .56
.29
.04
248..50
..50
.03
16.21
.61
.15
16.11
..34
.08
5.46
.20
.08
14.89
.29
.07
45.6.3
..39
.05
92.29
.30
.03
223.30
.32
.02
42.51
.21
.03
.35.46
..30
.05
160.63
.30
.02
1.32.56
.66
.05
Fig. 7. Correlation of dominant species of each zone from potholes and drainages with changes in elevation along
1,000 ft (925 m) bisect; 1 = Sporobolus heterolepis, 2 = Helianthus ^rosseserrottis. 3 = Calarnagrostis canadensis. 4
= Carex atherodes, 5 = Scirpus fluviatilis, 6 = Polygonum coccineum.
dominated chiefly by Polygonum coccineum
and Lijsimachia hijhrida. Zone 2 is found to
conipletelv encircle Zone 1 and is character-
ized by the dominants Polygonum coccineum
and Scirpus fluviatilis. Zone 3 is found as a
very narrow band that encircles Zone 2 or
occurs as rather wide patches in areas of
equivalent elevation. It is characterized
chiefly by Carex atherodes. Zones 4 and 5 are
best distinguished in potholes and drainage
ways that are somewhat shallow. Zone 4
most often occupies the center of these shal-
low depre.ssions surrounded by Zone 6. The
dominant species {Carex aquatilis) of Zone 5
is u.sually found as a subdominant of Zone 4
but at times appears as a dominant zone sur-
rounded bv Zone 6. Whether this relationship
is due to competition and/ or environmental
influences is unknown. Zone 4 is character-
ized by the species Spartina pectinata, Carex
aquatilis, and Calamogrostis canadensis.
Zone 6 is distinguished by the dominant spe-
cies Calamagrostis canadeiisis and a few
other participating species (i.e., Apocynum
sibiricum, Lycopus americanus, Teucrium
canadense, Carex meadii, and Iris virginica).
Zone 7 and column 6 of Table 1 represent
the prairie edge and are characterized pri-
marilv by the presence of Helianthus grosse-
serratus.
December 1980
Brotherson: Kalsow Prairie, Iowa
383
These zones appear to represent a succes-
sional sequence that is controlled basically by
the degree of pothole fill and consequently
by related moisture regimes. The successional
scheme (Fig. 5) parallels in many respects a
scheme proposed bv earlier authors (Sherff
1912, Schaffner 1926, Hayden 1943).
The actuality of this scheme is based on
the repeatability of the zonation pattern as
found within the potholes of Kalsow Prairie.
Evidence for change of fluctuations in pot-
hole vegetation paralleling this sequence will
depend on the results obtained from long-
term studies.
Literature Cited
AiKMAN, J. M. 1959. Prairie research in Iowa. American
Biology Teacher 21:7-8.
.\iKMAN. J. M., AND R. F. Thorne. 1956. Tlie (Javier
Prairie: an ecologic and taxonomic stuck' of a
northwest Iowa Prairie. Iowa Acad. .Sci.
ft3: 177-200.
Berry, W. J. 1927. The influence of natural environment
in north central Iowa. Iowa J. Historv and Poli-
tics 25:277-298.
Fig. 8. .\ssociated species of potholes and drainage,
Groups I and 2 (Table 1) as determined by Cole's (1949)
Index; (a) Ca at = Carex athcrodes. Ly hy = Lysi-
machia liyhrida, Po co = Polygonum coccineum, Sc fl
= Scirpns fluviatilis\ (b) El sp = Eleocharis sp., Ph ar
= Phalaris unindinacea.
Bhen.nan, K. M. 1969. Vertebrate fauna of Kalsow
Prairie. I'npublished thesis. Iowa State Univer-
sity.
Cole, L. C. 1949. The measurement of interspecific as-
sociation. Ecology .30:411-424.
Daibe.N'mire, R. 1959. A canopy-coverage method of
vegetational analvsis. Northwest Science
.33:4.3-66.
Dic:k-Peddie, VV. .\. 1955. Presettlement forest types in
Iowa. Unpublished thesis. Iowa State Univcrsitv.
Do.NDORE, D. \. 1926. The prairie and the making of
middle .America: four centuries of description.
Cedar Rapids, Iowa, Torch Press. 472 pp.
Ehrenreich, J. H. 1957. Management practices for
maintenance of native prairies in Iowa. Unpub-
lished dissertation, Iowa State l'niversit\'.
Esau, K. L. 1968. Carabidae (Coleoptcra) and other ar-
thropods collected in pitfall traps in Iowa corn-
fields, fencerows and prairies. Unpublished dis-
sertation, Iowa State University.
GiLLY, C. L. 1946. The Cvperacae of Iowa. Iowa State
Coll. J.Sci. 21:.5.5-151.
(Ileason, H. \. 1952. The new Britton and Brown illus-
trated flora of the northeastern United States and
adjacent Canada. Lancaster, Penns\lvania. Lan-
caster Press, Inc. 3 vols.
H.\yde.\, .\. 1943. .\ botanical survey in the Iowa lake
region of Clav and Palo .\lto Counties. Iowa
State Coll. }. 17:277-416.
1945. The selection of prairie areas in Iowa
which should be preserved. Proc. Iowa .\cad. Sci.
52:127-148.
1946. .\ progress report on the preservation of
prairie. Proc. Iowa .\cad. Sci. .53:45-82.
Hewes, L. 19.50. Some features of earlv woodland and
prairie settlement in a central Iowa count)'. .An-
nals .\ssoc. .\mer. Geog. 40:40-.57.
1951. The northern wet prairie of the United
States: nature, sources of information, and extent.
.\nnals .\ssoc. .\mer. Geog. 41:.307-.32.3.
Hewes, L., a.nh P. E. Fra.ndson. 19.52. Occupying the
wet prairie: the role of artificial drainage in Story
Countv, Iowa. .\nn. .\ssoc. .\mer. Geog.
42:24-.50.
Kaesler, R. L., a.nd J. Cair.ns. 1972. Cluster analysis of
data from limnological surveys of the upper Poto-
mac River. .\m. Midi. Nat. 88:56-67.
Kennedy, R. K. 1969. \x\ analysis of tall-grass prairie
vegetation relative to slope position, Shccder
Prairie, Iowa. Unpublished thesis, Iowa State
Universitv.
Landers, R. Q. 1966. Visit the \irgin prairie. Iowa Farm
Science 21:418-419.
Marean, H. VV., AND G. B. Jones. 1903. Soil survey of
Storv Countv, Iowa. Field Operations of the Bu-
reau of Soils 836.
MoYER. J. F. 1953. Ecologv' of native prairie in Iowa.
I'npublished dissertation, Iowa State University.
Norton, D. C.. and P. E. Ponchilla. 1968. Stylet-bear-
ing nematodes associated with plants in Iowa
prairies. J. Iowa .\cad. Sci. 75:32-35.
Oreoci. L. 1966. Geometric models in ccolog). I. The
theorv and application of some ordination meth-
ods, j'. Ecology .54:19.3-215.
PoHL, R. \\'. 1966. The grasses of Iowa. Iowa State J.
Science 40:341 -.566.
384
Great Basin Naturalist
Vol. 40, No. 4
Fig. 9. Associated species of potholes and drainage. Groups 3-6 (Table 1) as determined by Cole's (1949) Index,
the more lines between species, the greater the association; groups A, B, and C are basic clusters; .\n ge = Andwpo-
gon gerardi. An cv = Anemone eijlindnca, Ap si = Apoci/ntim sihiiicuni. As er = Aster ericoides. As si = Aster
simplex. Ca aq = ('arex cujuatilis, Ca ca = Cakimagrostis eanadensis, Ca la = Carex hisiocarpa, Ca me = Carex
meadii. Ca re = Carex retrorsa, De ca = Desmodiiim canadensis. El ca = Elymits canadensis, Er yu = Enjngium
i/iicci folium. Fr vi = Fragaria virginiana, Ga ob = Galium obtusum. He gr = Helianthus grosseserratus, Ir vr = Iris
virginica. La pa = Lathyrus paUistris, Li pii = Liatris pycnostachya, Ly al = Lythrum alatum. Pa vi = Panicum
virgatum. Po pr = Poa pratensis, Ra co = Ratibida columnifera, Ro su = Rosa suffulta, Se pa = Serteeio pauper-
culus. So ca = Solidago canadensis. So gy = Solidago gymnospermoides. Sp he = Sporobohis heterolepis, Sp pe =
Spaiiina pectinata. Te ca = Teucrium canadense, Tha da = Tlialictrum dasycarpum. Ve fa = Vernonia fasicukita.
Zi au = 7.iziii aurea.
Richards, M. S. 1969. Observations on responses of
prairie vegetation to an .\pril fire in central
Iowa. Unpublished thesis, Iowa State University.
RiHE, R. V. 1969. Quaternary landscapes in Iowa. .Ames,
Iowa, Iowa State University Press. 249 pp.
RuziCKA, M. 1958. Anwendung mathematish-statisticher
methoden in der Geobotanik (Synthetische bear-
bietung von aufnahmen). Biologia, Bratisl.
1.3:649-661.
Rydberg, p. \. 1931. A short phvtogeographv of the
prairies and great plants of Central North .\nieri-
ca. Brittonia 1:57-66.
ScHAFFNER, J. H. 1926. Grasslands of the Central United
States. Ohio State Universitv Studies Contr. Bot.
178:5-56.
Sc:hmitt, D. p. 1969. Plant parasitic nematodes and
nematode populations in the Kalsow Prairie. Un-
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Sherff, E. E. 1912. Tlie vegetation of Skokie marsh,
with special reference to subterranean organs
and their interrelationships. Bot. Gaz.
53:415-435.
Shi.mek, B. 1915. The plant geography of the lake Oko-
boji region. Bull. Lab. Nat. Hist., State Univ.
Iowa 7:1-90.
Sne.\th, p. H. a., and R. R. Sokal. 1973. Numerical ta.x-
onomy. W. H. Freeman. San Francisco, Califor-
nia. 573 pp.
Steiger, T. L. 1930. Structure of prairie vegetation.
Ecology 11:170-217.
Traucer, D. L. 1967. Habitat factors influencing duck
brood use of semi-permanent and permanent
prairie potholes in North Dakota. Unpublished
thesis, Iowa State University.
U.S. Government. 1868. First survey of the state of
Iowa. Plants deposited in the State House, Des
Moines.
Weaver, J. E., and F. E. Clements. 1938. Plant ecolo-
gy. 2d ed. McGraw-Hill, New York.
Weaver, J. E., and T. J. Fitzp.atrick. 1934. The prairie.
Ecol. Monogr. 4:109-295.
Yapp, R. H. 1909. On stratification in the vegetation of a
marsh, and its relations to evaporation and tem-
perature. Ann. Bot. 23:275-320.
PLANTS OF ANGEL ISLAND, MAIUN COUNTY, CALlFORNL\
J. D. Ripley'
Abstract.— The floristic composition of .\nge\ Island, Marin County, California, is presented. The vascular flora
consists of 416 native and naturalized plants, representing 2.52 genera and 7.J families. Eight) species of ornamental
plants persisting from cultivation, representing 64 genera and '39 families, are also identified. The nonvascular flora
consists of 42 species of fvuigi, 7 species of green algae, 8 species of brown algae, and 41 species of red algae. The
\ascular flora occurs in seven plant communities and is composed mainly of the following plant types: forbs (69
percent), grasses (1.3 percent), shnibs (8 percent), and trees (4 percent), .\pproximately 25 percent of the 1605 native
and naturalized vascular plants of Marin ('ounty occur on .Xngel Island.
Angel Island is situated in San Francisco Tiburon Peninsula, hv Raccoon Strait, which
Bav off the southern coast of Marin Countv, is about a mile in width at the narrowest
California. The island lies approximately 3 '2 point.
miles north of the city of San Francisco and The shape of the island, as shown in Figure
is separated from the nearest mainland, the 1. roughly resembles an equilateral triangle
10
12
13
15
16
17
19
20
CAMPBELL POINT
North Garrison
SIMPTON POINT
POINT lONE
STUART POINT
West Garrison
Camp Reynolds
KNOX POINT
LEGEND
aacz:^ Fire Road
= = = == Lower Road
Sunset Trail
North Ridge Trail
FT. Mc DOWELL
East Gorrison
Quorry Beach
SAN FRANCISCO BAY
BLUNT POINT
10
12
13
15
16
19
20
Fig. I. Mapot Augcl MaiKJ.
■Department of Biology, US.\F .\cademy. Colorado 80840. Present address: Departn.cnt of Botanv and PUnt Pathology. OreRon Stale University.
Corvallis, Oregon 973.30.
385
386
Great Basin Naturalist
Vol. 40, No. 4
with the sides measuring about VA miles in
length. Hence, the area of the island is
sliglitly larger than one square mile and equal
to approximately 740 acres.
The topography of the island is roughly
pyramidal in form, with the central peak, Mt.
Livermore (elevation 781 ft), forming the
central axis for various spurs that radiate out
to the perimeter of the island.
Numerous water seepages occur in the
canyons formed by the radiating ridges, but it
is only in the canyon extending from the
summit of Mt. Livermore to Perle's Beach
that any significant water flow occurs. The
damming of this small stream in two loca-
tions has resulted in the formation of two
small ponds.
With the exception of several miles of nar-
row, sandy beaches, the island is generally
hilly. In some areas, particularly near Simp-
son Point and Blunt Point, the cliffs are ex-
tremely steep.
Climate
Due to the steady sweep of air from the
Pacific Ocean, the climate of Angel Island is
characterized by few extremes of heat or
cold. Like other central coastal regions of
California, the winters are cool and wet and
the summers are warmer and dry. Until re-
cently, only intermittent precipitation and
temperature records were maintained on the
island. The limited data available indicate a
total annual precipitation ranging from a low
of 2L34 inches to a high of 34.53 inches,
with the average being 26.62 inches. These
data further reveal that 84 percent of the to-
tal average precipitation occurs during the
five-month period between November and
March and that measurable amounts of pre-
cipitation occur on the average less than 70
days a year. As in many coastal regions of
California, summer fog and low overcast play
an important role in tlie island's climate.
Summer mornings are frequently foggy, but
clearing usually begins early in the forenoon.
These low summer fogs no doubt contribute
soil moisture as condensation occurs on
leaves of trees and drips to the ground (Ober-
lander 1956).
The average annual temperature on Angel
Island is 56 F. The average monthly mini-
mum temperature occurs in January and is 42
F, and the average monthly maximum tem-
perature occurs in September and is 73 F.
The average monthly temperature ranges
from a low of 46 F in January to a high of 65
F in September.
Geology
Ransome (1894) conducted the first and
most extensive study of the island's geology.
This study has been supplemented and up-
dated by Bloxam (1960) and by Schlocker,
Bonilla, and Radbruch (1958), who compiled
an extensive geological map of the island.
The geological features of the island have
also provided source material for other stud-
ies of rock formations in the San Francisco
Bay area such as those of Bloxam (1956),
Schlocker (1961), and Bailey, Irwin, and
Jones (1964). Ransome (1894) describes the
structure of the island as consisting essen-
tially of a syncline trough with its axis plung-
ing to the northeast. The rocks of Angel Is-
land all belong to the Franciscan Formation
and consequently were formed during the Ju-
rassic approximately 100 million years ago.
Bloxam (1960) identified the following six
major rock groups on the island: (1) Francis-
can graywackes, conglomerates, and radio-
larian cherts; (2) jadeite-bearing metagray-
wackes derived from the Franciscan
graywackes; (3) glaucophane-schists derived
from graywackes, cherts, and basic igneous
rocks; (4) a large and apparently concordant
sheet of altered diabase intrusive into the
Franciscan sediments; (5) pillow-lavas at
Blunt Point; and (6) a vertical dikelike body
of serpentinite and pyroxenite near the west-
ern end of the island. Of these rock groups,
the near vertical serpentine dike extending
across the western portion of the island is the
predominant geological feature of the area.
Bloxam (1960) further points out that all the
rocks of the island are metamorphosed to
various degrees, with the exception of the
graywacke and shale at Quarry Point, which
are unmetamorphosed.
Vegetation Communities
Seven categories of vegetation cover are
recognized as occurring on Angel Island.
December 1980
Ripley: Angel Island Plants
387
S niXED EVERGRtEM FOREST
I 1 GRASSLAND
\M CHAPARRAL
H COASTAL SCRUB
H COASTAL 5TRAND
^ DISTURBED AREAS
FRESHWATER POND
Fi<i. 2. Map ot Angel Island showing location ot its se\("n tatcgories of vegetation cover.
Table 1. Tabulation of the native and naturalized vacular flora.
Division
Class
Family
Genus
Specific
and infraspec
ific taxa
Subclass
Native
Naturalized
Total
Calamophyta
Traeht'ophvta
I
1
2
0
2
Filicinae
3
7
7
0
7
Cyinnospermae
.\ngiosperiTiae
Monocotvledonae
2
10
2
45
0
60
2
30
2
90
Dicotyledoneae
57
197
213
102
315
Totals
73
252
282
134
lib
These include mixed evergreen forest (33
percent), grassland (28 percent), altered or
disturbed areas (20 percent), coastal scrub (10
percent), chaparral (4 percent), coa.stal strand
(4 percent), and freshwater ponds (less than
one percent). The location of each vegetation
type is indicated in Figure 2.
Flora of the Island
The native and naturalized vascular flora
of Angel Island consists of 73 families. 252
genera, and 416 species and infraspecific taxa
(Table 1). In addition, 80 species of culti-
vated plants have been identified (Table 2) as
388
Great Basin Naturalist
Vol. 40, No. 4
Table 2. Tabulation of the nonvascular flora identi-
fied.
Division
Genus Species
Eumycophyta
Chlorophyta
Phaeophyta
Rhodophyta
Totals
28
5
8
33
74
42
7
8
41
well as 42 species of higher fungi and 56 spe-
cies of marine algae (Table 3). The 10 largest
families are listed in Table 5 and contain 138
genera and 265 species, or 64 percent of the
total native and naturalized vascular flora.
Comparing the 416 species and infraspecific
taxa or native and naturalized vascular plants
with the 1605 species and infraspecific taxa
recognized by Howell (1970) for all Marin
County indicates that the flora of Angel Is-
land represents approximately 25 percent of
the total Marin County flora. No endemic
species occur on Angel Island nor is the is-
land the type locality for any plants.
Table 3. Tabulation of the cultivated plants identi
fied.
Class
Subclass
Family Genera Species
Gymnospermae 5 11 12
Angiospermae
Monocotyledoneae 5 8 10
Dicotyledoneae 29 45 58
Totals 39 64 80
Checklist
The relative inaccessibility and long mili-
tary history of Angel Island precluded a de-
tailed floristic examination of the area prior
to the study upon which the checklist below
is largely based (Ripley 1969). Early collec-
tors included Albert Kellogg (no date),
George Vasey (1876), T. S. Brandegee (no
date), Volney Rattan (1877), and Joseph
Burtt-Davy (no date). Only one or two Angel
Island collections exist from each of these
collectors. Collectors in the first half of the
Table 4. Plant characteristics of Angel Island native, introduced, and cultivated plants.
Percent of
Percent of
total flora
total native
(native.
Generalized life for
ni
Total
Percent of
individual
category
and
introduced
flora
introduced,
and
Growth habit
Annual
Biennial
Perennial
cultivated)
I. Native plants
Fems
0
0
7
7
3
1,7
1.3
Sedges
Rushes
1
1
0
0
5
5
6
6
2
2
1.5
1.5
1,2
1,2
Grasses
5
1
24
30
11
7.3
6,0
Forbs
73
8
110
191
67
46.0
.39.0
Shrubs
0
0
.30
30
11
7,2
6.0
Trees
0
0
10
10
4
2.4
2.0
Subtotals
80
9
191
280
100
67.3
56.4
II. Introduced plants
Grasses
17
0
8
25
19
6,0
5,0
Forbs
61
8
28
97
71
23,0
19,9
Shnibs
0
0
7
7
5
1,7
1,3
Trees
0
0
7
7
5
1,7
1,3
Subtotals
78
8
50
136
100
32.7
27,4
III. Cultivated
(ira.sses
Forbs
Shnibs
Trees
PLANTS
0
0
0
0
0
0
0
0
0
0
1
6
22
51
80
1
6
22
51
80
1,3
7.5
26.5
64.7
,20
1,2
4.4
10.0
Subtotals
100.0
16.2
Grand totals
158
17
320
493
December 1980
Ripley: Angel Island Plants
389
Table 5. The 10 largest families of vascular plants.
Species
and
infraspecific
Family
Genera
taxa
Compositae
47
76
Gramineae
25
61
Letjuniinosae
12
45
ScTophiilariaceae
7
16
Cniciterac
14
16
Rosaccac
10
12
I'mhelliferae
10
12
Car\()phyllaceae
6
11
Onagraceae
4
9
Cyperaceae
3
7
Totals
138
265
twentieth centurv included E. C. Sntliffe
(1920), Alice Eastwood (1925), and John
Thomas Howell (1946 and 1949). The 51
specimens collected by Howell are the most
important of these collections. In addition,
numerous plants were observed but not col-
lected by Howell on his two trips that
formed the basis for many citations in Marin
Flora (1970). The most recent collectors have
included Otto T. Solbrig (1956), Peter H.
Raven and Michael P. Johnson (1967), J.
Douglas Riplev (1967-1969), Gordon True
(1978), Alva Dav (1979-1980), and Catherine
G. Burke (1979-1980). Of these collections,
the most extensive are those of Ripley with
1074 and Raven and Johnson with 337.
The following abbreviations are used in
this checklist to indicate individual collec-
tors
B
B-D
D
(Catherine G. Burke
Joseph Bvirtt-Davy
Alva Dav
E Alice Eastwood
H John Thomas Howell
K Albert Keilogt;
R V'olnev Rattan
JDR J. Douglas Ripley
R & J Peter H. Raven and Michael P. Johnson
S E. G. Sutlifte
OS Otto T. Solbrig
T Gordon True
TB T. S. Brandegee
\' George Vasey
Certain plant characteristics are also in-
dicated in the checklist following the scien-
tific name for all vascular plants. The follow-
ing abbreviations are used for this purpose:
P perennial
B biennial
A annual
N native
I introduced
(' ornamental plant persisting from cultivation
T tree
S shrub
F fori)
Fr fern
G grass
R rush
Sd sedge
Finally, the approximate collection loca-
tion of each specimen is indicated bv map
grid coordinates from Figure 1 .
The checklist is arranged in alphabetical
order by families for ease of reference. Iden-
tification of the algal flora follows that of
Smith (1944) and Hollenberg and .\bbott
(1966). The vascular plant identification fol-
lows that of Munz (1973) and Howell (1970).
Division Eumycophyta— Higher Fimgi
Agrocybe pediades (Fr.) Favod.
JDR 1008, 2 Mar 1968, 'L-6; JDR 1048, 31
Mar 1968, G-12.
Aleuria atirantia (Fr.) Fuckel.
JDR 1000,2 Mar 1968, F- 10.
Amanita gemmata (Fr.) Gill.
JDR 1005, 2 Mar 1968, F- 10.
Amanita pantherina (DC. ex Fr.) Seer.
JDR 1656, 30 Dec 1968, F-10.
Amanita rubescens (Pers. ex Fr.) Gray
JDR 1704, 20 Apr 1969, E-Il.
Amanita vaginata (Fr.) Vitt.
JDR 998, 2 Mar 1968, F-10.
Bolbitius vitcllinus (Fr.) Fr.
JDR 975, 6 Feb 1968, HI 3; JDR 1002, 2
Mar 1968, D-11.
Boletus chrysenteron (Bull, ex Fr.) Q)uel.
JDR 1003, 2 Mar 1968, F-12.
Boletus subtomentosus (L. ex Fr.) Quel.
JDR 1662, 30 Dec 1968, D-10.
Chroogomphus rutilus (Fr.) Miller
JDR 974, 6 Feb 1968, J-9.
Clavariadelphus ligula (Fr.) Donk.
JDR 963, 6 Feb 1968, F-10.
Clavulina cristata (Fr.) Schroet.
JDR 1660, 30 Dec 1968. F-10.
390
Great Basin Naturalist
Vol. 40, No. 4
Laccaria striatula (Peck) Peck
JDR 1659, 30 Dec 1968, F-10.
Lactarius chrysorheus Fr.
JDR 964, 6 Feb 1968, F-10.
Lactarius suhdulcis (Bull, ex Fr.) Gray
JDR 965, 6 Feb 1968, F-10; JDR 1669, 30
Dec 1968, E-12.
Lepiota rachodes (Vitt.) Quel.
JDR 944, 27 Dec 1967, J-4.
Marasmius Candidas Fr.
JDR 999, 2 Mar 1968, F-10.
Marasmius plicatulus Peck
JDR 968, 6 Feb 1968, F-10; JDR 1004, 2
Mar 1968, F-12.
Nematolonia fascicidare (Fr.) Karst.
JDR 945, 27 Dec 1967, F-7; JDR 976, 6
Feb 1968, 1-13.
Peziza badia Merat.
JDR 480, 28 Apr 1967, J-9.
Pleuteits cervinus (Schaeff ex Seer.) Kummer
JDR 1667, 30 Dec 1968, F-12.
Ramariopsis kiinzei (Fr.) Donk.
JDR 1010, 2 Mar 1968, K-9.
Schizophylhim commune Fr.
JDR 1047, 31 Mar 1968, F-10.
Suillus pungens Thiers
JDR 938, 27 Dec 1967, L-5; JDR 939, 27
Dec 1967, J-9; JDR 1657, 30 Dec 1968, F-10.
Trernella mesenterica Fr.
JDR 970, 6 Feb 1968, E-11.
Clitocybe inversa (Fr.) Quelet
JDR 971, 6 Feb 1968, K-14.
Clitocybe ntida (Fr.) Cooke
JDR 969, 6 Feb 1968, F-10; JDR 1049, 31
Mar 1968, G-12.
Clitocybe robusta Peck
JDR 1661, 30 Dec 1968, F-12.
Cortinarius cinnamonieus Fr.
JDR 976, 6 Feb 1968, F-10.
Cortinarius croceofolius Peck
JDR 1007, 2 Mar 1968, J-9.
Entolonia serriceus (Fr.) Kuhn. & Romagn.
JDR 966, 6 Feb 1968, F-10.
Gyroniitra hicunosa Afx. ex Fr.
JDR 475, 28 Apr 1967, J-9.
Hebeloma crustiliniforme Quelet
JDR 476, 28 Apr 1967, J-9.
Hygrophorus ehurneus Fr.
JDR 1672, 30 Dec 1968, H-12.
Hygrophorus flavescens (Kauffmann) Smith &
Hesler
JDR 1009, 2 Mar 1968, K-9.
Hygrophorus hypothejus (Fr.) Fr.
JDR 1666, 30 Dec 1968, E-12.
Hygrophorus ponderatus Britz.
JDR 479, 28 Apr 1967, J-9.
Laccaris amethystina (Bolt, ex Fr.) B. & Br.
JDR 940, 27 Dec 1967, J-9; JDR 1671, 30
Dec 1968, H-12.
Tricholoma flavovirens (Fr.) Lundell
JDR 942, 27 Dec 1967, J-9; JDR 1644, 30
Dec 1968, F-12.
Tricholoma virgatum (Fr. ex Fr.) Kummer
JDR 1359, 29 Apr 1968, F-10.
Volvariella speciosa (Fr.) Sing.
JDR 1673, 30 Dec 1968,1-11.
Xylaria hypoxylon (Linn.) DuMortier
JDR 1673, 30 Dec 1968, Ml.
Division Chlorophyta— Green Algae
Bryopsis corticulans Setchell
JDR 1492, 10 Jun 1968, E-9.
Cladophora columbiana Coll.
JDR 1475, 10 Jun 1968, J-3.
Enteromorpha intestinalis (L.) Link.
JDR 1466, 10 Jun 1968, J-4.
Spongomorpha coalita (Ruprecht) Collins
JDR 1449, 10 Jun 1968, L-16.
Ulva angusta Setchell & Gardner
JDR 1471, 10 Jun 1968, J-3.
Ulva linza L.
JDR 1488, 10 Jun 1968, J-3.
Vha lobata (Kutzing) Setchell & Gardner
JDR 1472, 10 Jun 1968, J-3.
Division Phaeophyta— Brown Algae
Cystoseira osmundacea (Menzies) C.A.
Agardh
JDR 1531, 14 Jun 1968, C-13.
Desmarestia ligulata (Lightf.) Lamour
JDR 1456, 10 Jun 1968, N- 17.
December 1980
Ripley: Angel Island Plants
391
Ectocarpus acutus Setchell & Gardner
JDR 1456, 10 Jun 1968, J-3.
Egregia menziesii (Turn.) Aresch.
JDR 1483, 10 Jun 1968, J-3.
Fucus distichus L. ssp. cdcntatus (De la Py-
laise) Powell
JDR 1470, 10 Jun 1968, J-4.
Laminaria sinchiirii (Harvev) Farlow
JDR 1431, 10 Jun 1968,6-17.
Xcrcocystis luctkeana (Mertens) Ruprecht &
Postels
JDR 1532, 14 Jun 1968, K- 16.
Pelvctia fastigiata (J. G. A^ardli) De Toni
JDR 1477, 10 Jun 1968,^K-4.
Division Rhodophyta— Red Algae
Ahnfcltia pliaita (Hud.son) Fries.
JDR 1439a, 10 Jun 1968, 0-17.
AnfitluininioneUa glandulifera (Kvl.) Woll.
PR 1431, 10 Jun 1968,0-17. '
CaUithamnion pikcanum Harvey
JDR 1443, 10 Jun 1968, 0-17^
CaUoplu/Ilis obtusifolia J. G. Agardh
PR i437, 10 Jun 1968,0-17.
Ceramium eatonianum (Farlow) De Toni
JDR 1430, 10 Jun 1968, 0-17.
Ceramium gardneri Kvlin
JDR 1451, 10 Jun 1968,0-17.
Cnjptopleura violacea (J. G. Agardh) Kylin
JDR 1434, 10 Jun 1968, 0-17.
Cryptosiphonia woodii J. G. Agardh
JDR 1438, 10 Jun 1968, 0-17.
Endockidia murkata (Postels & Ruprecht) A.
G. Agardli
JDR 1480, 10 Jun 1968, L-5.
GelidUim coulteri Harvey
JDR 1440, 10 Jun 1968,0-17.
Gigartina agardhii Setchell & Gardner
JDR 1468, 10 Jun 1968, J-3.
Gigartina exasperata Harv. & Bail.
JDR 1478, 10 Jun 1968, J-3.
Gigartina canaliculata Harvey
JDR 1447, 10 Jun 1968, 0-i7.
Gigartina papillata (C. Ag.) J. Ag.
JDR 1479, 10 Jun 1968, J-4.
Gracihiriopsis sjocstcdtii (K\lin) Dawson
JDR 1460, 10 Jun 1968, k-16.
Grateloupia dorypfwra (Montagne) Howe
JDR 1491. lOJun 1968, E-8.
Gi/mnogongru.s Icptopln/llus J. G. .\gardh
JDR 1436, 10 Jun 1968,0-17.
Gymnogongnis linearis (Turner) J. G. Agardh
JDR 1446, 10 Jun 1968, N- 16.
Gymnogongnis plati/pliyllus Gardner
JDR 1432, 10 Jun 1968, O- 17.
Ihdosaccion glandiforme (Gmelin) Ruprecht
JDR 1530, 14 Jun 1968, K-4.
Leptocladia conferata Setchell
JDR 1464, 10 Jun 1968, N-17.
IJthophyllum decipiens (Foslie) Foslie
JDR 1490, 10 Jun 1968, J-4.
Iridaea heterocarpa Postels & Ruprecht
JDR 1445, 10 Jun 1968, 0-17.
Iridaea eordata S. & G. var. splendens (S. &
G.) Abb.
PR 1485, 10 Jun 1968. J-4.
Melobesia mediocris (Foslie) Setchell &: Ma-
zon
JDR 1428, 10 Jun 1968,0-17.
Microeladia borealis Ruprecht
JDR 1444, 10 Jun 1968, 0-17.
Microckidia coulteri Harvey
JDR 1453. 10 Jun 1968,0-17.
Odonthalia florcosa (Esper) Falkenberg
JDR 1433, 10 Jun 1968,0-17.
Petrocelis franciscana Setchell & Gardner
JDR 1487, 10 Jun 1968,0-17.
Pikea californica Harve\'
JDR 1441, 10 Jun 1968. O- 17.
Platyihamnion pectinatum Kylin
JDR 1462, 10 Jun 1968, J-3.
Polyneura latissima (Harvev) Kylin
JDR 1435, 10 Jun 1968,6-17.
Polysi})honia hcndn/i
JDR 1463, 10 Jun 1968, J-3.
Porphyra lanceolata (Setchell & Hus) G. .\1.
Smith
JDR 1482, 10 Jun 1968, J-4.
Pterosiphonia dcndroidca (Montagne) Falken-
berg
JDR 1439, 10 Jun 1968. O- 17.
392
Great Basin Naturalist
Vol. 40, No. 4
Ptilota filicina (Farlow) J. G. Agardh
JDR 1465, 10 Jun 1968, 0-17.
Rliodomeki larix (Turner) G. A. Agardli
JDR 1481, 10 Jun 1968, J-4.
Scfiizymenia pacifica Kylin
JDR 1457, 10 Jun 1968, 0-17.
Smithora naiadum (Anderson) Hollenberg
JDR 1427, 10 Jun 1968, 0-17.
Stenogramme interrupta (C. A. Agardli) Mon-
tague
JDR 1429, 10 Jun 1968, 0-17.
Division Calamophyta
Class Equisetinae
Equisetaceae
Equisetum hyemale L. var. affine (Engelm.)
A. A. Eat. PNF
JDR 703, 25 Jun 1967, L-16.
Equisetum telmateia Ehrh. var. braunii
Milde. PNF
JDR 386, 20 Apr 1967, L-7; JDR 625, 7
Jun 1967, K-8.
Division Pterophyta— Ferns
Class Filicinae
Dryopteridaceae
Dryopteris arguta (Kaulf.) Maxon. PNFr
PR 953, 27 Dec 1967, G-9.
Polystichum mimittim (Kaulf.) Presl. PNFr
JDR 563, 7 Jun 1967, 1-6; JDR 982, 6 Feb
1968, F-10.
Polypodiaceae
Polypodiiim californicum Kaulf. PNFr
JDR 564, 7 Jun 1967, E-7; JDR 981, 6 Feb
19681, F-10.
Pteridaceae
Adiantum jordanii C. Muell. PNFr
JDR 559, 2 May 1967, G-7; JDR 1191a, 11
Apr 1968, H-13; JDR 980, 6 Feb 1968, K-8.
Pellaea andromedifolia (Kaulf.) Fee. PNFr
JDR 1069, 31 Mar 1968, E-6; JDR 1176,
11 Apr 1968, K-9.
Pityrogramma triangularis (Kaulf.) Maxon.
PNFr
JDR 457, 28 Apr 1967, F-13; JDR 511, 2
May 1967, C-14; JDR 587, 7 Jun 1967, F-11.
Pteridiiim oquilimim (L.) Kuhn var. pub-
escens Underw. PNFr
JDR 565, 7 Jun 1967, F-7; JDR 588, 7 Jun
1967, E-12; JDR 1320, 12 Apr 1968, 1-9.
Division Coniferophyta
Araucariaceae
Araucaria bidwillii Hook. PCT
JDR 497, 2 May 1967, D-13.
Araucaria heterophylla (Salisb.) Franco.
PCT
JDR 499, 2 May 1967, D-13.
Cupressaceae
Chamaecyparis laivsoniana (A. Murr.) Pari.
PCT
JDR 749, 22 Sep 1967, F-14.
Cupressus macrocarpa Hartw. ex Gord. PIT
JDR 469, 28 Apr 1967, K-14.
Calocedrus dccurrens (Torr.) Florin. PCT
JDR 513, 2 May 1967, H-14.
Pinaceae
Abies concolor (Gord. & Glend.) Lindl. PCT
JDR 486, 2 May 1967, D-13.
Cedrus deodara (D. Don) G. Don. PCT
JDR 485, 2 May 1967, D-13; JDR 991, 6
Feb 1968, H-15.
Pinus monophxjUa Torr. & Frein. PCT
Observed by B, Jan 1980, F-8.
Pinus ponderosa Doiigl. ex P. & C. Lawson.
PCT
JDR 598, 7 Jun 1967, G-13.
Pinus radiata D. Don. PIT
JDR 470, 28 Apr 1967, J-9.
Pscudotsuga nienziesii (Mirbel) Franco. PCT
JDR 1365, 29 Apr 1968, J-13.
Taxaceae
Taxus baccata L. PCT
JDR 697, 25 Jun 1967, D13.
Taxodiaceae
Cryptonieria japonica D. Don. PCT
JDR 516, 2 May 1967, H-15.
December 1980
Ripley: Angel Island Plants
393
Seciiioia senipenirens (D. Don) Endl. PCT
JDR 568, 7 Jun 1967, 1-4; JDR 1251a, 12
Apr 1968, K-13.
Scijuoiadcndnm gigfintcum (Lindl.) Buchh.
PCT
JDR 514, 2 May 1967, J- 16.
Division Anthrophyta— Flowering Plants
Class Dicottjh'don cae
Aceraceae
Acer negundo L. ssp. ralifomicuni (T. & C.)
W'esniael. PCT
JDR 491, 2 Mav 1967, E-13; JDR 726, 25
Jun 1967, K-6.
Acer macrophijlhim Piirsh. PNT
JDR 597, 7 Jun 1967,0-12.
Aizoaceae
Mescmhrijanthemum chdense Mol. PNF
JDR 1533, 14 Jun 1968, J-4.
Mesemhnjandxeinwn ediile L. PIF
JDR 660, 20 Jun 1967, L- 16.
Mcscinhrijanthcmum floribiinduni Haw. PIF
JDR 535, 2 May 1967,1-17.
Tctragonia tetragonioides (Pall.) O. Kuntze.
PIF
JDR 715, 25 Jun 1967, K-4.
Anacardiaceae
Rhus divcrsdoha T. & C. PNS
JDR 410, 20 Apr 1967, 1-7; JDR 1224, 12
Apr 1968, L-6.
ScJiinus mollc L. PCT
JDR 515, 2 May 1967, H- 15.
Apocynaceae
Neriutn oleander L. PCS
JDR 756, 22 Sep 1967, E-14.
Vincd major L. PIF
JDR 441, 20 Apr 1967, J-5; JDR 1231, 12
Apr 1968, 1-4.
Aristolochiaeeae
Aristolochia californica Torr. PNF
K without number or date; R without num-
ber, 1877; JDR 812, 22 Sep 1967, F-9; JDR
1014, 2 Mav 1968, E-10.
Basellaceae
Andrcdera cordi folia (Ter.) Stennis. PCF
JDR 717, 22 Sep 1967,1-14.
Berberidaceae
Bcrbcris daruiiiii Hook. PCS
JDR 526, 2 May 1967, H-15.
Betulaceae
Corylus cornuta Marsh, var. californica (A.
DC.) Sharp. PNT
JDR 637, 20 Jun 1967, K-15; JDR 670a, 20
Jmi 1967, F-12; JDR 1364, 29 Apr 1968, H-
10.
Boraginaceae
Amsinckia intermedia F. & M. ANF
JDR 418, 20 Apr 1967. 1-5: JDR 1071. 31
Mar 1968, F-7.
Cynoglosstim grandc Dougl. ex Lehm. PNF
JDR 560, 2 Mav 1967, E-7; JDR 1013. 2
Mar 1968, F-11.
Echiiim fastuosum Jacq. PIS
JDR 385, 20 Apr 1967, J-4: JDR 1180. 11
Apr 1968, C- 11.
Myosotis latifolia Poir. PIF
JDR 524, 2 May 1967, E-14.
Callitrichaceae
Callitrichc marginata Torr. PNF
JDR 632, 20 Jun 1967, J-9.
Caprifoliaceae
Loniccra hispidula Dougl. var. lacillans
Crav. PNS
JDR 561, 7 Jun 1967, F-6.
Lonicera japonica Thunb. PIS
JDR 709, 25 Jun 1967, D-14.
Samhucus coerulca Raf. PNS
JDR 569, 7 Jun 1967, 1-6.
Symphoricarpos rivularis Suksdort. P.\S
JDR 763, 22 Sep 1967, E-Il.
Symphoricarpos mollis Nutt. PNS
R. 6c J. 21177,2 May 1967, J-12.
Car\'ophyIlaceae
Arcnaria douglasii Fen/.l ex T. 6i C. ANF"
JDR 401, 20 Apr 1967, Ml 6: JDH 595. 7
Jim 1967, E-12.
394
Great Basin Naturalist
Vol. 40, No. 4
Cerastium ghmeratum Thuill. AIF
JDR 1103, 11 Apr 1968, F-7; JDR 1271, 12
Apr 1968, E-9.
Pohjcarpon tetraphiiUum (L.) L. AIF
JDR 1409, 10 Jun 1968, 1-16.
Sagina apetala Ard. var. harbata Fenzl. AIF
R. & J. 21158, 2 May 1967, K-8.
Silene califomica Durand. PNF
JDR 1389, 8 Jun 1968, C-11.
Silene gallica L. AIF
JDR 423, 20 Apr 1967, 1-4; JDR 1149, 11
Apr 1968, J-14; JDR 1378, 29 Apr 1968, K-
15.
Spergularia bocconii (Scheele) Foucaud. AIF
JDR 1358, 29 Apr 1968, J-14.
Spergularia macrotheca (Hornem.) Heynh.
PNF
JDR 390, 20 Apr 1967, M-15; JDR 1311,
12 Apr 1968, K-9.
Spergularia rubra (L.) J. & C. Presl. PIF
JDR 647, 20 Jun 1967, H-13.
Spergularia villosa (Pers.) Canib. PIF
JDR 531, 2 May 1967, H-15; JDR 1108, 11
Apr 1968, F-7; JDR, 1051, 31 Mar 1968, G-8;
JDR 1221, 12 Apr 1968, K-6.
Stellaria media (L.) Vill. AIF
JDR 996, 6 Feb 1968, E-14; JDR 1017, 2
May 1968, F-10.
Casuarinaceae
Casuarina striata Dry. PCT
JDR 487, 2 May 1967, D-14.
Celastraceae
Maytenus boaria Mol. PCT
JDR 1407, 10 Jun 1968, G-14.
Chenopodiaceae
Atriplex leucophylla (Moq.) Dietr. PNF
Observed by H on 16 May 1946.
Chenopodium album L. AIF
JDR 667, 20 Jun 1967, L-6.
Chenopodium ambrosioides L. var. vagans
(Standi.) J. T. Howell. PIF
Observed by H on 16 May 1946.
Chenopodium californicum (Wats.) Wats.
PNF
JDR 1411, 10 Jun 1968, 0-16.
Chenopodium multifidum L. AIF
JDR 1395, 8 Jun 1968, N-16.
Salicornia virginica L. PNF
JDR 12.32, 12 Apr 1968, L-5.
Cistaceae
Helianthemum scoparium Nutt. var. vulgare
Jeps. PNF
JDR 1054, 31 Mar 1968, G-12.
Compositae
Achillea borealis Bong. ssp. califomica (Pol-
lard) Keck. PNF
JDR 408, 20 Apr 1967, 1-5; JDR 1115, 11
Apr 1968, J-5.
Achijrachaena mollis Schauer. ANF
Observed by H on 16 May 1946.
Agoseris apargioides (Less.) Greene. PNF
JDR 1297, 12 Apr 1968, H-8.
Agoseris grandiflora (Nutt.) Greene. PNF
JDR 580a, 7 Jun 1967, E-11.
Ambrosia chamissonis (Less.) Greene var. bi-
pinnatisecta J. T. Howell. PNF
JDR 664, 20 Jun 1967, K-5; JDR 720, 25
Jun 1967, N-15.
Anthemis cotula L. AIF
JDR 1706, 16 Jun 1969, E-12.
Artemisia califomica Less, in Hook. PNS
JDR 725, 25 Jun 1967, F-13; JDR 1505, 14
Jun 1968, L-5.
Artemisia douglasiana Bess. PNF
JDR 713, 25 Jun 1967, J-14.
Artemisia pijcnocephala DC. PNF
JDR 950, 27 Dec 1967, 1-9.
Aster chilensis Nees. PNF
JDR 814, 22 Sep 1967, J-9.
Baccharis pilularis DC. PNS
JDR 822, 22 Sep 1967, J-9.
Baccharis pilularis DC. var. consanguinea
(DC.) C. B. Wolf. PNS
JDR 708, 25 Jun 1967, H-7.
Bcllis perennis L. PNF
JDR 558, 2 May 1967, F-8.
Carduus pycnocephalus L. AIF
JDR 1373, 29 Apr 1968, J-12.
Carduus tenuiflorus Curt. AIF
JDR 521, 2 Mav 1967, L-16; JDR 438, 20
Apr 1967, J-8.
December 1980
Ripley: Angel Island Plants
395
Centaurea meUtensis L. AIF
JDR 656, 20 Jun 1967, J- 15; JDR 694, 25
fun 1967, D-14; JDR 1525, 14 Jun 1968, K-9.
Cirsiiun hrcvisti/lmu Cronq. PNF
JDR 623, 7 Jun 1967, K-9.
Cirsium occidentale (Nutt.) Jeps. P\F
JDR 696, 25 Jun 1967, D-13.
Cirsiimi piotcanuni J. T. Howell. P\F
R. & J. 21304, 2 May 1967, D-10.
Cirsium qucrcctorum (Grav) Jeps. P\F
JDR 616, 7 Jun 1967, K-6.
Cirsium rcmotifolium (Hook.) DC. P\F
H 21878, 16 May 1946, without location.
Cirsium vulgare (Savi) Ten. BIF
JDR 454, 28 Apr 1967, K-8.
Conijza bonariensis (L.) Cronq. AIF
Observed by H, 16 May 1946.
Comjza canadensis (L.) Cronq. AIF
Observed by H, 16 May 1946.
Cotula australis (Less.) Hook. AIF
JDR 435, 20 Apr 1967, 1-4; JDR 1074, 1 1
Apr 1968, F-8; JDR 1097, 11 Apr 1968, F-9.
Cotula coronopifulia L. PIF
JDR 432, 20 Apr 1967, 1-4.
Ci/nara scoh/mus L. PCS
JDR 807,22 Sep 1967, F-8.
Erechtites arguta (A. Rich.) DC. AIF
JDR 595a, 7 Jim 1967, G-13.
Erechtites prenanthoides (A. Rich.) DC. AIF
JDR 737, 22 Sep 1967, E-I4.
Erigeron glaucus Ker. PNF
JDR629, 7 Jun 1967, N- 17.
EriophiiUum confertiflorum (DC.) Grav.
PiNS
PR 589, 7 Jun 1967, J- 14.
EriopJu/lhim staechadifolium Lag. var. ar-
temisiifolium (Less.) Macbr. PNS
JDR 504, 2 May 1967, J-6.
Evax sparsiflora (Grav) Jeps. ANF
JDR 1353, 12 Apr 1968, K-9.
Fikigo gallica (L.) L. AIF
JDR 643, 20 Jun 1967, K-16; JDR 692, 25
Jun 1967, D-14.
Gnaphalium califurnicum D(>. BNF
JDR 574, 7 Jun 1967, H-6; JDR 1194, 12
Apr 1968, E-11.
Gnaphalium chilcnsc Spreng. B\F
JDR 673, 20 Jun 1967, K-15; JDR 1191, 11
Apr 1968, 1-I4; JDR 1399, 8 Jun 1968, H-16.
Gnaphalium luteo-alhum L. AIF
JDR 1192, 11 Apr 1968, 1-14; JDR 1094,
12 Apr 1968, G-7.
Gnaphalium purpureum L. BNF
JDR 1192a. 11 Apr 1968,1-14.
Grindclia campurum Cireene var. Daii/i
(Jeps.) Steverm. PNF
JDR588a, 7 Jun 1967, F- 13.
Grindelia hirsutula H. (be A. var. hrcvisquama
Steyerm. PNF
JDR 1337, 12 Apr 1968, 1-9.
Gutierrczia californica (DC.) T. & G. PNS
E without number, 3 Apr 1925; OS 2681,
Oct 1957; JDR 636, 20 Jun 1967, 1-6; JDR
716, 25 Jun 1967, J-7.
Haplopappus ericoides (Less.) Hook. 6c .Arn.
PNS
JDR 721, 25 Jun 1967, M-10.
Heterotheca bolanderi Grav. PNF
JDR 669, 20 Jun 1967, 'K-15; JDR 699, 25
Jun 1967, E-12.
Hijpochoeris glabra L. PIF
JDR 380, 20 Apr 1967, M-15; JDR 1233,
12 Apr 1968, L-5; JDR 1269, 12 Apr 1968, E-
9; JDR 1281, 12 Apr 1968, K-9.
Hijpochoeris radicata L. PIF
JDR 809, 22 Sep 1967, F-7; JDR 580, 7 Jun
1967, F-11.
Iva axillaris Pursh. PNF
JDR 661, 20 Jun 1967, L-15; JDR 765, 22
Sep 1967, K-9.
iMctuca saligna L. AIF
JDR 813, 22 Sep 1967, F-7.
Lasthenia chrysostoma (F. & W.) Greene.
ANF
JDR 1295, 12 Apr 1968. K-8.
Madia gracilis (Smith) Keck. PNF
JDR 553, 2 .May 1967, K-13.
Madia sativa .Mol. PNF
JDR 621, 7 Jun 1967, K-8.
Matricaria matricarioidcs (Less.) Porter. \\Y
JDR 435a, 20 April 1967, E-13; JDK 1067,
11 Apr 1968, G-10; IDH 1.357. 29 Apr 1968,
1-8.
396
Great Basin Naturalist
Vol. 40, No. 4
Micropus califomiciis F. & M. ANF
JDR 1225, 12 Apr 1968, J-5; JDR 1344, 12
Apr 1968, K-9.
Microseris higelovii (Gray) Sch.-Bip. ANF
H 21859, 16 May 1946.
Microseris douglasii (DC.) Sch.-Bip. ANF
JDR 1347, 12 Apr 1968, K-9.
Microseris heierocarpa (Niitt.) Chamb. ANF
JDR 1349, 12 Apr 1968, K-9.
Microseris linearifolia (Nutt.) Sch.-Bip. ANF
JDR 1310, 12 Apr 1968, K-9.
Pentachaeta alsinoides Greene. ANF
H 21865, 16 May 1946.
Picris echioides L. BIF
R. & J. 21032, 2 May 1967, K-9.
Psilocarphus teneUiis Nutt. ANF
Observed by H, 16 May 1946.
Rafinesqiiia californica Nutt. ANF
Observed by H, 16 May 1946.
Senecio aronicoides DC. PNF
E without number, 3 Apr 1925; JDR 466,
28 Apr 1967, E-11; JDR 1030, 2 Mar 1968,
G-13.
Senecio rnikanioides Otto. PIF
JDR 756a, 22 Sep 1968, K-16.
Senecio sylvaticiis L. AIF
JDR707, 25Jun 1967, L- 15.
Senecio vulgaris L. AIF
JDR 433, 20 Apr 1967, 1-4; JDR 1336, 12
Apr 1968, 1-8.
Silybum marianum (L.) Gaertn. BIF
JDR 581a, 7 Jun 1967, F-11; JDR 623, 7
Jun 1967, L-16; JDR 949, 27 Dec 1967, F-8.
Solidago californica Nutt. PNF
JDR 752, 22 Sep 1967, L-15.
SoHva daucifolia Nutt. ANF
JDR 663, 20 Jun 1967, L-7.
Soliva pterospertna (Juss.) Less. ANF
JDR 1096, 11 Apr 1968, J-5; JDR 1304, 12
Apr 1968, K-8.
Soliva sessilis R. & P. ANF
D 80-39, 29 May 1980, F-8.
Sonchus asper (L.) Hill. AIF
JDR 1.388, 29 Apr 1968, 1-6; JDR 1414, 10
Jun 1968, N-17.
Stephanonieria virgata Benth. ANF
Observed by H, 16 May 1946.
Stylocline amphihola (Gray) J. T. Howell.
ANF
R. & J. 21328, 2 May 1967, K-16.
Taraxacum officinale Weber in Wiggers.
PIF
JDR 508, 2 Mav 1967, D-14; JDR 1092, 11
Apr 1968, J-4.
Tragopogon porrifolius L. BIF
JDR 1423, 10 Jun 1968, H-16.
Wyethia angustifolia Nutt. PNF
JDR 594, 7 Jun 1967, H-7; JDR 1287, 12
Apr 1968, K-8.
Xanthium spinosum L. AIF
JDR 755, 22 Sep 1968, K-16.
Convolvulaceae
Calystegia polymorpha (Greene) Munz. PNF
Observed by H, 16 May 1946.
Calystegia purpurata (Greene) Bnnnmitt.
PNF
JDR 450, 28 Apr 1967, G-13; JDR 624, 7
Jun 1967, K-7; JDR 1117, 11 Apr 1968, J-9.
Calystegia suhacaulis H. & A. PNF
JDR 1305, 12 Apr 1968, K-8.
Convolvulus arvensis L. PIF
Observed by H, 16 May 1946.
Dichondra donneUiana Tharp & Johnston.
PNF
JDR 1103a, 11 Apr 1968, F-7; JDR 1187a,
11 Apr 1968,1-14.
Crassulaceae
Dudleya farinosa (Lindl.) Britt. & Rose.
PNF
R without number, 26 Mav 1877; JDR 570,
7 Jun 1967, J-4.
Sedum spathulifolium Hook. PNF
JDR 505, 2 May 1967, J-3.
Tillaea erecta H. & A. ANF
JDR 1037, 2 Mar 1968, K-8; JDR 1236, 12
Apr 1968, K-5.
Cruciferae
Arabis glabra (L.) Bernh. BNF
JDR 1302, 12 Apr 1968. K-8.
Brassica campestris L. AIF
JDR 1333, 12 Apr 1968, G-7.
Brassica geniculata (Desf.) J. Ball. BIF
JDR 626, 7 Jun 1967, 1-7; JDR 631, 20 Jun
1967, E-12.
December 1980
Ripley: Angel Island Plants
39-
Cakile maritima Scop. AIF
JDR 431, 20 Apr 1967, K-5; JDR 634, 20
Jun 1967, L-16.
CapscUa hursa-pastoris (L.) Medic. AIF
JDR 1129, 11 Apr 1968, F-8; JDR 1173, 11
Apr 1968, G-7.
Cardamine oligospenna Nutt. BNF
JDR 1012, 2 Mar 1968, E-10; JDR 1120,
11 Apr 1968, E-6.
Coronopus didymus (L.) Smith. BIF
Observed by H, 16 May 1946.
Dentaria integrifolia Nutt. var. californica
(Nutt.)Jeps. PNF
JDR 414, 20 Apr 1967, J-4; JDR 979, 6 Feb
1968, E-10.
Enjshmim capitatiun (Dougl.) Greene. BNF
JDR 502, 2 May 1967, D- 14.
Lepidium nitidum Nutt. ANF
JDR 1036, 2 May 1968, J-7; JDR 1518, 14
Jun 1968, K-9.
Lepidium strictum (Wats.) Rattan. AIF
D 80-44, 29 May 1980, F-9.
Lobidaria maritima (L.) Desv. PIF
JDR 409, 20 Apr 1967, J-5.
Raplianus sativus L. BIF
JDR 510, 2 May 1967, D-14.
Rorippa nasturtium-aquaticum (L.) Britt. &
Rendl. PIF
JDR 735, 22 Sep 1967, L-9; JDR 1415, 10
Jun 1968, N-16.
Thehjpodium lasiophyllum (H. & A.)
Greene. ANF
JDR 1503, 14 Jun 1968, K-9; JDR 1529, 14
Jun 1968, E-6.
Thysanocarpus curvipes Hook. ANF
JDR 1207, 12 Apr 196, E-6.
Cucurbitaceae
Marah fabaceus (Naud.) Dunn. PNF
JDR 389, 20 Apr 1967, L-6.
Marah oregonus (T. & G.) Howell. PNF
JDR 1330, 12 Apr 1968, 1-9.
Ericaceae
Arbutus rtienziesii Pursh. PNT
JDR 405, 20 Apr 1967, D-12; JDR 818, 22
Sep 1967, E-11.
Arctostaphylo.s glandulosa Eastw. var. Cush-
ingiana (Eastw.) .Adams ex McMinn. PNS
JDR 978, 6 Feb 1968, E-11.
Euphorbiaceae
Euphorbia pephis L. AIF
JDR 731, 22 Sep 1967, H-15.
Euphorbia spathulata Lamk. ANF
JDR 1351, 12 Apr 1968, K-9; JDR 1.384, 29
Apr 1968, K-15; JDR 1417. 10 Jun 1968, \-
17.
Fagaceae
Quercus agrifolia Nee. PNT
JDR 423, 20 Apr 1967, 1-4.
Quercus wislizeni A. DC. var. frutescens
Engelm. PNT
JDR 977, 6 Feb 1968, D- 11.
Quercus suber L. PCT
Observed by B, Jan 1980, K-14 and L-8.
Garryaceae
Garrya eUiptica Dougl. PNS
JE)R 566a, 7 Jun 1967, E-7.
Gentianaceae
Centauriitm muhlcnbergii (Griseb.) W.
Wight. ANF
H 21857, 16 May 1946.
Geraniaceae
Erodium botrys (Gav.) Bertol. AIF
JDR 1516, 14 Jim 1968, K-9.
Erodium cicutarium (L.) L'Her. .\IF
JDR .396, 20 Apr 1967, N-16; JDR 811, 22
Sep 1967, G-9; JDR 984, 6 Feb 1968, E-11.
Erodium moschatum (L.) L Her. .\IF
JDR 1170, 11 Apr 1968, G-7; JDR 1413, 10
Jun 1968, N-16.
Geranium carolinianum L. ANF
JDR 1106a, 11 Apr 1968, G-7.
Geranium dissectum L. .-MF
JDR 417, 20 Apr 1967, 1-5.
Geranium molle L. BIF
JDR 520, 2 May 1967, H-16; JDR 537, 2
May 1967, C-11.
Pelargonium zonale (L.) L'Her. PGS
JDR .530. 2 May 1967,1-16.
398
Great Basin Naturalist
Vol. 40, No. 4
Hippocastanaceae
Aesculus californica (Spach) Nutt. PNT
JDR602, 7Jiin 1967, H- 14.
Hydrophyllaceae
Eucnjpta chrifsanthetnifolia (Benth.)
Greene. ANF
TB without number or date.
Nemophila heterophyUa F. & M. ANF
JDR 1217, 12 Apr 1968, F-6.
Phacelia californica Cham. PNF
JDR 437, 20 Apr 1967, J-6; JDR 1158, 11
Apr 1968, F-12.
Phacelia malvifolia Cham. ANF
JDR 1393, 29 Apr 1968, C-11.
Phacelia nenioralis Greene. BNF
H 21892, 16 May 1946.
Pholistoma aiiritum (Lindl.) Lilja. ANF
Observed by H, 16 May 1946.
Hyperieaceae
Hypericum caneriensis L. PCS
JDR 1408, 10 Jun 1968, K-15.
Juglandaceae
Juglans hindsii (Jeps.) Jeps. PCX
JDR 730, 25 Jun 1967, K-15.
Labiatae
Lepechinia cahjcina (Benth.) Epl. PNS
JDR 472, 28 Apr 1967, H-8; JDR 583, 7
Jun 1967, K-7.
Mentha pulegium L. PIF
Observed by B, 1980, E- 11.
Monardella villosa Benth. var. franciscana
(Ehner)Jeps. PNF
JDR 615, 7 Jun 1967, K-7.
Rosmarinus officinalis L. PCS
JDR 1140, 11 Apr 1968, F-8.
Salvia leucantha Cav. PCS
JDR 962, 27 Dec 1967, J-6.
Stachys rigida Nutt. var. quercetorum (Hel.)
Epl. ANF
JDR 416, 20 Apr 1967, J-3; JDR 545, 2
May 1967, D-12; JDR 1109, 11 Apr 1968, G-
7.
Lauraceae
Umbellularia californica (H. & A.) Nutt.
PNT
JDR 413, 20 Apr 1967, 1-4; JDR 1019, 2
May 1968, F-10.
Leguminosae
Acacia deciirrens (Wendl.) Willd. PIT
JDR 494, 2 May 1967, E-13; JDR 1043, 2
May 1968, F-7.
Acacia longifolia Willd. PIT
Observed iDy B, 1980, H-15.
Albizia lophatha (Willd.) Benth. PIT
JDR 960, 6 Feb 1968, L-6.
Acacia melanoxylon R. Br. PIT
JDR 442, 20 Apr 1967, K-6.
Acacia verticillata (L'Her.) Willd. PCT
JDR 523, 2 May 1967, E-14.
Astragalus gambellianus Sheld. ANF
JDR 1511, 14 Jun 1968, K-9; JDR 1315, 12
Apr 1968, M-16.
Astragalus nuttalli (T. & G.) J. T. Howell var.
virgatus (Gray) Barneby. PNF
S without number, May 1920; H 21869, 16
May 1946.
Cytisus monspessulanus L. PIS
JDR 961, 27 Dec 1967, K-14.
Cytisus multiflorus (L'Her.) Sweet. PCS
JDR 1142, 11 Apr 1968,1-15.
Cytisus scoparius (L.) Link. PIS
JDR 542, 2 May 1967, C-10; JDR 1214, 12
Apr 1968, J-7; JDR 1334, 12 Apr 1968, MO.
Cytisus scoparius (L.) Link. var. andrianus
(Puiss.) Dippel. PIS
D 80-42, 29 May 1980, F-7.
Lathyrus latifolius L. PIF
JDR 1392, 8 Jun 1968, L-15.
Lathyrus vestitus Nutt. ex Torr. & Grav ssp.
Bolanderi (Wats.) C. L. Hitchc. PNF
JDR 612, 7 Jun 1967, F-12; JDR 1059, 31
Mar 1968, D-15; JDR 1133, 11 Apr 1968, E-
11.
Lotus scoparius (Nutt.) Ottlev. PNF
JDR 578, 7 Jun 1967, E-12.
Lotus strigosus (Nutt.) Greene. ANF
H 21869, 16 May 1946.
December 1980
Ripley: Angel Island Plants
399
Lotus siihpinnatus Lag. ANF
JDR 556, 2 Mav 1967, K-13; JDR 1223, 12
Apr 1968, L-6; JDR 1299, 12 Apr 1968, K-9;
JDR 1340, 12 Apr 1968, G-12.
Lupinus albifwns Benth. var. collinu.'i
Greene. PNF"
H 21873, 16 May 1946.
Lupinus arborcus Sims. PNS
JDR 603, 7 Jmi 1967, E-12; JDR 536, 2
May 1967, C-11.
Lupinus hicolor Lindl. var. umhcllatus
(Greene) C. P. Sm. ANF
JDR 379, 20 Apr 1967, N-15; JDR 1370, 29
Apr 1968, J- 11.
Lupinus densiflorus Benth. ANF
H 21863, 16 May 1946.
Lupinus fonnosus Greene. PNF
JDR 459, 28 Apr 1967, 1-9.
Lupinus latif alius Agardh. PNF
JDR 575, 7 Jun 1967, J-8.
Lupinus poll/corpus Greene. ANF
JDR 1510", 14 Jun 1968. K-9.
Lupinus rivularis Dougl. PNF
JDR 604, 7 Jun 1967, 1-7.
Lupinus succulentus Dougl. ANF
Observed by H, 16 May 1946.
Medicago arahica (L.) All. AIF
JDR 393, 20 Apr 1967, M-16; JDR 1376,
29 Apr 1968, H-14.
Medicago pohjmorpha L. AIF
JDR 1179, 11 Apr 1968, F-15; JDR 1335,
12 Apr 1968, I-IO; JDR 1521, 14 Jun 1968.
K-9.
Medicago poli/inorpha L. var. l)revispina
(Benth.) Hevn." AIF
JDR 1175, 11 Apr 1968, G-7; JDR 1247, 12
Apr 1968, E-10; JDR 1332, 12 Apr 1968, 1-8.
Melilotus indicus (L.) All. BIF
JDR 704, 25 Jun 1967, K-14; JDR 1401, 10
Jun 1968, J-16; JDR 1416, 10 Jun 1968, M-16.
Rohinid pseudo-acacia L. PCT
JDR 567, 7 Jun 1967, 1-6.
Sophora micropIu/Ua Aiton. P(>T
JDR 496, 2 May 1967, D- 14.
Thennopsis macrophijUa H. & A. PNF
JDR 1.387, 29 Apr" 1968, 1-6.
Trifoliwn amplectens T. & G. ANF
JDR 1296, 12 Apr 1968, K-9.
TrifoHum alhopurpurcum T. & G. .\NF
JDR 1293, 12 Apr 1968, K-9.
Trifoliutn hifidum Grav. .\NF
JDR 1151, 11 Apr 1968, H-13.
Trifoliuni duhiuin Sibth. .\IF
JDR 1080, 11 Apr 1968, G-13; JDR 1377,
29 Apr 1968, J-15; JDR 1418, 10 Jun 1968,
N-16.
Trifoliuni fucatuni Lindl. .\.\F
Observed by H, 16 May 1946.
TrifoHum gracilentum T. & G. .\NF
JDR 1135, 11 Apr 1968, F-11; JDR 1200,
12 Apr 1968, 1-14; JDR 1354, 12 Apr 1968,
K-9.
Trifoliuni macraci H. & .\. ANF
JDR 461, 28 Apr 1967, KIO; JDR 1316, 12
Apr 1968, K-8.
Trifoliuni niicroccphc.hnn Pursh. .ANF
JDR 1216, 12 Apr 1968, J-7; JDR 1265, 12
Apr 1968, J-5; JDR 1276. 12 Apr 1968. K-9;
JDR 1339, 12 Apr 1968, 1-9.
TrifoHum microdon H. & A. ANF
JDR 552, 2 May 1967, K-14; JDK 1121. II
Apr 1968, F-12.
TrifoHum repcns L. PIF
JDR 671, 20 Jun 1967, K-14.
TrifoHum tridentatum Lindl. .\NF
JDR 423a, 20 Apr 1967, J-5; JDR 1 166. 11
Apr 1968, 1-9; JDR 1254, 12 Apr 1968, D-IO.
TrifoHum variegatum Nutt. .\NF
JDR 391, 20 Apr 1967. M-15: JDR 555. 2
May 1967, C-11.
Vicia americana Muhl. var. orcgana (Nutt.)
Nels. PNF
R. & J. 21333, 2 May 1967, E-12.
Vicia angustifolia L. .\IF
JDR 1215. 12 Apr 1968. J-7.
Vicia cxigua Nutt. ANF
JDR 11.34, 11 Apr 1968, F-13; JDR 1250,
12 Apr 1968, D-10.
V'irjV; sativa L. .\IF
JDR 546, 2 May 1967. C-11.
Lvthraceae
Lijtlirum hyssopifoHa L. PNF
JDR 768, 22 Sep 1967, H-9; JDR 1523, 14
Jun 1968, K-9.
400
Great Basin Naturalist
Vol. 40, No. 4
Magnoliaceae
Magnolia grandi flora L. PCT
JDR 805, 22 Sep 1967, J-6.
Malvaceae
Lavatera cretica L. AIF
H 21899, 16 May 1946, H-16.
Malva parviflora L. AIF
JDR 668, 20 Jun 1967, 1-6.
Sidalcea malviflora (DC.) Gray. PNF
JDR 421, 20 Apr 1967, 1-4; JDR 1270, 12
Apr 1968, C-12.
Moraceae
Ficus carica L. PCT
PR 742, 22 Sep 1967, D-14.
Myoporaceae
Mijoporwn laetum Forst. PCS
Observed by B, 1980, E-9.
Myricaceae
Myrica califomica C. & S. PNT
JDR 1679, 16 May 1969, G-15.
Myrtaceae
Eucalyptus amygdalina Labill. PCT
JDR 488, 2 May 1967, D-13; JDR 702, 25
Jun 1967, 1-14.
Eucalyptus cornuta Labill. PCT
JDR 754, 22 Sep 1967, K-13.
Eucalyptus eugenioides Sieb. PCT
JDR 638, 20 Jun 1967,1-14.
Eucalyptus ficifolia F. J. Muell. PCT
JDR 744, 22 Sep 1967, D-13.
Eucalyptus globulus Labill. PIT
JDR 610, 7 Jun 1967, 1-15.
Eucalyptus goniocalyx F. J. Muell. PCT
JDR 525, 2 May 1967, D-13.
Eucalyptus leucoxylon F. J. Muell. PCT
JDR 819, 22 Sep 1967, 1-15.
Eucalyptus polyanthemos Schau. PCT
JDR 823, 22 Sep 1967, 1-14.
Eucalyptus pulverulenta Sims. PCT
JDR 714, 25 Jun 1967,1-16.
Melaleuca nesophila F. J. Muell. PCS
JDR 748, 22 Sep 1967, D-14.
Syzygium paniculatum Gaertner. PCT
JDR 729, 25 Jun 1967, G-15.
Nyctaginaceae
Ahronia htifolia Esch. PNF
JDR 534, 2 May 1967, J- 16.
Abronia umbellata Lamk. PNF
JDR 533, 2 May 1967, J-16.
Bougainvillea spectabilis Willd. PCS
JDR 723, 25 Jun 1967, 1-15.
Oleaceae
Ligustrum ovalifolium Hassk. PCS
JDR 609, 7 Jun 1967,1-16.
Syringa vulgaris L. PCS
JDR 495, 2 May 1967, E-14.
Onagraceae
Clarkia amoena (Lehm.) Nels. & Macbr.
ANF
JDR605, 7 Jun 1967, G- 14.
Clarkia concinna (F. & M.) Greene. ANF
JDR586, 7 Jun 1967, E-11.
Clarkia unguiculata Lindl. ANF
JDR 760, 22 Sep 1967, J-15.
Epilobium adenocaulon Hausskn. var. Parishii
(Trel.) Munz. PNF
JDR 1524, 14 Jun 1968, K-9.
Epilobium brachycarpum Presl. ANF
R. & J. 21064, 2 May 1967, K-9.
Epilobium minutum Lindl. ANF
JDR 1356, 12 Apr 1968, K-9.
Epilobium watsonii Barbey var. francisca-
num (Barbey) Jeps. PNF
Observed by H, 16 May 1946.
Fuchsia magellanica Lam. PCS
JDR 600, 20 Jun 1967, H-15.
Oenothera ovata Nutt. PNF
JDR 458, 28 Apr 1967, J-9.
Zauschneria califomica Presl. PNF
JDR 718, 22 Sep 1967, J-6; JDR 946, 27
Dec 1967, K-14.
Orobanchaceae
Orobanche fasciculata Nutt. var. franciscana
Achey. ANF
JDR 1703, 20 Apr 1969, K-8.
Oxalidaceae
Oxalis pes-caprae L. PIF
JDR 403, 20 Apr 1967, J-16; JDR 1169, 11
Apr 1968, F-8.
December 1980
Ripley: Angel Island Plants
401
Oxalis pilosa Nutt. PNF
JDR995, 6Feb 1968, D-13.
Oxalis rubra St. Hil. PIF
JDR 404, 20 Apr 1967, J- 16.
Papaveraceae
Eschscholzia caUfornica Cham. ANF
JDR .383, 20 Apr 1967. M-14; JDR 601, 7
Jun 1967, H-14; JDR 1078, 11 Apr 1968, F-
14; JDR 1182, 11 Apr 1968, J-15.
Stylomecon heterophylla (Benth.) G. Tavl.
ANF
JDR 1068, 11 Apr 1968, E-7.
Pittosporaceae
Pittosponini crassifoUiim Banks & Soland.
PCX
JDR 408, 20 Apr 1967, D-14.
Pittosponini eugenioidcs A. Cunn. PCT
JDR 490, 2 May 1967, J-6.
Pittosporum rhomhifolium A. Cunn. PCT
JDR 743, 22 Sep 1967, D-14.
Pittosporum undulatwn Vent. PCT
JDR 449, 20 Apr 1967. K-6.
Pittosporum viridiflorum Sims. PCT
JDR 712. 25 Jmi 1967, K-6.
Plantaginaceae
Plantago hookeriana F. & M. var. caUfornica
(Greene) Poe. ANF
JDR 1034, 31 Mar 1968, H-12; JDR 1161.
11 Apr 1968, G- 13.
Plantago lanceolata L. PIF
JDR 607, 7 ]nn 1967, G-I3; JDR 1193, 11
Apr 1968, F-14; JDR 1227, 12 Apr 1968, K-6.
Polemoniaceae
Cilia achilleijolia Benth. ssp. multicaulis
(Benth.) V. & A. Grant. ANF
JDR 1089, 11 Apr 1968, F-6.
Cilia capitata Dougl. ANF
JDR 382, 20 Apr 1967, M-14.
Cilia clivorum (Jeps.) V. Grant. ANF
JDR 1355, 12 Apr 1968, K-9.
Navarretia squarrosa (Esch.) H. & A. ANF
JDR 624, 7 Jun 1967, K-14.
Polemonium carneum Gray. PNF
V without number, 1876.
Folygalaceae
Poll/gala \ dalmaisina Baile\ . PCS
JDR 1406, 10 Jun 1968, E-13.
Polygonaceae
Eriogonum latifoliuin Smith in Rees. PNF
JDR 654, 2 May 1967, F-7.
Poli/gonum avicularc L. ,\IF
JDR 764, 22 Sep 1967, F- 12.
Pterostegia drymarioides F. 6c .M. .\NF
JDR 1038, 2 May 1968, K-9.
Rumex acetosella L. PIF
JDR 635, 20 Jun 1967, L-16; JDR 666. 20
Jun 1967, M-16; JDR 1199, 12 Apr 1968, H-
14.
Rumex crispus L. PIF
JDR 665, 20 Jun 1967, L-16.
Rumex pulclwr L. PIF
JDR 766, 22 Sep 1967, E-12.
Portulacaceae
Calandrinia ciliata (R. 6c P.) DC. var. men-
ziesii (Hook.) Macbr. ANF
JDR 1055, 31 Mar 1968, E-12.
Montia perfoliata (Donn) Howell. .\NF
JDR 388, 20 Apr 1967, L-5; JDR 1015, 2
Mar 1968. E-10.
Primulaceae
Anagallis arvensis L. .\IF
JDR 462. 28 Apr 1967, F-7.
Anagallis arvensis L. forma caerulea
(Schreb.) Baiung. .\IF
Observed by R. & J., 2 May 1967.
Dodecatheon hendersonii Grav. PNF
JDR 1024, 2 Mav 1968, E-'lO; JDR 1031. 2
May 1968, G-I2.
Proteaceae
Grevillea robusta \. Cunn. PCT
JDR 7.39. 22 .Sep 1967, D-14.
Ranunculaceae
Delphinium califoniicum T. & G. PNF
JDR 613, 7 Jun 1967, J-9.
Ranunculus califomicus Benth. PNF
JDR 415, 20 Apr 1967. 1-4: JDR 10.32, 2
Mav 1968, 1-9.
402
Great Basin Naturalist
Vol. 40, No. 4
Ranunculus niuricatus L. AIF
JDR 658, 20 Jim 1967, J-9; JDR 1705, 20
Apr 1969, F-7.
Thalictrum pohjcarpum (Torr.) Wats. PNF
JDR 1057, 31 Mar 1968, E-10; JDR 1184,
11 Apr 1968, G-12.
Rhamnaceae
Ceanothus thyrsiflorus Esch. PNS
JDR 406, 20 Apr 1967, D-13; JDR 706, 25
Jun 1967, 1-4.
Rhamnus crocea Nutt. PNS
JDR 948, 6 Feb 1968, J-8; JDR 983, 6 Feb
1968, E-12.
Rosaceae
Acaena californica Bitt. PNF
R. & J. 21187, 2 May 1967, J-11.
Adenostoma fascicuhtum H. & A. PNS
JDR 569, 7 Jun 1967, E-12.
Cotoneaster franchetti Bois. PCS
JDR 483, 2 May 1967, E-14.
Cotoneaster pannosus Franch. PIS
D 79-40, 6 July 1979, G-8.
Crataegus monogyna Jacq. PCT
JDR 492, 2 May 1967, E-14.
Eriobotnja japonica (Thunb.) Lindl. PCT
JDR 7.38, 22 Sep 1967, E-14.
Heteromeles arhutifoUa (Ait.) M. Roem. PNS
JDR 492a, 2 May 1967, G-14; JDR 611, 7
Jun 1967, K-15; JDR 951, 27 Dec 1967, L-15.
Holodiscus discolor (Pursh) Maxim. PNS
JDR 562, 7 Jun 1967, G-8; JDR 701, 25 Jun
1967, D-10.
Horkelia californica Cham. & Schlecht. PNF
R. & J. 21194, 2 May 1967, H-9.
Mains sijlvestris Mill. PCT
JDR 493, 2 May 1967, E-14.
Oernleria cerasifortnis (T. & G.) Landon.
PNS
JDR 412, 20 Apr 1967, G-7; JDR 1683, 30
Dec 1968, L-7.
Potentilla glandulosa Lindl. PNF
Observed by H, 16 May 1946.
Prunus amieniaca L. PCT
JDR 754, 22 Sep 1967, E- 13.
Prunus avium L. PCT
JDR 566, 7 Jun 1967, F-8.
Prunus cerasifera Ehrh. 'atropurpurea'. PCT
JDR 484, 2 May 1967, E-13.
Pyracantha angustifolia (Franch.) Schneid.
PCS
JDR 946, 27 Dec 1967, K-15.
Pyrus communis L. PCT
JDR 1374, 29 Apr 1968, K-15.
Rosa californica C. & S. PNS
JDR 1390, 8 Jun 1968, 1-16.
Rosa gymnocarpa Nutt. PNS
JDR 576, 7 Jun 1967, J-4.
Rubus parviflorus Nutt. var. velutinus (H. &
A.) Greene. PNS
JDR 590, 7 Jun 1967, F-U; JDR 1114, 11
Apr 1968, 1-8.
Rubus procerus P. J. Muell. PNS
JDR 1070, 11 Apr 1968, F-7.
Rubus ursinus C. & S. PNS
JDR 453, 28 Apr 1967, L-7; JDR 1072, 11
Apr 1968, G-7.
Rubiaceae
Coprosma repens A. Rich. PCS
JDR 740, 22 Sep 1967, D-13.
Galium aparine L. AIF
JDR 1264, 12 Apr 1968, C-12; JDR 1286,
12 Apr 1968, K-9.
Galium nuttallii Grav. PNF
JDR 463. 28 Apr 1967, F-11; JDR 1267, 12
Apr 1968, D-12; JDR 1298, 12 Apr 1968, K-
9; JDR 1329, 12 Apr 1968, G-7.
Sherardia arvensis L. AIF
JDR 1421, 10 Jun 1968, F-8; JDR 1106, 11
Apr 1968, G-7.
Salicaceae
Populus alba L. PCT
JDR 473, 28 Apr 1967, J-6.
Populus nigra L. var. italica Du Roi. PCT
JDR 734, 22 Sep 1967, J-5.
Salix babylonica L. PCT
JDR 557, 2 May 1967, F-8.
Salix lasiolepis Benth. PNT
JDR 719, 22 Sep 1967, L-11; JDR 987, 6
Feb 1968, J-15; JDR 988. 6 Feb 1968, G-15.
Salix lasiolepis Benth. var. bigelovii (Torr.)
Bebb. PNT
JDR 989, 6 Feb 1968, L-8.
December 1980
Ripley: Angel Island Plants
403
Saxifragaceae
Heuchera niicranlha Doiigl. PNF
JDR 577, 7 Jun 1967, H-7; JDR 1528, 14
Jun 1968, E-7.
Lithophra^nia hctvwplujUum (H. & A.) T. &
G. PNF
JDR 547, 2 Mav 1967, D-11; JDR 1058, 31
Mar 1968, F-IO.
Rihes cahfornicuin (H. 6c A.) PNS
JDR 1243, 12 Apr 1968, L-6.
Saxifrag^a califomica Greene. PNS
JDR 1016,2 May 1968, F- 10.
Scrophulariaceae
Castilleja affinis H. & A. PNF
JDR593", 7 Jun 1967, F-10.
Castilleja foliolosa H. & A. PNF
JDR 1052,31 Mar 1968, E-11.
Castilleja franciscana Penn. PNF
R. & J. 21310, 2 May 1967, D-10.
Castilleja latifolia H. & A. PNF
JDR 1150, 11 Apr 1968, J-15; JDR 1258,
12 Apr 1968, D-11; JDR 1322, 12 Apr 1968,
G-12.
CoUinsia heterophylla Buist ex Grah. ANF
JDR 538, 2 May 1967, D-11; JDR 1212, 12
Apr 1968, G-7; JDR 1346, 12 Apr 1968, K-9.
CoUinsia multicolor Lindl. & Paxt. ANF
R. & J. 21312, 2 May 1967, D-10.
Linaria canadensis (L.) Dum. -Coins, var. tex-
ana (Scheele) Penn. BNF
JDR 381, 20 Apr 1967, M-12.
Linaria cipnbalaria (L.) Mill. PIF
JDR 1502, 14 Jun 1968, L-5.
Mimuhts aurantiacus Curt. PNS
JDR 468, 28 Apr 1967, K-15; JDR 461, 28
Apr 1967, C- 11.
Mimnlus guttatus Fisch. ex DC. PNF
JDR 655, 20 Jun 1967, K-16.
Mitnulus guttatus Fisch. ex DC. var. arvensis
(Greene) Munz. PNF
JDR 1400, 10 Jun 1968, N- 15.
Orthocarf)us attenuatus Gray. ANF
JDR 1322, 12 Apr 1968, 1-8.
Otihocarpus densiflorns Benth. ANF
JDR 456, 28 Apr 1967, F-12; JDR 584, 7
Jun 1967, J-14; JDR 1244, 12 Apr 1968, D-
13.
Orthocarpus pusillus Benth. .\NF
JDR 426, 20 Apr 1967, 1-4; JDR 1147, 11
Apr 1968, F-8; JDR 1245, 12 Apr 1968, D-
10.
Scropliularia califomica ChauL PNF
JDR 591, 7 Jun 1967, E-11.
Veronica arvensis L. .AIF
JDR 1098, 11 Apr 1968, F-8.
Simarubaceae
Ailanthus altissinia (Mill.) Swingle. PCX
JDR 648, 20 Jun 1967, L-16.
Solanaceae
Nicotiana glauca Grah. PCS
Observed by B, 1980,1-15.
Solanum nigriwi L. AIF
Observed by H, 16 .May 1946.
Tropaeolaceae
Tropaeoluui niajiis L. .\IF
JDR 1424. 10 Jun 1968, N-16.
Umbelliferae
Angelica tomentosa Wats. PNF
Observed by H, 16 May 1946.
Daucus ptisilhis Michx. ANF
JDR 644, 7 Jun 1967, F-7.
Foeniculum vulgare Mill. PNF
JDR 722, 25 Jun 1967. E- 13.
Ileracleum lanatum Michx. PNF
JDR 503, 2 May 1967, D-12; JDR 1209, 12
Apr 1968, F-7.
Lomatiiim dusycarpuni (T. ik G.) C. & R.
PNF
JDR 467, 28 Apr 1967, 1-8.
Osniorhiza chilensis H. & A. PNF
JDR 617, 7 Jun 1967, J-8.
Perideridia kelloggii (Grav) Mathias. PNF
R. & J. 21 123, 2 May i967, K-9.
Sanicula hipinnatifida Dougl. P.NF
JDR 420, 20 Apr 1967, 1-9; JDH 670. 7 Jun
1967, K-15; JDR 1273, 12 Apr 196S, F in
JDR 1212, 12 Apr 1968, G-7.
Sanicida crassicaulis Poepp. PNF
JDR 436, 20 Apr 1967, 1-4; JDR 118.3. 11
Apr 1968, 1-15; JDR 1282, 12 Apr 1968, K-9.
Scandix pecten-veneris L. \\F
JDR 518, 2 May 1967, H-15.
404
Great Basin Naturalist
Vol. 40, No. 4
Torilis arvensis (Huds.) Link ssp. purpurea
(Ten.) Hayek. AIF
D 80-39, 29 May 1980, F-8.
Torilis nodosa L. AIF
JDR 663, 20 Jun 1967, L-6; JDR 1160, 11
Apr 1968, F-14; JDR 1348, 12 Apr 1968, K-9.
Urticaceae
Urtica holosericea Nutt. PNF
JDR 736, 22 Sep 1967, G-15.
Valerianaceae
Centranthus ruber (L.) DC. PIF
JDR 528, 2 May 1967, H-17.
Plectritis macrocera T. & G. ANF
JDR 434, 20 Apr 1967, E-14.
Verbenaceae
Verbena robusta Greene. PNF
Observed by H, 16 May 1946.
Violaceae
Viola pedunculata T. & G. PNF
JDR 419, 20 Apr 1967, 1-4; JDR 1268, 12
Apr 1968, D-12.
Vitaceae
Parthenocissus tricuspidata (Sieb. & Zucc.)
Planch. PCF
JDR 724, 25 Jun 1967, G-14.
Class Monocotyledoneae
Agavaceae
Agave americana L. PCS
JDR 803, 22 Sep 1967, K-4.
Agave americana L. 'variegata\ PCS
JDR 959, 27 Dec 1967, K-6.
Cordyline striata Endl. PCS
JDR 990, 6 Feb 1968, H-15.
Amaryllidaceae
Allium dichlamijdeum Greene. PNF
JDR 622, 7 Jun 1967, K-8.
Amaryllis belUidonna L. PCF
JDR 725, 25 Jun 1967, J-6.
Brodiaea lam (Benth.) Wats. PNF
JDR 592, 7 Jun 1967, F-11.
Brodiaea pulchella (Salisb.) Greene. PNF
JDR 440, 20 Apr 1967, J-6; JDR 1190, 11
Apr 1968, 1-14; JDR 1237, 12 Apr 1968, K-5.
Narcissus poetaz Hort. PCF
JDR 992, 6 Feb 1968, E-13.
Narcissus pseudo-narcissus L. PCF
JDR 1042, 2 Mar 1968, 1-7.
Araceae
Zantedeschia aethiopica (L.) Spreng. PIF
JDR 806, 22 Sep 1967, 1-6.
Cyperaceae
Carex barbarae Dewey. PNSd
JDR 700, 25 Jun 1967, F-12; JDR 1189a,
11 Apr 1968, G-14; JDR 1367, 29 Apr 1968,
D-14.
Carex brevicaulis Mkze. PNSd
H 21850, 16 May 1946.
Carex gracilior Mkze. PNSd
JDR 1288, 12 Apr 1968, K-9.
Carex tumulicola Mkze. PNSd
JDR 400, 20 Apr 1967, M-15; JDR 1093,
11 Apr 1968, M-16; JDR 1347, 12 Apr 1968,
K-9.
Cyperus eragrostis Lanik. PNSd
JDR 750, 22 Sep 1967, G-15.
Scirpus cernuus Vahl. var. californicus (Torr.)
Beetle. ANSd
JDR 1402, 10 Jun 1968, N-15.
Gramineae
Agrostis diegoensis Vasey. PNG
JDR 619, 7 Jun 1967, 'K-8.
Agrostis exarata Trin. var. pacifica Vasev.
PNG
JDR 672, 20 Jun 1967, L-6.
Agrostis hall a Vasey. PNG
Observed by H, 16 May 1946.
Agrostis setniverticillata (Forsk.) C. Chr.
PIG
JDR 769, 22 Sep 1967, M-15; JDR 747, 22
Sep 1967, D-11.
Aira caryophyllea L. AIG
JDR 628, 7 Jun 1967, K-8; JDR 1164, 11
Apr 1968, F-14; JDR 1314, 12 Apr 1968, K-9.
Ammophila breviligulata Fernald. PIG
JDR 947, 27 Dec 1967, K- 16.
Avena barbata Brot. AIG
JDR 395, 20 Apr 1967, M-I4; JDR 1145,
11 Apr 1968, 1-8; JDR 1263, 12 Apr 1968, C-
12.
December 1980
Ripley: Angel Island Plants
405
Aiena fotua L. AIG
JDR 572, 7 Jun 1967, K-8; JDR 1382, 29
Apr 1968, K-15.
Briza maxima L. AIG
JDR 1701, 20 Apr 1969, 1-7; JDR 1707, 20
Apr 1969, E-U.
Briza minor L. AIG
JDR 572, 7 Jun 1967, H-7; JDR 1382, 29
Apr 1968, J-15.
Brumus carinattis H. & A. BNG
JDR 447a, 20 Apr 1967, 1-7; JDR 1171, 11
Apr 1968, G-7; JDR 1284, 12 Apr 1968, K-9.
BroDius diandrus Roth. AIG
JDR 394, 20 Apr 1967, M-15; JDR 447, 20
Apr 1967, J-7; JDR 509, 2 May 1967, D-14;
JDR 527, 2 May 1967, 1-16: JDR 1086. 11
Apr 1968, F-14. '
Bromus madritensis L. AIG
JDR 446, 20 Apr 1967, J-6: JDR 646, 20
Jun 1967, L-5.
Bromus marginatus Nees + PNG
JDR 1324, 12 Apr 1968, 1-8.
Bromus mollis L. AIG
JDR 427, 20 Apr 1967, J-5; JDR 1372, 29
Apr 1968, J-13; JDR 1404, 10 Jun 1968, N-
16.
Bromus racemosus L. .\IG
R. & J. 21330, 2 May 1967, D-10.
Bromus ruhens L. AIG
JDR 1124, 11 Apr 1968, F-12; JDR 1338,
29 Apr 1968, 1-9.
Cortaderia selloana (Schult.) Aschers. &
Graebn. PCG
JDR 532, 2 May 1967, K-16; JDR 633, 20
Jun 1967, H-16; JDR 957, 27 Dec 1967, L-15.
Cynodon dactijlon (L.) Pers. PIG
JDR 1522, 14 Jun 1968, K-9.
Cynosurus erhinatus L. AIG
D 79-41, 6 Jul 1979, E- 12.
Dactylis glomerata L. PIG
Observed by H, 16 May 1946.
Danthonia californica Roland, var. amcricana
(Scribn.) Hitchc. PNG
JDR 1188, 11 Apr 1968, F-14.
Danthonia pilosa R. Br. PIG
T8468, 16 Jun 1978, H-7.
Distichlh spicata (L.) Greene. PNG
JDR 728, 25 Jun 1967, K-5.
Elymus glaucuso Buckl. PNG
JDR 460, 28 Apr 1967, 1-9; JDR 548, 2
May 1967, K-8; JDR 1280, 12 Apr 1968, K-9.
Ehjmus triticoidcs Buckl. PNG
Observed by H, 16 May 1946.
Elymus triticoides Buckl. ssp. multif torus
Gould. ANG
JDR 705, 25 Jun 1967, G-8.
Festuca californica Vasey. PNG
JDR 551, 2 May 1967, K-8; JDR 1023, 2
Mar 1968, 1-9; JDR 1077, 11 Apr 1968, E-9;
JDR 1123, 11 Apr 1968,0-12.
Festuca drrtonensis (All.) .\schers. &
Graebn. AIG
JDR 541, 2 May 1967, G-11; JDR 1104, 11
Apr 1968, G-7; JDR 1285. 12 Apr 1968. K-9:
JDR 1231, 12 Apr 1968, K-6.
Festuca idahoensis Elmer. PNG
JDR 1146, 11 Apr 1968,1-9.
Festuca megalura Nutt. .ANG
JDR 1082, 11 Apr 1968, G-14; JDR 1022, 2
Mar 1968, E-11; JDR 1260, 12 Apr 1968. D-
10; JDR 1303, 12 Apr 1968, K-9.
Festuca myuros L. AIG
JDR 507, 2 May 1967, G-13; JDR 550, 2
May 1967, D-11; JDR 1262, 12 Apr 1968, D-
10.'
Festuca pacifica Piper. ANG
JDR 1187, 11 Apr 1968,1-14.
Festuca reflexa Buckl. ANG
JDR 1420, 10 Jun 1968, K-9.
Festuca rubra L. PNG
Observed by H, 16 May 1946.
Gastridium ventricosum iCiouan) Schin/ 6c
Thell. ANG
JDR 652, 20 Jun 1967, K-15; JDR 762, 22
Sep 1967, M-15; JDR 1506. 14 Jun 1968. L-6.
Holcus lanatus L. PIG
JDR 1708, 16 Jun 1969, F-7.
Hordeum hrachyantherum Nevski. PNG
JDR 582, 7 jun 1967, E-U; JDR 693, 25
Jun 1967, D-13; JDR 986, 6 Feb 1968, D-13.
Hordeum glaucum Steud. AIG
JDR 1095, 11 Apr 1968, G-7; JDR 1128, 11
Apr 1968, F-12.
Hordeum leporinum Link. MC
JDR 443, 20 Apr 1967, H-6: JDH 1101. 11
Apr 1968, J-7; JDR 1363. 29 Apr 1968, 1-9;
JDR 1383, 29 Apr 1968. J- 16.
406
Great Basin Naturalist
Vol. 40, No. 4
Hordeum vulgare L. AIG
Observed by H, 16 May 1946.
Koeleria macrantha (Ledeb.) Spreng. PNG
JDR 554, 2 May 1967, K-14; JDR 1307, 12
Apr 1968, K-9; JDR 1324a, 12 Apr 1968, 1-9.
Lolium multiflorum Lamk. var. muticum
DC. PIG
JDR 651, 20 Jun 1967, J-15; JDR 1369, 29
Apr 1968, J- 13.
Lolium perenne L. PIG
R. & J. 21040, 2 May 1967, K-9.
Melica californica Scribn. PNG
JDR 1203, 12 Apr 1968, G-15; JDR 1381,
29 Apr 1968, K-15.
Melica torreijana Scribn. PNG
JDR 549, 2 May 1967, K-8; JDR 1249, 12
Apr 1968, D-12; JDR 1163, 11 Apr 1968, G-
15; JDR 1157, 11 Apr 1968, G-12.
Monenna cylindrica (Willd.) Coss. & Dur.
AIG
Observed by H, 16 May 1946.
Phahris californica H. & A. PNG
JDR 464, 28 Apr 1967, 1-9; JDR 1143, 11
Apr 1968, H-10; JDR 1331, 12 Apr 1968, G-
12.
Plialaris minor Retz. AIG
JDR 618a, 7 Jun 1967, K-8.
Poa annua L. AIG
JDR 985, 6 Feb 1968, D-11; JDR 1020, 2
Mar 1968, F-13; JDR 1100, 11 Apr 1968, F-7.
Poa bolanderi Vasey ssp. howellii (Vasey &
Scribn.) Keck. ANG
JDR 1186, 11 Apr 1968, F-14.
Poa scahrella (Thiirb.) Benth. PNG
JDR 1085, 11 Apr 1968, F-14; JDR 1148a,
11 Apr 1968, 1-14; JDR 1240, 12 Apr 1968,
D-12.
Poa unilateralis Scribn. PNG
JDR 1087, 11 Apr 1968, F-14; JDR 1317,
12 Apr 1968, K-9; JDR 1321, 12 Apr 1968, I-
9.
Polypogpn interrnptiis H. B. K. PIG
JDR 1396, 8 Jun 1968, N-17; JDR 1403, 10
Jun 1968, M-14; JDR 1526, 14 Jun 1968, N-
16.
Pohjpogon monspeliensis (L.) Desf. AIG
JDR 653, 20 Jun 1967, L-15; JDR 758, 22
Sep 1967, N-16.
Sitanion x Jianscnii (Scribn.) J. G. Smith.
PNG
JDR 1371a, 29 Apr 1968, D-14.
Sitanion jubatum J. G. Smith. PNG
JDR 767, 22 Sep 1967, J-15; JDR 1381a, 29
Apr 1968, L-15.
Stipa lepida Hitchc. PNG
JDR 1198, 12 Apr 1968, F-13.
Stipa pulchra Hitchc. PNG
JDR 539, 2 May 1967, D-11; JDR 1127, 11
Apr 1968, H-11; JDR 1131, 11 Apr 1968, E-
11; JDR 1371, 29 Apr 1968, C-14.
Iridaceae
Iris longipetala Herbert. PNG
JDR 429, 20 Apr 1967, J-7.
Sisijrinchium helium Wats. PNG
JDR 387, 20 Apr 1967, L-6; JDR 1118, 11
Apr 1968, G-9; JDR 1246, 12 Apr 1968, C-
11.
Juncaceae
Juncus balticus Willd. PNR
JDR 1292, 12 Apr 1968, K-9.
Juncus bufonis L. ANR
JDR 640, 20 Jun 1967, J-8.
Juncus effusus L. var. pacificus Fern. &
Wieg. PNR
JDR 751, 22 Sep 1967, G-15.
Juncus patens E. Mey. PNR
JDR 397, 20 Apr 1967, M-16; JDR 579, 7
Jun 1967, F-12.
Juncus tenuis Willd. var. congestus Engelm.
PNR
JDR 657a, 20 Jun 1967, K-16; JDR 1291,
12 Apr 1967, K-9.
Luzula subsesilis (Wats.) Buch. PNR
B-D 6912 without date; JDR 540, 2 May
1967, D-11; JDR 1035, 2 Mar 1968, F-12;
JDR 1075, 11 Apr 1968, J-7.
Liliaceae
Aloe saponaria (Ait.) Haw. PIS
JDR 517, 2 May 1967, G-14.
Calochortus luteus Dougl. ex Lindl. PNF
D 80-47, 18 Jun 1980, 1-12.
Chlorogalum pomcridianum (DC.) Kunth var.
divaricatum (Lindl.) Hoover. PNF
JDR 630, 20 Jun 1967, 1-8.
December 1980
Ripley: An(;el Island Plants
407
Fritillarid hmceohta Pursh. PNF
JDR 1206, 12 Apr 1968, E-9; JDR 1681, 16
Mar 1969, L-7.
Knaphofia uvaria (L.) Okeii. PCS
JDR 439, 20 Apr 1967, J-5.
S7nilacina raccmosa (L.) Desf. PNF
JDR 1116, 11 Apr 1968, G-9.
Smilacina stellata (L.) Desf. var. sessilifolia
(Baker) Renders. PNF
R. &J. 21111, 2 May 1967, L-9.
Trilliutu chloropctdluiu (Torr.) Howell. PNF
JDR 1680, 16 Mar 1969, K-7.
Zigadcnus fremontii (Torr.) Torr. PNF
JDR 465, 28 Apr 1967, 1-9; JDR 1033, 2
Mar 1968, G-12.
Orchidaceae
Hahernaria elegans (Lindl.) Roland. PNF
JDR698, 25Jnn 1967, F-12.
Palmae
Phoenix canariensis Chaboud. PCT
JDR 512, 2 May 1967, C- 13.
Typhaceae
Typlia angustifoUa L. PNF
JDR 824, 22 Sep 1967, J-9.
Zosteraceae
Zostcra marina L. var. latifolia Morong.
PNF
JDR 1391, 8 Jun 1968, F-8; JDR 1426, 10
Jun 1968, G-6.
PhijUospadix torrciji Wats. PNF
JDR 1425, 10 Jun 1968, N-17.
ACKNOWLEDG.VIENTS
It is my pleasure to acknowledge the in-
valuable a.ssistance of the following individ-
uals who generously assited in the prepara-
tion of this paper: J. T. Howell, H. D. Thiers,
M. Josselvn, C. Best, C. H. True, A. Day, and
CO. Burke.
LlTEHATl HE ClTEl)
Bailey, K. li., W . I', liuviv. and D. L. Jones. Ujf>4.
Franciscan and related rocks and their signifi-
cance in the neolojfy of western California. Cali-
fornia Division of .Mines and Geologv, Bull. 18.3.
Blo.vvm, T. W. 19.56. Jadeite-hearing metagravwackes in
California, .\merican .Mineralogist 41:488-496.
1960. Jadite-rocks and glaucophane-schists from
.\ngel Island, San Francisco Bav. California. Am.
Jour. Sci. 258:.55.5-.57.3.
HouLE.NBERG, C. S., A.ND I. .\. .\bbott. 1966. Supplement
to Smith's marine algae of the Monterev Penin-
sula. Stanford University Press, Stanford, Califor-
nia.
HowEi.i., J. T. 1970. .Marin flora and supplement. Uni-
versity of California Press. Berkelev.
-Mc.NZ, P. A. 1973. .\ California flora and supplement.
University of California Press, BerkeleN'.
Obkki.ander, C T. 19.56. Simimer fog precipitation on
the San Francisco Peninsula. Ecology
37(4):851-852.
Ra.nso.me, F. L. 1894. The geolog\ of .\ngel Island. Uni-
versity of California Pub. Dept. Geology
1:19.3-240.
Ripley, J. D. 1969. .\ floristic and ecological study of
.\ngel Island State Park, .Marin County, Califor-
nia. Unpublished thesis, San Francisco Slate Uni-
versity, San Francisco. California.
ScHLOCKEH. J. M. 1961. Podingite from .\ngel Island.
San Francisco Bav. California. U.S. Geological
Survey Prof. Paper 400-B, p. B311-B312.
Scni.ocKER, J. M., M. G. Bomlla. ano D. H. Radhiuch.
1958. CJeology of the San Francisco north quad-
rangle. California. U.S. Geological Survey Mi.sc.
(;eol. Inv. .Map 1-272.
Smith. G. .M. 1944. .Marine algae of the Monterey Penin-
sula, (California. Stanford Uni\ersit\' Press. Stan-
ford, California.
ADDITIONS TO THE VASCULAR FLORA OF TETON COUNTY, WYOMING
Ronald L. Hartman' and Roliert W. Lichvar^-^
Abstract.— An annotated list of 125 taxa new to the flora of Teton County is presented, increasing the number of
known species to 1043.
Recently, Shaw (1976) published a "Field
Guide to the Vascular Plants of Grand Teton
National Park and Teton County, Wyom-
ing," which was said to be based in part on
the holdings of the Rocky Mountain Herbar-
ium. Unfortruiately, that herbarium was con-
sulted mainly for the taxonomic groups
treated by Dr. Robert D. Dorn, including the
Poaceae and the genera Artemisia and Salix.
Consequently, many of the 125 taxa listed be-
low were omitted. Additionally, recent in-
tensive collecting by the junior author in the
Gros Ventre Mountains resulted in the dis-
covery of 20 species apparently new to the
flora of Teton County. This paper updates
the list of known taxa for this much-visited
area and emphasizes the richness of its flora
(1043 species). The nomenclature and tax-
onomy follow that of Hitchcock and
Cronquist (1973), unless otherwise indicated
by synonymy. The senior author has checked
determinations on those specimens not anno-
tated by specialists in their respective groups.
Dr. Dorn is acknowledged for assistance with
the manuscript.
Apiaceae
Angelica roseana Hand. Teton Mts., 16
Aug 1899, A. Nelson 6 E. Nelson 6500, anno-
tation by M. E. Mathias & L. Constance,
1940.
Cicuta douglasii (DC.) Coult. & Rose.
Kent's Corner (SI T41N R115W), elev. 6700
ft, 10 Aug 1977, R. Lichvar 1235.
Cymopterus terebinthinus (Hook.) T. & G.
var. calcareus (M. E. Jones) Cronq. Teton
Pass Mts., elev. 7500 and 9000 ft, 22 Jul
1920, E. B. 6 L. B. Paijson 2073, annotation
by M. E. Mathias, 1930, as Pteryxia t. var. c.
Ligusticum porteri Coult. & Rose. 7.7 mi
W of Flagg Ranch, elev. 7600 ft, 14 Jul 1956,
W. G. Solheim 4571; Whetstone Creek, 24
Jul 1929, O. /. Murie 86.
Apocynaceae
Apocynum cannabinum L. var. glaberri-
muni DC. Hot Springs Bar, 20 mi S of Jack-
son, 19 Jul 1901, E. D. Merrill 6 E. N. Wilcox
1042, cited by Woodson (1930).
ASTERACEAE
Antennaria anaphaloides Rydb. Gros
Ventre River, Jackson Hole, 14 Jul 1901, E.
D. Merrill 6 E. N. Wilcox 986; Sheep Creek
(S16 T41N R115W), elev. 6900 ft, 5 Jun
1977, R. Lichvar 242.
Arnica sororia Greene. Near Moose (SWi/4
S16 T43N R115W), 4 Jul 1970, D. W. Sa-
binske 13C.
Aster eatonii (Gray) Howell. Along Tag-
gert Creek, elev. 7500 ft, 14 Aug 1932, L.
Williams 1041, annotation by M. L. Dean,
1963; Gros Ventre River (SI T42N R115W),
elev. 6900 ft, 21 Aug 1977, R. Lichvar 1285.
Chrysopsis horrida Rydb. Gros Ventre
River, 16 Aug 1894, A. Nelson 1084, annota-
tion by V. L. Harms, 1963, as Heterotheca
horrida (Rydb.) Harms.
Crepis runcinata (James) T. & G. ssp.
glauca (Nutt.) Babe. & Stebb. Gros Ventre
Mts., E of Gros Ventre Slide, elev. 7300 ft, 28
Aug 1949, W. G. Solheim 2733.
Erigeron corymbosus Nutt. Jackson Hole,
10 Jul 1931, L. Williams 290, annotation by
A. Cronquist, 1943-44; Crvstal Creek (S28
T42N R113W), elev. 7100 ft, 7 Jul 1977, R.
Lichvar 655.
'Rocky Mountain Herbarium, Department of Botany. University of Wyoming. Laramie, Wyoming 82071.
■Present address: Wyoming Natural Heritage Program, The Nature Conservancy, 1603 Capitol Avenue, No. .325, Cheyenne, Wyoming 82001.
408
December 1980
Hartman, Lichvau: V asci lah Flora
409
Erigeron flagellaris Gray. Glacier Creek,
Jackson Hole, elev. 7500 ft, 9 Aug 1920, E. B.
6 L. B. Paijson 2258.
Erigcron rydbergii Cronq. Sheep Mt. (S34
T42N H114\V), elev. 9800 ft, 1 Autr 1977, R.
Liclnar 1015.
Erigeron simplex Greene. Cirque basin
near Amphitheater and Surprise Lakes, elev.
9750 ft, 31 Jul 1962, /. Merklc 62-36- Snake
River, 22 Aug 1894, A. Xel.son 96.9, annota-
tions by S. Spongberg, 1970; Sheep Mt. (S34
T42N R114W), elev. 9800 ft, 13 Jul 1977, R.
Lichvar 804.
Erigeron tirsiniis D.C. Eat. Two-gwo-tee
Pass, elev. 10,500 ft, 27 Jul 1932 L. Williams
947; Alaska Ba.sin, W slope Teton Range,
elev. 9500-10.000 ft, 14 Aug 1965. /. Merkle
65-44.
Gaillarida aristata Pursh. Kellv Post Of-
fice (SI T42N R115W), elev. 6900'ft, 18 Aug
1977, R. Lichvar 1278.
Gnaphalium chilense Spreng. Along Snake
River, 20 mi S of Jackson, 19 Jul 1901, E. D.
Merrill ir E. N. Wilcox 949.
Gnaphalium viscosiim H.B.K. Cascade
Canyon, elev. 7500 ft, 30 Jul 1934, L. Wil-
liams 1684.
Haplopappus lyallii Gray. Two Ocean
Mt., Two-gwo-tee Pass, elev. 10,000 ft, 1 Aug
1933, L. Williams 1366.
Microseris nigrescens Hend. Near Two-
gwo-tee Pass, elev. 9650 ft, W. G. 6 R. Sol-
Iwim 4031.
Senecio debilis Nutt. Gros Ventre Mts.,
elev. 7100 ft, 28 Aug 1949, T. F. 6 M. S.
Reed 2702, annotation bv T. M. Barkley,
1960.
Senecio fremontii T. & G. var. fremontii.
Near Leighs Lake, Teton Mts., 26 Jul 1901,
E. D. Merrill 6 E. N. Wilcox 1047; Gros
Ventre Mts., 10 mi N of Bondurant, 15 Aug
1922, £. B. 6 L. B. Payson .3022.
Senecio werneriaefolius Gray. Teton Mts.,
elev. 9000 ft, 21 Aug 1894, A. Xelson 979.
Solidago gigantea Ait. var. serotina
(Kuntze) Cronq. Double Diamond Ranch,
elev. 7000 ft, 31 Jul 1932, L. Williams 966.
Townsendia florifer (Hook.) Gray. Snake
River, 29 May 1892, F. McCullough s.n., an-
notation by J. H. Beaman, 1956.
Townsendia hookeri Beaman rather than
T. exscapa as reported in Shaw (1976) ac-
cording to R. Dorn (pers. conun.).
Townsendia leptotes (Gray) Osterh. Sheep
Mt., ca. 14 mi NE of Jackson, elev. 11,190 ft,
30 Jul 1957, J. //. Beaman 6 K. J. Stone 1487.
Toivnsendia parryi D.C. Eat. Road above
Upper Slide Lake, (iros Ventre River, elev.
9300 ft, 27 Jul 1955, W. G. 6 R. Solheim
4143; Sheep Mt. (S33 T42N R115W), elev.
11,100 ft, 13 Jul 1977, R. Lichvar 824.
Boraginaceae
Cryptantha amhigua (Grav) Greene. Near
Moose iSE'4, S27 T43N R115\V), 1 Jul 1971,
D. W. Sahinske B3.
Brassicaceae
Arabis lemmonii Wats. Teton Pass .\lts.,
elev. 10,100 ft, 25 Jul 1920, E. B. 6 L. B.
Payson 2135.
Arabis microphylla Nutt. var. micro-
phylla. Treasure Mt. Scout Camp, Teton
Can) on, 25 Jun 1955, L. C. Anderson 109.
Barbarea vulgaris R. Br. Elk Ranch, elev.
6900 ft, 13 Jun 1948. /. F. 6 M. S. Reed 2318.
Berteroa incana (L.) DC. Alta. Jun 1969.
//. P. Alley s.n.
Descurainia californica (Gray) Schultz.
Snow King Mt. (S7 T41\ R116W)', elev. 7700
ft. 22 Jun 1977, /^. Lichvar 407.
Draba lanceolata Rovle. Sheep Mt. (S16
T41N R114W), elev. lO.'oOO ft, 23 Jun 1977.
R. Lichvar 462.
Draba praealta Greene. Hoback Canvon,
elev. 7500 ft, 23 Jun 1933, L. Williams 1143.
annotation by C. L. Hitchcock, 1939.
Erysimum inconspicuum (Wats.) MacM.
Bacon Creek, 15 Aug 1894, A. Xelson 916.
Hesperis matronalis L. Snow King Mt. (S7
T41N RI 16W), elev. 6700 ft. 22 Jun 1977. R.
Lichvar 392.
Physaria didymocarpa (Hook.) Gray var.
integrifolia Rollins. Near .\dams Ranch,
Jackson, 15 Jul 1901, E. D. Merrill 6 E. N.
Wilcox 965; Gros Ventre River, 16 Aug 1894,
A. Selson 927.
Rorippa curvipes Greene var. curvipes {R.
obtusa (Nutt.) Britt. var. obtusa). Lizard
Point, N end Jackson Lake, elev. 66(K) ft. 15
Aug 1961, C. L. 6 M. Porter 8829a. duplicate
annotated by R. L. Stuckey, s.d.
Rorippa curvipes Greene var. alpina
(Wats.) Stuckey {R. obtusa (Nutt.) Britt. \ar.
410
Great Basin Naturalist
Vol. 40, No. 4
alpina (Wats.) Britt.). E slope Grand Teton,
elev. 8500 ft, 6 Aug 1920, E. B ^ L. B. Pay-
son 2210, duplicate annotated by R. L.
Stuckey, s.d.
Sisymbrium altissimum L. Gros Ventre
River (SI T42N R115W), elev. 6800 ft, 27
Jun 1977, R. Lichvar 497.
Subularia aquatica L. Near Togwotee
Pass, elev. 9600 ft, 28 Aug 1952, C. L. Porter
6211.
Thelypodium paniculatum A. Nels. Bacon
Creek, 15 Aug 1894, A. Nelson 922.
Callitrichaceae
Callitriche hermaphroditica L. Near Tog-
wotee Pass, elev. 9000 ft, 4 Aug 1953, C. L.
Porter 6373.
Caryophyllaceae •
Arenaria stricta Michx. ssp. dawsonensis
(Britt.) Maguire. Vicinity of Hoback Canyon,
elev. 7500 ft, 24 Jun 1932, L. Williams 6 R.
Pierson 718.
Dianthus armeria L. Near Pilgrim Creek,
17 Aug 1965, A. A. Beetle s.n.
Stellaria jamesiana Torr. Hoback Canyon,
6 mi SE of Hoback Junction, elev. ca. 7000
ft, 7 Jun 1969, R. L. Hartman 2845.
Chenopodiaceae
Chenopodium glaucum L. var. salinum
(Standi.) Boiv. Near Gros Ventre River, 9 or
10 mi E of the slide, elev. 7100 ft, 28 Aug
1949, /. F. ir M. S. Reed 2700; Jackson Lake,
24 Aug 1922, E. B. ir L. B. Payson 3087, an-
notations by H. A. Wahl, 1960.
Chenopodium leptophyllum (Moq.) Wats.
Gros Ventre Road (S4 T42N R115W), elev.
7100 ft, 18 Aug 1977, R. Lichvar 1281.
Cyperaceae
Carex backii Boott. Treasure Mt. Scout
Camp, Teton Canyon, 25 Jun 1956, L. C. An-
derson 359, annotation by F. J. Hermann,
1958, as C. saximontana Mack.
Carex buxbaumii Wahl. E shore String
Lake, elev. ca. 6000 ft, 29 Jul 1964, W. M.
Johnson 465, annotation by F. J. Hermann,
1964.
Carex illota Bailey. Alaska Basin, W slope
Teton Range, elev. 9500-10,000 ft, 14 Aug
1965, /. Merkle 65-62.
Carex muricata L. Jackson Hole, 24 Jul
1901, E. D. Merrill 6 E. N. Wilcox 907.
Carex narditia Fries. Summit of Table Mt.,
elev. 11,100 ft, 17 Jul 1956, L. C. Anderson
532; Head of S Fork Cascade Canyon, elev.
10,500 ft, 1 Aug 1949, H. b V. Bailey 4936,
annotations by F. J. Hermann, 1958 as C.
hepburnii Boott.
Carex norvegica Retz. Just E of Treasure
Lake, Teton Canyon, 22 Jun 1956, L. C. An-
derson 347, annotation by F. J. Hermann,
1958, as C. media R. Br.
Carex nova Bailey. Teton Mts., 16 Aug
1899, A. Nelson <b E. Nelson 6527, annotation
by F. J. Hermann, 1958, as C. pelocarpa Her-
mann.
Carex praticola Rydb. Grassy Lake road
W of Flagg Ranch, elev. 6600 ft, 9 Jul 1959,
C. L. 6 M. W. Porter 7876, annotation by F.
J. Hermann, 1959.
Eleocharis flavescens (Poir.) Urban, Flagg
Ranch, Jul 1954, A. A. Beetle 16409.
Eleocharis rostellata Torr. Huckleberry
Hot Springs, 12 Aug 1974, A. A. Beetle s.n.
Eriophorum polystachion L. Grand Teton
National Park, elev. 7000 ft, 1 Aug 1932, L.
Williams 989.
Dipsacaceae
Knautia arvensis (L.) Coult. Jackson, Jun
1969, H. P. Alley s.n.
Ericaceae
Vaccinium globulure Rvdb. Along Taggart
Creek, elev. 7500 ft, 15 'jun 1933, L. Wil-
liams 1123, cited by Camp (1942).
Fabaceae
Astragalus argophyllus Nutt. Near Snake
River at Simon's, E of Moran, elev. 6750 ft, 2
Jun 1948, /. F. 6 M. S. Reed 1833, annotation
by R. C. Banieby, 1960.
Astragalus diversifolius Gray. Gros Ventre
River, 16 Aug 1894, A. Nelson 1086.
Astragalus eucosmus Robins. Spread
Creek, E side of Jackson Hole, elev. 7200 ft,
13 Jun 1948, /. F. 6 M. S. Reed 2261, annota-
tion by R. C. Barneby, 1960.
December 1980
Hartmax, Lichvar: Vascular Flora
411
Astragalus miser Dougl. ex Hook. var.
tenuifolius (Nutt.) Banieby. V^icinitv of Ho-
back Canyon, elev. 8000 ft, 24 Jun'l932, L.
Williams ir R. Pierson 721, annotation bv D.
Isely, 1976.
Astragalus teiwUus Pursh. Near Adams
Ranch, Jackson Hole, 15 Jnl 1901, E. D. Mer-
rill 6 E. N. Wileox 964.
Lupinus tiyethii Wats. Grand Teton Na-
tional Park. elev. 7000 ft, 21 Jun 1932, L.
Willidtiis 6801); Two-gwo-tee Pass, elev.
10,500 ft, 27 Jul 1932, L. Williams 940, anno-
tations by B. J. Cox, 1969.
Oxytropis sericea Nutt. var. spicata
(Hook.) Barneby. Near mouth of Fish Creek,
Gros Ventre drainage, 28 Jun 1958, D. E.
Wilbeii 5.
Oxytropis viscida Nutt. Gros Ventre Riv-
er, 16 Aug 1894, A. Nelson 928; Sportsmans
Ridge (SIO T40N R112W), elev. 9200 ft, 21
Jul 1977, R. Liehvar 980.
Juncus torreyi Gov. Teton Mts., 21 .\ug
1895, A. Selwn 956, annotation by F. J. Her-
mann, 1957.
Juncaginaceae
Triglochin palustris L. 2 mi SW of Jack-
son, elev. 2000 m, 30 Aug 1949, J. F. Reed
2726; Near Gros Ventre River, ca. 9 mi
above the slide, elev. 7100 ft, 28 Aug 1949. /.
F. 6 M. S. Reed 2697.
Lamlaceae
Nepeta cataria L, Kelh Warm Springs (SI
T42N R115W), elev. 6800 ft, 10 Aug 1977,
R. Lichvar 1240.
Stachys palustris L. var. pilosa (Nutt.)
Fern. 20 mi S of Jackson, 19 Julv 1901, E. D.
Merrill 6 E. N. Wilcox 953.
Gentianaceae
Gentiana barbellata Engelm. Crvstal
Creek divide, 25 Aug 1933, O. J. Murie 1052.
Gentiana prostrata Haenke. E of the slide,
Ciros Ventre Mts., elev. 7300 ft, 28 Aug 1949,
/. F. 6 A/. S. Reed 2701.
Grossul.ariaceae
Ribes aureum Pursh. Snake River bottom,
6 mi SW of Jackson, elev. 6000 ft, 29 Jun
1933, L. Williams 1185.
Hydroch.aritaceae
Elodea longivaginata St. John. Two-Ocean
Lake, elev. 6900 ft, 4 Aug 1953, C. L. Porter
6365.
Hydrophyllaceae
Phacelia linearis (Pursh) Holz. Snake Riv-
er, 29 May 1892, F. McCoullough s.n., anno-
tation bv G. W. Gillett, 1958.
JUNCACEAE
Juncus nodosus L. 2 mi SW of Jackson,
elev. 2000 m, 30 Aug 1949, /. F. Reed 2727.
Lemnaceae
Lemna minuta H.B.K. Third Creek near
Swan Lake, 10 Jun 1959,/. Wetherell 40.
LiLIACEAE
Allium cernuum Roth. Big Cow Creek (S3
T41N R112W), elev. 8100 ft, 30 Jul 1977, R.
Lichvar 993.
Asparagus officinalis L. Gros Ventre Ris-
er at Kellv (SI T42\ R115W). elev. 6700 ft.
27 Jun 1977, R. Lichvar 496.
Calochortus nuttallii T. & G. Teton Pass
Mts., elev. 7000 ft, 22 Jul 1920. E. B. 6 L. B.
Payson 2093, annotation by M. Ownbey,
1939.
Malvaceae
Sidalcea oregana (Nutt.) Gray var. orc-
gana. Granite Creek, vicinity of Hoback
C:anyon, elev. 8000 ft, 10 Jul 1932, /.. Wil-
liams 836.
Najaoaceae
Najas guadalupensis (Spreng.) Morong.
Kellv Warm Springs, elev. 6700 ft. 17 Aug
1971. K. D. Dorn 1424.
412
Great Basin Naturalist
Vol. 40, No. 4
Onagraceae
Clarkia pulchella Pursh. Snake River, 29
May 1892, F. McCouUough s.n., annotation
by M. & H. Lewis, 1951.
Orchidaceae
Habenaria saccata Greene. Jackson Hole,
elev. 6700 ft, 8 Aug 1920, E. B. 6 L. B. Pay-
son 2248, annotation by D. S. Correll, 1946;
Granite Creek (S5 T39N R113W), elev. 8200
ft, 10 Jul 1977, R. Liclwar 721.
Orobanchaceae
Orobanche corymbosa (Rydb.) Ferris.
Near Cliff Creek, Hoback Canyon, 19 Aug
1922, E. B. ir L. B. Paijson 3078, annotation
by L. R. Heckard, s.d.
Plant AGiNACEAE
Plantago tweedyi Gray. Trail Creek, vicin-
ity of Teton Pass, elev. 7500 ft, 1 Jul 1932, L.
Williams 790.
POLEMONIACEAE
Gilia spicata Nutt. Sportsmans Ridge (SIO
T40N R112W), elev. 9200 ft, 21 Jul 1977, R.
Lichvar ,977; Curtis Canyon Campground
(S16 T41N R115W), elev. 8000 ft, 5 Jun
1977, R. Lichvar 263.
Gilia tenerrima Gray. Near Moose (NWVi,
S12 T44N R115W), 3 Jul 1970, D. W. Sa-
binske 12A; Teton Canyon, 11 mi E of
Driggs, Idaho, 2 Jun 1956, L. C. Anderson
264.
Navarretia breweri (Gray) Greene. Gros
Ventre River, 18 Aug 1894, A. Nekon 1094.
POLYGONACEAE
Erigonum brevicaule Nutt. var. laxifolium
(T. & G.) Reveal (£. chrysocephalum Gray).
Bacon Creek, 15 Aug 1894, A. Nelson 903;
Gros Ventre slide area, elev. 7000 ft, 10 Jul
1959, C. L. 6 M. W. Porter 7892, annotations
by J. L. Reveal, 1971; Soda Creek (S14 T41N
R112W), elev. 7200 ft, 21 Jul 1977, R. Lich-
var 941.
Polygonum achoreum Blake. Jackson Hole
Wildlife Park, elev. 6750 ft, 25 Aug 1949, /.
F. 6 M. Reed 2674.
Polygonum confertiflorum Nutt. Near
Jackson Lake, elev. 6750 ft, 21 Aug 1935, M.
Ownbey 976; Snake River, 15 Aug. 1899, A.
Nelson b- E. Nelson 6463, annotations by J.
O. Coolidge, 1963.
Polygonum saivatchense Small. Two-gwo-
tee Pass, 25 Jul 1939, Miss Gooding 39-139.
Polygonum watsonii Small. Jackson Lake,
21 Aug 1899, A. Nelson 6 E. Nelson 6556;
Hwy. 89, 2 mi S of Moran, 27 Jul 1939, /. F.
Brenckle 6- O. A. Stevens 44, annotations by
J. O. Coolidge, 1963.
Primulaceae
Primula incana Jones. Hot Spring Bar, 20
mi S of Jackson, 19 Jul 1901, E. D. Merrill b
E. N. Wilcox 1039; Adams Ranch, Jackson
Hole, 14 Jul 1901, E. D. Merrill 6 E. N. Wil-
cox 990; Soda Lake (Sll T41N R112W), elev.
7100 ft, 21 Jul 1977, R. Lichvar 937.
Potamogetonaceae
Potamogeton amplifolius Tuckerman. In
Lake of the Woods, near the Continental Di-
vide, elev. 9400 ft, 8 Aug 1956, C. L. Porter
7196.
Potamogeton epihydrus Raf. Signal Mt.,
elev. 6800 ft, 19 Jul 1963, C. L. 6 M. W. Por-
ter 9401.
Ranunculaceae
Anemone parviflora Michx. Teton Pass
Mts., elev. 9500 ft, 25 Jul 1920, E. B. 6 L. B.
Paijson 2134.
Anemone nuttalliana DC. Snow King Mt.,
(S35 T41N R116W), elev. 6500 ft, 25 Mav
1978, R. Lichvar 1446.
Delphinium burkei Greene. N drainage of
Pilgrim Creek, E of Colter Bay, 20 Jul 1957,
A. A. Beetle 13974, annotation bv R. J. Tay-
lor, 1960.
Ratiuticulus gmelinii DC. Lower Slide
Lake (S5 T42N Rl 14W), elev. 6900 ft, 4 Jul
1977, fl. Lichvar 572.
Ranuticulus uncinatus D. Don var. parvif-
lorus (Torr.) Benson. Snake River bottom,
elev. 6500 ft, 28 Jul 1932, L. Williams 754,
annotations by L. Benson, 1945.
December 1980
Hartman, Lichvar: Vascular Flora
413
Rosaceae
Potentilla ovina Macoun. Summit of
Table Mt., elev. 11,100 ft, 7 Jul 1956, L. C.
Anderson 429; Togwotee Pass, 8 Jul 1940, L.
E. Wehmei/er ct al. 5322 (NY), annotations bv
B. C.Johnston, 1979.
Potentilla hookeriana Lchm. Hoback Can-
yon, elev. 7500 ft, 24 Jun 1933, L. Williams
1164, annotation by B. C. Johnston, 1978.
Pyrtis mains L. Kellv Warm Springs (SI
T42N R115\V), elev. 6800 ft, 15 Aug 1977,
R. Lichvar 1237.
Salicaceae
Poptilus acuminata Rvdb. Gros Ventre
River (SI T42N R115W),' elev. 6800 ft, 27
Jun 1977, R. Lichvar 524.
Saxifragaceae
Conimitella williamsii (D.C. Eat.) Rvdb.
Hoback Canyon, elev. 7500 ft, 23 Jun 1933,
L. Williams 1144.
Saxifraga adscendens L. var. oregonensis
(Raf.) Breit. N side of the Grand Teton, elev.
11,000 ft, 10 Aug 1932, L. Williams 1016.
Saxifraga caespitosa L. var. minima
Blake. Single-Shot Mt., 4 Jul 1897, R. S. Wil-
liajns s.n., cited by Hitchcock et al. (1961).
Saxifraga cernua L. Sheep Mt. (S3 T41N
R114W), elev. 10,500 ft, 1 Aug 1977, R.
Lichvar 1028.
Scrophularl\ceae
Castilleja gracillima Rvdb. Blackrock
Creek, Two-gwo-tee Pass, elev. 9500 ft, 29
Jul 1932, L. Williams ,960, annotation by M.
Ownbey.
Mimulus tilingii Regel. Grand Teton Na-
tional Park, elev. 7000 ft, 22 Jul 1933, L.
Williams 1322.
Orthocarpus tolmiei H. & A. Teton Pass
Mts., elev. 8000 ft, 22 Jul 1920, E. B. b L. B.
Payson 2072, annotation by D. D. Keck,
1926.
Penstemon glaher Pursh ssp. glaber. Gros
Ventre slide area, elev 7000 ft, 10 Jul 1959,
C. L. 6 .\/. \V. Porter 7S95, annotation bv F.
S. Crosswhite, 1969.
Penstemon rydbergii A. Nels. var. ryd-
bergii. Grand Teton National Park, elev.
8000 ft, 19 Jul 1932, L. Williams S92, annota-
tion by D. V.Clark. 1971.
Synthyris pinnatifida Wats. Teton Pass
Mts., elev. 9200 ft, 22 Jul 1920, £. B. 6 /.. B.
Payson 2079, annotation by C. G. Schaack,
1975; Cache Divide (S14 T40N R115W),
elev. 9200 ft, 4 Aug 1977, R. Lichvar 1166.
Veronica scutellata L. Jackson Lake, 12
Aug 1899, A. Xelson 6 E. Selson 6561, anno-
tation by F. W. Pennell, 1920.
Solanace.ae
Solanum dulcamara L. Spalding Bav (S34
T44N R114W), elev 6400 ft, 1 Aug 1978, R.
Lichvar 1604.
Violaceae
Viola canadensis L. var. canadensis. Vi-
cinitv of Hoback Canvon, elev. 7000 ft. 25
Jun 1932, /.. Williams'{:r R. Pier.wn 731, an-
notation bv N. H. Russell, 1967.
Viola nephrophylla Greene. Hot Spring
Bar, 20 mi S of Jackson, 19 Jul 1901. E. D.
Merrill 6 E. X. Wilcox 1040, N'icinitv of .Mo-
ran, 7500-8500 ft, 22-30 Jun 1935, T. G.
Yuncker 5291, annotations bv N. H. Russell,
1967.
Literature Cited
lIiTfncocK, C. L.. A. Cronqiist. \t. Ownbev. and J.
W. Thompson. 1961. Vascular plants of the Pacif-
ic Northwest. I'nivcrsitv of W'ashiiiiitoii Press,
Seattle, vol. .3.614 pp.
HiT( IK ocK, C. L., A.ND .\. Cronqiist. 1973. Flora of the
Pacific Northwest. University of Washington
Press, Seattle, xix -f- 730 pp.
Shaw, R. J. (with major contributions by R. D. Dorn).
1976. Field piide of the vascular plants of Grand
Teton National Park and Teton Co\inty, Wyom-
ing. Utah State University Press. Logan, xvi -1-
301 pp.
INDEX TO VOLUME 40
The genera and species decrihed as new to science in this vohiine appear in hold t\ pe in
this index.
Ahronia argillosa, p. 78.
Additions to the vascular Oora of Teton
County, Wyoming, p. 408.
Allred, Dorald M., articles by, pp. 116. 165.
Anipldcninus spectus, p. 358.
Anaxylebonis, p. 90.
Andersen, Ferron L., and Peter M. Schantz,
article by, p. 216.
Andersen, William R., and Jack D. Broth-
erson, article bv, p. 121.
Anderson, Loran C, articles bv, pp. 73, 117,
351.
Andre, John B., and James A. MacMahon, ar-
ticle bv, p. 68.
Androsace alaskana, var. reedae, p. 80.
Apoxyleborus, p. 90.
Araptus speciosus, p. 357.
Atwood, N. Duane, Clyde L. Pritchett, Rich-
ard D. Porter, and Benjamin W. Wood, ar-
ticle by, p. 303.
Bacterium Thioploca ingrica on wet walls in
Zion National Park, Utah, The, p. 98.
Bateman, Lucinda, article by, p. 268.
Baugh, Thomas M., article by, p. 139.
Baugh, Thomas M., and Bnice C. Brown, ar-
ticle by, p. 359.
Baumann, Richard W., and Bill P. Stark, ar-
ticle by, p. 63.
Breinholt, J. Craig, and Richard A. Heck-
mann, article by, p. 149.
Brotherson, Jack D., article by, p. 372.
Brotherson, Jack D., and William R. Ander-
sen, article bv, p. 121.
Brotherson, Jack D., William E. Evenson,
and Richard B. Wilcox, article by, p. 167.
Brotherson, Jack D., Lee A. Szyska, and Wil-
liam E. Evenson, article bv, p. 229.
Brown, Bruce C, and Thomas M. Baugh. ar-
ticle by, p. 359.
Bryan, James A., and Ronald \l. Lanner, ar-
ticle by, p. 190.
Burton, Sheril D., Sanniel R. Rushforth, Jef-
frey R. Johansen, and fuciith A. Grimes, ar-
ticle by, p. 98.
Campbell III, Thomas M., and Tim W.
Clark, article by, p. 183.
Chiracdnthiiini sj^idcr ])ite. A. p. 116.
Cltnisi>th(iinnus pulchelloides, p. 351.
Clark, Tim W., and Thomas M. Campbell
111, article by, p. 183.
Collin.s, Nicholas C, and Grav Stirling, ar-
ticle by, p. 131.
Comparative floral biology of Pcnstrmon ca-
tonii and Pcnstemon cyananthiis in central
Utah: a preliminarv studv, p. 268.
Comparison of epiph\ tic diatom assemblages
on living and dead stems of the conunon
grass Plinif^niitcs (iiistnili.s, .\, p. 223.
Cono))htlu)nis michoacanae, p. 354.
Cou()j)lithoni.s teocotum, p. 354.
Cryphalogenes, p. 91.
Ciyphal()<gcncs cuphorbiae, p. 91.
CrypluiloigciU's exiguus, p. 92.
Cushing, Bruce S., and .\nne Matherne. ar-
ticle by, p. 193.
Differential habitat utilization l)\ the sexes of
mule deer, p. 273.
Dog owners and hydatid disease in Sanpete
County, Utah, p. 216.
Egoscue, Harold J., article bv, p. 361.
Elliott, Charles L., and Jerran T. Flinders, ar-
ticles by, pp. 175, 362.
Ernocladiu.s, p. 93.
Evenson, William E., Jack D. Brotherson,
and Lee \. Szvska, article by, p. 229.
Evenson, \\'illiam E., Jack D. Brotherson,
and Richard B. Wilcox, article by. p. 167.
Feeding ecology of Gila honixohitts (Os-
teichthves: Cvprinidae) endemic to a ther-
mal lake in southeastern Oregon, p. 101.
Ferrington, Leonard C, Jr., and Bernard C
Swegman, article by, p. 287.
Field observations on the response of the
Railroad Valley springfish [Crcnichtliijs nc-
vadae) to temperature, p. 359.
First record of the pallid bat iAutroznits piil-
lidus) from .Montana, p. 115.
Flath, Dennis L., and Jeff Shryer, article l)\.
p. 115.
Flinders, Jerran T., and Charles L. Elliott, ar-
ticles b\ . pp. 175, 362.
415
416
Great Basin Naturalist
Vol. 40, No. 4
Flood frequency and the assemblage of dis-
persal types in hanging gardens of the
Narrows, Zion National Park, Utah, p.
365.
Garcia, James R., Clive D. Jorgensen, and H.
Duane Smith, article by, p. 282.
Genus Eriogoniim Michx. (Polygonaceae) and
Michel Gandoger, The, p. 143.
Goodrich, Sherel, and Stanley L. Welsh, ar-
ticle by, p. 78.
Grimes, Judith A., Samuel R. Rushforth, She-
ril D. Burton, Jeffrey R. Johansen, article
by, p. 98.
Grimes, Judith A., Larry L. St. Clair, and
Samuel R. Rushforth, article by, p. 223.
Habitat and plant distributions in hanging
gardens of the Narrows, Zion National
Park, Utah, p. 178.
Hadrodemius, p. 94.
Haplopappits alpinus, p. 73.
Haplopappus alpinus (Asteraceae): a new
species from Nevada, p. 73.
Harper, K. T., and L. M. Kunzler, article by,
p. 127.
Hartman, Ronald L., and Robert W. Lichvar,
article by, p. 408.
Hatch, Stephan L., article by, p. 221.
Heckmann, Richard A., and J. Craig Brein-
holt, article by, p. 149.
Henderson, Jan A., David J. Schimpf, and
James A. MacMahon, article by, p. 1.
Hesperoperla hoguei, p. 63.
Hesperoperki hoguei, a new species of stone-
fly from California (Plecoptera: Perlidae),
p. 63.
Hijlesinus aztecus, p. 354.
Hijlurdrectonus corticinus, p. 94.
Identity of narrow-leaved CJinjsothamniis
viscidiflonis (Asteraceae), p. 117.
Impact of the 1975 Wallsburg fire on ante-
lope bitterbrush {Purshia tridentata), p.
299.
Johansen, Jeffrey R., Samuel R. Rushforth,
Sheril D. Burton, and Judith A. Grimes,
article by, p. 98.
Jorgensen, Clive D., H. Duane Smith, and
James R. Garcia, article by, p. 282.
Kay, Jeanne, and George P. Malanson, article
by, p. 365.
Keiss, Robert E., Patrick W. Roberts, and
Donald J. Nash, article by, p. 141.
King, Michael M., and H. Duane Smith, ar-
ticle by, p. 273.
Kunzler, L. M., and K. T. Harper, article by,
p. 127.
Lanner, Ronald M., article by, p. 265.
Lanner, Ronald M., and James A. Bryan, ar-
ticle by, p. 190.
Lepidium ostleri, p. 80.
Leptoxyleborus, p. 94.
Lichvar, Robert W., and Ronald L. Hartman,
article by, p. 408.
Lygodesmia entrada, p. 83.
MacMahon, James A., and John B. Andre, ar-
ticle by, p. 68.
MacMahon, James A., David J. Schimpf, and
Jan A. Henderson, article by, p. 1.
Malanson, George P., article by, p. 178.
Malanson, George P., and Jeanne Kay, article
by, p. 365.
Matherne, Anne, and Bruce S. Gushing, ar-
ticle by, p. 193.
Microperus, p. 94.
Miscellaneous plant novelties from Alaska,
Nevada, and Utah, p. 78.
Murphy, Robert W., John R. Ottley, and
Geoffrey V. Smith, article by, p. 59.
Nash, Donald J., Patrick W. Roberts, and
Robert E. Keiss, article by, p. 141.
New American bark beetles (Coleoptera:
Scolytidae), with two recently introduced
species, p. 353.
New genera and new generic synonymy in
Scolytidae (Coleoptera), p. 89.
New grass distribution records for Arizona,
New Mexico, and Texas, p. 221.
New records of western Trichoptera with
notes on their biology, p. 287.
New species of fossil Chrysothamnus (Aste-
raceae) from New Mexico, A, p. 351.
Observations on seasonal variation in desert
arthropods in central Nevada, p. 292.
Ottley, John R., Robert W. Murphy, and
Geoffrey V. Smith, article by, p. 59.
Parasites from two species of suckers (Ca-
tostomidae) from southern Utah, p. 149.
Parker, Albert J., article by, p. 254.
Pediococtus despainii, p. 83.
Phloeocleptus punctatus, p. 355.
Pietruszka, Robert D., article by, p. 292.
Pityogenes mexicanus, p. 356.
Poisonous plants of Utah, p. 229.
Porter, Richard D., N. Duane Atwood, Clyde
L. Pritchett, and Benjamin W. Wood, ar-
ticle by, p. 303.
December 1980
Index
417
Postemergence development and interyear
residence of juvenile Columbian groimd
squirrels in the Idaho primitive area, p.
362.
Plants of Angel Island, Marin ('ountv, C^ali-
foniia, p. 385.
Pritchett, Clyde L., N. Duane Atwood, Rich-
ard D. Porter, and Benjamin W. Wood, ar-
ticle by, p. 303.
Psetidotliifsanocs atonius, p. 355.
Pscudothi/sanocs leechi, p. 356.
Recovery of Gambel oak after fire in central
Utah,' p. 127.
Relationship between environmental and
vegetational parameters for luiderstorv
and open-area commimities, p. 167.
Relationships among total dissolved solids,
conductivitv, and osmositv for five Ar-
temia habitats (Anostraca: Artemiidae), p.
131.
Reproduction in three sympatric lizard spe-
cies from west-central Utah, p. 68.
Reveal, James L., article by, p. 143.
Ribulose diphosphate carboxylase activities
in cold-resistant common mallow, Malvo
neglecta Wallr. and a cold-sensitive to-
mato, Lycopersicon esculciituin L., Ace 55
var., p. 121.
Ripley, J. D., article by, p. 385.
Roberts, Patrick W., Donald J. Nash, and
Robert E. Keiss, article by, p. 141.
Rushforth, Samuel R., Sheril D. Burton, Jef-
frev R. Johansen, and Judith A. (wimes, ar-
ticle by, p. 98.
Rushforth, Samuel R., Judith A. Grimes, and
Larry L. St. Clair, article by, p. 223.
St. Clair, Larry L., Judith A. Crimes, and
Samuel R. Rushforth, article by, p. 223.
Schantz, Peter M., and Perron L. Andersen,
article by, p. 216.
Schimpf, David J., Jan A. Henderson, and
James A. MacMahon, article by, p. 1.
Seasonal activity pattern of Columbian
ground squirrels in the Idaho primitive
area, p. 175.
Self-pollination experiment in Piiuis ('dulis,
A, p. 265.
Senecio toiyabensis, p. 86.
Short-term effects of logging on red-backed
voles and deer mice, p. 183.
Shryer, Jeff, and Dennis L. Flath, article by,
p. 115.
Smith, Geoffrey V.. John R. Ottley, and Rob-
ert W. Murphy, article bv, p. 59.
Smith, H. Duane, Clive D. Jorgensen. and
James R. Garcia, article by, p. 282.
Smith, H. Duane, and Michael M. King, ar-
ticle by, p. 273.
Soil water withdrawal and root distribution
under grubbed, sprayed, and undisturbed
big sagebrush vegetation, p. 157.
Some aspects of succession in the spruce-fir
forest zone of northern Utah, p. 1.
Spatiotemporal variation in phenology and
abundance of floral resources on .shortgrass
prairie, p. 197.
Spawning of the least cinib i/o/ir/i/Zii/.s
pJiIegetJwntis), p. 139.
Sphaeralcea grossularufoHa. \'ar. moorei, p.
35.
Sphaeralcea Icptopln/lla, var. janeae, p. 36.
Sphaeralcea psoraloides, p. 36.
Stanton, N. L., and V. J. Tepedino, article by,
p. 197.
Stark, Bill P., and Richard W. Bauniann. ar-
ticle bv, p. 63.
Stinger utilization and predation in the scor-
pion Paruroctonus horeus, p. 193.
Stirling, Gray, and Xicliolas C. Collins, ar-
ticle by, p! 131.
Sturges, David L., article by, p. 157.
Successional status of Cupressus arizonica.
The, p. 254.
Swarming of the western harvester ant, Pu-
gonoDiijmiex occidentalis, p. 165.
Swegman, Bernard G., and Leonard C.
Ferrington, Jr., article by, p. 287.
Szyska, Lee A., Jack D. Brotherson, and
William E. Evenson, article by. p. 229.
Taphrodasus, p. 95.
Taurodemus, p. 96.
Taxonomic status of the rosy boa Lichonura
roseofusca (Serpentes: Boidae), The, p. 59.
Temporal activity patterns of a Dipodonuis
ordii population, p. 282.
Tepedino, V. J., and N. L. Stanton, article by,
p. 197.
Terminal bud formation in linilHT piiK-. p.
190.
Terrestrial vertebrate fauna of llu- Kaiparo-
wits Basin, p. 303.
Transferrin polymorphism in bighorn sheep.
Otis canadensh, in Colorado, p. 141.
Utah flora: Malvaceae, p. 27.
Utah flora: miscellaneous families, p. 38.
418 Great Basin Naturalist Vol. 40, No. 4
Waejstaff, Fred J., article by, p. 299. Wood, Benjamin W., N. Duane Atwood,
Wetsh, Stanley L., articles by, pp. 27, 38. Clyde L. Pritchett, and Richard D. Porter,
Welsh, Stanley L., and Sherel Goodrich, ar- article by, p. 303.
tide by, p. 78. Wood, Stephen L., articles by, pp. 89, 353.
Wilcox, Richard B., William E. Evenson, and Woodrat nest flea Anamiopsi/Uus amphibolus
Tack D. Brotherson, article by, p. 167. . ^u 4- r>, ^ oa^
„ ^ , ^ -, -r 1 V; ^^r.ii. in southeastern Oregon, p. 361.
Williams, Cvnthia D., and Jack E. Williams, ^, , , ^^'/
article by, p. 101. Xyleborus praestans, p. 358.
Williams, Jack E., and Cynthia D. Williams, Zonation patterns in the potholes of Kalsow
article by, p. 101. Prairie, Iowa, p. 372.
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TABLE OF CONTENTS
Impact of the 1975 Wallsburg fire on antelope Ijitterbnish (Purshia tiidcntata). Fred
J. Wagstaff 299
Terrestrial vertebrate fauna of the Kaiparowits Basin. N. Duane Atwood, Clyde L.
Pritchett, Richard D. Porter, and Benjamin W. Wood 303
A new species of fossil ChrysotJiamnus (Asteraceae) from New Mexico. Loran C.
Anderson 3.51
New American bark beetles (Coleoptera: Scolytidae), with two recently introduced
species. Stephen L. Wood 353
Field observations on the respon.se of the Railroad Valley springfish (Crenichthijs
nevadae) to temperature. Thomas M. Baugh and Bruce G. Brown 359
Woodrat nest flea Anomiopsyllus atnphibolus in southeastern Oregon. Harold J.
Egoscue 361
Postemergence development and interyear residence of juvenile Columbian groimd
squirrels in the Idaho primitive area. Charles L. Elliott and Jerran T.
Flinders .362
Flood frequency and the assemblage of dispersal types in hanging gardens of the
Narrows, Zion National Park, Utah. George P. Malanson and Jeanne Kay 365
Zonatioii patterns in the potholes of Kalsow Prairie, Iowa. Jack D. Brotherson 372
Plants of Angel Island, Marin County, California. J. D. Ripley .385
Additions to the vascular flora of Teton County, Wyoming. Ronald L. Hartman and
Robert W. Lichvar .". '. '. 408
Index to Volume 40 414
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