HARVARD UNIVERSITY
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X Jl g J'J'^ 5 1992
HA RV A R g
UMVERSrEy
GREAT BASIN
MURALIST
VOLUME 52 NO 1 - MARCH 1992
BRIGHAM YOUNG UNIVERSITY
GREAT BASIN NATURALIST
Editor
James R Barnes
290 MLBM
BrighaiTi Young University
Provo, Utali 84602
MichaklA howKHs
Blandy ExperiuKMital F"anii
University of N'irginia
Box 175
Boyce, Virginia 22620
Pai'lC Marsh
Center for Environmental Stndies
Arizona State University
Tempe, Arizona 85287
Associate Editors
Jeanne C. Chambers
USDA Forest Service Research
860 North 12th East
Loiran, Utah 84322-8000
Brian A MAifRER
Pepartnient of Zoology
Brigham Yonng University
Fro\o, Utah 84602
Jeffrey R. Johansen
Department of Biology
John Carroll University
Cleveland, Ohio 441 18
JimmieR Parrish
BIO-WEST, Inc.
1063 West 1400 North
Logan, Utah 84321
Editorial Board. Richard W. Baumann, Chairman, Zoology; H. Duane Smith, Zoology;
Clavton M. White, Zoology; Jerran T. Flinders, Botany and Range Science; William Hess,
Botany and Range Science. All are at Brigham Yovmg University. Ex Officio Editorial Board
members include Clayton S. Huber, Dean, College of Biological and Agricultural Sciences;
Norman A. Darais, University Editor, University Publications; James R. Barnes, Editor, Great
Basin Wituralist.
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Copyriylit © 1^W2 l)y BriKliam Yoiini; Univt-rsitv
Ofllci.il piililication dati; 22 Mav 1992
ISSN 0017-3614
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LIBRARY
JUN 5 1992
The Great Basin Natiiralist
Published at Prono. Utah, by
Brigham Young Uni\ kusiit
ISSN 00 17-36 14
" I r
Volume 52
Margh 1992
No. 1
Great Ba.sin Natiiridi.st 52( 1 ). 1992. pp
IN MEMORIAM— A. PERRY PLUMMER (1911-1991
TEACHER, NATURALIST, RANGE SCIENTIST
E. Duiaiit McAitlm
A. Pern' Pliininier died in tlie Gunnison \ixllev
Ho.spitiil, Gunni.son, Utah, on October 3, 1991,
after several years of iU heiiltli. His piissing
deserves comment because he was a mixn who
made a difference in natin'al re.source manage-
ment luid research in the Intermountain area. He
spent his professional career (1936-1979) with the
Intermountain Research Station (INT, formerK'
the Intermountain Forest iuid Range E.xperiment
Station) of the Forest Senice, U.S. Department
of Agriculture, at duh' stations in Utaii near Mil-
ford and in Ogden, Ephraim, and Pion'o.
Teagiier .\nd Mentor
Perrv was a caring, effecti\e mentor and
teacher. His assignment witli the Forest Service
was research and research administration,
which he did w(^ll; but his professional lo\ e was
teaching, especialK' small groups and indixidu-
als. His formal teaching was limited to a couple
semesters at Brigham Young Universit)' (BYU)
shortl\- after the 1975 establishment of INT's
Shnib Sciences Laboratory on that campus. He
e.stal)lished a wildland shrub biologv class that
remains a part of the BYU curriculum, in addi-
tion, he instnict(nl numerous workshops at the
Great Basin Experimental liange (Ephraim
Canvon) and conducted man\' field tours at out-
planting, common garden, range rehabilitation,
and other research sites throughout Utiili and the
Intermountain area. Under these ci renin stance \s
he was a master teacher whose points mad(" lasting
impressions on whoever was there — agencx land
manager, private landowner, public school
teacher, Washington Office Forest Senice
research administrator, politician, junior col-
league, or uni\ ersit\ professor.
Perry had a rare gift of integrating in his mind
the potential vegetative states of degraded lands
because he knew soil t\pes, compatible plant
associations, plant adaptations, planting e(|uip-
inent, and seedb(nl re(juir(Miients. Becau.se of
this gift and his willingness to share it, he was
often called on to consult those n'sponsible for
rehabilitating degraded huids. Txpically. he
would visit potential rehabilitation sites and
folkm- up bv providing detailed w iitt(^n recom-
mendations. He completed well over one hun-
dred careful, thoughthil consultations lor tlie
good of tlu^ laud, for those who manage it, and
for its human and other occupants. He was a
mentor to others wlio continue on in this tradi-
tion: I think csnccialK of Steve \h)iisen of our
' Slinib Sciences Liihoniton,, IiiliriiioMTil.un Kesearcli Slalion. Kore.st Semce, U.S. Department of Agricnllure. Provo. Utah S4(t()6.
Great Basin iNatuiullst
[\ohinie
laborator)' and Richard Stexens of the Utah
Division Or W'ildHfe Resources (DWR) in
Ephraini.
I illustrate Pern's teaching st)le with a ])er-
sonal example. In May 1972 I had been working
for INT for four months when Perrv' took me on
a field trip to the Brown's Park area of northeast-
em Utah to exaluate the results of some earlier
work (he took or sent me on monthly field trips
those first two or three years). At one stop I saw
a patch of green in the distance at a spring. I
suspected monkey flowers {Miniiiltis sp. — the
subject of m\- Ph.D. degree research a few years
earlier) would be growing there. I hustled over
and confirmed mv suspicion. Perry ambled up
and said, 'It's nice to appreciate these monkey
flowers the wa\ \ on do, but look back toward the
tnick. What else (k) \on see? There aie lots of
other plant species and plant communities
between here and there. You can learn a lot by
looking at the whole plant communit)." He
laughed in his characteristic \\'a\', and we dis-
cussed the \arious plant species present and
their habitat requirements. A lasting lesson to
me. it is similar to other Perrv teaching
moments shared bv \n\ colleagues.
Back(;rou\d, Education, Work
Ethic:, and Honors
Arthur Pern Plummer (Hg. I ) was bom on a
farm in Daniel, Wasatcli Count\', Utah, on April
10, 1911. His mother died when he was young;
he and his siblings had a resourceful, indepen-
dent upbringing with their \\i(k)wer father. He
was educated in the Wasatch Count\ public
schools, at East High School in Salt Lake Cit)',
and at the University' of UtiJi. Peny received a
B.S. degree (1935) in botany from the U, began
his INT career (1936), married Blanche Swin-
dle of Monroe (1938), and completed his M.S.
degree also in botany at the U (1939) in a busy
h)ur \(*ars. He enjoyed his universitv' davs and
called on that background and experience
throughout his career. Notable among his pro-
fessors were Kim Newby Walter Cottam, Ralph
Chamberlain, Fayette Stephens, and Angus
Woodburx. He and Doc" Cottam continu(>d a
producti\ (â– interchange of ideas and shared field
trips into the mid-]97()s.
Perrv was a doer. He performed and worked
hard. He didn't just a.sk his subordinates to get
souK^thing done— he did it with them. As a new
Ph.D., I didn't e.xpect to be on the bu.siness end
of a hoe for .several hours a dav, but then 1 didn't
expect mv boss to be in that situation either. He
would show up anywhere a work crew was,
reach' to help with \1gor and energv', and he
expected anyone working to do the same. It
wasn't uncommon for Perr)^ to show up at these
sites at 11:30 a.m. or 4:30 p.m., seemingly
unaware of the impending lunch hour or (quit-
ting time.
Perrv's record of accomplishment was noted
by several organizations. In 1965 INT recog-
nized him with a certificate of merit and a sub-
stantial cash award for outstanding performance
in wildlife habitat research and application f)f
that research. Also in 1965 the Utah Wildlife
Federation honored him as Consen'ationist of
the Year. In 1973 the Utah Chapter of the Soil
Consenation Societ}' of America gaxe him their
Chapter Recognition Award. He received a
USD A Superior Seivice Award in 1969 for
implementing and luaking successful the coop-
eratixe work between INT and DWR. Pern', a
1949 charter member of the Societx' for Raiiiie
Management (SRM), was president of the Utah
Section and received SRMs Outstanding
Achievement Award (1974), the premier Fred-
eric G. Renner Award (1976), and the Fellow
Award (1977). He was president of the Utah
Chapter of the Soil Consenation Societv during
the early 197()s.
Scientific Contributions
In this section 1 comment not onl\ on Pern's
direct contributions but also on work that he
stimulated and inspired. Pern's contril)utions
were not limited to those he personalK' made;
but, like those of many great teachers, his
achievements have been enhanced aiul
expanded b\' those who came after and built
upon the foundation he laid.
('onsidering Pern's later contriliutions to
shmb biologx, it is of interest that his first pub-
lication was on de\ eloping a techuicjue for prep-
aration of microscopic sections of stems and
roots of shrubs (Newby and Pluuuuer 1936). His
master's degree thesis (1939), published in
1943, d{\ilt with germination and seedling
development of range grasses. He continued his
interest in seed germination, (jualitv, storage,
and processing, and in seedling de\elopment,
on a wick' range of plants throughout his career,
and his successors have continued this work
(Rudolf et al. 1974, Stein et al. 1974, Plummer
19921
In Mkmouiam — A. Pkkhy Pia \imkh
Fig. 1. A. Pern,^ Pliimiiier in his office about 1975.
and Jorgensen 1978, Stexeiis ct al. 1981, Meyer opment of procedures for revegetating degraded
et al. 1989, Ste\ens and Me\er 1990. \Ie\er and lands, including plant materials and operational
Monsen 1991). c(]uipnicnt infonnation and answers to liow.
Pern's greatest contributions iuNolwd tlc\el- wlicn. \\li\. and where. He was priuian author
Great Basin Natuiialist
[Volume
of three "how to" publications that have been
broacllv accepted and applied (Plumnier et al.
1955, i96S, Plummer 1977). The 1968 publica-
tion. Restoring Big Game Range in Utah,
became a classic; it has been used extensively in
the cKussroom and in the field and is now out ol
print after several press runs. It is serving as the
foundation of a new compendium for western
wildland rehabilitation techniques (Monsen
and Ste\ens, in press).
Other publications of note for general and
specific re\egetation applications include
Phunmer et al. (1943), Stewart and Plummer
(1947), Plummer and Fenlev (1950), Plummer
(1959, 1970), Plunnner and Stapley (1960), Ste-
vens et al. (1974), Hamer and Haq^er (1976),
Giunta et al. (197Sa), McArthur et al. (1978b),
Monsen and Phunmer (1978), Stevens et al.
(1981), Mon.sen and Shaw (1983), Monsen and
McArthur (1985), Da\is (1987), and Blauer
et al. (in press).
His earl\' rexegetation work led to a coopera-
tive research and application xenture bet\veen
INT and the Utiili Dixision of Wildlife Resouces
(knowTi then as the Utali Department of Fish
and Game) under Perry's direction. This effort
was stinmlated bv big game winter range prob-
lems brought on b> the [)artial urbanization of
those ranges, large deer populations, and the
heaxA' snowfalls of the late 1940s and earlx'
1950s. The program began in 1 954 at the behest
of the directors of INT and DWR. It is the most
extensive and longest running such arrange-
ment in the countrx'. He and his colleagues from
DWR produced 11 substautixe reports betxx^een
1956 and 1971 detailing their findings and rec-
ommendations in revegetation science ( Plum-
mer etal. 1956-1971).These reports, published
by DWR, xvere sought out and used xx'idelx^ bx
land management professionals.
Perrx- had a particular interest in and impact
on plant materials development including
exploration, collection, evaluation, adaptation,
culture, genetic xariation, hybridization, and
breeding systems. In this area he read carefulK'
and folloxx'etl the xxorks of Luther Burbank
(wide and unusual hvbridizations, see Kraft and
Kraft 1973), N. I. N'axilox- and E. V. Wulff (ori-
gins and dexelopment of related plant groups,
Wulff 1943, \'ax-ilov 1951 ), Jens Clausen, David
Keck, and William Hicsev' (accessional or pop-
ulational compari.sons in common gardens and
reciprocal transplantations, Clausen et al.
1940), and G. L. Stebbins (natural hybridization
and intraspecific variation, Stebbins 1950,
1959). He xvas particularly interested in applx-
ing these concepts to xvestem shnib species,
xx'hich had received little prior attention despite
their obvious ecological importance.
He spelled out his dream of a regional
common garden testing scheme (LeGrande,
Oregon; Boise, Idaho; Ephraim, Utah; and
Reno, Nevada) in a 1972 document (Plummer
1972a). Although this dream was not fully
implemented because of funding problems,
several useful and interesting studies resulted —
e.g.. Van Epps (1975), McArthur and Plummer
(1978), McArthur et al. (1978c, 1979, 1981),
Welch and McArthur (1979, 1981), Welch and
Monsen (1981), McArthur and Welch (1982),
Edgerton et al. (1983), Welch et al. ( 1983), Geist
and Edgerton (1984), Hegerhorst et al. (1987).
His specific interests in h\l)ridization, breed-
ing systems, and genetic xariation and selection
hav'e been addressed in a series of publications
specific to certain shrub taxa (Plummer et al.
1966, Nord et al. 1969, Hanks et al. 1971, 1973,
1975, Plummer 1974b, Blauer et al. 1975, 1976,
McArthur 1977, Stevens et al. 1977, Giunta et
al. 1978b, McArthur et al. 1978a, 1978c, 1979.
1988, in press, Welch et al. 1981, 1987, 1991,
McArthur and Freeman 1982, Davis 1983,
Freeman et al. 1984, 1991, Davis and Welch
1985, Welch and McArthur 1986, Pendleton et al.
1988, Welch and Jacobsen 1988, Wagstaff and
Welch 1991) and in more general terms (Drobnick
and Plummer 1966, Plunnner 1972b, 1974a,
Monsen 1975, Monsen and Christensen 1975,
Ciu-lson and McArthur 1985, McArthur 1989).
He had a keen eye for recognizing unusual
and/or superior plant populations occurring nat-
urally and in test plantings and in enhancing
tliose materials for improved productivity and
esthetics of degraded and badlv disturbed lands.
Several of these collections have been given
distinctive 'cultivar' or source identified names
and released for commercial propagation and
use by his associates since his retirement. These
includ(^ "Appar" Lewis flax {Liniini pcrcnne).
"Cedar" Palmer penstemon {Pcnstcnioii pal-
nieri), 'Rincon' founving Sixltbush [Ati'iplrx
canesrcii.s), "Hatch" xxinterfiit {Ccratoidcs laiidta),
"Hobble Creek" mountain bigsagebnish (Aiiciiiisia
tii(h'iit(ita ssj). vasci/ana), 'bnmignuit' forage
kochia {Kochia prosinitit), "Lassen" antelope
bitterbmsh (Piirsliia trident at a), "Ephraim"
crested wheatgnuss {A^ropi/roii cristatnni), and
"Paiute" orchardgrass {Dactijlis ^lonwrata)
19921
I\ MKMOHI AM — A. Pehhy Fiammkh
5
(McArtliur et A. 1984, Monsen and Ste\(^n.s
1985, Stevens and Monsen 1985, 1988a, 19881).
Stevens et al. 1985, Shaw and Monsen 1986.
Welch et al. 1986, McArthnr 1988). Othei .spe-
cies and populations were not released but iiax'e
had their usefulness documented and lia\(^
become axailable in the revegetation species
repertoire.
Perr\' Plunniier sened lor man\ \ears as the
Forest Ser\ice technical representatix'e to the
Western Regional Plant Introduction Couuiiit-
tee (W-6). His plant materials expertise was put
to use as a member of 1976 and 1977 plant
collection and e.xplo ration teams in the So\iet
Union (Dewey and Plummer 1980) and in 1980
as an on-site consultant in a New Zealand range
rehabilitation program. He also stimulated
interest in shnib disease and microbial and
entomological relationships (Tiernan 1978,
Nelson and Krebill 1981, Moore et al. 1982,
Nelson 1983, Nelson and Tiernan 1983, Nelson
and Schuttler 1984, Haws et al. 1988, Nelson
and Lopez 1989).
Aspects oi Pern's kne of plants can be high-
lighted bv two that were named after him: (1)
'Appar' Lewis flax was the first of se\eral plant
releases effected b\ INT, DWR, USD A Soil
Conser\ ation Service, and sex'eral state agricul-
tural experiment stations (the "App" in Appar is
for his initials); and (2) Gravia brandegei ssp.
phimnieri is a \\ide-lea\'ed tetraploid \ariet\' of
spineless hopsage that Howard Stutz named in
honor of its di.sco\erer (Stutz et al. 1987). These
tvvo plants illustrate the poles of Perry's work:
one is a show\' revegetation and horticultuial
cultivar; the other a restricted edaphic endemic,
new to science.
Perr\ helped develop and refine equipment
and techniques including anchor chaining, seed
dribblers, scalpers, seed collection and process-
ing, rangeland drills, and transplantation and
interseeding equipment (Plummer et al. 1956-
1971. 1968).
Lecacy
Manx of FeriA "s 80+ |)ubHcations are listed in
the Literature Cited section. .\si(k^ from these,
I see the following components of his legacy: ( 1 )
with Blanche, a fine family of seven children, (2)
an expanded scientific foundation that he and
his disciples ha\e laid for wildland reclamation
(see recent examples documented in the Liter-
ature Cited section) and for the incipient dis-
cipline oi shiub science, (3) hundreds of thou-
sands ol acres of successfullv rehabilitated
wildlands that retain sufficient plant diversitvto
supj)ort a rich native fauna, and (4) a native
wildland plant industrx' (.several seed companies
in Sanpete Counts' alone owe their existence, at
least in part, to Perrv and his team for back-
ground information, collecting and processing
techni(jues, and (k'velopment of a market for
products). I will acklress onl\- item 2.
Perrv beam his can^'r with the seeding, eval-
nation, and development of range grasses
(Plummer 1944, 1946, Plunnner and Stewart
1944, Plummer and Frischknecht 1952.
Frischknecht and Plummer 1955). He was
sinmltaneousK" involved in range management
research (Rotli and Plummer 1942, Phunmer et
al. 1943, Bleak and Phnnmer 1954) and sagebrusli
control work (Pehmiec et al. 1944, 1954, 1965).
Later, he managed the Great Basin E.xperimental
Range in Ephraim Canyon (Keck 1972).
When his assignment changed to restoration
of wildlife habitat in 1954, he quickk' became
conxertetl to the value of shnibs on wildlands.
Perrv liked to recount his subsequent attempts
to convert others to the value of shrubs, even the
heretofore "weed " sagebnish, by recalling an
anecdote. In the late 195()s he was with a crew
on a vegetative rehabilitation project above a
central Utah tovvni. The local Forest Service
district ranger came bv' to see what thev were
doing. Perr\' pointed out the v arious seeds in the
seed mix — crested wheatgrass, orchard gniss,
alfalfa, fourwing saltbush, Lewis flax, small bur-
nett, etc. The ranger wanted to know what one
particular small black seed was. When Pern-
answered tliat it was sagebnish, the ranger took
him to task for planting a weed. Perrv acknowl-
edged that he, himself, had spent much of his
career tning to rid western lands of that plant
but pointed out that it was neeck'd for v\ikllife
food and habitat. Thev were on a bciuli above
a vallev. Below them was recentiv cleared land
that had been choked with a thick stand of
sagebnish. Pern- pointed out that there were
good HMsons to do both: thin sagebrush stands
and plant sagebrush.
Pern had the vision to understand the useful-
ness of all plants v\ithin acommunitv. He .sought
to include the use of less common but important
taxa, including buckwheat, globemallow, and
smooth aster. He understood that plants sene
main- important functions in addition to forage.
He stronglv supported management and resto-
Great Basin Naturalist
[Volui
ration efforts needed to improve disturbed sites.
His standing, knowledge, and ahilit)' to work
witli different people were extremely helpful to
federal and state land management agencies as
the\ attempted to balance livestock grazing
pressure with earning capacity- of rangelands.
He was particularh' interested in presenation
and stud\- of natural plant communities. He
worked to maintain the exclosure facilities of the
Great Basin E\i)erimcntal Range and provided
numerous plant vouchers for herbaria.
His work with shnib management and values
was important in garnering support for constmc-
tion of the Shnib Scic^nces Laboratoiy. V. L.
Haiper, retired Depuh (^hief for Research,
Forest Service, sent me a letter in 1985:
... I wa.s dding ;i Rcsearcli In.spcction of the Iiiter-
moiiiitaiii Station (about 1960) . . . One of the cen-
ters Director Joe Peclianec and I \isited was the
work on shrub rescarcli. After listenint^ to the Project
Leader's {Perrs's) presentation and \'iewing some of
the Held experiments, 1 turned to Joe and said
"mavbe we ought to amend the Ten-year Reseaich
Program to include a new laboratorv' at Provo . . .
featuring shnib research including genetics, etc."
Joe grinni'd broadk and said "I hoped von would see
this need." He then produced a menx) outlining the
justification for such a laborator\ to be located on the
grounds of Brigham Young Uuixersitv. He further
remarked, "I ha\'e outlined a speech which I can now
cut sliort. <ji\in<i a big yiitvli for the lab."
Tlic laboratorv was completed in 1975 (Stutz
1975). FeriA and his colleagues saw great oppor-
tunities and benehts in v\ ildland shnib research
(Van Epps et al. 1 971 , McKell et al. 1972). Some
of their vision has been realized (McKell 1989),
one piece of evidence being a viable Shrul)
Research ('onsortium (Tiedeman 1984) head-
quartered at tlie Shnib Sciences LaboratoiAand
involved with v ital ongoing activ ities ( McArthur
1990).
I was fortunate to v isit PeriA about two vv(>eks
b<4bre he died. He was at home between hospital
stavs. It was pleasant to update him on lab
activities. He talked about his friends and col-
leagues who had gone on before and e.\press(>d
the view that his time was near. Later, as I drove
home, I reflected through mistv^ eves the good
fortune I had of knowing and being mentored
bv the man. .\hmv share this view.
AcKNow i,Ki)(;\n:\Ts
1 thank Clyde Blaucr, Kim Ilaiper, Steve
Mon.sen, Blanche Plummer. and Rich Stevens
for u.seful comments on an earli(>r version of this
memoriain.
Literature Cited
Bi.AUKH. A. C>"., E. D. McAhtiu H. R. Stevens, tmd S. D.
Nelson In press. E\tJnation of roadside stabilization
and beautification plantings in south centriJ Utah.
US DA Forest Senice Research Paper. Intermountain
Research Station, Ogden, Utah.
Blai KH. A. C, A. P. Pllmmeh. E. D. McAirrm. h, R.
Stem-ins. tuid B. C. Giln ta 1975. Characteristics and
hybridization of important Intermounttun shrubs. L
i^ose familv. USDA Forest Service Research Paper
INT-169. Intermountain Forest and Range Experi-
ment Station, Ogden, Utah. 36 pp.
. 1976. Characteristics iuid h\iiridizati()n of impor-
tant Intermountain shrubs. II. Chenopod famil\'.
USDA Forest Sendee Research Paper INT-177. Inter-
mountain Forest and Range Experiment Station.
Ogden, Utah. 42 pp.
Bleak, A. T, and A. P. Plimmkr 1954. Grazing crested
wheatgrass bv sheep, [ournal of Range Management 7:
63-6s'.
Cahlson. J. R., and E. D. McAUTiiUK, EDS. 1985. S\mpo-
sium on range plant improvement. Pages 107-220 in
Proceedings: selected papers presented at the 3Sth
Annual Meeting of the Society for Range Manage-
ment. Society for Range Management, Demer, Colo-
rado.
Claiskn. J., D. D. Keck and W. M. Hiesev 1940. Exper-
imentiJ studies on the nature of species. I. Effect of
\aried environments on western North Americiui
plants. Cai'uegie Institution of W'ashington Publication
520. W'ashington. D.C. 452 pp.
Da\ IS. ). N. 1983. Performance comparison among popula-
tions of bitterbnish, cliffrose, and bitterbrush-cliffrose
h\brid crosses on study sites throughout Utah. Pages
38-44 in A. R. Tiedemann and K. L. |ohn.son, compil-
ers. Proceedings — research anil management of
bitterbnish and cliffrose in western North .America.
USDA Forest Senice General Technical Report INT-
152. Intermountain Forest and Range Experiment Sta-
tion, Ogden, UtiJi.
. 1987. SeecUingestablishmentliiologv and patterns
of interspecific association among established seeded
antl nonseeded species on a chained juniper-pinvon
woodland in central Utah. Unpublished doctoral tlis-
seitation, Brigham Young Unixersitv Pro\o, Utali. 80 pp.
Dams J. N., and B. L. Welch 1985. Winter preference,
nutritive \alue, and other range use characteristics of
Kocliid prostnitd (L. ) Schrad. Great Basin Naturalist
45: 778-783.
Di;w EV. D. R., and A. R Plimmeh 1980. New collections
ol range plants from the Soviet Union. )ournal of Range
Management 33: 89-94.
Dkohnk K R., and A. P. Pllmmki! 1966. Progress in
browse h\bridi/,ati()n in Utah. Proceedings, .Annual
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tion of mountain big sagebmsh enhanced through
short-term protection from heav)- browsing, (ournal of
Range .Management 44: 72-74.
WV.i.cii. B. L,, and T I,. C. Jacohsox 198S. Root growth
oi Artemisia thdciilalii ]onrnal oi Range Managcnient
41:332-3.34,
Welch. B. L., and E, D. McAkthuk 1979. Variation in
winter leyels of cnide protein among Artemisia
fridentafa subspecies. Joumd of Range Management
.32: 467-469.
. 1981. Variation of monoterpenoid content among
subspecies and accessions of Artemisia tridentata
grown in a uniform garden. Journal of Range Manage-
ment 34: 380-384. '
1986. Wintering nuile deer preference for 21
accessions of big sagebmsh. CJreat Basin Naturalist 46:
281-286.
Welch. B. L., E. D. McArthuk and J. N. D.wis 1981.
Differential preference of wintering mule deer for
accessions of big sagebmsh and black sagebmsh. Jour-
nal of Range Mimagement 34: 409-411.
. 1983. Mule deer preference and monoterpenoids
(essential oils). Journal of Range Management .36: 48.5-
487.
Welch, B. L., E. D. McArthuk. D. L. Nelson. J. C.
Pederson, and J. N. Davis. 1986. 'Hobble Creek'— a
STiperior selection of low elevation mountain big sage-
bmsh. USDA Forest Service Reseaich Paper INT-370.
Intermountain Research Station, Ogden, Utah, 10 pp.
Welch, B. L., E. D. McArthur. luid R. L. Rodriguez
1987. Variation in utilization of big sagebrush acces-
sions b\ wintering sheep. Journal of Range Manage-
ment 40: 11.3-11.5,
Welch, B, L,, and S, B, Mcjnsen 1981. Winter crude
protein among accessions of founving saltbush grown
in a uniform garden. Great B;isin Naturixlist 41: .â– 34.'3-.346.
Welch, B. L., F. J, Wac;stafe, tuid J, A, Roberson 1991,
Preference of wintering sage grouse for big sagebmsh.
Journal of Range Management 44: 462-465.
WuLFF, E, V, 194.3, An introduction to historical plant
geography. Chronica Botanica 10: 1-223. (translated
from the Russian bv E. Brissenden).
Received 24 Febnian/ 1992
Accepted 3 March 1992
Great Basin Naturalist 52( 1). 1992, pp. 1 1-24
SECONDARY PRODUCTION ESTIMATES OF BENTIIIC INSECTS
IN THREE COLD DESERT STREAMS
1.2
W. L. Gaines ' ", C. E. Cushintr' . and S. D. Siiiitl
Abstiuct. — ^We studied aquatic in.sect production in three cold desert streams in soutlieastem Washington. Tlie
Size-Frequenc\' (SF) and P/B methods were usetl to assess production, wiiich is expressed h\- taxon. functional trroup. and
trophic le\el.
Diptenuis (midges anil black tlies' were the most productivx' taxa, accounting lor 4()-7()'f oltlic total insect ])roduction.
Production b\ collectors and detiitixores was the greatest oi all functional groups and trophic le\ els, respecti\eK', in all stud\'
streams.
bisects with rapid development times and multiple cohorts are \en important in cold desert streams; they were major
contributors to the total insect production. Total insect production rates in our stuil\ streams (14—23 g DW-m'-AT" ) were
greater thiui diose found in Deep Creek, Idalio ( 1.2 g DW-m" yr" ), the onlv other cold desert stream for which production
data are axailable. Our values also were generall)' greater than published data for most cold/mesic (3-27 g DW-m'^-vr" )
and humid/mesic (3-25 g DW-m'"yr' ) streams, but lower than in Sonoran Desert Streams (>120 g DW-m""-\T" ) or New
Zealand streams (—40 g D\\'ni'"\T" ).
Our data support the contention of othcis that production, rather than tlensitv or bioniass, is the most accurate^ and
meaningful wax to assess die role of these organisms in lotic ecosystems.
Kc'tj words: pwdnctiiity, benthos. sprin(^-streaiu.s. cold dcscii. fmictioitnl 'groups, trophic levels, Dijrtera. Tiiehopteni.
Coleoptera. Epiieineroptera, Odonata. Plecoptera.
Coniinunit\-le\el production of iiisect.s has
been assessed in relatively few stream types, and
of all niacroinxertebrates in exen fewer. Partic-
ularh; little is known about secondan' produc-
tion in arid region streams. The only studies of
secondar\- production in arid region streams
that we are aware of are those of Minshall et al.
(1973) in Deep Creek, Idaho, in the cold desert
proxince, and Fisher and Gra\- ( 1983) and lack-
son and Fisher (1986) in S\'camore Creek, Ari-
zona, in the hot desert region.
Secondar\ production is the rate of animal
tissue elaboration over time regardless of the
fate (e.g., cannvorx; emergence) of that produc-
tion (Benke and Wallace 1980). Estimating sec-
ondary' production in a stream provides one
assessment of the role of animals in the ecosvs-
tem (Benke and Wallace 1980) as well as insight
into ecoswstem dxnamics. Estimating onl\' den-
-sity- and biomass. regardless of time, ma\' not
accurately describe the role of organisms in the
stream. For instance, the role of gathering-col-
lector imertebrates was underestimated 1)\ bio-
mass anaK'sis and o\erestimated 1)\ nunuMJcal
analysis in a southeastern stream (Benke et al.
1984). Waters (1977) states that production is
important to imderstanding ecoswstem d\ nam-
ics because it is the means bv which cnergx is
made a\ailable to higher trophic le\els.
While most secondan production studies
ha\ e focu.sed on one or a few species in a stream
(Benke and Wallace 1980, Waters and
Hokenstrom 1980. O'Hop et al. 1984). more
recent studies have estimated secondan- pro-
duction of the entire macrobenthic fauna
(Kmeger and \\aters 1983, Benke et al. 1984.
Smock et al. 1985, Ilumi and \\al lace 1987).
Yet to be integrated into c()niiuuiiit\ -Icnx'I anal-
\-.ses are the Inporheic fauna, proto/oa. and
other microiuNfrtebrates. Thec()nnnunit\-le\el
apjiroach proxides a mon^ integrated insight
into the ecoIogN' of stream ecosvstenis.
11ie purpo.se of this study was to measure the
secondarN' production of insects in three streams
located in the cold desert physiographic pro\-
ince of .southeiisteni Washington. We emphasize
^ Department ol Biolof^cul Sciences. Central Wa.shington Uni\ersit>\ Ellensbnri;, Wiusliini^on 9S926.
"Present addres.s: U.S. Forest Service, l^>a\en\v()rtli Ranger District, Lea\en\v<)rtli, Wiusliinnlon 9SS26.
En\ironniental Sciences Department. Pacific Nortliwesl Laliorator) , Ricliland. Washington 99.3.52.
11
12
Great Basin Naturalist
[\
oiunie oz
TaBI.K 1. Plivsical and chemical cliaracteristics of" stiicK ivaclus in Don^las Cn-i^k, SnKely Springs, and Rattle-snake
Springs, July 19S5 to June 1986.
Stream
I^onglas C-'reek
Snivelv Springs
Rattlesnake Springs
A\'erag('
widtli
(m)
4.0
1.3
1.7
Axfragc
de]itii
0.31
0.10
0.05
Axi'iagc
discliargc
invVs)
0.6
0.04
0.05
i)lSS()Kt'd()2
(nig/L)
9.6-14
8.6-12
8.2-10
T.Mil.K 2. Percent snitstiatnni t\pes in stnd\ reaches of Donglas Creek, Snively Springs, and Rattlesn;xke Springs, July
1985 to June 1986.
Substratum type
Stream
Boulder
(>256 nnn)
Cobble
( 64-225 mm)
Pebble
(16-64 nnn)
(Jraxel
(2-16 nnn)
Sand/silt
{<2 mm)
Douglas Creek
SniveK Springs
Rattlesnake Springs
21
7
29
20
1
24
25
7
16
11
11
10
37
81
(hat the estimates [)ul)li.shed here are, in sexeral
cases, l)a.se(l on assnmptions that we have
explained (see Methods). Given the choices to
which we could devote the available resomx'es,
we chose to prochice an estimate of total insect
production in the.se spring-streams rather than
detailed data on a few taxa. We hope futme
studies will proxide data on growth, CPIs, etc.,
for all taxa in tlu^se spring-streams which we can
then use to refine tlu^ initial estimates presented
heri'.
Study Sitks
This shnih-.steppe region is characterized bv
a climax conuiiunitx' consisting ofbig sage (Aiie-
misia tridentata) and hluebunch wheat<irass
{Aoropijron spicatuDi). Mean aimual precipita-
tion in the area is about 14 cm. The study
stnnuns were Douglas Greek {r>C), SniveK
Springs (SS), and Rattlesnake Springs (RS) (Fig.
1 ). The axerage width, depth, discharge, and
dissoKed oxygen concentration for each stud\
reach are shown in Table f , and the substratum
composition is gi\en in Table 2. Figure 2 shows
the daily and seasonal temperature rang(\s.
Douglas Greek
DG is a spring-fed stream located in Douglas
Gouutx; \\'ashington. It is the largest ofthe three
streams studied, the stream it.self draining an
area of 530 km". Our studv sites were located in
the upper reaches where flow is permanent and
not affected bv irrigation withdrawal. Riparian
vegetation is dominated bv water birch (Bctitld
occidentalis) and peachleaf wallow {Salix
anii/c^daloicles).
Sni\el\' Springs
SS is a small spring-stream located on the U.S.
Department of Energy's Hanford Site, Wash-
ington. It drains an area of approximately 40
km". The lower reaches ofthe spring-stream drv
up during the summer, leaxing about 3.6 km of
perennial flow (Gushing 1988). Riparian vege-
tation is dominated bv cattails (TijpJui kit i folia)
along the upper and lower reaches, and willow
{Salix sp.) and wild rose (Rosa sp.) along the
mid- reaches, where it flows through a canyon.
Watercress {Nastuiiiimi officinale = Rorippa
nastiii'titun-acjuatiniui) grows extensivelv
within the s[)ring-stream.
l^attlesnake Springs
RS is a small spring-stream also located on the
Hanford Site. It drains an area of 350 km"
(Crushing et al. 1980). Portions of the lower
reaches diA up during the summer, leaxing
about 2.5 km of perennial flow. Mean annual
total alkalinit\ (as C^aGO.^) is 127 ppm, and the
spring-str(>am is subject to periodic severe
spates in winter (Gushing and Wolf 1982, Gush-
ing and (xaines 1989). Riparian vegetation is
dominated b\- peachleaf willow and cattails.
19921
Insect Pkoduc:ti\ity in Spkinc;-Stkkams
13
Fig. 1. Stiulx ivachfs: A. noiujas Creek; H. Siiiwly Springs; C. Rattlesnake Springs
14
Great Basin Naturalist
[\blunie 52
O 15-
0)
I 10
CD
Q.
E
I- 5
Y Snively Springs
J I I I L
J I L
A S O N D
1985
F M A M J
1986
Fig. 2. ATimiiil water teiniXTdtiire regimes: Douglas Creek, Sniwly Springs, and Rattlesnake Springs, |nl\ 19S5 to June 1986.
\\atfi-cTes,s is presentl)' the cloniiiiaiit in-.sti-eaiii Mkti K ) DS
autotroph, altlioiigh periph\ton primary pro-
chictioii exceeded that of watereres.s in 1969-70 We sampled seo;iiu^nts of eacli stream repre-
(Ciishiiig and Wolf 1 984 ). senting the various hal)itats that were present.
1992]
Insect Phodi (;ri\ irv in Simunc-S riiKAMs
15
One study reach was sampled in SS and one in
RS, and three reaches were saniphnl in the
larger DC. Samples were taken to calculate an
average standing stock lor each stream to he
used to calculate production estimates. The
sampling scheme was not designed to allow
intrastream comparisons ot production esti-
mates hetween dilTerent hahitats, hut rather to
pro\ide representatixe production estimates ol
the entire stream.
Samples were collected monthly from lul\
1985 through June 19S6. We collected three
samples during each visit. A Portable Inxerte-
brate Box Sampler (PIBS) (0.1 m", mesh size
350 ^.m) was used in DC. A Surber sampler
(0.09 m~, mesh size 350 |xm) was used in SS and
RS because these spring-streams are too slial-
low for a PIBS. Samples were taken to a depth
of 10 cm and presened in 70% eth\l alcohol.
Insects were separated from organic debris b\
sugar flotation (Anderson 1959) and sorted by
taxa. Insects were identified to the lowest taxo-
nomic level possible and counted, and bod\
length was measured to the nearest 1 mm using
a microscope and ocular micrometer. The tro-
phic status of each taxon was determined bv
examining gut contents (Gaines et al. 1989) or
b\- reference to Merritt and Cummins (1984).
Biomass was determined as dn' weight (DW)
for all size classes after dning at 60 C for 24 h
and weighing to the nearest 0. 1 mg.
The Size-Frequency (SF) method (Hviies and
Coleman 1968, Hamilton 1969, Hynes 1980,
Waters and Hokenstrom 1980) was used to
(estimate secondare production of the most
common taxa. An average SF distribution was
determined from montliK' sample sets; these
represented the sunixorship cune of an "axer-
age cohort" (Hamilton 1969, Benke and Waide
1977); "zero" xalues xx'ere included xx'hen calcu-
lating densities. Production xxas estimated bx
calculating the loss between succ(\ssix-e size
classes and then multiplving the loss bx the
number of size classes using the etjuation gixen
bx Hamilton ( 1969). Production estimates xx'cre
rehned by multiplying by 365/CPI (Cohort Pro-
duction Interval; Benke 1979).
We fovmd that conducting groxxth studies lor
all taxa present xxithin each of the streams xxas
not practicable. To establish reasonable (\sti-
mates of larxal dexelopment times and CPIs, xxe
followed the example of Benke et al. (1984),
xvho u.sed axailable life-histon- data and field
data to estimate CPIs. We used three major
sources of information to estimate CPIs for each
taxon in our study streams. First, xve surxeyed
[\\r ax ailable life-histor)' data gathered from lit-
erature reviexx's and extrapolated the results to
applx' to our situations. Second, xxe made field
obsen'ations to dctcruiiue presence/absence of
taxa and collected size-lre(juencv information
for each taxon to estimate larval development
times and (>PIs. Lastlx; xve conducted in situ
groxxth studies for Bactis sp., Clicuiiuifopsi/che
sp., and Sintulijini s[). to alloxx fuiilici- refine-
ment of our CPI estimates. These groxxth stud-
ies inxolxed placing insects xxithin groxx'th
chambers in RS. Chambers xx'ere constructed
xxitli mesh netting on each end to alloxv water
and food material to pass through. Measure-
ments xx'cre taken and dexelopment times
recorded to estimate CPIs. Using the combina-
tion of all these data sources, we feel confident
that our CPI estimates are reasonable apj'jroxi-
mations.
Production/Biomass (P/B) ratios (Waters
1977) xxere used to estimate secondan produc-
tion for less-abundant taxa. These P/B ratios
xx'ere either ta.x()n-specific xalues derixed from
the study streams or an assumed cohort P/B
xalue of 5 (Waters 1977, Benke et al. 1984).
These taxa xx'ere not present in sufficient num-
bers to proxide an accurate SF distribution
cune that is necessan to compute SF produc-
tion estimates.
RKS LILTS
Production calculations for DC, SS, and RS
are gixen in Tables 3, 4, and 5. respcH'tixclx-. The
folloxving text describes some ot the assmnj)-
tions xve u.sed in our calculations, data support-
ing the.se assumptions, and other information
relexant to the production calculations. .All pro-
duction estimates, unless noted otheivxise, are
gixen in units ol iiig DW-m" xr .
Douglas ( ,'rcek
Fpih:MEROFT1:u.\. — Maxilies txpically exhibit
xxidelx- xaried laival dexelopment times (Clif-
ford i982). Clifford (1982) examined life-cycle
data of 85 species of Heptageniidae and found
that >909f had at least one unixoltine cycle.
Field data for Baetis sp. in DC proxided little
clarification of the CPI. Based upon field data
oi' Baetis sp. from RS and SS, and agroxvth study
in RS, xx'e estimated a CPI of 60 d. Similar
temperature regimes in DC and RS support this
16
(;heat Basin Natuhalist
[Volume 52
TaHLK .3. Annual production ofinsects in Douglas Creek, JuK 19S5 to June UlSfi.
(.'alculation
365/C:Pr' method X/in"
B
Annual
production
SE CV (mcrDW/m-) SE C\' (lus; DW/nr
Ephemcroptera
Bart is sp. (jjc. D)''
F(ir(ilq)toplilehi(i sp. (gc, D)
U'ucwctita sp. (g. ID
Tricon/tluxlcs sp. {gc, D)
TOT.M.
Odonata
An^id tibialis (,p, (;)
Plecoptera
Isopcrlii sp. (p, C')
IVichoptcra
lli/dropsi/cltc sp. (fc, D)
Chatmatopsrjchc sp. (fc, D)
LcucDtriclua pictipcs (g, H)
TOT.M,
Coleoptera
OpfiosciTus sp. (g, II)
Diplera
Cliiniiioinus sp. (gc, D)
Siinitliuiu sp. (fc, D)
ParautcthiHiH'inns sp. (gc, D,
Chdctodadius sp. (gc, D)
Hcloiiclla sp. (gc, D)
Tipulidae (s, D)
Pluiciiospcctrd sp. (g, ID
Poh/fK'diluiii sp. (s, II)
Tahanidae (p, C)
Tlii('itcnuimiii)ii/ia sp. (p, C)
Brillia flaiifrotis (s, D)
Enipididae (p, (>)
ToT.M.
Gk.wo Total
6°
r
9°
r
1.5°
12°
15°
1.5'^^
1.5'^
r
9°
l.S°
1°
15°
1.5°
15"
.SF'
SF
SF
PBd
PB
SF
SF
SF
SF
SF
PB
PB
SF
SF
SF
PB
PB
SF
PB
PB
PB
PB
2416 0.41 92.4
225 0.35 7S.5
IfiO 0.47 104.0
(i 0.80 1.59.2
2S()7
.30 0.46 103.9
77 0.5S 129.4
445
1.56
95
696
753
41
196
115
141
37
60
1451
9.3S3
0.57
0.53
0.63
127.1
118.3
139.7
0.71
0.75
0.44
0.57
()..52
0.37
0.07
0.69
0.48
0.81
0.25
0.22
1.52.3
168.6
98.0
127.8
116.4
82.5
15.5
1.54.5
106.6
180.5
.55.0
50.0
263.7
48.1
51.4
1.7
364.9
8.9
42.8
413.5
84.1
7.7
505.3
4.322 0.37 83.5 606.7
60.7
31.2
10.4
3.5
4.5
82.1
4.9
2.2
27.8
0.9
0.9
0.1
229.2
17.57.8
0.41 91.9
0.38 85.4
0.51 104.0
0.67 151.0
0.49 1 10.3
()..58 113.9
0.65 145.8
0.60 1.35.0
0.68 153.2
0..36 80.0
0.69
0.72
0.46
0.66
()..54
0.48
0.07
0.78
0.48
0.83
0.26
0.18
1.53.8
1.36.1
101.9
129.4
1 16.5
103.1
15.0
129.1
107.5
185.4
.57.4
40.0
S320
249
238
884
44
183
1700
818
32
2550
2160
4920
1680
875
426
423
411
221
161
1.30
75
68
8
9358
2,3219
Annual
P/B
31.5
5.2
4.6
45.0e
5.0''
4.3
4.1
9.7
4.2
3.6
81. If
54.0'
84.1
121.7
94.0
5.0"
45.0''
73.1
S.O''
83.6*
75.0''
75.0"
'Sdurcc of (.-pi used; ° = <l,-n\.-(l fiuni ..^nmlli sliuliis, + =
otlitT S(>iirce.s \vf re not iivailal)If)
's = shredder. i;c = j;allifriiii;-collector; Ic = lilli'nnij-collrctc
'SK = ])r(xliicli()ii Ciilc:ilalcd 1>\ llic Si/r-Krci|iiciic\ iiietliod
■'I'B = prodiKlion calculated 1» an ,i.sMiiii<-d IVH n.lio
'.•VssiuiiedLoliort P/Holo.
'.VsMiined .iniiu.il I'/H is tlie same as dcnved In SF l,,i diis 1,
d.ila.nidSKdlstnlniti.i.is: .. = liti-r.itiiiv.
= Sra/er/scnii.cT; p = prrdatni' II = lierln
111 ciiii-r.rtlie ntluT si ud\ streams
lusi-dupc.iiCPl t,
ic; D = detntiNore;
iilar eited insects (used when
e.stiiiiatc. Paralcptophlchia .sp. i.s geiu'ralK iiiii-
voltine, haxiug either suninier or winter cycles
(Ciiriord 1982). In DC, however, sea.sonal cycles
coukl not be distinguished. Pairileptopltlchia
were present in DC throughout the studv vear,
and we assumed a CPl of 1 yr. Because of low
numbers of" Tricon/tluxle.s sp., field data pro-
vided little indication of their CPI. McCullough
et al. (1979) reported a 34-d laival development
time for T. iiiiiiittn.s grown in the field at ISC;
therefore, we estimated a CPI of 40 d for
Triconjthodcs sp. because of lower stream tem-
peratures in DC.
OUONATA. — The dam,selfly AroUi tibialis is
inii\-oltine.
Pi .Fcx )PTE RA.— A CPI estimate for hoperla .sp.
could not be made from Held data. Sexeral stud-
ies (Macka\ 1969, Haiper 1973, Barton 1980)
o( Isoperhi sp. showed seasonal variation in growth
rate, but generally their development time was
about 1 yr. Therefore, we assumed a CPI of 1 \t.
TlUCHOPTERA. — Lcticotrichio pictipcs was
uni\()ltin{\ and as SF distributions and field data
indicated, the lanae oxei-wintered as late instars
and emerged in spring. This obsenation is sup-
ported by studi(\s on L. pictipcs in Owl Creek,
Montana (McAuliffe 1982).
COLEOi'TERA. — An accurate CPI estimate for
the riffle beetle Optiosctxiis sp. was difficult to
estimate because few data are axailabie con-
cerning their development times. W'e tlius
assumed a CPI of 1 yr.
19921
Insect PHoniuTiN ity i\ Si'hi\(;-Sthkams
17
Tahi.I-: 4. Ainnial protliiftioii ol insects lidiii Siii\cl\ Spriiuj;s. |uK 19S5 to |mic 19.S(i.
.\iiiiual
(
'alciilatioi
1
B
procliictioii
.\iiiiual
.m5/c;pr'
inctliiKl
Wiii-
SF
(:\
ingDW/iii-
) SF
C\
(mg DWVin")
P/B
Ephemcioptera
B(icti.ss\\ (jic D)''
f-.=
SFc
I.ISS
0.fi2
104,7
1 S5.4
0.55
96.3
7010
37.8
F(iral('j)t(>])lil('hin sp. (gc, D)
V
SF
5-1
0.27
47.5
15.5
0.28
48.2
67
4.3
TOIAI,
1442
200.9
7077
OHonata
.\r<^i/i lihialis (p, C)
r
PB''
22
0.(il
1 06.6
27.8
0.(iS
118.6
139
5.0''
Trichoplera
('liciiiiiiitojisiichc sp. (fe. D)
2+-0
SF
433
0.41
83.0
200.9
0.51
86.9
1300
6.5
Dipltia
SiinulitiDi sp. (fc, D)
12+,°
SF
27fi
0.70
121.3
34.3
0.82
142.6
1880
54. S
Cliironoiniis sp. (gc, D)
15°
SF
412
0.54
93.2
17.1
0.58
99.8
1390
81.1
Tipulidac (s. D)
1°
I'B
25
0.60
103.8
219.2
0.50
87.4
1100
5.0e
Hi'lciiiclla sp. {gc, D)
15°
SF
381
0.40
69.2
9.2
0.37
64.7
.550
60.3
PoUjpcdihini sp. (s, H)
18°
SF
123
0.56
96.2
3.2
0.52
89.1
220
68.6
Cluictochidiiis sp. (gc, D)
15°
SF
92
0.63
108.3
2.7
0.69
120.2
210
77.8
DLxicIae (gc, D)
15"
PB
21
()..55
95.9
1.3
0.(i5
1 1 1 .5
98
75.0*'
Thieii('iiwintii)u/i(i sp. (p, C)
15°
PB
18
0.42
72.3
1.1
o..3;5
57.. 3
92
S3.6''
Talianidae (p, C)
1°
PB
52
0.47
81.5
10.5
0.50
86.4
53
s.o'-
Enipiilitlae (p, C)
15
PB
4
0.15
26.6
0.6
0.12
32.1
45
75.0'^
T( ) I'A! .
1404
299.2
5638
Chand Total
3301
728.8
14,154
.)t(;l>Illsecl:
Ml,
â– 's
other sources uere not a\'ailalile I-
's = slireclder; gc = gathering-collector; fc = niteriiig-collector: [^ -
'.SF = production calculated In the Size-Frequenc\' method
' I'B = ])rothiction calculated In an ;i5sunied 1V15 ratio
' Assunu'd cohort IVB o(5.
Assumed annual IVB is tlie same as ileri\ed In ,SF tor this taxon ii
L.i.i
IS|-,l,sl,,l.„l,.,ns ,
/scraper: II = lierl.i
i)t the other stud\str<-ai
n- - r Ims..I m|...ii( I'I r..r
detriti\(ire; (^ = camix'ore.
DiPTFB.X. — Simiiliiim sp. were not present in
sufficient numbers in DC to calculate an SF
production (\stiinate. The P/B ratio was calcu-
lated 1)\ axeratfing the P/B ratios obtained for
Siinuliiiiit sp. in SS and RS b\- the SF method.
Accurate CPI estimates for (^hironomidae
could not be obtiiined from field obsenations or
SF distribution. Therefore, we derived CPI esti-
mates, as did Benke et al. (1984), and u.sed
growth data from Macke\ (1977). Macke\
(1977) reported lanal development times of 21
d for Chiroiioiniis sp., 13 d for Poli/pcdihim
com idiiin. and 36 d lor Phaenospectra jlavipcs
at 15 (;. CPIs were compensated for slightK
lowx^r a\'era(2;e temperatures in D(> (13 (^) and
eii\irouuienta] stress (e.g., food axailabilitA',
competition, etc.). These P/B ratios seem high
but are comparable to other data \vher(> short
CPIs were used to estimate P/B ratios (Benke et
al. 1984, Jackson and Fisher 1986). Tabanidae
and Tipulidae were assumed to bc^ unixoltine
with a dexelopment time of 1 \r (Knieger and
Cook 1984). This is consistent with the estimate
of a 1-yr development time for Tahaiiiis dorsifcr
in S\camore Creek, Arizona ((wax 1981).
Empididae grew to a maximum .size similar to
nuun of the midges; therefore, a CPI of 25 d was
Snix'cK Springs
EPIIFMFHOITFHA. — (irax' (1981) reported a
lanal dexclopnu^ut time of 20 d lor Bactis
(jiiiUch in Sxcamore Creek, Arizona. Because of
knxer stream temperatures, howexer, Bactis sp.
dex(^lope(l more sloxvlx in all streams in this
studx'. We assumedaCTT of 6()d. ParaJcpto))lilc-
hia sp. xxas present oulx' during the sununer;
thus, xx'e used oulx summer data to ciilculate
production because annual P xxas essentially
e(jual to sinnmer P.
OdonaPA. — Ar<^ia tibkilis was not present in
suffici(>nt numbers to make an SF production
estimate.
TUK.llOPTFHA. — Field data and SF data indi-
cated a bixoltine life ex cle and a CPI of 6 mo for
Chen mat opsijche .sp., the only caddisflx in SS.
Dli'TFIVV. — Becker (1973) reported a lanal
dex clopment time of 13 d for .S. vittatum grown
in the laboratorx' at 17 C. A 30-d CPI xx'as esti-
mated considering loxx-er stream temperatures
18
Great Basin Naturalist
[Volume 52
Tahi.f: 5. AnniiiJ prochiction of insects from Rattlesnake Springs, July 19S5 to June 1986.
Calculation
365/CPr' method N/m
B
SE (:\' (mgDW/m-) SE
Annual
production
CV (maDW/m2)
Ephemeroptera
Bactis sp. (gc. D)'
TricDnjtluxIcs sp. (gc, D)
TOIAI.
Odonata
.Ari^fV/ tibial is {p, C)
Trichoptera
Clicitmatopsijclw sp. (fc, D)
Parapsijclie sp. (fc, D)
LimncphiUis sp. (s, D)
ToiAl.
Cole<»ptera
Hi/ddticiis sp. (p, C)
IKdropliilidae (p, C)
ToTM,
Diptera
Siiniiliitin sp. (fc, D)
Chin)ii(»nus sp. (gc, D)
Helcnielld sp. (gc, D)
Tlii('iicm(tiiiii)nt/i(i sp. (p, (J)
Tahauidae (p, C.)
Misc. C'hironomidae (gc, D)
Polijpcdiliim sp. (s, II)
Cliactorladiii.s sp. (gc, D)
Empididae (p, C)
TipuIidae(s,D)
Di,\idae(gc. D)
TOTAI.
Grand ToiAi.
Annual
P/B
go,.,o
SEc
1336
0.61
107.2
47.3
0.58
104.0
2540
53.8
9"
BB''
1
1.337
0.05
8.3
0.3
47.6
0.07
12.2
14
2554
45.0''
r
BB
67
0.72
124.1
74.3
0.78
134.9
372
5.0''
20. + .0
SF
140
0.69
118.9
48.6
0.78
134.5
486
10.0
1-
PB
10
0.24
41.7
26.8
0.25
43.4
134
5.0"
1
PB
52
202
0.45
76.9
22.0
97.4
0.38
66.3
115
735
5.0''
r
PB
4
0.50
87.4
1.2
()..35
60.1
6
5.0''
r
PB
1
5
0.27
47.6
0.3
0.25
43.1
2
S
5.0"
12°"°
SF
1777
0.73
125.8
212.3
0.73
127.5
11,180
52.6
15°
SF
192
0.50
87.3
7.0
0.58
IOCS
489
69.9
15°
SF
352
0.51
89.0
5.4
0.51
88. 4
480
88.9
15°
SF
114
0.55
94.9
3.3
0.55
95.2
279
83.6
1°
PB
34
0.51
85.6
15.9
0.64
111.0
80
5.()e
15°
PB
IS
0.29
50.1
0.8
0.38
66.3
60
75.0"
1S°
PB
13
0.62
108.2
0.6
0.46
78.9
41
68.6*
15°
SF
59
0.73
126.4
0.4
0.56
97.7
30
75.0
15"
PB
8
0.39
68.3
0.4
0.23
39.8
30
75.0"
1°
PB
3
0.21
35.9
2.0
0.26
44.3
10
5.0"
15-
PB
2
2572
4183
0.28
64.7
0.1
248.2
469.0
0.29
50.0
8
12,687
16,356
75.0"
mlli Nludiis
: + = l!.-|(l ,
;lat.iamlSF,lisliil
.uho„s:„ =
lilrralniv - =
:lMM..l„pn
11 CI'I lors
innUutcanivt
â– 1^ "l-i-
= tilti-ring-o
.>ll<-ct<,r, n
= gruzer/scraper; p
1 = predatiir
■:II =lK-rl,lv„
„v: D = (let
ntnort-: C;
= carnivore.
'Source of CI'I used: " = derived troiii
other sources were not availalile).
s = shredder: gc = gathering-collector;
'SF = prcxliiction calculated l)y the Size-Fretjuencv method
' PB = production calculated by an assumed P/B ratio
'Assumed cohort P/B of 5.
Assumed annual P/B is the same a.s derived by SK for this taxon in oni- ol the otiii'r study streams
and en\'iron mental stress. CPIs of C^hironom-
idae in SS were estimated as thev were in DC.
We iLsed Grays (1981) estimateOf a 1-yr CPI
and nnivoltinism for Tabanidae and Tipulidae.
Dixidae and Empididae reached ma.xinnmi
sizes similar to manv of the midges, and a (>PI
of 25 d was assumed.
Rattlesnake Springs
Ephemeroptera.— We isolated several
Bncti.s sp. lar\ae in growth chambers in RS to
estimate lanal development time. These data
and field data indicated a CPI of 60 d.
Tricon/tlKulcs sp. were not present in sufficient
numbers for an SF production estimate.
OiX)\ATA.— Field data for Arg/V/ tibialis indi-
cated a CPI of 1 vr.
Trichoptera.— We isolated several Chcumafo-
psijclw sp. lan'ae in growth chambers in RS to
estimate lanal development time. These data
indicated a bivoltine life cvcle and a CPI of 6
mo. Because of low densities, field data ga\e no
indication of the CPIs of LiinncpJiihis sp. or
Farapsijchc sp.
COI.EOPTERA. — Field data pnnided little
indication of the (]PIs of beetles because of low
numbers.
Diptera. — Several Siinulimit sp. lanae were
isolated in growth chambers in RS to estimate
lanal development time. As in SS, we used
(irays (1981) estimate of a 1-vr CPI and uni-
Noltinism lor Tabanidae ami Tipulidae. Dixidae
and I'jupididac^ grew to maximum sizes similar
to main ol the midges, and C>PIs of 25 d were
assumed.
19921
InsectPiu)i:)U(:ti\ rn i\ Sfhixc-Sti^kams
19
TvHl,! (i. \iiiiual production (P. nuj; l)\\ -in x r- 1 ' and ])iit<-nt production ol insect tnnctional <4ronps in Douglas C.Vcek,
Sni\rl\ Springs, and Rattlesnake Springs, )nl\ 1SIS5 to |nnc 19S(i.
Functional
''roup
Douglas (Jrcek
Sni\fl\ Springs
7f
Rattlesnake Springs
(;r;i/.i'r/scraper
Collector
(Jatlierer
Filterer
(Totd)
Slui'dder
I'lcdator
(;i{\\i)i()r\i.
2(i51
11.4
0.0
15.2S2
65. ,S
9332
65.9
.3621
22.2
4198
18.1
3177
22.5
11.800
72.1
(19,4.S0)
(83.9)
(12.509)
(88.4)
(15,421)
(94.3)
639
2.S
1316
9.3
166
1.0
449
1.9
329
2.3
769
4.7
23,219
100.0
14.1.54
lOO.O
16..356
1(K).0
TaHI I 7. Annual production ( F, nig DW'in "-nt-D and percent production ol insect trophic le\els in Douglas C^reek.
Sni\c'K .Springs, anil Rattlesntiki' Springs. |uK 1985 to |une 1986.
Trophic
IcN-el
Douiilas CJreek
'Tf
SnixeK Sprinjj
<-/<
Rattlesnake Springs
^f
iirrliixorr
Detritixore
( 'aniix'ore
ToTvl.
2SI2 121
19.967 Sfi.O
440 1 .9
2:>.2I9 1 ()().(>
220 1.6
13.605 96.1
.329 2.3
14.154 100,
4 1 0.3
15.546 95.0
769 4.7
16.356 100.
Functional (yi-oiip Production
Production In collectors Wius greatest of all func-
tional groups in all stud\ streams. ( Collector pro-
duction was highest in DC, 19.5 gin'~\T' ,
accoiuiting for 83.9% of the total annual produc-
tion of insects. In SS and RS. collector production
was 12.5 gaud 15.4 g, representing 88.4 and 94.3'/f
ot the total aruiual production, re.spectixeK . The
annual pioduction of ;i]l ftiuctional groups in each
stud\ stream is sliowu in Table 6.
IVopliic Ije\('l Production
Heihixores and detritixores are both second-
an producers at the same trophic le\el; carui-
xores are teitiarx producers. Fortliis discussion,
we address them .separateK. Detritixore pro-
duction was greatest of all trophic lexels in each
stuck stream. In DC, detiitixore production was
about 20.0 g in'~\T' , accoimting lor 86.()9( of
the total annual insect production. In SS andHS,
detritixore production xxas 13.6 g and 15.5 g,
rejn-esentiug9rs.l and 95. 09^ of"th(^ total amuial
insect production. Herbixores contributed
12.]'^^ ol the productixitx' in DC", but no other
tropliic lexel in anx of the three streams x\as an
important contributor to secondaiA j)roductiou.
The annual production of all trophic lexcls in
each stream is i£ixen in Table 7.
Discussion
Interstream (Comparisons
DC x\as clearlx the most productixe of the
three streams studied (Table 6), and this is prob-
ablx' related to the xaiietx' of substratum (Tal)le
2) and resulting increase in microhabitat diver-
sit)'. Minshall (1984) thoroughlx rexiexxed the
importance of substratum heterogeneitx' and its
influence on insect abundance and distiibution.
SS and HS xxere similar in size and had similar
total productixit\- estimates (Table 6), although
im[)ortant differences existed among the biotic
coniponeMits.
In t(Mnis of hmctional group productixitx', col-
lectors dominated in each of the streams. Gath-
erers xxere more important in DC and SS, and
lilterers in HS. The greater filterer/gatherer
ratio in US is probablx related to the shifting
nature of the sandx' substratum (Table 2) and
resulting absence of areas lor detritus to collect
and be hancsted. The filtering sinuiliids
occurred on the abmidant xxatercress plants.
The scarcitx of solid snbstratimi for periphxton
dext'lopment in HS also explains the absence of
grazers in this stream. Htnxexer, substratum
composition does not explain a lack of grazers in
SS, xx'here solid substratum is present (Table 2).
20
Ghkat Basin Naturalist
[Voluni
In SS, tlie dense riparian canopy almost coni-
pleteK' sliaded and obscured the stream. This
proliahK pre\ented the development ol a sub-
stantial periplutic food base (or grazers. In DC,
which had both solid snbstratnm and unshaded
stream bottom, a significant grazer commnnitx
was present (Table 6).
Comparing die prodnctixit) of taxa common
to all three streams shows some differences that
are difficult to (^xplain (Table 8). For example,
Si)miliitiit sp. production was similar in DC and
SS, but was an order of magnitude greater in RS.
This nia\ indicate a richer source ol suspended
food in RS; howexer, comparatix (^ measure-
ments of this resource were not made, (wishing
and Wolf (1982) report a \alue of L513
Kcal !n'~\r" of suspended POM in RS, but
comparable data are not available for DC and
SS. This value is much less than diat reported
In iMinshall ( 1978) for Deep Creek, a small, cold
desert stream in .southeastern Idaho. Since
SiimtUum sp. production far exceeded that of
auN- other iu.sect in RS (Table 5), competitive
exclusion (Hemphill and C'ooper 1983) max
make it more sncc("sshil in competing for the
limited attachment sites. CJ}eiinuttoj)si/clie sp.
and Paraj)si/c)i(' sp., two filtering Triclioptera in
RS, had a combined production of 620 mg as
compared xxitli Sintiiliinii sp. production of
> 1 1,000 mg. This is a 20-foId difference for
organisms of the same functional group. Except
for Siinnliiun sp., dipteran production xvas high-
est in D(" for Chiroiioiims sp. and Tabanidae,
xvhile in SS. production oi' PoltfpediliDit sp. and
Tipnlidae xxas highest. Tipniidae j)rodnction
increased bx' an order of magnitude from RS to
DC to SS. This max be relatcnl to the relatively
high amounts of particulate organic matter
(POM) found in the study .section of SS (Cush-
iug 1988). Production of Bactis sp. is three to
four times loxx-er in RS than in the other txxo
streams (Table 8).
A likely explanation lor some of the difler-
ences shoxxii in Table 8 is the xxinter spates lliat
occur in RS, but not in SS or DC. These spates,
described by Cushing and (;aines (1989), .scour
die entire streambed, flushing out accumulated
POM and much of the fauna. They occur about
exerx three xears and act as a "reset" mecha-
nism. Because they occur in xxinter xx'hen there
are no oxipositing adults, and because they
scour and eliminate sources for both upstream
migration and doxxTistream drift, thex- must
T.ARi.K (S. (Comparative annual production (mg DWin^-yr-
I ) of taxa common to Douglas Creek, Sni\'el\- Springs, and
Hattlesnake Springs, |ul\ UiS5 to |une ]9Sfi.
Douglas
Sni\'el\
Rattlesnake
TiLxon
Creek
Springs
Springs
Ephemeroptera
Bdclis sp.
8317
7012
2542
Odonatii
.4rg(V/ tibialis
44
1.39
372
Trichoptera
Cliniiii<ifoi>si/clu- sp.
SIS
1298
486
Diptera
Siiiiii/iinii sp.
IfiSO
1879
11.175
Cltinnioiiins sp.
4920
1386
489
Poh/jx'dihiin sp.
161
220
41
Tabanidae
130
.53
80
Tipulidae
411
1096
10
severely limit the potential productixitx of RS.
It is notable that the dominant secondarx* pro-
ducers in RS are the black flies, organisms that
are found in abundance soon after discharge
diminishes (Cushing and Cxaines 1989).
Intrastream Comparisons
DouCiLAS Creek. — Secondan production in
DC xx'as spread over a wider varietv of fimctional
groups (Table 6) and trophic lex'els (Table 7),
exen though it xx'as dominated bx' detritus-feed-
ing collector-gatherers. Cliirononiiis sp. and
Baeth sp. xx'ere the dominant secondan produc-
ers in the stream.
Snix'ELY Sprincs. — In SS, about 50% of the
secondan- production xvas due to Baetis sp., a
tletritus-feeding collector-gatherer; and, as
mentioned aboxe, the grazing component xx'as
absent. Total dipteran production xxas of the
same order of magnitude as that for Bactis sp.
but xx'as spread out among several organisms,
notablx Siinulitdii sp., Cliiro)wmtis sp., and
Tipulidae (Table 4).
RaTTEESXAKK SPRIXCS. — Secondan pro-
duction in RS xxas less dixerse than in the otlnM-
studx' streams, xxith oxer 68% of the production
due totlu^ filtering detritixore Sinuiliiiin sp. The
second liiglu\st produc(M' xxas Bactis sp., but
production was lai' loxxer than the black (lies
(Table 5). The high [)r()duction ol simuliids in
RS can be attributed to the presence of midtiple
coliorts xxith short dexelopment times. Cirax'
(1981) suggested that rajMcl dexelopment max'
be adxantageous in streams subject to spates.
19921
iNsi'Xrr l'iu)i)r(Ti\ iTvix Sphixc-Sti^IvWi.s
21
Tahi.i: 9. (loiiiparatix-e whole stream .secoiulaiA production ol inscct.s (\\ <; l)\\'-m'~\T-l), e.xcept as indicated, in l'i\e
i^eoc-liniatic resiion.s. Streams CTroiiped In' '^eograjihical region, not 1)\ temperature rep;inies.
.Stream
(;c Cr/.sc I'red
Sonrc(
Cold/mcsic
Unnamed, Quebec
Facton' Br., Maine
Sand H., Alheita
Caribou H.. Minnc^sota
BlackliooC R.. Minnesota
No. Branch C^r., Minnesota
Fort R., Massachusetts
Bear Br., Massachusetts
L'Anee (hi Nord, France
Bisbalh" Iniek, Denmark
Huiiiicl/inesic
.Satilla K. Ci-orgia'
Snag substrate'
SancK' substrate*^
Mud substrate'
Cedar R., So. Carolina
Lower Shope Fk., No. Carolina
Upper Ball Cr., No. Carolina
Bedrock-outcrop
Riffle
I'ool
Hot de.seii
S\camore Cr. .Xrizona
New Zealand
Hinau R.
Horokiwi R.
Cold desert
DeepCr., Sta. 1. Iddio
Dougkis Cr.. Wasliington
Sni\el\ Spr.. Washington
Rattlesniike Spr., Washington
5.8"
Haqierl978
12.2
Nexes 1979
O.S"
Soluk 19S5
3.54
().S3 ().fS2
1.3fS
0.14
0.59
Kruegerand Waters 19S3
7.13
1.00 3.53
1,15
0.37
l.O.S
Knieger and Waters 1983
13.23
0.73 5.33
9.43
1.00
2.07
Knieger and Waters 1983
3.3
Fisher 1977
4.8
Fisher and Likens 1973
12.5
(Total detriti\ore
P=P
- Fred.)
2.0
Maslin and Pattee 1981
26.7
1 .3
.Mortensen and Sinionsen 198^3
25.2
64.8
2.9 18.0
49.3 8.1
4.3
21.0
17.9
3. 1
17.9
0.2
8.6
9.2
3.0
0.1
1.0
1.3
0.02
1.4
0.6
6.1
0.6
2.1
2.1
0.6
0.7
5.6
1.4
0.3
1.8
1.0
1.1
7.6
2.4
0.03
3.0
o.:'^
1.9
120.9
38.2
Hopkins 1976
41.5
Hopkins 1976
1.2
Minshalletal.
23.2
0.6
4.2
15.3
2.7
0.4
This stuck
14.2
1.3
3.2
9.3
0.3
This stuck
16.4
0.2
3.6
lis
O.S
Thisstu(k
Benkeet al. 1984
Smoeketai. 19S5
Ceorgian and W'lJkice 1983
Humi and Wallace 1987
Jackson and Fisher 1986
197.3
'S = sliredtlen Fc = filterins-cx)llecf()r; (..l =
'Kmergers onlv.
'Only two species of cliironomicLs.
' F.xprcs.scil per iinil .lira of total stream bott
'T,\prcssi'il pir ijiiil an-., oflialiilal.
Hatlu
C'oniparisons with ()th(>r Strc^im.s
Annual IVB latios langcd Iroiii .3.6 to 121.7 lor
insects from the studvstrcaiii.s. The high animal
P/R ratios are attiibntetl to insects with rapitl
(le\('lo])nient and multiple cohorts (e.ii;., main
(."hironomidae). The annual P/B ratios lomid in
thes(^ cold desert spring-streams arc generalK
lower than those reported 1)\ fackson and I'^isher
( 1986) for Sonoran Desert stream insects and 1)\
Benke et al. (1984) for southeastcM-n hlackwater
stream iiisc^cts. The .Sonoran and hhickwatcr
streams are warmer and insect (k'xclopnient is
faster, resulting in a greater iiumher of cohorts.
Our annual P/B ratios wen^ geneialK hi'^hcr
than reported for northern temperate streams
(Knieger and Waters 198.3). wlien^ cooler
streams result in insect dexelopment at slower
rates with fewer cohorts.
Total ins(>ct i)roduction rates in this stud\-
ranged Irom 14 to 23 g DWuf-N r' and are
compared with \alues for other streams
grouped In geographical region (Table 9). Pro-
duction rates in cold desert streams are well
below the higher \alues found in New Zealand
streams, the ricluM" areas (snags) of humid/mesic
streams in the soiitheasteni United States, and
Sonoran hot ck'sert streams. Howexer, produc-
tion rates in cold desert streams are higher than
those in stre.niis in cold/mesic areas of the
I iiited Stales. These rankings relate to the
int(Maclioii among stream water temperature,
insect deNclopuKMit, cohort production inter-
\als, and other factors. Howexer, it should be
kept in mind that other factors, e.g., geochem-
istn, ma\ be influential in goxerning production
as well as temperature. Production \alues in
oo
Ghkat Basin Naturalist
[y^
52
Rattlesnake Springs, which has a sandy substra-
tum, are comparable to the sandy areas of the
Satilla Hi\er in Georgia (16.4 vs 13.1 g
DW ni'"\r"\ respecti\eK); production of col-
lector-gatherers was identical.
Benke et al. (1984) stated that measurement
ofsecondan' productivit)' ofbenthic organisms
pnnides a tnier indication of their importance
in lotic ecosNstems than does measurement of
either den.sit\- or biomass. This is intuitively
rea.sonable since measurement of P, a rate,
includes consideration of both biomass and den-
s\t\: Our results support the validit)' of Benke et
al.s (1984) cf)ntention. (>learly, our data reveal
that collectors are the dominant hmctional
group, and detiitixores the dominant trophic
le\el in terms of die secondan producti\it) of
insects in the.se three streams (Tables 6 and 7).
If onK biomass or (k'nsit>' data are evaluated
from these streams (Tables 3, 4, and 5; Gaines
et al. 1 989 ), anomalies become evident. Density-
data in DC re\eal that herbivores are ecjualK' as
numerous as detntivor(\s, but biomass data
re\eal that detritixores are about two times
greater than herbivores. Conversely, when the
insects are separated into functional groups, the
bicnnass of grazer/scrapers (herbivores) exceeds
that of collectors in D(] h\ a factor of two.
Further, collector-filterers in DC; represent
18% of the production and 30% of tlie biomass,
but onl\- 7% of the densit\'. In SS, trophic level
comparisons reveal that detritix'ores dominate
production, biomass, and densit); but if hmc-
tional groups are compared, biomass data would
oxereniphasize the importance of shredders
(30%), wliich form onl\ 5% of the densit)- and
9% of total production. In HS, the largest anom-
aly appears when comparing functional groups.
.Although collector-filterers represent 72% of
the total production and 61% of the biomass,
tlu^ir densit)' is similar to the collector-gathenMs.
In c-onclu.sion, we ha\e found that taxaw id I short
(k'x-elopment times and multiple cohorts, sucli as
midges and black flies, are important to cold
desert .spring-stream production. Pre\-ious studies
ha\-e addressed the difficulties in obtaining accu-
rate field estimates of Simuliidae (black flv) and
( :liironomi(kie ( midge) lanae CPIs, and duis pro-
duc-tiou estimates (BcMikeetal. 1984, Beginner and
Hawkins 1986. Stites and Benke 1989). Their
small si/.e, rapid turnoxer rate, high densitx, and
dixei-sit)' make accurate species-.specific CPI esti-
mates difficult. These same characteristics, how-
e\er, make midges antl black flies vetv importtmt
to stream communities in terms of production.
In nianv streams, thev contribute a large per-
centage of the total community production
because of their rapid development and liigh
timiover rates. We found high P/B ratios for
siniuliids and chironomids, but other inxestiga-
tors luue reported similar results (Fisher and
Gray 1983, Benke et al. 1984, Stites mid Benke
1989). This life-liistory strateg\- is particularK-
advanta<2;eous for insects inhabitins; the streams
that are subjected to severe spates.
Detritus is the major food resource in these
small streams; collector-gatherers predominate
where there is more substratum diversit\- (DC
and SS), and filterers in svstems more prone to
the effects of spates (RS). Grazer/scrapers are
present whenever suitable substratum and suf-
ficient sunlight are available for development of
a peripli)ton crop. Shredders, surprisingh-, are
not well represented in these small headwater
streams. This may be related to the flushing of
the systems b\' the spates and/or the low
amounts of allochthonous detritus reaching the
streams (Gushing 1988). Secondaiy productiv-
ity of these cold desert spring-streams was less
than that of streams in hot deserts, but generally
higher than that in most cold/mesic and
humid/mesic .streams. FinalK', our results
underscore the contentions of Benke et al. (1984)
that measuring the secondan production of
in.sects in streams piTnides a better iissessment of
their role than densitv or bioniiiss, but the anom-
alies described abo\-e argue for care in appKing
this genenilization to all streams.
Ac K N ( ) \ V L E D C; M E N T S
This paper represents a portion of the thesis
submitted b\-WLG to Central Washington Uni-
\'ersit\- for the M.S. degree. The research was
pcM-formed at Pacific Northwest Laboraton-
during a North.west C^ollege and Uni\-ersit\-
Association for Science (NORGUS) Fellowship
(Unixersitv of W'ashington) to WLG. It was
funded mider Contract DE-AM06-76-
KL()2225 and was supported b)- the U.S.
Department of Energv' (DOE) under Contract
DE-AC()6-76RLO 1830 bet^veen DOE and
Battell(> Memorial In.stitute.
We would lik(^ to thank Dr. William Coffman
for identif\ing the chironomids, and Dr. Pat
Scliefter for identifN-ing the caddisflies. The
manuscript was impro\ed b\' comments from
three anonxnious rexiewers; our thanks to them.
1992]
Insect Phoductin ity in Sphin(;-Sthi£ams
23
LlTERATURK ClTKI)
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budget of Rattlesuiike Springs. Washington, .\iiicrican
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. 1984. Piimarv production in Rattlesnake Springs,
a cold-desert spring-stream. Hydrobiologia 114: 229-
236.
GisiiiNc;, G. E., C. D. Mclntiue. J. R. Sedfli. )>:. W.
GUMMINS. G. W. MiNSHALL. R. G. PETERSEN, and R.
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Fisher. S. G. 1977. Organic matter processing by a stream-
segment ecos\stem: Fort Ri\er, Massachusetts. U.S.A.
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701-727.
Fisiii i; S. (;.. and 1,. (.',. GnAV 198.3. SecondarvproductioTi
and organic matter processing l)y collector macro-
invertebrates in a desert stream. EcologN' 64: 1217-
1224.
Fisher, S. G.,andG. E. Likens 1973. Energv- flow in Bear
Brook, New Hampshire: an integrati\e approach to
stream eco.svstem metabolism. Ecological Monographs
4.3: 421-439.
Gaines, \V. L.. C. E. G\siiiN(;. and S. D. Smith 1989.
Trophic relations and functional group composition of
bentliic insects in three cold desert streams. South-
western Naturalist 34: 478-482.
(Jl.oHci \N r.. and |. 15. WALLACE 1983. Seasonal produc-
tion (Knaniics in a guild of periph\ton-grazing insects
in a southern Appalachian stream. l']colo_g\' 64: 12.36-
124S.
(;i;\v L j. 1981. Species comjxjsition and life histories of
aquatic insects in a lowland Sonoran desert stream.
\merican Midland Naturalist 106: 229-242.
11 win.roN, A. L. 1969. On estimating annual production.
Linmol<)g\' and Oceanograph\' 14: 771-782.
IIaiU'ER. p. p. 197.3. Emergence, reproduction, and growth
of setipalpian Plecoptera in southern Ontario. Oikos
24:94-107.
. 1978. Variations in the production of emerging
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Intemationalen Wninigun fiir Limnologie 20: 1.317-
1.32.3.
llFMi'iULi. N., and S. D. GoooPER 1983. The effect of
ph\ sical disturbance on the relatixe abundances of two
filter-feeding insects in a small stream. Oecologia .58:
37,8-.382.
Hopkins. G. L. 1976. Estimate of biological production in
some stream inxertebrates. New Zealand Journal of
Marine and Freshwater Research 10: 629-640.
Hi HVN. A. D., iuidj. B. Wallace 1987. Local gcomorphol-
ogy as a determinant of macrofaunal production in a
mountain stream. Ecokjg)' 68: 19.32-1942.
Hynfs, H. B. N. 1980. A name change in the sec-ondary
production business. Limnolog\- and Oceanography
25:778.
IIVNES, H. B. N., iuid M. J. Goi.E.MAN 1968. A simple
method of assessing the aimual production of stream
benthos. Limnologv- and Ocetuiography 13: .569-573.
J.vcKsoN |. K., andS. C FisiiER 1986. SeconcLuy produc-
tion, emergence, and export of aquatic insects of a
Sonoriui desert stream. Ecolog\ 67: 629-638.
Krlfc;er, G. G., and E. F. GooK. 1984. Lifecycles, stanckng
stocks, and drift of some Megaloptera, Ephemerop-
tera, and Diptera from streams in Minnesota. U.S.A.
.\quatic Insects 6: 101-108.
Ki4i FCFH, G. G.. and T. F. Waters. 198.3. Annual produc-
tion of macroinxfrtebrates in three streams of different
water qualitA. Ecolog\' 64: 840-8.50.
Mack.w, R. |. 1969. Aijuatic insect communities of a small
stream on Mont St. Ilihiire, Quebec. Journal of the
Fisheries Research Board of C;anada 26: 1157-11(8:3.
Mackfv a. p. 1977. Growth and dexelopment of lanal
(;hironomidae. Oikos 28: 270-275.
Maslin, J-L., and E. P.vitee 1981. La prockiction du
peuplenient benthique dune petite ri\iere: son e\alu-
ation par la mc'-thode de Hviies. Goleman et Hamilton.
Archiv fiir IIy(lrol)iobgie.'92: .321-;34.5.
.\I( AiLiEEE. J. R. 1982. Behaxior and life histon of
Lcucotrichia pictipes (Banks) (Trichoptera: Hydroptil-
idae) with spi-cial emphasison ca.se reoccupancv Gana-
dian Journal of Z(x)log\ 60: 1.557-1.561.
M( ci LLOLcai. D. A., G. W. Minsiiall. and G. E. Gt'sii-
INC 1979. Bioenergetics of a stream "collector" organ-
ism, Trinmithodcs miinttus (Insecta: Ephemeroptera).
Linuiol()g\ and Oceiuiography 24: 4.5^58.
Merri'it. R. \V., luid K. W. GXm.shns. eds. 1984. An intro-
duction to the acjuatic insects of North America. 2nd
ed. Kendall/Hunt Publishing Gomp;un. Dubuque,
Iowa.
Minsiiall. G. W. 1978. Autotrophy in stream ecosystems.
BioScience 28: 767-771.
24
Gkeat Basin Naturalist
[N'olume 52
. 19S4. Acjiiatic- iiisc'ct-siil)stratiiin rclalionsliips.
Pages 358-400 in V. II. Hc-sh and D. M. Koseiiherg,
eds.. The ecwlog\- of aquatic insects. Praeger Puhlisli-
ers. New York.
MiNsii.M.i.. G. W', D. A. Andkku s. F. L. RosK D. W. Shaw
and R. L. Ni;\\ KLL 1973. Validation studies at Deep
Creek, Curlew \ alley. Idaho State University Research
Memorandum No. 73-48.
MoKTKNsr.N. E., iuid j. L. Simonskn 1983. Pnuluctioii
estimates of the henthic invertebrate conniiunity in a
small Danish stream. Hvdrobiologia 102: 155-162.
Nk\ Hs R. J. 1979. Second;ir\- production of epilithic fauna
in a woodland stream. American Midland Naturalist
102: 209-224.
O'lIoR J.. J. B. \\'ai.1..\(.f.. and J. 1). Haki'NKH 1984.
Production of a stream shredder, Pcltopcrla inaria
(Plecoptera: Peltoperlidae) in disturbed and undis-
turbed hardwood catchments. Freshwater Biologv 14:
13-21.
Smock. L. A., F. (Jilinskv, and D. L. Stoxkbl h\7-.k 1985.
Macroinvertebrate production in a southeastern
United States blackAvater stream. Ecolo'^v fi6: 1491-
1503.
SoiA K D. A. 198.5. Macroimertebrati' abundance and pro-
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Fisheries and Aquatic Sciences 42: 1296-1.302.
SniKs. D. L., and A. C. Bf.nkk 1989. Rapid growth rates
of chironomids in three habitats of a subtropical black-
water river and their implications for P:B ratios. Lini-
nolog)' and Oceiuiograpln- 34: 1278-1289.
\V/\TF,HS. T. F. 1977. Secondarv' production in inland waters.
Advances in Ecological Research 10: 91-164.
Watkks, T F., and J. C. Hokenstrom. 1980. Annual pro-
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og\' and Oceanograph\' 25: 700-710.
Received 1 ]nnc 1991
Revised 1 December 1991
Accepted 10 Jannan/ 1992
CJreat Basin Naturalist 52;
m)-i
25-28
EFFECT OF REARING METHOD OX CIIUKAR SI :R\'1\ AL
BartclT. Slaiidi
laii \. I* lin
|a\ A. Roherson' , and X. I'nil |(
AUSTHACT, — Sun i\al nl adult cliukar-iuiprintcd. >j;anic tarui isil)liii<j;/liiiiuaii-inipiiutcd L and wild ihnkais was c-oniparcd
in three releases (two sites), (.'(jinhiiied results iuilicate similar i/' < .05) sniAJxal lor adult-iniprinti-d and wild cliukars. hut
lower rates (P < .05) for ijanie farm elmkars. With early l)ilia\ ioral conditioning, some potential exists for using captive-
icarcd iluikars to estalilisli new populations.
Kci/ tcoiil.s: cltiikdi: clinkiir rcariiiti. piirlri(hj(\ iinjtiiiitiit'j^. hiluii ior. iii'i}j}(i'^(iti(ni. suiriidi
Captixe-reared game liirds ixdea.sed in the
wild geiieralK liaxe poor .siinixal (CsermeK' et
al. 19(S3, Krauss et al. 1987). A probable reason
is beha\aoral deficiency (Hessler et al. 1970,
HoseberrN' et al. 1987). Hess (1973) reported
that imprinting is indispensable for surx-ixal of
an animal nnder natnral conditions. Tlialer
(1986) and IDowell (1989) obsened imprtned
pr(xlator-a\()idance behavior ot "properK"
imprinted game birds. Postnatal \isnal imprint-
ing as well as embrvonic anditon" imprinting
( Baile\ and Ralph 1975) appear to be important.
Om" objectixe was to e\ahiate snni\al ot cap-
tive-reared (adult chnkar-imprinted \s. conven-
tional game farm-reared) and wild chnkars
(Alcctohs chiikar).
Mi<:tii()13s .\xd Stuidv .\he.\s
.Adnlt-im printed C'hnkars
(;hukar (;ggs were expensed dming the final
week of incubation to recorded adult chiikar
xocalizations. The recordings, from the (Cornell
LaboratoiA of ()rnitliolog\ Libran of Natural
Sounds, appeared to fit the descri])tion of (he
"rally call" described 1 )\- Stokes ( 1 96 1 ) ( rec( )\d(. â– ( 1
\ ocalizations of incubating or brooding hen chn-
kars were not a\ailablc).
The brooding facilitA was a 6.1 x 15.2x2.1-m
room at tlie Brigham Young llui\ersit\' (BYU)
F()ultr\ He.search Unit (Proxo, Utah). Fecnl and
watei" were provided tiiioiigh automatic s\s-
tems, and cliukar habitat was mimicked b\ co\-
cMTUg the floor with gra\('l. small shrubs, grass,
and rocks.
(Jhicks were removed troiu the iucubatoi-
within 5 h after hatching and transferred to the
brooding facilitv' without allowing exposure to
humans. Six adult cliukars were released so that
the chicks could \istiall\ imprint on th(Mu.
When four weeks old, the chicks were allowed
to access a 5.6 x 22.9 x 2-m outdoor pen. Tlic
outdoor pen was xisualK isolated because of its
solid walls and the netting-cox ered top. ("oxer
xx'as proxided bx' grass, small shnibs, and txxo
deciduous trees.
A haxx'k mod(d was passed (ropc/pullex'
.sxstem) ox-erthepen and a dog introduced twice
xxeeklx so chicks could as.sociate adults" alarm
calls xxitli predator pre.stMice.
Came Farm (>'hnkars
(Ihnkars (same genetic stock as the adult-
imprinted birds) xxere rai.sed at the Utah Dixi-
sion of W ildlife Resources (DWR) C^ame Farm
in Springxille, Utali, under conxentional meth-
ods (broock'd in l)ox-tx]-)e brooders, fed and
watered xxith humau conlacl [sibling/hnman-
imprinted], antl moxcd into ni<j;lit pens at lour
xxeeks of age).
Wild Cliukars
W ild chukars xxere trapped iu the Dugwax'
and Hiomas ranges, Utah. 3-5 .August 1989.
^Dcpartiiienl otBotam anil H.iiip' Scirncc. Bns^h.ini Voiins; UniM-rsit) , I'nno. llali S4(i()2.
"Author to whom com'spiiii(lciicc slioiild he ail<h-csst'(l.
â– ^Utah i:)ivisioii ofWildhfV' Hcsourtcs. 1.596 UVsl North Temple. Salt L^ike City. Utah 841 16.
Department of Animal Science, Brigham X'onng University. Pro\'0. Utah 84602.
25
26
GuKAT Basin Naturalist
[Volume 52
Release Site I
Antelope Island, located in the Great Salt
Lake in Da\'is Count); Utah, varies in elevation
from 1282 m to 2010 m. In size it is 24 x 8 km
and co\ers 10,409 ha. Rock)' slopes and grass-
land are the dominant ecological t)'pes. Average
\earl\- high and low temperatures are 38.9 and
-12.2 C, respecti\elv (Jones 1985). Antelope
Island had self-peipetuating and self-sustaining
chukar populations until the severe winter of
1983-84, after which no chukars were obsened.
On 8 August 1989 (release I), 80 chukars from
each group were released, 13 ol which were
equipped with haclqxick-mouut radio transmit-
ters (Slaugh et al. 1989, 1990). On 2 May 1990
(release 111) 65 adult-imprinted, 65 game farm,
and 4 wild chukars were released; 9 chukars in
each captive- reared grf)up and all 4 of the wild
group were fitted with radio transmitters.
Radios were attached to even' fifth bird cap-
tured from the capti\'e-reared groups to reduce
bias from ease (jf capture. All birds were fitted
with patagial tags and legbands. Captive-reared
chukars were 14 weeks old in release I and 22
weeks old in release III. Wild chukars in all
releases were trapped 3-5 August 1989.
Eighteen coyotes (Canis latrans) were
remox'cd from site I preceding the 1990 release.
MortalitN data were recorded dail\ during the
first two weeks, tlu^u weekK thereafter.
Release Site II
Th(> second studv site was the Sterling
IIollowA\ind Rock Ridge area of Spanish Fork
Canyon. This area ranges in elevation from
1470 m to 3057 m, and the dominant ecological
t)pe is mountain brush. Annual precipitation
a\erages between 38.8 cm and 52 cm. Average
yearly high and low temperatures are 40 C and
-30 C, respecti\el\.
On 25 September 1989 (release II), 1 1 birds
Ironi each group were radio-marked and
released at site II. Captive-reared groups were
21 weeks old. Mortalit) was recorded daiK for
t^\'() weeks, then weekK thereafter.
Statistical AuaUsis
Data were anaKy.ed using a Product limit
(Kaplan-Meier) estimator; a k)g rank test was
used to compare sunixal cui-ves (Pollock et al.
1989). Onl) radio-markt>d birds were compared
since their obsenation was not biased b\ ea.se of
approach and proximit)- to release site.
Results
Release 1
All adult-imprinted and game farm chukars
(both radio and patagial tagged) died within
three weeks of release (Fig. 1) with no differ-
ences between groups (P < .05). Wild birds
decreased in number shortly thereafter but
experienced higher sunival rates (F < .05) than
captive-reared groups. Coxote predation was
the principal cause of mortality.
Release II
There were no significant (F < .05) differ-
ences (Fig. 1).
Release III
Mortality was similar (F < .05) for the adult-
imprinted and wild groups but higher (F < .05)
for game farm chukars (Fig. 1).
All Releases
Combined data for releases 1, 11, and III indi-
cate similar (F < .05) suni\ al for wild and adult-
imprinted groups, both having higher (F < .05)
\alues than game farm birds (Fig. 1).
Discussion
During relciLse 1, wild birds mo\ed (juickh' to
high, I'ockA areas, whereas captive -reared birds
remained at lower elexations and sought co\'er in
the sp(U\se vegetation, where they suffered liigh
mortalits'. Immediatelv following demise of cap-
ti\e-reared birds, wild birds began to be killed.
Adult-imprinted and wild birds demonstrated
the greatest fear response to human presence,
whereas game farm birds tolerated approach.
These findings correspond with those of
CsenneK' et al. (1983), who found that red-
legged partridges {Alectoris nifa) displaved
greater fear response toward humans when iso-
lated from them during imprinting. The flight-
ier behaxior of the adult-imprinted chukars
would likeK' proxide more hunting sport than
game farm birds but did not offer sufficient
suivix al ad\ antage under the existing predator
j)r("ssur('.
Adult-imprinted birds appareiitK had a
behaxioral adxantage over the game farm birds
tliat was not ex|oressed in release 1 but was
demonstrat(Hl at release II, apparentK due to
lower prcxlator pressure. Wild chukar m()rtalit^•
was similar for releases I and II.
19921
CiiikAH 1U:ari.\(;
27
RELEASE
RELEASE II
â– Adult imprinted
O Game farm
A Wild
RELEASE
ALL RELEASES
Fig. 1. Chiikar sunival prohahilitN cuncs: i 1 i release I ( Aiitt-lope Islainl. S August-15 Noveniher 19S9) — no difference
(P < 0.5) between game larni and adult-imprinted elmkar.s. hut botli group.s are lower than wild ehukar.s; (2) release II
(Spanish Fork Canyon, 5 Septemher-12 December 1989) — no differences (P < .05) between gronps; (3) relea.se III
(Antelope Island. 2 May-S Augnst 1989) — no differences (P < .05) between adult-imprinted and wild, but lioth groups are
higherthiui game form chukars; (4) all releases — no differences (F < .05) between adult iiiiiiriiited and wild, but k)wer for
iiame farm chukars.
Re.sult.s irom rcle;i.se III iiulicated tliat .sur-
vival on Antelope Island for all groups was
greater than in the prexious \'ear, especially for
the a(lult-ini[)rintecl group. The iinproxcnu^nt
was attributed to predator remoxaf wliieli nia\
he heneficial e\en in establishing transj)huit(Hl
wild birds in good habitat. Season ot the year
ina\ ha\'e affected sunixal as altematixe pre\'
abunchmce and predator location on the island
nia\ ha\e \aried. |()nkel (1934), however,
obsened little difference in chiikar siuAival
related to sea.son of release.
Combined data from all releasees suggest that
captixe-reared chukars can be used to establish
wild populations if gixcn properearlvbehaxioral
conditioning. This stiuK, howe\er, does notpro-
\ ide intorniation on reproductive success.
ACKNOW LFDCMF.XTS
We express appreciation to the Utah Dixision
of Wildlife Resources for project hmding, also
to M. A. Lar.s.son and J. Fillpot (Utah Dixision
of Parks and Recreation — Antelope Island State
Park) and BYU and DWR personnel who
as.sisted with the project, and to C;. C. Pi.\ton
(BYU Statistics Department) for statistical
assistance.
IJTFHATI'I^K ClTFD
H\ii i;v K. D., and K. M. Rm.imi. 1975. The effects of
embiAonic exposure to pheasant \ociJi7,ations in later
call identification bv chicks. Canadian |ournal of Z(X)I-
og\ 53: 1028-1038.'
28
Ghkat Basin Naturalist
[N'olunie 52
CSF.HMKLV. D., D. Mainakdi. andS. Si'ano. I9S3. Escape-
reaction of captixc \oung recl-Iej;j;ecl partridges
{Ah'ctoris nifa) reared with or without \isual contact
witli man. Applied Animal Etholog) II: 177-1S2.
DouKl.l. S. 19S9. Hearing and prcdation. The Game ('on-
senancN Amiiial Ri'\iew 20: <S.'>-<S8.
Hkss. E. H. 1973. Imprinting: earh experience and tlie
developmental ps\chohiology of attachment. \an
N'ostrand Heinholcl Company New York. 472 pp.
IlKssLKU. E.. J. R. Tkstkk, D. B. SiMFF. and M. M.
Nlll.SON 1970. A biotelemetr\- study of sunixal of
pen-reared pheasants released in .selected habitats.
Journal of Wildlife Management ;34: 267-274.
Jones. C. D. 1985. A manual of the vasculiu- flora ol Ante-
lope Island State Park. Da\is Co., Utah. Unpublished
master's thesis, Brighani Young Universits, Provo,
Utiili. 101 pp.
JoNKKL. C. M. 1954. A comparative stuck oi I'all and spring
rele;ised chukar partridges (Alcctoris 'graced (â– hitkarl
Unpublished master's thesis, Montana State Unixer-
sit\. Bo/.eman.
Kkaiss. G. D., II. B. (;ha\ KS and S. M. Zf,h\ anos I9S7.
Sur\ival of wild and game-farm cock pheasants
releiised in Pennsvl\ania. Journal of Wildlife Manage-
mt'nt 51: 55.5-559.
F()l,l.()( K K. II., S. R. WiNTFHSTFIN. C. M. BUNCK, and
P. D. Gl inis 19S9. Sur\i\al imalvsis in telemetn' stud-
ies: the staggered entn' design, foumal of Wildlife
Mimagement 53: 7-15.
RosKBKHHV. J. L.,D. L. ELF.swouTii.aiKl W. D. Kli.mstka
1987. Comparative post-release behavior and survival
of wild, semi-wild, and game fiu^m boliwhites. Wildlife
Society Bulletin 15: 449-455.
SiAi c;ii. B. T, J. T Flindfhs. J. A. Rohfkson. iuid N. R
Johnston 1990. Effect of backpack radio trimsmitter
attachment on chukar mating. Great Basin Naturalist
50: 379^80.
Slai(;if B. T, J. T Flinufhs. J. A. Robekson M. R.
Olson. ;md N. P. Johnston 1989. Radio transmitter
attachment for chukars. Great Basin Naturalist 49:
632-636.
Stokfs. a. W. 1961. N'oice antl social behaviour of the
chukar partridge. Condor 63: 111-127.
Thalfi^ E. 1986. Studies on the behavior of some
Plia.sianiddc chicks at the Alpenzoo — Innsbnick. Pro-
ceedings of the III" International Symposium on
Pheasants in Asia 1-12.
Received 12 June 1991
Accepted 14 Jdniuin/ 1992
Cicat Basin Naturalist .52; 1 i. 1992. pp. 29-,34
DNA EXTRACTION FROM PRESERVED TROUT TISSUES
D. K. ,Slii<)/a\\a'. j. Kudo'. H. 1'. Evans', S. K. Wocxiwaid-. am! li. \. Williams'
Absth.act — \\V ha\t' ailaptcil t('cliiii(|iics cli'vclopt^d lor the cNtrai-tioii ol l)\,A iroin toniialin-rixcd. paiairiii-iiiihcddi'd
liuinan tissues tor use on presened fisli tissues. DNA was successfullv extracted and tlu' d-loop region ol niitochonilriai
]^N.\ was amplified with the poKinenise chain reaction (PCR). The setjuences ofthe amplified DN.A from preserved and
inockru sainplts wen- identical. These teclinicjues were also applied t(j lin tissue treated with a \ariet\' of preser\ati\es.
Ivxtractiou ol !).\.\ irom ethyl alcohol and air-dried fin tissues gave vields e(jui\;ilent to those from frozen tissues. Extraction
of DNA from presen'ed museum specimens of rare or extinct taxa could significantK' increase the scope of s\ stematic and
jilnlogeuetic studies. Similarlw extraction of DNA from tin tissues proxides a nonlethal sampling strategv allowing
InoiheuiiciJ s\ stematic anaKses ol rare or endangered taxa.
Kcii uord.s: I)\A s(vy//(7ir/;(i^. jHtlijincnisc cIkiui nddioii. \(vy;((7/r/;/g, rntthnxit Iroiit. I )nr()rli\ uehus.
As a part ol our onu;oiiitf .stiulie.s ot the .s\steiii-
atics of western salnionicls, niainK' cutthroat
trout (Oncorl}ijiicluis clarki), we were inter-
ested in extracting DNA from presence! fish
tissues. Museum collections contain man\ pre-
sened specimens, usnalK stored in alcohol hut
originallv fixed in formalin. These could repre-
sent a significant resene of information for s\s-
tematics research if the DN.\ could be
successfulK' extracted. In addition, mau\- popu-
lations of western trout are in such low numbers
that collecting fish for systematic studies could
seriouslv jeopardize their sunival. For this
reason we also wanted to e\aluate the applica-
l)ilit\ of presened-tissue DNA extraction tech-
ni(|U('s to samples of fin tissue. Fin samples
could be taken rapidK' in the field with minimal
str("ss to the fish. These samples could (lien be
])re.seiA('d lor later I^N.\ extraction.
Medical researchers lia\c developed tech-
iiicjues for the extraction of DNA from forma-
lin-fixed, paraffin-imbedded tissues (Coet/. et
al. 1985, Debeau et al. 19S6). The DNA
(extracted from these tissues was of sufficient
qualitNthat restriction cutting and sou tluM'u blot
aiuiKsis were possible (Debeau et al. l9S(ii.
DN.\ has also been successhdK' extracted bom
biiils held in miiseum collections, both tliied
andpre.senedin 7()9( etlnl alcohol (Iloudeanil
Hraun 1988). The DNA extracted from alcohol-
[)resened birds was signilicantK degraded
(maximmn size, 200 ba.se pairs), while that from
the dried tissues contained fragments 9-20 kb
in length. But exen if the DNA obtainetl with
these procedures was degraded, the recent
de\ elopment of the poKmerase chain reaction
procedure (PCR) (Saikietal. 1985, 1988, Mullis
et al. 1986, Mullis and Faloona 1987, Wong et
al. 1987, White etal. 1989) pro\ides a technique
to amplifv specific fragments of DN.\ as small
as 200 bas(^ pairs. Tlu\se amplificxl fragments
can then be se(|uenced to decipher genetic rela-
tionships (Saiki et al. 1985. WVischnik et al.
1987, Kocher ct al. 1989, Thomas and
Beckenbach 1989).
Mati;hi.m.s \m) Mithods
Arcliixcd Specimens
(jdthroat trout collected between 1926 and
1982 and archix (^d in the fish range at the .\h)nte
L. Bean Life Science Museum. Brigham Young
Universitx', were n.sed to determine the uselul-
n(\ss ofthe formalin-extraction techni(|ue when
a[)pliedt() nniseum specimens. Samples of fixer,
nuiscle, or gut were taken from .specimens rep-
lesenting a range of preserxation times (Table
1 ). Tissues were renioxed from the specimens
and placed in 20 xolumes of TE9 buffer (500mM
Tris, 20 mM E13TA, 10 inM NaCl. pll 9.0: Coetz
^ Depart tiieiil olZooloi^', Brii;liaiii X'oniis; Uiiiversilv. I'rovo. IJtali
"Departmenl c)IMien)l>iol()i^ . IJriijhani Yoiini; Uni\i-rsih'. rro\(>. Ulali.
â– Departiiieiil of Biolog). Boisi- State Uiii\crsit\ Biiise. Idaho.
29
30
Great Basin Naturalist
[X'olume 52
TMil.K 1 DNA viekls froin fbmialin-fixed musetim spednieiis ofcuttliroat trout {Oncorhijnclius riarki). DNA \ields were
tlcteriiiiiH'd iisinij l)\' spcctroiiieter ahsorliance readings at 260 niii.
Saiuple
Total
DNA
tissue
weight
DNA
vie Id
Sill
)S[X'C'ic's
^ear
Location
Museum No.
t\pe
(g)
(f-g)
(|jig/mgtis.sue)
o.
c. hoiivirh
1926
Snake R.. ID
BYU #26792
li\c'r
0.13
77.5
0.59(1
o
c. ntali
1927
Utah L.. UT
BYU #26755
ii\(T
0.64
567.5
0.887
o
c. iitali
1940
Utali L.. UT
BYU #26756
liver
0.65
310.0
0.477
o
r. iittilt
1982
Deaf Smith, UT
BYU #176896
uiusele
0.24
147.5
0.615
o.
r. iildli
1982
Deaf Smith. UT
BYU #176890
gut
0.42
965.0
2.298
o.
c. Utah
1928
Trout Cr.UT
BYU #26858
li\-er
0.07
51.0
0.728
o
t: Utah
1981
DeepCr. UT
BYU #176793
muscle
0.11
57.5
0.523
et al. 19S5). The bufft^- was changed twice oxer
24 hours.
Fin Tissues
Fin tissues were taken from anesthetized
hatcheiA rainbow trout {Oncorlu/nchtis i)u/kiss)
that ranged in length from 15 to 25 cm. Samples
were taken from all (ins hut were restricted to
the outer edges of the fins to more accnrateK
represent the region that would he sampled in
the field. ApproximateK 1 cm" of fin was
remo\ed for each sample. These were placed in
labeled 1.8-ml poKetlnlene tubes with gas-
keted screw caps. Four sample's were taken for
each of si.x treatments applied to the fins. These
were (a) 10% formalin, (b) 40% isopropvl alco-
hol, (c) .storage in a standard freezer at -20 C,
(d) storage in an ultracold freezer set at -80 C,
(e) 70% ethyl alcohol ( Et( )H ), and (0 air-dning.
The samples were held in the tubes for 45 da\s,
after which the presenatives were decanted and
the tissues soaked in TE9 for 24 hours, with no
change in the buffer. The frozen and air-dried
.samples were not soaked in buffer piior to
extraction. One sample stored at -20 (] was lost
during storage.
Extraction Pn )cedme
Tissue samples were minced with a clean
razor blade (to 2 mm or less in cross section) and
placed in 15-ml centrifuge tubes with 10 ml of
TEyandO.I gof SDS. Fixe mgofproteina.se K
xvas ackied to each sample, and the tubes were
cappi'd and incubated in a shaking water bath
lor 24 hours at 55 C. An additicmal 5 mg of
proteinase K and 0.1 mg SDS xxere added to
each .sample and the tubes returned to tlu> shak-
ing water liath for 50 hours at 55 C to remoxc
residual undigesteil tissue. The samples xvere
transferred to 30-ml tubes, and an equal xolume
of phenol-chloroform xxas added to each. The
tubes were inxerted sexeral times to mix and
then centrif uged in an SS-34 rotor at 10,000 ipni
for 10 minutes. The aqueous phase from each
sample xxas remoxed xxith an inverted glass
pipette and placed into clean 30-ml tubes and
the procedure repeated. A final extraction of the
acjueous phase xx'as made xvith one xolume of
chloroform and centrifused as liefore. The
aqueous phase from each sample xx^as trans-
ferred to a new tube and .1 xolume of 3 M
sodimii acetate solution added. The mixtures
xvere precipitated xx'ith one xohmie of 95%
EtOH and .stored at -20 C oxemight (12 hours
minimum). Each sample xvas centrifuged at
10,000 ipm for 10 minutes and the supernatant
carefullx poiu'ed off, leaxing a DNA pellet. The
pellets XX ere xxashed xxith 70%- ethvl alcohol and
centrifuged again for 10 minutes at 10,000 ipni.
The alcohol xvas poured off and the samples
alloxx'ed to air drx; The pellets xx^ere resuspeuded
in a 3 mM Tri.s, 0.2 niM EDTA solution (pH
7.2). RNase xxas added to a final concentration
of 20 |jLg/ml.
Results and Discussion
.Archixed Specimens
Mu.scle and lixer tissues xielded comparalile
amounts of l^NA, and exceptionallx high xields
xvere obtained from the sample of gut tissue
(Table 1 ). Because the gut tissue xx'as xx'ashed in
buffer immediatelx after remoxal from the pre-
.serxed specimen, contamination from items in
tlu> alimentaiy- canal should haxe been minimal,
(wit tissue xxas easilx' digested, indicating a rel-
atixely rapid relea.se of DNA (Diibeau et al.
1986), and this coidd haxe been associated xxith
the high xields. DNA samples (20 |xl) from the
museum specimens xxere electrophoresed on a
1992]
D\A FROM Phi:sfr\ei:) Thout
31
B
m % ^ s 9^ fli
Fig. 1. DNA eleetrophoresed on 1% agarose gels after being extracted (Fig. lA) from formalin-presen'ed innsenm
specimens and following PCR amplification ( Fig. IB). The DNA from the trout collected in 192fi ( liver) is only faintk' visible
(lane 1, Fig. lA). The DNA from 1927 (liver), 1940 (liver), 19S2 (mnscle), luid 1982 (gnt) are in lanes 2-5, re.specti\el\-. The
DN.\ in huie 6 was extracted from a contemporary frozen liver sample. The PC'R prodncts are shown in Figure IB. Lanes
1-6 in Figure IB correspond to the D\'.\ tc^nplates shown in lanes 1-fi in Figure l.\.
TvHi.K 2. .\ comparison of the nucleotide sequence (120 ba.se pairs) from the SD-1 region oi the mitochondrial DN.A
il-loop. The DNA was amplified with the polvmerase chain reaction. The top row represents the base sequence from
frozen-tissue DNA, and the lower row represents the sequence from a formalin-preser\ed specimen. The frozen-tissue
specimen (BYU #90621) is O. r. ittah. from McKinzie (]reek, IT, collected S-I7-S8. The preser\ed-tissue specimen ( BYU
#26755) is O. c. utah, from Utah L., UT collected in 1927. Both vouchers are aicliivcd in tin- fish range at the Monte L.
Bean Life Science Museum.
l'"ro/en
l^reservcd
A A c; c; c TAT c; c:
A A G G C T A T C C
A c; c c G A A c; T A
A G C C G A A G T A
C A A T C T T A T T
G A A T C: T T A T T
GGGTTGTGTT
GGGTTGTGTT
T T \ .\ C; A A A G G
T T \ \ G A A A C C
A A(;G ATGTGG
A A G G A T G T G G
C; G G G G T T A G C:
C; G G G G T T A G C:
A TAT C; A G T A C;
A T A T (; A G T A C;
A c; G c; c: g t c a a 30
a g g g g g t c; a a
ttaatg(;tgt 6o
ttaatg(;tgt
gaggaag(:g(; 90
g agg aagggg
ggggtgtggg 120
c; c, G c: T c: T G G G
\7c agarose gel containing etiiidinni hromidc
( Fig. lA) to verify extraction. The DNA samples
extracted from fresh and presened tissne sam-
ples were nsed in a P(>H reaction (25 jxl total
\()hnne) nsing primers for the d-loop region ol
front mitochondrial DNA dexeloped b)- K.
Thomas (Universit)' of California, Berkeley),
with standard conditions (Perkin Elmer Cetns.
Non\alk. (lonnecticnt). C>\cle times and tem-
peratnres wtM-e I iniinite at 92 ( ,', 1 minute at 53
(>. and 2 minntes at 72 C, for 35 c\cles. PCI^
products are showni in Figure IB. DNA extrac-
tion controls containing no fish tissue did not
\ield PCR products under identical conditions
(data not shown). Subsamples of the PCH prod-
ucts from preserved and fresh tissue samples
were secjuenced (Fig. 2) and compared with
contempoiaiA secjuence data from cutthroat
trout (Table 2). Tlie .sequence data were identi-
cal, indicatingthatwithin the amplified segment
no base niodilicafions had occurred in the for-
malin-present hI samjile.
Fin (;lij)s
We obtained DNA from all fin clips regardless
of presenation method. Mean \ields ranged
from a low of 0.40 [xg/mg of tissue from forma-
lin-preser\ed fin clips to a high of 1.104 |JLg/mg
in air-dried samples (Table 3). The treatment
effects were examined with anak sis of \ariance
( Table 4), and a highly significant difference was
found bt>t\\e(Mi the treatments. Fishers least
significant difference multiple comparison pro-
cedure w as applied to separate those treatment
32
Great Baslx Naturalist
[V'«
olunie o'l
B
Fig. 2 (at left). Sequence gel from a portion of the mito-
cliondrial l^NA tl-Ioop. (Joluuin A i.s the .sequence for a
conteinporaiT sample of trout DNA (BYU #90621) and
coluum B is the ,se(juence from a preser\-ed trout specimen
I BYU #26755) collected in 1927. The sequence ge! is read
from the hottoni up, and the colunms represent guanine (G),
adenine (A), tliNininc (T), and c\tosine (C), respectix-elv.
Q.
O
Q.
O
W3
:CM
O
^o O^
00 p
T3
CO
— r"
0.50
0.25
0.75
1.00
1.25
mean DNA yield
{^ig / mg)
Fig. .3. Multiple comparisons of the means of the six fin
tissue treatments, using Fisher's leiLst significant difference
test (alplia = 0.01 ). Lines connect means tliat do not differ
siiruilicautK from one another.
Tabi.K .'3. DN.^ \ields Irom fui tissue presened with dif-
ferent methods. The lin clips, approxiniateh 1 cm" each.
were taken from hatchen -reared rainhow trout
{Onctirhi/iicliiis im/kiss). D\\ \ields were determined
using U\' spt'ctrometer alisorliance readings at 260 um.
I'resen atiou
N
Mean
Stantlard
metliod
\ield
( (JLg/uig)
deviation
formalin
4
0.402
0.15743
40';^ isopn)p\l
4
0.569
0.19111
-20 c:
â– 3
0,644
0.10016
SOC
4
0.740
0.06295
70'7r KtOlf
4
0.S22
0.07964
air-dried
4
1.104
0.13443
a
i;;r()ui),s that clifFei-ccl significantK From one
another. Tho.se compansons (Fig. 3) indicate
that the air-(hi(xl treatment ga\e \ields signifi-
eantlx higliei- than the other treatment.s.
Becan.se the weights used in ealenhiting the
DNA yi(^hls were the preextraetion \ahies and
not the pretreatnient weights, the initial weights
(pre(hAing) of the air-ch-ied samples are not
known. I lowexer, ha.sed on the initial si7,e of the
tin cli})s, tliey are assnuied to \\a\c heen similar".
WTiile air-dnini: \ields ar(> nmch better tlian
19921
DNA Fwnw PRKSEn\i:D Troit
33
T\Hl.l'4. ()iic-\\a\ aiial\sis ol \ariaiuc ot tlic Ihi clip ticatinciit clictt on DNA \i('l(l.
Source
Degrees of
freecloiu
Sum of
scjuiU'es
Mean sfiuare
Prob. > F
iVcatuient
Error
Total! ad j)
17
9.1
1.14512
0.2891 1
1.43424
0.22902
0.01700
I3.4';
O.OOCX)
tlio.si" resiiltiiiti; Iroiii other prcsenatioii iiiclli-
(xls. the lack ol preseniitixes could allow
socoiulaiA foiitaniiuation of samples through
l)aet(Mial or luugal colonizatiou, aud air-dning
prohahK should not be used in collecting sani-
j)les in humid areas or where adequate storage
is not possible. The yields obtained from ethyl
alc-ohol presi^iAation are equal to those from
hozen tissues and superior to both isopropxl
alcohol and formalin presenation. Of the pre-
senati\"es examined in this studx; eth\'l alcohol
would appear to be the preservative of choice in
most field situations. This eliminates the neces-
sit\- of earning drv ice or lic|uid nitrogen into the
field to presene tissues. Other presenative solu-
tions should be considered; for instance, Seutin,
W'liite, and Boag (1991) reported successful DNA
extraction from a\ian tissues presened in a mix-
ture of EDTA, NaCl, and DMSO.
Conclusions
The abilit\ to extract, amplif\; and sequence
D\,\ from formaliu-presened museum .speci-
mens increa.s(^s the inloriuation value of mu.seum
holdings. In addition tol)eingarecordof moipho-
logical and meristic information, the specimens
can l)e u.sed in biochemical studies. Because
museum collections include hpe specimens, rare
spcx'ies, and representatives of now extinct fonus,
many ke>' phylogenetic relationships can be reex-
amined. The extraction techni(|ues can be applied
to contemporan pr(\s(M-\ed tissues as well. Fin
tissues gi\e ade(juate \ields with this techni(jne for
1 )oth restriction enz)'me digestion and P( A\ ampli-
tication. Fin samples, which can be taken nonleth-
alK. present opportunities to examine fish
populations that would othenxi.se be inaccessi-
ble to tissue collection becau.se of management
considerations.
LlTKKATliHK CiTKD
Di:hi:ai L., L. A. Cii wdi.kh J. H. CiUAi.ow. I^. R. Nu.ii-
()l,s. and P. A. Jonks 1986. .Soutliern hlot analysis of
DNA extracted from fonualin-lixed patliologs' speci-
mens. Ciuicer Research 46: 2964-2969.
CoK 1/ S. E., S. R. ri\.\iii.T()N. and B. \'()c;ki.stkin 198.5.
Purification of DN.\ from fornialdelnde fixed and par-
affin embedded human tissue. Biochemical and Bio-
ph\sical Research Conununications 1.30: 11 8-126.
Iloi l)i: P. and M.J. Bk.M N 1988. Museum collections as
a source of DN.V for studies of a\i;ui phxiogein. ,\uk
10.5: 77:^776.
Kociii.ii T D.. W. K. TiioM.vs. A. Mf.ykh. S. \'. Eowahds,
S. I'wHo. F. X. \ ii.i.ABi.ANCA, and A. C. Wn.soN
1989. D\namics ol mitochondrial DNA e\<)hition in
animals: amplification and sefjuencingwith con.served
primers. Proceeding ol the National .Acadenn of Sci-
ence 86: 6196-620().
.Ml l.i.is. K. B., luid F. A. Fai.oona 1987. Specihcs\nthesis
of DNA in vitro \ia a poKinenuse-cataK zed chain reac-
tion. Methods in Enz\inolog)' 1.55: 3.3.5-.3.5().
.Ml 1.1 IS K. B., F. a. Fai.oona, S. Sciiahk. R. Saiki C.
lloHX, and n. A. ElU.lcil. 1986. Specific enzxniatic
amplilication of DN.A in ritro: the poKinera.se chain
reaction. Cokl Springs Harbor ,S\mposinm on Qiianti-
tati\ e Biolog) 5 1 : 262-273.
S.Mki R. K., D. II. Oi'.i.ANn S. Srcnri;, S. |. Sciiakf R.
IlKaciu G. T. IIOKN K. B. Mllijs. and II. A.
Ehi.ICII 1988. Primer-directed enz\niatic amplilica-
tion of" DNA with tlu-nnostable l^N.A polvmerase. Sci-
ence 2.39: 487-49 1 .
S\iKi. R. K., S. Sciiakf. F. Fai.oona. K. B. Mi i.i.is. C.
IIoHN. II. A. Eklicii. and N. Ahnhf IM 198.5. Enz\-
matic amplification of B-globin genomic secjnences
and restriction site aiiaK sis of sickle cell anemia. Sci-
ence 2.30: 1350-1.354.
Skitin. C, B. N.Whitk. and P. T. Boac; 1991. Presi-rva-
tion ola\ ian blood and tissue samples for DN.V analysis.
(Canadian Journal of Zoolog\' 69: 82-90.
Thomas. W. K.. and A. T Bfckfnhacii 1989. N'ariation in
salmonid mitochondrial DN.A: exoltitionan constraints
and mechiuiisms of substitnticjn. Journal ol .Molecular
Exolution 29: 2.3.3-245.
WiiiTF. T. J., N. Aknmki.m. and II. A. Eklicii. 1989. The
poKinerase chain reaction. Trends in (Jenetics 5: 18.5-
189.
W'oNc.C. C. E. Dow I, INC li. K.Smki R. C;. Hick hi II.
A. ElU.lCll. and II. II. Ka/.a/ian 1987, Oharacteri/ii-
tion of B-thalassaemia mutations using diri'ct genomic
secjuencing of amplified single c-op\ DN.\. Nature. 3.30:
3S4-386. '
34
Grkat Basin Naturalist
WiuscnN.K, L. A., R. G. H.glchi M. Stonek.ng, H. A.
EHI..CH, N. AHNHKiM. and A. C. Wilson. 198/.
Length mutations in hum^ mitochondnd DNA:
direct sequencing ol enz)'matically aniplitied DNA.
Nucleic Acids Research L5: .529-542.
[Volume 52
Received 27 ] tine 1991
Revised 10 Febnianj 1992
Accepted 20 Febnianj 1992
Crcat Hasin Naturalist 52( 1 1. 1992, pp. 35^40
RELATIXC; soil. CIIKMISTHY AND PLANT RELATIONSHIPS IN
\\ OODED DRAW S OE THE NORTHERN CiREAT PLAINS
Mumierite E. Nborliees and Daniel W. Urcsk
\.-2
Ahsthact — Soils of till' ijrccn asli/c'liokcclicrn liahitat t\pc in iioitliwcstnii South Dakota were cxaluatcd lor 22
properties to deterniine whether an\ could he correlated with densit\ ol chokeeherr\ il'miiiis vin^iiiiana) ami siiowhern'
iSiiiitplioricdrihts occidcntalis). Siirfaee soils were moderateK teitile, with liiiili levels ol all elements except phosphorus
and nitToij;eu. Soils wfre tine textiH'ed, with uioderateKhigh cation exchange capaeit\' anil saturation percentages. Ilowex'cr,
soils \MH' nonsaline-nonalkaline with low amounts ol exchangeable sodium. None of the soil properties showed good
eonclation w ith ehokeeliern and snow hern densities. (Greatest correlations were loiind between each of the shrub species
Kci/ U(ir(l\: uixxlrd (Imws. <^rccii ash. slinihs. i^runus \irginiana, Sxniphoiieaipos oet-identalis. '^raziiit.
Wooded draws constitute a Naluahic liahitat
(\ |K^ ill the northern Great Plains. The\ pro\ide
shelter from wind and weather and contain
L;;reater moisture than surrounding areas, result-
ing in an abundance of plant life and forage. An
understanding of soil-plant relationships of
tiiese wooded draws has become more critical
since these areas ha\e been obsei^ved to be in
decline (Boldt et al. 1978) for a \"ariet\" of rea-
sons (Girard et al. 1987).
Studies that correlate habitat t\pe with soil
properties are particularl)' useful in efforts to
manage these systems. Knowledge gained from
such studies might help managers determine
(he potential habitat t\pe of a site after \egeta-
tioii decimation. Pfforts and limited resources
could then be concentrated on sit(\s with the
greatest potential for rehabilitation.
This studx' was conducted to characterize the
surface soil chemistiA' of the grecMi ash/choke-
cheriT (Fraxiiiiis pcnnsi/lcanica/pmniis rif^iiii-
(iHd) habitat tA'pe in northwestern South Dakota
and to n^latc^ these soil properties as well as grass
co\cr to (leiisitx ol chokechei'n and snowbern
iSiiinplioiicaiyos occidcntalis). This habitat
type is considered a topographic climax
(Hansen, Hoffman, and Steinauer 19S4.
Hansen and Hoffman 1988) and is one of the
most important in the northern Great Plains.
Si IDY .\Hi: A
The stud\ areaisap])ro\imatel\ 5 miles north-
west of Bison, South Dakota, in Perkins Count\'
on lands administered b\' the USDA Forest
Senice, Custer National Forest. Geologx of the
area has been described In I lansen (1985). The
topography is rolling to stec^p plains dissected b\-
streams and drainagewaws. The climate of the
area is characterized b\ warm summers and \er>'
cold winters. Annual ])recii)itati()n axerages .36
cm, witli most receixcd in the spring and
sunniuM".
The habitat txpes ol the area ha\e been
described l)\ Peterson (1987). The green
asli/chok(X'hern habitat t\pe was found on shal-
low to moderateK dee[). well-drained, Cabba-
Lantn loam soils of upland ridges and the sides
of steep drainagewa\s with slopes of 159^ to
40%.
Mktiiods
Gollection ol Samples
Soil samples were colKx'ted during the
summer of 1986 from 24 green ash/chokechern'
diaw s spaced oxer a 2769-ha pasture. The \eg-
etation ol (he 24 wooded draws ranged from few
trees and shrubs (o a dense ox crstoiy and under-
stonol trees and shnibs. Sampling was conducted
L'SD.X Forest Senice. HockN Mi
:it),, Soudi Dakota .5770 1.
"Corresponding aiitlior.
and Kans;e F.\periMienl Station, Soulli Dakota Seli(K)l ol'Mines and Teclinoloirv . .501 P.. St. Joseph St.. Kapid
35
36
Cheat Basin Natuhalist
[Volume 52
TaBI.K 1. Cheinic-al nrop-itii-s of soil samples collected from ijrceii asii/cliokecliern liahitat h pe near Bison. Sontli Dakota
(n = 72).
Soil
pll
IX'. (mmiios/cni)
Ori^aiiic matter {%)
N0.5-N ((xg/g)
P(m.,u;/U)
Zn (ML,n/g)
Fe (jtg/g)
Mn(fjLg/g)
Cii(|j.g/g)
Ca(meq/I)
Mg (meq/1)
Na (iiieq/l)
SAR
Saturation i%)
CEC(me(i/l(X)kg)
Ext.'Ca(mg/kg)'
Ext. Mg(ing/kg)
Ext. Na(mg/kg)
SaiulC/f)
SiltC/f)
Clav(7f)
Meiui
7.3
0.6
9.1
3.1
2.5
321
3.4
21.2
7.6
2.1
4.5
2.1
0.2
0.1
72.9
45.2
4311
684
15.2
32.9
40.8
26.3
l^aMtre
6..3-7.S
0.4-2.6
4.2-19.8
1.0-17.0
0.1-10.5
202-491
0.9-9.2
6.9-268.0
3.2-24.1
0.,8-5.6
2.0-20.8
1.0-12.5
0.1-0.9
0. 1-0.2
48.8-106.5
29.9-62.4
2580-6830
90-987
1.8-57.5
20-67
21-51
11-40
Standard deviation
0.3
0.3
3.3
2.6
2 2
67
2.0
31.5
3.4
0.8
2.3
1.4
0.1
0.1
11.2
7.6
937
171
I . I
9.1
5.4
6.2
K\tr;iclal>ltc.i(i(
at tliree locations in each draw. At each location
(approxiniatek' 250 m"" in area), three frames
(20 X 50 cm) were randomly located. Stem den-
sities of chokechern at tlu^se locations ranged
from low (0-2 stems/frame), to medium (3-6
stem.s/frame). and high (greater than 8
stem.s/trame). All stems were counted within a
frame and the three \alues axeraged for each
location. Canopy cover of grass was estimated in
each frame (Daubenmin* 1959). One soil
sample was collected within each frame to a
depth of 10 cm. The t]\wc soil samples from
each location were comhiiunl lor chemical anal-
ysis, xielding a total of 72 samples.
Soil .\nal\ses
Amounts of 'soil elements (R K, Zn, Fe, Mn,
(Ju) were determined In' using the annnonium
hicadionate-diethylenetriamine pentaacetic
acid (AB-DTPA) extract (Soltanpour and
Schwab 1977) and iuducti\el\- coupled plasma
atomic emission spectrometr\- (ICP-AES)
(Jones 1977). The AB-DTPA procedure was
de\-eloped and is used by the C:()lorado State
Unixersit) Soil Testing Laboratory An ecjual
amount ol pota.ssium is extracted as with the
ammoniuui acetate test (Knudsen et al. 1982),
antl the same amount of iron is extractcnl as with
the standard DTPA test (Haxlin and Soltanpoiu-
1981). Half as much phosphonis is extracted
using AB-DTPA as in the sodium bicarbonate
extract (Olsen et al. 1954), and slightly less zinc
is extracted than in the standard DTPA test
(Ilavlin and Soltanpour 1981). AB-DTPA
extractable copper and manganese are highly
correlated with DTPA-extractable le\els of
these elements (/•" = .75 and .86, respecti\'ely)
(Soltanpour and Schwab 1977).
The pH was measmed with a pH meter that
used a combination electrode on a saturated
past(\ Sodium adsoiption ratio (SAR) was esti-
mated from lexels of soluble calcium, magne-
siiun, and sodium measured in a saturation
extract In means of ICP-AES. Total soluble salts
were nunisured on the filtered extract with a
solubridge.
Organic matter was (U^ermined b\ wet oxida-
tion with spontaneous heat of reaction. Potas-
sium dichromate and concentrated sulfuric acid
were us(>d lor organic matter, and results were
determintxl calotim(4ricalI\. Nitrate nitrogen
was determined In the chromotropic acid
method. Le\els of extractable Ca, Mo and Na
w ere measured In using ICP-AES on an annno-
niiun acetate extract. Cation exchange capacity'
was determined b\ the .sodium satiuation
method (Page 1982)'.
[992]
Soil. ClIKMlS Tin \\n Fl.AXT Rklatioxships
37
Statistical AnaKses
Simple linear regression was nsed to relate soil
clieniistn \ariahl(^s to cliokecliern and snow-
l)err\' densities; the points were plotted to clieek
tor nonlinear relationships. Stepwise regression
was nsed to test relationships between soil
eheniistn; canop\ eo\(^r of grass, and densit\ ol
each shnil). The regression model Y = a + 1)\'^
pi-o\ided the best fit in relating chokechern and
snow bern densities with canop\- eo\"er of grass.
Soil \ ariables and densities of both shrnbs were
subjected to a nonliierarchieal cluster analvsis
(ISODATA) to group the sites (Ball and Hall
1967). Stepwise^ disciiminant anaKses were
nsed to estimate compactness of clusters and
identifv the ke\ xariables that accounted for
their differences. However, cluster anaKses and
discriminant anaKses and simple correlation
plots did not pro\ide an\- meaningful results.
KHsri;rs .wd Discussion
Nitrate nitrogen lexels averaged 3.0 fxg/g and
ranged from 1.0 to 17.0 |xg/g (Table 1). Soil
organic matter ranged from about 4% to nearlv
2()7c. These \alues compare well with values
tiom surface soil samples from hardwood forest
on fine-textured .soils (Charle\' 1977). Organic
matter le\els ranged substantiallv higher than
tho.se from soils from similar sites in North
i^akota (Han.sen, Hoffman, and Bjugstad 1984),
Montana, and South Dakota (Hansen and Hoff-
man 1988). Nitrate le\els appeared ade(juate
lor growth of rangeland plants ( Soltanpour et al.
1979).
Soils were near neutral in pH (Table 1) and
similar to other sites in Montana, North Dakota,
and Soutli Dakota (Han.sen, HolTman. and
Bjugstad 1984. Hansen and Hoffman 1988).
A\ailabilit\ of nutrients at this pH is near maxi-
mum except for Fe, Mn, Zn, and i'.w. which
l)ecome less a\ailable alxne pH 7.0 (Brad\
1974). Plants nsnalK' grow well bet\veen pH 5
and 8.5 ( Donahue et al. 1977) if no other growth
factor is limiting. Phosphoins and potassimn
content a\ eraged 2.5 jJ-g/g and 321 |i.g/g, respec-
ti\('l\. Thus, phosphorus le\els were low,
whereas potassium, /iuc, copper, and manga-
nese levels were high (both generallv and rela-
ti\e to similar sites in the northern Hi";h Plains
[Hansen. Hoffman, and Bjugstad 1984, Han.sen
and Hoffman 1988]). Iron Itnels a\ eraged 21.2
M-g/g and were fairl\- high.
The cation exchange capacitx (CEC) was
rather high at 45.2 meq/100 kg (Tiible 1 ). Cla\s
in these .soils are likelv to ha\e high adsorptixc^
capacities since organic matter content and cla\
content did not fulK account for the high (>EC
(BracK 1974). The sodium adsorption ratio
(SAB) indic-ated iiiiiiinial saturation ol (he
exchange c-omplex In .sodium. Electrical con-
ductixity was low at 0.6 mmho.s/cm. The.se soils
woukl be classed as nonsaline-nonalkaline with
low ek'ctiical conducti\it\' and exchangeable
sodium percentage. The saturation percentage
at 72.9 was somewhat higher than othcM" nonsa-
line-nonalkaline^ soils in this classification ( Rich-
ards 1954). The soil moistun^ percentage at 15
MPa, which is approximateK* equivalent to the
wilting percentage, was 18%. These soils are
thus relati\el\- fine textured on average. Sand,
silt, and cla\' averaged 33%, 41%, and 26%,
respectiveK'.
Soluble Ca, Mg, and Na were 4.5, 2. 1, and 0.2
me(|/l, respectively (Table 1). Extractable (]a,
Mg, and Na averaged about 431 1. 684, and 15
mg/kg, respectively. These con-e.sponded to 10.8,
5.7, and 0.065 meq/100 g soil luid exchangeable
percentages of 23.8, 12.6, and 0.1, re.spec-tix'elv
Thus, of the.se elements, (Ja wiis predominant on
the exchange complex, and exchtuigeable Na was
\ei"\' low. Howe\er, calcium was low relatixe to
comparable sites of \egf4ation and landsc-ajx\s
(Hansen, Hoffman, and l^jugstad 1984. Hansen
and Hoffman 1988).
Simple correlation coefficients for densitxol
either chokechern' (r = .26 to -.18) or snow-
berr\' (r = .36 to -.20) with various soil proper-
ties were low (Table 2). TweKe soil properties
were negatixcK associated with chokechera'
d(^iisit\. Phosphoins showi^d the greatest posi-
tive relationship with chokechern densitx (/" =
.26). OnK four soil xariables (pH, P. extractable
(>a, and (JEC) were negati\eK correlated with
snowbern' densitv Magnesium showed the
highest coriclation with snowbern densit\' (r =
.36). Soil properties \aried some tor both spe-
cies at the microsite le\el but were not statisti-
calK different (/; < .10). For example, when
densit\ ofchokechern w'iushigh (no snow bern),
phosphonis was somewhat greater than phos-
[)li()rus on sites with high snowbern densities
(no chokecherpy), and thus, a positive correla-
tion.
St(>j)wi.se nmltiple regression using all soil
properties with either chokecheny or snow-
bern- stem densitx did not pnnide meaningful
results. Howexer, a good relationship wa.s found
38
c;heat Basin Natuiulist
[Volume 52
Taui.k 2. Simple correlation coefficients for densities of
chokeclierr\- luid snowbi-ra witli chemical properties of soil
of green ash/eliokeclierr\ habitat t\pe near liison. South
Dakota (n = 72).
Soil
Chf)kechern
Snowbern
pll
KC
Orgiuiic matter
NO:vN
P
K
Zn
Fe
Mn
c:u
C.'a
\a
SAK
Satn ration
Ext.'Ca
Ext. Mg
Ext. Na
CEC(meq/l(X)kg)
0.1 9°
-O.Hi
-0.17
-0.03
0.26°
O.M
-0.13
-0.11
-0.03
0.07
-O.IS
().]7
- 0.00
-O.OS
-0.10
0.02
0.0]
-0.13
0.04
-().20°
0.2S°°
0.15
0.10
O.Ofi
O.IS
0.23"
0.03
0.23"
0.09
0.25"
()..3ft"
0.30"
0.08
0.10
-0.16
0.23"
0.17
-0.02
•Sinniricaiit ill a =0.5.
°°Sij;niricaiil al a = .()l.
Extrattahle cation
tor [)ro(liclino; chokccliern den.sitrv using snow-
hern' tlensih and cauopx' eo\er of grass (Table
3). Predicting snowheny stem density using
choked lern densit\ and grass cover similarly
showed a good relationshij) (r~ = .50). When
snow'hern' stem density was high, chokecherry
.stem densitv was low and \ice versa (Fig. 1).
Chokecherrv densitv' showed a good relation-
ship (r" = .48) with canopy co\'er of grass (Fig.
1 ). Stem densities of chok(^chenv were greatest
when canop\ coxcr of grass was k)w\
Oxcrall. soil properties were not highK' corre-
lated with either chokechern or snowbern'
stem densits'. Each shrub was more infhienced
by the densit\of the otheror the amount of grass
co\er. Factors such as other shrubs, trees, dis-
ease, fire, .soil compaction, and grazing ma\ also
inlhience stem densit)'of"both chokechern and
snowbertA (Boldt et al. f97(S, Se\erson and
Boldt 1978, Uresk and Paintner f985, Uresk
and Boldt 1986, Uresk 1987), but these factors
were not considered in the present study.
Summary
Surface soils of the gnx-n ash/chokecheny
woodland in northwestern South Dakota near
Bison were found to be moderateh' fertile with
CO
z
HI
Q
>-
cr
LU
m
O
z
15
12
-♦
_
FITTED
â– X
ACTUAL
â– *
*
â– * +*
** *
**
1 . . 1
7r**^
3 6 9 12 15
CHOKECHERRY DENSITY
15
W 12
LU
Q
>- 9
CC
LU
I
O 6
LU
o
^ 3
— FITTED
* ACTUAL
-\
* *
* *
\
c * *
*
â–
\.
i
-
*s<
* *
. ♦--<
* + ^\^
* *
* ^s,
>^* *
*
.
* *
^<
*
â–
*
">
s.-
Ij_
20 40 60 80
% GRASS COVER
100
Fig. 1. Snowheny stem densitv (stems/0. 1 \u~) is greatest
wiien chokecherry stem densitv is the least, but decreases
as chokechern densit\^ increases. CliokecheriA stem densitv
is greatest wlien grass co\'er is the least, ami densih'
decreases as grass ccner increases.
fairh- high lexeLs of nutrients except phospho-
rus, which was low, and nitrogen, which was
uioderateK low. Organic matter ranged from
about 47( to 20%. These soils were fine textured
with UioderateK' high cation exchange capacitv'
and saturation percentages. The\' were classed
as non.saliue-nonalkaliiK^ with low amounts of
exchangeable sodium.
Soil jirojierties showed low correlation rela-
tionships with chokechern' or snowbern stem
densit\. A good relationship was found lietween
the t^v() species of shrubs and grass. Additional
factors such as d(^usit\ of other shrubs or trees,
di.sease, lire, soil compaction, and grazing may
also infhience densities of chokechern or snow-
bern and interact with soil surface properties.
1992]
Soil Chemistry and Plant Rklationsiiifs
39
Tahlk 3. Coefficients (a. b, aiul c), standard error of the estimate (SE), and correlation (r ) describing relationsbips of"
cbokecherfN' (C), snowberrv (S), luid grass (C) in green ash/chokecherr\- habitat t\pe (n = 72).
Densih(Y)
SE
r\pe
Cliokt'c hern'
9.651
-().48SS
-().().S2(;
I.<S4
0.72
S
Snowlx'rn
1 1 .694
-L()76C:
-().()76(;
2.66
0.50
S
Snowbern
1L75.S
-6.266(:
0.197
2.51
0.55
E
C'hokechern
9.32,3
-().555C;
0.620
2.53
0.4S
E
'S = .sl.-purs,-rcgn.ssion(V =
= a+lK' + l«-); E = c
xponciitial rcijri'ssi
i()n(Y = a + lKM.
Ac:kn()\vledgments
Thanks are extended to Custer National
Forest for providing partial funding and study
areas. Appreciation is extended to Robert
Hordorff and Steve Denison for assisting witli
data collection.
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for summarizing nuilti\ariate data. Beha\ionil Science
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BoLDT, C. E., D. W. Urksk. and K. E. Sknekson 1978.
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Bhaov N. C. 1974. The natnre and properties of soils. Sth
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Cl I AH LEY, J. L. 1977. Mineral cycling in rangeland ecosys-
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IIxN.SEN, P L., cindC. R. Hofeman 1988. The yegetation
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of ,^^erto,V ret <>nundcrstor^â– Lod^ plants. 1'^ treatments on regeneration of ,u.t.v.woodIanck^
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bird. Utah. USDA Forest Senice Gener.J Technical Journal ol Range Management oS: 44(^2.
lk'i-K)rt INT-222. Intermountain Forest and Range
Exiwrimcnt Station, Ogden, Utah. 179 pp. Received 1 November 1991
Accepted 16 Jamianj 1992
(;R-at Basin NatiinJist 52^ 1 i. 1992, pp. 41-.52
THE GENUS AK/ST/D.A (GRAMINEAE) IN CALIFORNIA
KclK \\. Allrcd'
Arstuact. — Till' t;L\()iioi)i\ ol Aiistidd ( Crainiiicac ' in ( .'aliloniia is revised. Tlie liciiiis is ri'presented in tlie state 1)\ six
species and 1 1 ta\a. Identification ke\s, descriptions, selected s\ non\ in\, dislrilmtion records, and illnstiations are prox ided.
Kct/ uords: .\ristida. //()C/.s7/r.s, Ciilifonna.
As part oltlu" current rexision of Willis L\"nn
jepson's .\ Manual of the Flowerino; Plant.s of
(-"alifornia ( 1923), ,spon,sorecl l>\the Jep,son Iler-
hariuni ot the Unixersitvof California at Berke-
le\. an (\\ainination of the taxononn,
nouienclature, antUlistrihution of the California
sp(^cie.s of Aristida was undertaken. Jepson
( 1 923) originalK li.sted 10 .species oi'Aiistida for
California, and subsequent floristic endeaxors
increased this number to 12, reported by Munz
and Keck (196(S). This work treats si.x species
ap[)orti()ned to 1 1 total ta.\a.
Aristida are peculiar in the de\tdopnient ol the
iusilonii, indurate floret. The lemma (in North
.Ameiican species) is convolute iuid conipleteK'
encloses the palea and flower, forming a rather
firm anthoecinm. or flower casing. This configu-
ration customariK prexents the exsertion of
anthers and stigmas, resulting in cleistogamons
(and st^lf-pollinated) reproduction. Howe\er, in
souie spikelets of A. pmyurea Nuttall, A.
diiaricata Humb. & Bonpl. ex Wilk^now, and
other species, swelling of the lodicules will often
spread the lemma and palea, and the antheis and
stigi 1 uis are commoi \\\ e.x.serted from tl \v an tl k k'c-
ium during and afteranthesis,e\idence of possible
cros.s-pollination. In A. dicJiotonui Michaux of
ceutnil and ea.steni United States, two kiiuls of
flowers ai-e de\ eloped: one with three anthers
each 2-3 nun k)ng, presumabK adapted for
chasmogamous reproduction, and the other with
a single anther less dian 0.3 mm long (j)ersonal
ob.senation). The smaller anther is alwa\s found
retainetlwithiu the floret and aj^paRMitK functions
ill clcMstogamous n^production. I'his condition is
LiLso reported for A. oli^aiUha Michaax (Uenrard
1929).
The tip of the leuuna often bears a column or
beaklike structure in species ol Aristida, and tw o
terms describe this condition. An awii column is
formed b\ the couni\ent or coalescent. often
twistetl bases of the awns alxne the lemnui. This
is a relati\el\ unconnuon arrangement but is
seen in Aristida califoniica Thurber. A beak of
the lemma, howexer, is sometimes formed b\
the lennna apex. It is often narrow and twisted,
as in A. divaricata and A. pinyurca. The term
(iwn. as used luM'ein, refers to the free portion
onK and is measured from the summit of the
beak or awni coluum to the tip ol the awn.
North American Anstida have been classified
in three different sections of the gemis:
ArthradwrunL Sircptachnc, and Aristida
(Chactaria) (Uenrard 1929, Cla\ton and
Renxoi/.e 19Sfi). In section A/t/jraf/irn/;;;. the
lennna bodx is terminated by an awn column
that disartic-ulates from th(> rest of the floret.
This section is represenletl in California by A.
califoniica. The section Strcptacltnc is charac-
terized b\- the extn^ne reduction of the lateral
awns, illustrated consistently in A. ternipe.sCiXv-
auill(\s. but also found in other species that are
not usualK placcnl in this section, such as
A. adsccnsioiiis Linnaeus. In a study of Amf/f/r/
species affiliated with A. divaricata, Trent
(1985) found that some degree of reduction of
the lateral awns was a couunon occurrence in
numerous sjiecies, and concluded that this f(^a-
Inrc was often not a good indicator of biologic
relationship. The \alidit\ of the section
Stn'ptachnc ba.sed on this criterion is doubtful.
.Section Aristida comprises the remaining (Cali-
fornia species without articulation in the lennna
or consistent reduction of lateral awns.
' Dipartiiieiit ot Animal and Kange- Sciences. Bon .3-1. New Mexict) State University. 1-ls C:nKvs. New Mexic-o SS(K)3.
41
42
Great Basin Naturalist
[Volume 52
Because the sectional classification of the
genus remains lari^cly unexamined and imsatis-
factorv', for this re[)ort the California species are
sorted into informal "groups." These groups do
not necessariK correspond to any formal rank
hut parallel those used b\ Ilitclicock and (>hase
(1951) and Allred (1986).
Group ADSCENSIONES. — Ah.sfida ad.scen-
sionis; characterized h\ the annual habit,
branching at the upper nodes, and erect awns.
Group DiciiOTOMAE. — Aristkla oligantha;
characterized by the annual habit, branching at
the upper nodes, and a tendency for the central
awn to coil.
Group DixakiCATAE. — Ahstkia d'waricata
tmd A. temipes; chtiracterized by tlie stiffly spread-
ing piiman' (and often secondary) bnmches wdth
a\illan [)ul\ini. These two species are usuiJly
placed in different sections of the genus (Aristkla
and Streptachne, respectiveK).
Group PurPUREAE. — Aristkla puiyitrea,
including sexen \arieties; characterized by gen-
eralK unecjnal glumes, a narrowed beak of the
lenuna, and generally erect branches; merges
with the Divaricatae through A. purpurea van
parishii (Hitchcock) Allred, as well as A. pansa
Wboton & Standle\'of the Chihuahuan Desert.
( ;h{ )U 1' Tu B !■: 1k;u LOS a E . — Arisi kla califor-
iik-a; characterized by the disaiticulation of the
awnis and awn column from the l)od\- of the
lennua.
Following are identification keys to till taxa,
descriptions based on Cialifornia specimens,
counties of occurrence in California, lists of
selected specimens examined, and an illustra-
tion of each taxon. Herbaria arc^ abbreviated
according to Holmgren et al. (19(S1). Updated
information on the distribution of Aristkla in
Cialifoniia will be welcomed by the author.
Aristkla Limiaeus, Sp. Pi. (S2. 1753.
Tufted annuals or perennials; ailms generalK
erect, the internodes mostly semisolid. Sheatlis
open. Uiudes a ring of hairs. Blades flat to in\o
lute, lacking auricles. Injlorescence generalK a
panicle, occasionally racemose or spicate.
Sj)ikclrts 1 -flowered, di.sarticulating above the
glumes. Chinws etjual to \er\' unequal, thin,
membranous, 1- to 7-nened, often as k)ng as
the floret or longer. Lenuna 3-ner\'ed, terete,
indurate at maturity and enveloping the palea
and flower; eallus oblicjue, usuall)' sharp-
jiointed and bearded; aicns 3 in number, termi-
nal on the lenuna, the lateral awns sometimes
reduced or obsolete. Falea 2-nerved, thin,
shorter than the lemma. Lodicules 2. Stamens 1
or 3. Carijopsis enclosed in the anthoecium,
hisiform, the hilum scar linear, the embryo
.small. X= 11.
Key to the Genus Aiistkld
I . Culm internodes and nodes eonspicuously hairy
A. califonuca var. califomica
Cuhn internodes and nodes glabrous 2
2(1). Plants annual, generally much branched above
the base 3
Plants perennial, simple or onl\ \\ eaklv branched
above the base 4
3(2). Central awns mostly 3-7 cm long ... A. oligontJia
Central awais mostly 0.7-2 cm long . A. adsccnsionis
4(2). Primary panicle branches erect to spreading or
diooping, but at least the bases of the branches
appressed to the main iixis, without pulvini in the
branch axils A. ptiqnnva
Prinii\r)' panicle branches abniptlv spreading from
the main axis with pulvini in the branch axils ... 5
5(4). Lower panicle branches ascending, the upper
branches appressed .... A. pinjjurca vm. pari.sliii
Lower and upper panicle branches spreading ... 6
6(.5). Anthers O.S-l mm long; summit of lemma twisted
at maturitv; base of blade glabrous abo\ ethe ligule
A. dhurkdtd
Anthers L2-^3 nun long; sunnnit of lemma not or
onK slightK- twisted at maturity; base of bladewith
scatteied pilose hairs above the ligule A. tcntipcs
Aristida adscensionis Linnaeus, Sp. Pi. 82.
1753. Six weeks threeawn (Fig. 1) [A.
adscenswnis var. ahortiva Beetle, A. adscen-
sionis var. decolorata (Founiier) Beetle, A.
adscensionis var. niodesta Hackel]. Tufted and
generally annual, but e.xtremelv variable in size,
growth habit, and longevit)'; culms erect to
geniculate, simple to much-branched, (3)1()-
50(80) cm tall; internodes glabrous. Sheaths
generally shorter than the internodes. Li^jides
0.4-1 nun long. Blades flat to involute, 2-14 cm
long, 1-2.5 mm wide. Panicle narrow and con-
tracted, 5-15(20) cm long, often internipted
below, tlie spikelets aggregated on short
branches. CUumes unequal, 1-nerved, the first
4-8 mm long, the second 6-11 mm long.
Lcnufuis 6-9 mm long, slightly flattened, sca-
brous on th(^ midneiAe; awns flattened at the
base, .spreading, the central awii 7-18(23) mm
long, the lateral awns somewhat shorter, rarely
0-2 mm long. Palea 0.5-1 mm long, hvaline,
blunt, fan-shaped. Anthers 0.3-0.7 nuii long.
Can/opsis somewhat shorter than the lenuna.
2)1 = 22. Diy, open places and rocky hills below
19921
GKNUS/\/i/.S7V/A\ IN CJ.M.IFOHMA
43
Fi>4. I. Ari^tida (uiscciisioiiis. inflorescence and spikelet.
1 ()()() 111. COUNTIKS: Imperial Inyo, Los Angeles,
Hixerside, San Bernardino, San Diego, San Luis
Obispo, Santa Barbara.
Aristkla adscensionis ranges in liabit troin
small, unbranched plants scarcely 3 cm tall with
onK one or t\\T) spikelets to large, mnch-
branclied clumps SO cm tall witli immerous
branches and spikelets. Sexeral \arieties liaxc
been named based on differences in plant and
[xmicle size, degree of branching, and the devel-
opment of the awns. N'ariation in size and
robustness seems related to precipitation, and
populations at the same site max \ an drasticall\'
troni \('art()\ear. The\alidit\ ()l nnluced lat(M-al
awns as a taxonomic character is also (jiiestion-
able. Most species o{ Aristida haw forms with
the lateral awns reduced, and this seems to
occur almost indiscriminatek and without any
correlation with other features.
Selected specimens. — Imperial Co: rd
from Ogillix to Bhthe, 17 Feb 1958, Bacigalupi,
H. 6136 [|EPS]; Carriso Mts, Painted C;orge, 17
Mav 1938, Ferris, R. S. 9622 [UC]; near Dixie-
land, 13 Oct 1912, Parish, S. B. 8239 [JEPSf
Inyo Co: Panamint Mts, Deadi Valley, 18 Apr
1978, Dedecker4541 [UC]; 11 mi W of Death
Valley, 28 Mar 1947, Keck, D. 5847 [UC]. Los
Angeles Co: Pasafk'ua, 27 Feb 1882, Jones,
M. E. s.n. [CMl; San Clemente Island, 8 Mav
1962, Raven, P M. 17609 lUC]. Riverside Co:
9.4 mi N of BK-the, 19 Feb 1958, Bacigalupi, R.
6188 [JEPS];' Marshall Canyon, 10 mi W of
Coachella, 16 Apr 1905, Hall,' II. M. 5797 [UC];
near Mecca, 28 Jun 1902, Parish, S. B. 8122
[UCJ; S end of Coxcomb Mts, 27 Mar 1941.
Wiggins, I. L. 966 [UC]. San Bernardino Co:
NW side of Coi)per Basin, 6 Ma\ 1939, Alexan-
der 710 [UC]; Sheep Mole Mts, 25 Apr 1932,
Ferris, R. S. 8020 [UC]; Needles, 12 Mar 1919,
Tidestrom, I. 8556 [UC]. San Diego Co: San
Diego, 29 Apr 1902, Brandegee 832 [UC]; 6 mi
NW of Agua Caliente, 5 Apr 1960. Everett
24075 [UC]; 1.5 mi E ofWillecitos, 28 Jan 1940,
Munz, P A. 15856 [UC]; Borrego Springs, 18
Mar 1976, Schroeder 51 [UC]. San Luis
Obispo Co: San Luis Obispo, 9 Ma\ 1882,
Jones, M. E. 3245 [UC]. Santa Barbara Co:
Santa Ynez Mts, 9 May 1954, Pollard [ UC].
Aristida californica Thurber in S. Watson,
Bot. Calif 2:289. 1880. Tufted, slightly bush\
perennial; culms erect, much-branched, gener-
all\- 10-40 cm tall; inicrnodes glabrous or pubes-
cent. Sheaths much shorter than the intemodes,
pubescent at the throat and on the collar. L/g-
tdes about 0.5 mm long. Blades mo.stK' folded to
in\ olute, occasionalK' flat, stiffly .spreading, 2-.5
cm long, inostK' less than 1 mm wide, scabrous
to hispid-pnbenilent. Inflorescence few-flow-
ered, 2-6 cm long, the terminal ones paniculate,
the axillan- oiu\s racemose. Chimes unecjual,
l-nen(Hl. Lenniia with a narrow column at the
tip formed b\' the twisting and fusing of the awn
bases; awns nearly ecjual, breaking from the
lemma, the zone of articulation at the ba.se of
the awn column. 2n = 22.
var. californica. CxilFOHMA TIIKEPIWN
(Fig. 2). Iittenuxh's pubescent, the hairs pilose
to sublanose. Clluines \c\\ unequal, the first 4-8
mm louiiand the .second 9-12 mm Icmg. Lemma
bod\ 5 7 mm long when mature, the awn
coluum S 26 mm long; awns 2-4.5 cm long.
Diy, sancK, desert areas. Coi'NTIES: Imperial,
Riverside, San Bernardino, San Diego.
The other \ariet\- of this species is \ar.
olahrala \'ase\-, known principally at the species
Ie\el as Aristida <ihd>rata (Vasey) Hitchcock.
This varietx- differs from \ar. caUfonuca primariK'
in having glabrous, rather than pubescent,
44
(;hkat Basin Naturalist
[\\)luine 52
inteniodes and octiu-s in die slighdy higher ele-
sations oldie deserts to die east of the range of
\ar. calijomica. Both taxa are cbploids {2n = 22),
and the)- oxt-rlap considerably in spikelet
dimensions (Keeder and P\dger 1989). Variety
^lahrata is not knowni from ('alifoniia.
SELKCTLD SFECIMKNS. — Inipei-ial Co:
Signal Mt, 2 Apr 1903, Abrams, Ci. D. s.n. [DS-
] 86664] [ DS]; 8 mi E of El Centro, among larrea
bnshes. 22 Apr 1942, Beetle, A. A. 3172
[AllUC]; Bard, near Arizona line, 22 Sep 1912,
Thornber, J. J. s.n. [ARIZ], a few mi E of Holt-
xille, Jun 1951, Tofsrnd. H. s.n. [AHUC]. Riv-
erside Co: nearTlionsand Palms, rockv desert
slopes, 27 Apr 1943. Beetle, A. A. 1938
[AHUC]; Pinto Basin, 16 mi from Cottonwood
Springs, 15 May 1938, Ferris, R. S. 9522 [DS];
canxons along Colorado River, 1 May 1905,
Hall. H. M. 5963 [ARIZ, POM, UC]; Coachella
\'alle\, 6 mi SE of Caniet Station, sand dunes,
ca 500 ft, 1 1 Mar 1928, Howell, J. T. 3443 [DS,
CAS, AHU(]]. San Bernardino Co: Joshua
Tree National Monument, 1700 ft, north ledge,
TIS RIOE, 18 May 1941, Cole, J. E. 734 [UC];
Baxter, S of Mojave River, 23 May 1915, Parish,
S. B. 9886 [UC, DS]; Dale Lake Valley (W of
lake), 13 mi E of 29 Palms, sun-dn' sand flats,
abundant. 29 Max 1941, Wolf, C. B. 10876
[RSA, DS, CAS]. San Diej^o Co: San Felipe
Narrows, ca 350 ft, 20 Apr 1935, Jepson, W. L.
17101 [J EPS]; canvon W of Borrego Spring,
1.500 ft. 19 Apr I9()6. jcmes, M. E. s.n. [POM-
I 1 700 1 I 1 POM I; Colorado De.sert, clay hills, 25
jun 1SS8, Orcutt, C. R. 1486 [DS].
Aristida divaricata Ilumb. & Bonpl. ex
W'illck'now, Enum. Pi. 1:99. 1809. PON'KKTY
TiJKliEAWN (Fig. 3). Tufted perennials; ciihits
erect, mo.stly unbranched, 25-70 cm tall; iiitcr-
nodes glabrous. SJieatlis longer than the inter-
nodes. iJffih's 0.5-1 mm long. Blades looscK
inxolute, glabrous, 5-20 cm long, 1-2 mm wide.
Fnniclc open, 10-30 cm long, 6-25 cm wide;
priniaiy branches stifflx spreading from the
main axis. axillanpuKini present, 2-12 cm long,
generally naked on the lower portion. Brditch-
lels and s))ikclct.s general!)- appressed along the
branches, but .sometimes si)reading. Chimes
nearl) etjual, l-ner\ed, 8-12 mm long, acumi-
nate-aristate. Lemma (S-13 mm long to base of
awns, the terminal 2-3 mm narrowed ami geii-
erall) twisted fonror more turns; <'/u/j.s subecjual
to une(jual, (7)10-22 mm long, the lateral awns
at least slightl)- shorter than the central. Anthers
0.8- 1 nun long. 2n = 22. To be k)oked for on d\\
Ply;. 2. Ah.stiild ccdijontica, iiillorescenee, spikelt't, and
ck'tail of hranchiiii';.
slopes below 150 m elevation. COUNTIES: San
Diego.
It is doubtful that Arisiida divaricata cur-
rently occurs in (>alifornia. Most reports are
based on collections of C. R. Orcutt in 1884, and
no knowni specimens haxe been collected from
the state since that time. In addition, it is possi-
ble that Orcutt's labels are in error, because on
at least one specimen of A. divaricata he located
Hansen's Ranch, which is in Raja California, in
San Diego Countx'.
A similar species, Aristi(hi orciiftiana N'asev,
also supposetlK was collected from southern
Calilornia in 1884 b\ (]. R. Orcutt, and hvo
specimens are hou.sed at US. The labels
(k'scribe San Diego as the collection 1ocalit\'.
and these specimens are apparently the basis for
reports ol either A. orctittiana or A. scJiiedeana
Trinius & Ruprect from California (Abrams
1923, Jepson 1923, Hitchcock 1924, Munz &
Keck 1968). Coincidentally, the t\pe locality of
/\. orciitiiana is again Hansen's Ranch in Baja
California, mentioned abo\e. It is possible that
neither. A. divaricata nor A. orcitttiaiui was e\er
collected from California b\- Orcutt, but from
19921
CiENUS/\/^/.S77/;.A IN C^ALIFOHNIA
45
Fi'j;. 3. :\risli(l(i diiaricafa. innoresceiicc. spikek't, and
hasc ol i)laiit.
Baja ( iaiitornia. Arisfida orciiUidna rcsciiibles
A. (livdi'icald in tlit^ stiifl\ sprcadiiiu; panicle
hranclu's, hnt the lateral awnis are \eiy short or
absent, and the blades are cjeneralh' flat and
somewhat cm ling in orcutfiaiui.
Sl'i:(:iMi;\S KXAMINED. — Withont detiiiite
loealitx but recorded as California: Santa ( ^ata-
hna Mts [Santa Catalina Island?], in 18S4,
Orcutt, C. H. 2 [US]; Santa Clara Mountains
Ipo.ssibly Arizona?], in 1SS4, Orcntt, C. 1^ 2
|US[. San Diego Co: San Diego. Orcutt. C. H,
s.n. |\Y, US].
Aristida oU'^iintha Michanx, Fl. Bor. Ainer.
1:41. ISO,). OLDFIELD TIIREKAWN (Fig. 4) |A.
oli'j^tmllia \ar. nervata Real]. Tufted auuuals;
culms win, 3()-7() cm tall, mucii-branclied, the
iunoxations extraxaginal: iiifcniodcs glabrous,
pith\. SJwatlis nio.stlv shorter than the inter-
nodes. Ligules 0.1-0.5 mm long. Blades flat to
in\ olute, 3-22 cm long, 1-2 mm wide, reduced
Fig. 4. Ahstid/i olifiantliti. inllorL'scciici', spikclft, aiitl
detail ol hraiK-liiii''.
upwards. Injlorcsccncc few -flowered, race-
mo.se, the spikcdets nearh' sessile. GliiDics sub-
cHjual or the second longer, awn-tipped, most!)
(12)18-34 nun long, the hrst 3- to 5(7)-neived
and shoi-t-awTied, the second 1 - to 3-nened with
an awn S-13 nun long. Lciniua (10)13-20 mm
long to base of awns; cciitrdl (iwii (2)3.5-7 cm
long, the lateral awns generally somewhat
shortc-r. 2// = 22. Dn hills and fields, bare
ground, scrub land, 90-1000 m elevation.
C()L\Tli;S; .Vniador, Butte, El Dorado, Hvun-
boldt. Imperial, Lake, Madera, Mendocino.
Merced, Modoc, Nevada, Placer, Redding, Sac-
ramento, San Joacjuin, Shasta, Siskiyou, Solano,
Sonoma, Stanislaus, Tehama, Tuolumne, Yuba.
46
Great Basin Natuhai.ist
[\ blunie 52
Some specimens of Arisfidn oJi<uinth(i from
northern California (Lake and Modoc counties)
and adjacent areas of southern Oregon exhibit
smaller glumes, lemmas, and awns than are top-
ical and ha\e been segregated as either A.
rainosis.siina Engelmann var. chaseana Ileiuard
or A. oli'^aiitha \ar. iwrvatn Beal. In addition,
the central awii in these plants in sometimes
acuteK reflexed and the florets darkened. Tliis
configuration is intermediate between A.
oli^aiitlia and A. raniosissiDia.
Selected specimens: Butte Co: Chico, 27
Jul 1903, Copeland 3488 [US, WIS]; volcanic
uplands between Pent/ and Dn Creek, 15 Jul
1914. Heller, A. 1 1576 1 UC]; 2.5 mi S of Wyan-
dotte, 28 Nov 1933, Jensen 367 [UC]. Hum-
boldt Co: Cottrell Ranch, 17 Sep 1955, Mallory
122 [U(>]; Trinitv' River near mouth of Willow
Creek. 15 Sep 1919, Tracv 5222 [UC]; vicinity
of Carbenille, 27 Aug 1933, Tracy 13()()() [UC]';
Dobbyn Creek, 9 Juri934, Tracy 13341 [UC].
Lake Co: dn hills between Upper Lake and
Scott \alle\-, 17 Aug 1905, Tracy, J. P. 2365 [UC]
(\ar. nervata). Madera Co: Mintum, 1 Oct
1936, Hoover, R. F. 1618 [JEPS, UC]. Merced
Co: Tuttle, 17 Jul 1936, Hoover, R. F. 1580
[JEPS, UC]. Modoe Co: 19 Aug 1935, Whitnex,
L. 3627 [UC]; M(4cher Creek, 6 Sep 1935,
Wheeler, L. C. 3959 [US] (\ar. nenata).
Nevada Co: Talioe Natl Forest, S of Grass
\ alley, Aug 1931, Smith 2638 [JEPS, UC]. Sac-
ramento Co: 5 mi SE of Folsom, Yates, H. S.
5953 [UC]. Shasta Co: Redding, 21 Jun 1909,
Blankinship [JEPS]; 1 mi N of Anderson, 21 Jul
1932, Long 190a [UC]. Stanislaus Co: vicinit\
of La Grange, 30 Sep 1961, Allen [JEPS];
bet\\'een Knight's Fern- and Wanienille, 1 Sep
1941, Hoover, R. K 5582 [UC]; 1 mi NW of
Waterford, Yates, H. S. 6858 [UC]. Tehama
Co: 9.7 mi N of Red Bluff, 14 .Aug 1954. Bac-
igalupi. R. 4808 [JEPS]; \blcanic Plateau NE of
Red Bluff, 22 Sep 1940, Hoover. R. R 4617
[UC]. Tuolumne Co: near Kevstone, Yates
H.S.6148[UC].
Aristida purpurea Nuttall, Trans. Amer.
Philos. Soc. 5:145. 1837. Tufted perennials;
culim erect and general!)- unbranched, 10-80
cm tall; i)ifcnioclcs glabrous. Sheaths longer
than the inteniodes. Li<iidcs 0.1-0.5 mm long.
Bhulcs mostly in\-olute. PanicU' \ariable, con-
tracted and spikelike to open and fle.xuous, the
branches without puKini in the axils (except \ ar.
parishii). Chimes mostly unecjual (except xar.
pahshii), the first about half the length of the
.second, l(3)-nened, acuminate. Aiois about
e(|ual or the central slightK' longer. Because of
intergradation among forms (Allred 1984), the
taxa of this complex are recognized as varieties
within Arisfida piiffnirea.
1. I'riiiian panicle l)ranc-lies, at least the lower, with
a\illar\ piiKini and usually stifll\ spn^uliuo; to
iuscending from the main axis \ar. parishii
I'riiiiaiA panicle branches lacking a\illan piiKini,
tlie spikelets variously disposed but at least the
bases of the bnuiches appressed to the axis .... 2
2(1). Awns 4-10 cm long 3
Awns l-.'j..5cin long 4
3(2) Sunnnit of lenuna 0.1-0. .3 mm wide; awns rather
delicate, mostly 0.2 mm or less wide at the base,
4—6 cm long; second glume mostly shorter than
16 mm xdv. puqturca
Summit of lemma 0..3-0.8 nnn wide; awns usu;ilK
.stout, more than 0.2 nnn wide at the base. 4—10 cm
long; second glume 16-2.5 mm long . . \ar. low^iscfd
4(2). Snnmiit of lemma mostK' less than 0.2 nnn witle;
awns delicate, mostK less than 0.2 nnn wide at the
base .5
Sunnnit of leiMiiia mostK more thiin 0.2 mm wide;
awns stout, mostK 0.2 nnn or more wide at the
base 6
5(4). Panicle branches and pedicels erect, stiff", occa-
siomJK' spreading or flexuous var. iicallctji
Panicle branches and pedicels drooping to flexoious
\ai: ptu^jiirea
6(4). Panicles mostK 3-14 cm long; blades mostly basal
and less tluui 10 cm long yar.fcmlleiiana
Panicles mostly 15-30 cm long; blades mostly
canliue arul more than 10 cm long . . . \ar. ivii<^litii
Mil. fendleriana (Steudel) \'ase\', Contr. U.S.
Natl. Herb. 3:46. 1892. FENDLER THREEAWN
(Fig. 5) [A.fendh'riaita Steudel, Svn. Pi. Glum.
1:420. 1855]. Cuhns 10-40 cm tall Blades invo-
lute, mostly less than 10 cm long, usualK basal
but occasionally cauline. Pan'wk' 3-14 cm long,
narrow. Chimes unequal, the first 5-8 mm long,
the second 10-15 mm long. Lemma 8-14 mm
long;c/(r/;,v generallv 1.8—4 cm long, 0.2-0.3 mm
wide at the base. 2n = 22, 44. Diy, often rocky
slopes and hills, 1000-2000 m elevation. COUN-
TIES: Im-o, liixerside, San Bernardino, San
Diego.
Selected specimens.— Imo Co: Dexil's
Kitchen C\-n, SE V4, Sec 7, T22S R39E, 21 .\hi\
1978, Zembal, R. L. 531 [RSA/POM]. River-
side Co: 20 Jul 1905, Griffiths, D. 8008 [MO];
San Jacinto Mts, Pinvon Flats, 18 .Ma\' 1958,
Rawn. P. H. 13003 [RSA/l^OM]. San Bernar-
dino Co: near Jupiter Mine, Kingston Range,
1992]
Genus. \/i/.s77/;.\ i\ Califouma
47
Fig. 5. Aristida puqntrca \ar. fcndlcriana. inflorescence,
spikelet, and base of plant.
30 May 1980, de Nexers, G. 348 [RSA/POM];
SW New York Mts, 5.5 mi E of Ciina in Gottoii-
\\()()(l (Jan\'on near Cottonwood Spring, 2 |nn
1973, Hem-ickson, J. 10339 [RSA/POM];
I\ anpah Mts, Kessler Peak, 2 Jun 1931, Jep.son,
W. L. 15825 [lEPS]; San Bernardino Mt.s, 15
|un 1895, Pari.sh, S. B. [UC]; Budweiser Wasli,
near 35d 46m N, 115d 44m W, Granite Mts, 28
Oct 1977, Prigge, B. A. et al. 2320 [RSA/POM];
Caruthers Cyn, New York Mts, 30 Ma\ 1973.
Tliorne, R. F. 43639 [RS/V/POM]. San Diego
Co: 3 mi W'NW of |acnmba, Yates, H. S. 6805
[UC]; 5 mi ENE of'jacnmba, Yates, H. S. 6808
[UC].
var. longiseta (Steuck4) Vasey in Utjtlirock.
U.S. Suney W. lOOth Merid. Rpt. 6:286.1855.
RE15 THHFFAW \ (Fig. 6) [A. longiseta Stenck'k
S\ii. PL C;kim. 1:420. 1855, A. lonoiscfa \ar.
rohusta Merrill]. Culms 10^0 cm tall, delicate
or stout. Blades 4-16 cm long, mostly involute,
basal or cauline. Panicle 5-15 cm long, the
branches stout and erect to delicate and droop-
ing, but usuall)- not ver)' flexuous or tangled.
Fig, 6, Ari.stidii purpurea \ar. lon'^iiseta, inflorescence and
.spikelet.
Glumes unecjual, the first 8-12 nnn long, the
second 16-25 nnn k)ng, sometimes shorter.
Lemma 12-16 nun k)ng, 0.4-0.8 mm wide ju.st
below the awiis; awns stout, 4-10 cm long, 0.2-
0.5 mm \\ade at the base. 2n - 22, 44, 66, 88.
Dry, desert hills and plains, 300-1500 m ele\a-
tion. COUNTIES: Mono, Riverside, San Bernar-
dino, San Diego.
The \ari('ties lon<iiscta and fcndlcriana are
often contused, but are most easik distin-
guished b\ the width of the awnis and lemma
apices, and not b\ whether the lea\es are basal
or cauline.
SELE(:TE13 si'KCl.MEXS: Riverside Co:
Joshua Tree National Monument, 1 Ma\- 1942.
Roos 1153 [US]; Deep Can\on. T7S R5E. 27
Jun 1937. Yates, H. S. 6722'[RS.VPOM]. San
Bernardino Co: E New York Mts, W of Castle
Buttes between Corral and Do\e Spring. 12
May 1974, Henrickson, J. 13933 [RS.VPOMj:
Rock Springs, Palmer, E. 537 [UC]; plains near
Lea.stalk, 3 Jun 1915, Parish, S. B. 10329 [UC];
2.2 mi ESE of Brant on N side range of New
York Mts, 8 Ma\- 1978. Prigge, B. A. et al. 2905
[RS.Vl'OM]; San Bernardino Natl Forest,
48
Great Basin Naturalist
[Volume 52
above Cactus Flat ^^' of Hwv' 18 N of Baldwin
Lake, 2-3 Jun 1980, Thorne, R. F. 54375
[RS A/POM]. San Diego Co: head of Box
Canyon near Mason Vallev, 12 May 1932,
Duran, V. 3208 [WIS].
van nealleyi (Vasey in Coulter) Allred,
Brittonia 36:391. 1984. NEALLEY THREEAVVN
(Fig. 7) [A. glaiica (Nees) Walpers, A. stricta
Michaux van ncalletji Vasev in (Joulter, Contr.
U.S. Nad. Herb. 1:55. 189()]. Cxihm 20-45 cm
tall, tightly clustered. Blades generally basal,
involute, curving in age, 5-15 cm long. Panicle
narrow, spikelike, light brown, 8-18 cm long,
the branches mostK' erect-appressed. Glumes
mostly unequal, the first 4-7 mm long, the
second 8-14 mm long. Lemma 7-13 mn^ long,
0.1-0.2 mm wide just below the awns; awns
delicate, 1.5-2.5 cm long, mostly 0.1 mm wide
at the base. 2n - 22, 44. Drv; desert plains and
slopes, 200-1200 m elevation. COUNTIES:
Imperial, Inyo, Riverside, San Bernardino, San
Diego.
V^ariet)' nealleyi grades into \ an pitqutrca with
flexuous branches, and into van wrightii with
more robust panicles and broadc^r lemma apices
and awns.
Selected specimens. — Imperial Co:
Piiinted (iorge, Carisso Mts, 17 Ma)' 1938,
Ferris, R. S. 9623 [UC]. Inyo Co: john.son
Creek, Death N'allev, 28 Apr 1940, Gilman,
M. F 4190 [RSA/POM]; Cave Springs Wash, 25
Apr 1930. Hoffman, R. [US]; Funeral Mts, 2
May 1917, Jepson, W. L. 6907 [JEPS];
Titanothere Cyn, Grapevine Mts, E side of
Death Vallev, 26 Mar 1947. Wiggins, I. L. 11566
[RSA/POM', UC]. Riverside Co: Cottonwood
Spring, 30 Mar 1940, Hitchcock, C. L. 5871
[MO, RSA/POM, UC]; Eagle Mts, Cottonwood
Springs, 25 Apr 1928, Jep.son, W. L. 12585
[JEPS]; mouth of Andreas Canvon, 4-6 Aj^ril
1917, Johnston, I. M. 1010 [RSA/POM]; E of
Hemet, along San Jacinto Ri\en 7 Aug 1938,
Roos, J. C. 582 [RSA/POM]. San Bernardino
Co: Proxidence Mts, Fountain Cauxon, 15 Ma\
1937, Real .30] [JEPS]; route 95, 18 mi N of
Travis, 23 Apr 1942, Beetle, A. A. 3193 [WIS]:
39 mi from Needles on Parker Road, 24 Apr
1928, Ferri.s, R. S. 7226 [RSA/POM]. San
Diego Co: San Felipe, 16 Apr 1895, Brandegee
[UC]; San Felipe C;ap, 6 Apr 1901, Brandegee
[UC]; head of Fox C^anyon near Mason Vallev,
12 May 1932, Duran, '\'. 3208 [MICH, MO,
RSA/POM, UC]; Yaqui Well. 22 Apr 1928
Jepson, W. L. 12516 [JEPS].
Fig. 7. Aristkia purpurcd \ar. iwallet/i. inflorescence and
spikelet.
var. parishii (A. S. Hitchcock in Jepson)
Allred, Brittonia 36:392. 1984. PARISH'S
THREEAVVN (Fig. 8) [A. pariskU A. S. Hitchcock
in Jepson, Fl. Cahf 1:101. 1912, A. wiightii
Ndnh var. parishii (Hitchcock in Jepson) Gould].
Culms thick, stout, erect. Blades niostlx' flat,
longer than 10 cm. Panicle narrow, spikelike or
the lower branches with axillan" puKini and
spreading at al)out a 45-degree angle, 15-24 cm
long, reddish when \'oung. Glumes unequal to
e(|ual, the first 7-1 1 mm long, the second 10-15
mm long. Lemma 10-13 mm long, 0.2-0.3 mm
wide just below the awais; awns 2-3 cm long,
0.2-0.3 nun wide at the base. Chromosome
number not reported. Dn' hills and plains, 300-
]()()() 111 (-lexation. COUNTIES; Imperial, Imo.
Los Angeles, Rixerside, San Bernardino. Sail
Diego.
Vdm'ty ))arishii is very' similar to \an uri^zlif''
but differs most strikingly in the sometimes
spreading primary branches, the reddish color
of the panicle when young, and tlu^ more clus-
tered arrangement of die spikelets. It iilso
19921
GESLsAnisTin.\ i\ (;\i,ii"()i{\i.\
49
F\<^. S. Aiislida j)itif)nn'(i \'ar. parisltii. iiillorcscciK'c and
sjiikclct.
flowers earlier, mostK' March througli Maw
\\ liile \"ar. »;r/<^/(/// flowers iiiostK' Ma\ tIiroii<i;li
Octolx'i". Parisli s tlirecawn also resenil)l(\s some
nu^mhers of the Dixaricatae group because of
its spreadiug priuian' l)raucli{\s aud geuerallx
sul)equal gluuies.
SKLECTED specimens. — Imperial Co: 9.2
uiiles NE of Glamis, 18 Mar 1962. Ilitclicoclv,
C. L. 2225 [F]: Palo Wrde Mts. 8 Apr 1949.
Koos. }. (>'. 419S 1 US|. Inyo Co: spc'ciuieu with-
out lo(alit\ at KS.VPOM. Riverside Co:
(.'huckawalla Spriugs, 15 uii SE of (luiladax, 9
ful 1957. Crauiptou. H. s.u. [AIIUCl; Palui
(;auyou,4 Apr 191 7, johustou. 1. \1. 1008 [US.
MI(>H]; Rix'crside aud \iciuit\ ol upper fork of
Salt Creek Wash. 19 Mar 1927. Heed. E M.
5440 [AIIUC, HS.VPOM]: betweeu Marcli
AEB aud Lake\iew.29 Apr 1966. Koos. ]. C. s.u.
[RS.VPOMl. San Bernardino Co: 2 uii NE of
Eifteeun-iile Poiut. 3()()() ft, 28 Apr 1935.
Axelrod, D. 321 [AHUC, UC]; behveeu Bulliou
aud Sheep Hole Mts. 7 Apr 1940, Muuz. P A.
16568 [RS.Vl^OMl; Budweiser Wash. u(^u-35d
Fi'j;. 9. . \;-/.s7(r/c/ piiqiitrca \ ar. piiq)urc(i. iiilloivsctMice and
S|likrl('t.
46ui X, 1 15d 44iu W, (;rauite Mts, 28 Oct 1977.
Prigge, B. A. et al. 2320 [RS A/POM]. San Diego
Co: 0.5 uii N of Mirauiar Resenoir cla\ soil. 4 Mar
1981, Re\e;J, ]. s.u. [AHUC]; Auza Cauvou E of
juliau. 3 Apr 1940. W'ilsou. E. s.u. [AHUC].
var. purpurea. Pi Hl'LE THREEAWN (Fig. 9)
[/\. })iir})iir(ii \ar. raJifornicn Vase\]. Culms 2.5-
60 ciu tall. Blades flat to iu\olute, uiostly cau-
liue. .3-17 cui loug, 1-2 luui wide. Panicle
puiplisli. often uoddiug. 10-25 cui loug. tlie
brauches usualK delicate, droopiugor llexuous.
Chimes uue(jual. the hrst 4-9 luui long, tiie
second 7-16 uuu loug. Lemma 6-12 uiiu. 0.1-
0.3 luiu wide just below the awiis; aivns 2-3(4)
CUI loug. 0.2-0..) uiin wide at the ba.se. '2n = 22.
44. 66. 88. DiA, gra.s.sy hills, scrublands. 2.5()-S()0
ui elexatiou. COUNTIE.S: Mono, Rixenside, San
Bernardino, San Diego.
This is a beautiful grass, with its droojiing. red-
di.sh.plnnielike panicles. It conuuouly intergrades
w ith the varietes m'allei/i, lon^seta, iuid wn<ilifii.
SELECTED SPEC;E\IENS: Mono Co: McAfee
Creek, White Mts, Fishlake \alle\ drainage,
6 Aug 1984. Mor(fi(4d. ]. D. jbM-24S0(e)
f RSA/l^OMf Ri\er.side Co: 1 mile E of Banning,
50
Gkeat Basin Naturalist
[Volume 52
Fig. 10. Aii.stichi pui'j)iirca VAV. uri<s,lttii. intlorcsccncc and
.spikclct
20 Jul 1905, C;rifTith.s, D. 8007 [MO]; Palm
Canyon, 4 Apr 1917, jolmston, 1. M. 1008 [US,
MKJH]; ha.se ol San |acinto Mountain, fune
1882, Parish, S. B. et al." 1549 [F, MICH]; Lower
San Jacinto River Canvon, Yates, H. S. 6711
[UC]. San Bernardino C><): road from High-
land to Huiiniug Springs, 1 nil Irom valley floor,
26 Jun 1942, Beetle, B. A. 3644 [F, WIS]; near
Upland, 7 Nov 1916. lohnston, I. M. 1120
iMICHj; San Bernardino N'allev, 2 jun 1906,
Parish. S. B. 5783 [NMCR]; Clark Mts, 5 Aug
1950. Boos, j. C. et al. 4906 [BSA/I^OM, UC].
San Diego Co: 6 mi N of Ocean Side Ranch,
coast hills in chaparral, 21 Apr 1942. Beetl(\
A. A. 3145 [TAES]; near Vallecitos Station,
2 Apr 1939, Gander, F 7142 [MICH]; Ilarhi.son
C;an\T)n, 19 Jun 1938, C;ander, F F 5999
[BS.WOMl.
van wrightii (Nash in Small) Allrcd,
Brittonia 36:393. 1984. Whiciits thhki:a\\\
(Fig. 10) [A. wii<ihtii Nash in Small, Fl. South-
ea.st. U.S. 1 16. 1903]. Ciihiis erect, to 80 cm tall.
Blades involute to flat, cauline, 10-25 cm long,
1-3 nun wide. Panicle narrow, spikelike, 14-30
cm long, the branches erect-appres.sed. Gliiines
unefjual, the first 5-10 mm long, the second
Fig. 11. Aristida tcniipcs \ar. hdinulDsa. innorescence,
.spikelet, and detail of ligiilar region.
10-16 mm long. LeniDia 8-14 nuu long, 0.2-0.3
mm wdde just below the awais; awns mostly
2-3.5 cm long, 0.2-0.3 mm wide at the base. 2n
= 22, 44, 66. Sandv or rocky hills and plains,
500-1500 m elevation. COUNTIES: Ri\erside,
San Bernardino, San Diego.
Wright's threeawn intergrades with the \arie-
ties piiq)iirea,Jcn(Ueriana, and parishii.
Selected specimens. — San Bernardino
Co: Slo\er Mts, 14 Aug 1907, Reed F. M. 1307
[WIS]; 2.5 mi SE of Kingston Peak, T19N
RIOE, Sec 34-27. 23 Oct 1977, Ilenrickson, J.
16321 [RSA/l^OM]; rocky can\•()nbet^veen Bul-
lion and Sheep Holt Mts. 7 Apr 1940, Munz,
P. A. 16568 [UC]. San Diego Co: 3 mi WNW
ol'lacumba, T18S R8E, 3 Sep 1937, Yates, H. S.
68()5[RSA/POM].
Aristida ternipes Caxanilles, Icon. Pi. 5:46.
1799. Tufti'd pcMcmiials; minis few, erect to
s[)ra\\ ling, simple or ouK weakK branched, 2.5-
80 cm tall; internodes glabrous. Slieatlis mostly
longer than the internodes. Li^idcs 0.2-0.5 mm
long. Blades (Lit to inxolute, 5-40 cm long, 1-2
nun wide, with scattered long hairs above the
ligule. Panicle 15-40 cm long, open, the
branches widelv spreading from the main axis
and naked ;it the base, axillan ]iul\ini present.
19921
Genus, Ay>'/.s/7/)\ i\ (:\i,ii-()1{\ia
51
Spikclcts oppressed or sprcadinsj; Iroiii the
hranclu's. GliiDics about e(jual, l-nciAcd. 9-15
nun long. Lcinnia 10-15 mm long. nsnalK not
twisted at the ape.\; aivns e(|nal to \ci\ nnc(|iial.
Anthers 1.2-3 mm long.
var. hamuloHii (llenrard) Trent, Sida
14(2):26(). 1990. HooKTllHKK WW (Fig. 1 1) [.A.
hdinulosa llenrard, Med. Hijk.s Herb. Leiden
.54:219. 1926]. Central awn 10-25 mm long.
Ltifcral aicits mostly 6-23 mm long, .sometimes
shorter. 2// = 44. Dw hills and slopes. lOO-SOO
ni ele\ation. COUNTIK.S: Butte, Colusa, Fresno,
(ilenn. Kern, Los Angeles, Madera, Kixerside.
San Bernardino, San Diego, Santa Barbara.
Sonoma, Stanislaus. Sutter. Tehama. Tulare.
Wntura, Yolo.
Trent and Allred (1990) doeumented the
moiphologie \ariation and similarit\()LA/7.sf/r/c/
Irrnipcs and .A. Iianuilosa. eoneluding that tlu^
lunniilosa taxon should be treated as a \ariet\ ol
fcrnipcs. \'ariet\' ternipcs does not oceur in Cal-
iloniia and differs oiiK in the length of the
lateial aw lis. \'ixnct\luniinli>s(i also resembles A.
(lit tiricala. which diffeis most eonsistently in
liaxing shorter anthers and lacking pilose hairs
ab()\e the ligule. Based on numbers of speci-
mens in California herbaria. \ar. hainulosd is
unusualK' common.
Selected specimens. — Butte Co: Soudi
Butte, 10 Sep 1981, Ahart 1535 [UCJ; along
lIwT 32, 1 mi E of Chico, 16 Aug 1983, Ahart.
L. 4277 [TAES]. Colusa Co: 10 mi W of Wil-
liams, 5 Jul 1955, Burcham, L. T. 317 [AllUC,
TAES, UC]; 10.7 mi SE of Leesville, 19 May
1 958, Crampton, B. 4789 [AHUC]. Fresno Co:
Citnis Grove, 11 May 1940, Hoover, K. F 4385
[UC]; 8 mi N of Orange Cove, 8 |nn 1960,
Howell, J. T. 35481 [ISC]. Glenn Co: 5.5 mi S
ofOrland, 29 May 1942. Beetle. A. A. etal. 3353
[AHUC]; 5 mi \\' of OHand on the XewAJllc
road, 27 May 1914, Heller, A. A. 114.32 [US|.
Kern Co: lowest slopes of the Tehachapi Mts.
15 mi S of Bakersheld. 14 Apr 1942. Beetle.
A. A. .3017 [AHUCJ: 15 nn S of Bakersfiekl
7 |un 1946, Beede, A. A. 4679 [UC]. Lo.s Ange-
les Co: Alta Dena, 2 Apr 1905. Grant 66-64.59
[ARIZ, BS.ATOM, UC]; Pomona, 1 Jul 1937.
I lorton 448 [UC]; Li\eoak (^an\-on, San (iabriel
Mts. 15 Apr 1934, Wheeler, L. C. 2525 [ A H UC | .
Madera Co: near Raviiiond. on sheep lancli.
II Nhiv 1934, Wikson.'E. s.n. [AHUC]. River-
side Co: 10 mi N of Pala, 17 Way 1964. Hitch-
cock. C. L. et al. 23113 [NY]; lower San Jacinto
Rixer Canyon. Yates, H. S. 6710 [UC]. San
Bernardino Co: near Upland. 7 Xo\ 1916,
John.ston, I. 1121 [ARIZ]; nie.sa near Rialto. 20
May 1888, Parish, S. B. [UC]; Granite Nh)un-
tains, Budwei.ser Wash, 28 Oct 1977. Prigg(\
B. A. et al. 2321 [RS.VPOM]. San Diego Co:
Rolando. 14 |an 1938, Gander, F F 4936 [SD];
San [amento'. 4 [nl 1890. Hasse, H. E. s.n. [NY];
Escondido. 10 "Aug 1928, Meyer 652 [JEPSJ.
Santa Barbara Co: Santa ('ni/ island. X of
biological station in central \alley, 23 Apr 1979,
Thorne. R. F et al. .52466 [RSA/POM].
Sonoma Co: Little CyeNsers, 1 mi E of Big
Sulpliur Creek. 10 Aug 1984. Leitner [UC].
Stanislaus Co: \ icinitx of La Grange, 30 Sep
1961, Allen, P s.n. | AHUC, JEPS]. Sutter Co:
Sutter Buttes. 10 Sep 1981, Ahart L. 3129 [NY].
Tehama Co: about 5 km N of Black Butte
Resenoir and about 17 km N\\' ofOrland, 26
.Mar 1990. Buck. R. 1469 [JEPS]; Jelly's Fenv
Rd. 0.5 mi from 1-5 exit. 16 Aug 1991. Allred
K. \V. 5467 [NMCR|. Tulare Co: Three Rixers.
24 Aug 1905, Brandegee s.n. [UC]; 10 mi SE of
Portenille on Tule Indian Resen'ation Rd, 28
Dec 1964, (;uthrie. L. 66 [AHUC]; Fountain
Springs Rd. 6.3 mi W ol (California Hot Springs.
25 Jnn 1966. Twisselmann, E. C. 12537
[AHUC]. Ventura Co: Upper Santa Ana
Creek, Santa Ynez footliills, 13 fun 1957. Pol-
lard. H. M. s.n. [TAES]. Yolo Co:" foothills, open
slope. 2 mi W ol Winters. 24 Aug 19-53. (^ramp-
ton, B. 1600 [AHUC].
ACKNOW I.KDCMENTS
I am grateful lo the Friends of the Jepson
Ilerliarium. who proxided traxel funds for stuck
in Caliloiiiia. to an anon\uious rexiewcM' lor a
meticulous criti([ue, and to the curators of the
foHowing herbaria for their hcdpful cooperation
and n.se of plant materials: AHUC, ARIZ. DA\'.
JEPS. RS.\/I^()M. TAES, UC, and US. Geoffivx
Le\in of the San Diego Natural Histon
Mu.seum })r()\ ided \aluable assistance by track-
ing down pertinent collection information. Paul
Peterson of the Smithsonian Institution and
[ames P. Smith of Humboldt State Unixersity
look time to locate specimens and information.
and John W. Reeder and Richard Ledger ol the
Unixcrsitx ol Arizona generously shared with
mc^ before publication their obsenations on
Aristida californica. The illustrations were
expertK- rendered by Robert DeWitt Ley. Tliis
is [oumal Article No. 1583. New .Mexico Agri-
cultural Experiment Station.
52
(;hkat Basin N atuhalist
[N'oli
LiTKRATURK CiTKD
AlJKAMS. L. 1923. Illustrated flora of the i^icific States. X'ol.
I. Stanlord University Press, Stanford. California.
Ali.KI;1). K. W. 19(S4. Morphologic \ariatioii and elassihta-
tion of the North Auwrican Arislkla inirpiirca complex
(Gramineae). Rrittonia 36; 382-395.
. 1986. Studies in the Aii.stidfi ((Jraniineaei oi the
southeasteni United States. I\' Ke\ and conspectus.
Rhodora 88(855 ): 367-^387.
Cl.ayton. W. D., and S. A. Hiwoizk 1986. (;enera
graniinnni: grasses ol the world. Kew Bulletin Addi-
tional Ser XIII.
IIl".Mi\HD. J. T. 1929. .A monograph ot the genus Aristida.
I. Mededeelingen \'an"s Rijks Ilerharinm Leiden .No.
58.
Hitchcock. A. S. 1924. The North .American species of
Ari.slkhi. (Jontrihutions of the United States National
Herbarium 22:517-586.
IllK M< i)( K .\. S.. and A ClI.ASK 1951. Mainial of the
grasses of the United States. United States Depart-
ment of .Agriculture MiscelKuieous Publication No.
200.
HOLMCHKN. P K., W. KELKf'.N AND E. K. SCIIOFIFI.D
1981. Inde.x Herbariomm, Pt. I. Holm. Scheltema, and
Holkema, Utrecht, Netherlands.
|i;rs()\ W. L. 1923. A manual of the ilouering plants ot
California. University of California Press, Berkelew
.Ml :\/ P A., and D. D.' Kfck 1968. A CaHfornia flora.
Uni\ersit\()t {California Press, Berkeley. 1681 pp.
Rkkdkh J. li, and R. S. Fkl<;KH 1989. The Arislida
ralifi>niic(i-^lahr(itfi complex i (Iramiiieae). Madrono
36;' 187-197.
Thkn'I'. J. S. 1985. .A studv of moqihological variabilitv in
divaricate Aristicia of the southwestern United States.
Unpublished masters thesis. New Mexico State Uni-
\ ersih. Las Cnices. 90 pp.
Tlu;\T J. S., and K. \V. Allhkd 1990. A taxonomic com-
parison oiAiisfida tcniipcs Cav. und Ari.stida luiinuhmi
Ilenr Sida 14: 251-261.
Rccriicd loMati njyi
Rciisrd21 Jaiuian/ 1992
Accepted 1 Fehnian/ 1992
Cicat Basin Naturalist 52(1), 1992, pp. 53-5S
TEMPERATURE-MEDIATED CHANGES IN SEED DORMANCY AND LIGHT
REQUIREMENT FOR PENSTEMON FALMERI (SCROlTiULARI.ACEAE)
StanlcN (;. Kittlu'ii aiul Susan K. McNcr
Abstract. — Pciistciumt pdlmcri is a sli(irt-Ii\ccl prrcnnial Iit-rl) coloni/iiiii distmiu-d sites in sciiiiarid liahitats in iIr'
western USA. In this stuck .seeil was liarxcsted lioni si.\ nati\e ami ionr seetled p()|)nlali()iis dnriniJ \\\o conseciitixe \c"ars.
In lali(irat(>i\ t;einiination trials at eonstant 15 (', considerable between-lot \ariati()n in prinian' dormancy ;uid light
icijuirenii-nt wasohsened. Fonrwet'ksol moist chilling ( 1 (-) indnccdsccondar\ilormanc\ at 15C. Cold-induced secondan'
donnainA was rexersed 1)\ one wt-ek oltlark incubation at 30 C. This warm incubation treatment also reduced tlu' light
requirement of unchilled. after-ripened seed. Fluctuations in dorinancN and light reijuirement ol buried seeds haw been
linki'd to seasonal chtuiges in soil temperatin-e. Pcnstcinou palmcri germination responses to temperature ap[X"ar to be
similar to those ol lacnltati\e winter annuals.
Kiij words: seed 'ji'iin'uiatiou. P(diiur jxitstciuou. seed hduk. induced doiiiunuij. heardtoiiinir. Fenstemon palmeri.
Seed dorniancx iiiechanisms function to
ensure that germination i.s postponed until con-
ditions are favorable tor seedling suiAi\al
(Fenner 1985). The le\el ol donnanc\' of an
imbibed seed is dependent upon its dormanc\'
jc\ ("1 prior t( ) imbibition and on the enxironmen-
tal conditions to which it has been exposed in
the imbibed state (Bewley and Black 1982).
C'hilling, es.sential for breaking dormancN' in
seeds of nian\' temperate species, induces \aiA-
inii decrees of secondan' dormanc\ in others
iBaskin and Baskin 1985). Conxer.seK, warm
temperatures increase and diminish dormanc\'
in other species. These temperature-mediated
changes in seed dormancy are related to tlie
s(>ason in whicli seeds undergo germination and
cmergenc(\ Thus, spring and fall germinators
tend to ha\e opposite responses to chilling and
warm-temperatures regimes.
Poisteinon palmeri Gnw is a short-lixcd
perennial lierb nati\e to the southern half of the
Cireat Basin and adjoining regions of the west-
em United States (Cronciuist et al. 1984). It
occurs across a fairh' broad range in elexation
(8(){)-275() m), colonizing n^latixch ojM'U. carK
successional sites such as roadcuts and washes.
Indixidual plants produce large (juautitics ol
seed tliat remain \ial)le for several vears in stor-
age (Stevens et al. 1981). Numerous popula-
tions ha\"e been successtulK established
through artificial seeding on a \ariet\' of sites
outside its natixe range (Stexens and Monsen
1 988 ). This \ersatilit\' raises questions about the
establishment strateg\' of this species. In this
stud\ the effects of moist chilling and warm
incubation on seetl germinabilits' were deter-
mined under controlled laboratoiA' conditions.
The results are suf licientK clear to permit spec-
ulation about seedbed ecolog\ and ha\e led to
the fieldwork necessan to confirm the conc-lu-
sions drawn herein.
In laborator\ trials on F. paluicri. Young and
Exans (unpublished data. C.reat Basin Experi-
mental Range, Ephraim, Utah) demon.strated
tliat germination at a constant 15 C was not
significantK lower than at an\- other constant or
alternating temperature regime. Germination
o\er a 28-da\- period was suppres.sed at mean
temperatures Inflow 10 and abo\e 25 C. Allen
and Me\(M- (1990) reported similar results in a
stnd\ of three Penstemon .species and suggested
the p()ssibilit\- of cold-induced secondaiy dor-
iiiancx in P fxihiicri. Field sowing of this species
is usualK ( allied out in late fall and is based on
tlu^ assumption that acoid treatment is required
to break dormancv (Stexens and Monsen 1988).
' IS. D<-partiiieiit oi Ai;rii iilture-. Kort-st St-niix-. IntirMiouiit.iiii Kcsi-artli Station. Slinib Stiencrs Uilxiraton., Provo. Ctali S4fi(l6.
53
54
GwEA'Y Basin Naturalist
[\ oluiiie 52
Mktiiods
Seed Ac([iiisiti()n
Ripened seeds were harvested frf)ni nine poj)-
nlations in 1986. Collections wen^ made from
eight ot the original and one n(n\ population in
19S7 (Table 1). Four of the populations were
from roadside seedings outside the native range of
this species. The genetic origin of the aitifici;illy
seeded populations is unknown. Eacli collection
was clean(^d using standard tec-hni(|ues and stored
in envelopes at 20 (' (room temp(^rattn'e).
\'iabilit\ l^etermination
An estimate of viahilitv for each 1986 collec-
tion was obtained using a tetrazolium chloride
(TZ) t(>st. Four replications of 25 seeds from
each collection were imbibed overnight. Each
.seed was pierced and placcnlin a \% TZ solution
at room tempcM'ature for 24 liours. Embnos
were then evaluated for xiabilitv using estab-
lished procedures (Grabe 1970).
Gibberellic acid (CiA,3) effectivelv' breaks dor-
mancv in F. pal inch .seeds (Young and Evans,
unpublished data. Great Basin Experimental
Range, Epln-aim, Utah). Four replications of 25
seeds for each 1986 collection were imbibed in
250 mg L" (».*\.s. Germination temperature was
a constant 15 G. Germination percentages,
determined after 2 1 davs, showed no significant
differences betwec^i TZ estimates of \iabilitv
and genninalion percentages in GA.3. Hence,
germination in (iA^ was the onlv measure of
\ial)ilit\' en'.ploved with 1987 seixl.
FAperiment I
Experiment I was started on 1 |une 1987.
-Mean time after harvest date was a])proximatelv
nine months (Table 1). The experiment was
designed to ck'termine the effect of thn^e teni-
p(M-ature pretreatnients on germination of seed
from the nine 1986 collections under two light
regimes. Pretreatnients inchuk'd: (1) chilling
for 28 days at 1 G, (2) incubation for 7 davs at .'^O
G, (3) chilling lor 28 davs at 1 G followed bv
incubation for 7 davs at .'30 G. and (4) no pre-
treatnient. (termination temp(Matm-e and dura-
tion following pretreatment was a constant 15 ( '.
for 21 days. The light regimes were a 12-hr
photoperiod and constant darkness.
Each pretri'atment/light regime combination
was replicated fovu" times for each of the nine
collections. Replicates consisted of 25 seeds
placed on top of two germination blotters in a
100 X 15-nnn petri dish. Blotters were moist-
ened to saturation with deionized water.
Experimental units assigned the same pretreat-
ment and light regime were randomized in stacks
of 10. .'\ blank dish (blotters but no seeds) was
placed on top of each stack that would receive
light, ensuring that all seeds would receive light
throuiih the sides of the dish onlv. Litiht intensity
inside the dishes was 25 microein.steins m" sec'
PAR. Each stack was enck)sed in a plastic bag and
looselv sealed with a nibber band to retain mois-
ture and facilitate handling.
Dniing pretreatment, stacks were placed in
cardboard boxes, each of which was enclosed in
an additional plastic bag. After pretreatment,
stacks assigned the light regime were removed
from their boxes and randomly arranged in the
growth chamber directl)' beneath fluorescent
lights. The remaining boxes were placed in the
growth chamber and were not opened imtil the
ei^.d of their germination period.
Seeds with radicle extension >1 mm were
counted as germinated. Experience wath this
and other penstemon species has shovvni this to
be a clear indicator of the initiation of seedlins
development. A germination percentage was
determined for each replicate (dish). Germina-
tion percentages were arcsine transformed for
statistical analvsis. Experimental results were
subjected to analvsis of variance procedures
appropriate to the completelv randomized
design, l^ecanse of the collection X treatment
interaction in the analvsis of variance, each col-
k^ction and treatment was analvzed indepen-
dentlv. Significant differences among treatment
and colk^ction means were determined using
the Stndent-Neuman-Keul (SXK) method.
Experiment II
\ second (^\p(.Miment was started on 14 Octo-
ber 1987 using nine fresh (1987) collections
(Table I ). Mean time from hanest was approx-
imatcK one month. The objective was to deter-
mine the ellec't of .30 (1 (imbibed' on prinian-
dormancv and light recjuinMuent of fresh seed,
'fhe methods w(>re the same as those used in the
first experiment w ith tluee exceptions: onlv one
pretn^atment was used (30 C]), the length of the
preticatment was 14 davs, and the length of
germination v\as 28 davs. Light and dark con-
trols (no warm incubation) were a<iain included.
19921
ri:\srFMO\ PALMi.ni Skkd C'.i:rxMi\ vnoN
55
Tablk 1. Location and harvest dates tor 10 populations ( IS colk'ctions duruiii twoxcars' ol P pahncri. All populations are
n Utah except the Mountain Home pojiulation in Idaho.
Lat (N)
Long (W)
Ele\ation (ni)
11;
;ir\ est date
Collection
1986
1987
Snow's ("an\()n
37 12'
1 1:5 39'
loso
S/14
Urowse
37°21'
n3°L5'
1350
8/22
8/14
Let'ds
37°14'
1L3°2U
]()50
8/8
8/14
Zion
37°14'
112°54'
1740
8/22
9/14
Kolol) lioad
37°16'
n3°()fV
1410
8/8
9/13
Utah Hill
37°08'
1 13°47'
13S0
8/8
Mountain Home''
42°57'
1 15°()5'
9:)()
8/13
8/27
Mercur Canxon''
40°25'
112°10'
1650
12/15
9/22
Salt (;reek ("auNon''
39°42'
111°45'
1740
9/10
10/10
NeI)o IjOop'
39°52'
nr4()'
2100
1 0, 2fi
10/10
'ArtilkulK
.utslde tlu- n.aural la.
RESULTS
Experiment I
Four weeks of chillino; redueed "â– eniiinati(Jii
ill light significantly below the level of controls
lor six of the nine collections (Table 2). Incuba-
tion at 30 C caused no significant change for
germination in light when compared to the con-
trol. When the four-week chill was followed bv
one week at 30 C, mean germination percentage
was onl\' slightK lower than that of the control.
This indicates that incubation at 30 C effecti\el\
reversed the secondan dornianc\' induced bv
chilling. \n addition, incubation at 30 C' substan-
tialK increased the dark germination ptM'cent-
age over the dark control (Table 3). The 30 (>
warm incubation was much less effectixe in
HMuoxing the light requirement when preceded
b\ chilling.
CTermination rate at 15 C was onl\' slightlv
accelerated b\ chilling and warm incubation
pretreatments (data not shown). Mean gcMiiii-
nation for the light control treatment after se\(Mi
da\s was 15%, indicating that most essentialK
nondonnant seeds recpiired a considerable
period of imbibition before germination was
possible. Foiu' weeks of chilling and one week
of warm incubation increascxl the piojjortion of
seeds that germinated b\ da\ 7 to 24 and 28%,
respecti\el\ . Howexer, a major fraction of the
seeds still required more than one week of con-
stant imbibition at 15 C to cerminate.
Experiment II
In the first experiment there was a slight trend
in the more dormant lots for germination to be
higher after warm incubation relatixe to the
control. The second experiment was conducted
to determine if warm incubation could break
the priman- dormancv of fresh seeds.
Contran to what was expected for fresh seed,
onlv t\vo of the nine 1987 collections showed
significant priman' dormancx' (Table 4). The
increase in germination percentage following
warm incubation was significant when com-
pared to the nonincubat(>d light control for one
of these collections. In the remaining collec-
tions, neither the light control nor the light,
warm-incubated germination percentages were
significantK' different from total \iabilit\- esti-
mates determincnl In germination in GA3.
The xariation in dark germination was similar
to that obser\'ed in the first experiment with
after-ripened seed (Table 4). The effect of warm
incubation on dark germination was not as clear
as in the initial experiment. Germination of the
warm-incubated secnls resulted in a mean net
increase oxer iioiiinciibated, dark controls of
onl\- 11%. Fotn-of the nin(> collections showed
significant increases, whili' one showed a
decR^i.se.
Discussion
Moist chilling for four weeks caused vaning
degrees of secondan dormancy in P. palnwri
seed collections. Incubation at 30 C clearl)
56 Gii MAT Basin Natl'ivxlist [\bliiiiie52
TaBI.K 2. CkM-mination response ofniiie after-ripc'iied collections of'/' jxihiwri seed to moist cliilliiiti; ( 1 (^ lor 2S days) and
warm incnhation (30 C for 7 davs). Tlie germination period was for 21 days at a constant 15 ( .' witli a 12-hr photoperiod.
Cermination in 250 mg L (lAr; was nsed as an estimate of total \ial)ilit\ lor cacli collection.
Mean germination percentage'
Pretreatment
Collection
(.'ontrol
Browse
yoa
Li'cds
S9a
Zion
72a
K.0I0I) Hoad
95a
Utah Hill
S9a
Moimtain \\t
)me
S<Sal)
Mercnr (Jan\()n
S61)
Salt Creek C;
an\'on
5SI.
Neho l,oo[)
75a
Means
S2I.
1 c 30 c 1 c/3() c: c;a3
411)
38c
73a
63b
39b
65b
21c
55b
38b
4S,I
92a
Sda
91a
92a
73b
93a
SOa
71a
81a
90a
86a
97a
.S8a
78a
82a
89al)
S7al.
92a
S71)
81b
99a
SOal)
72b
92a
S4a
SOa
89a
S7I)
79c
91a
broke cold-iiuhiced secoiidaiA d()inuuic\ in siil)se(|uentl\, light seiisitixih" is sti'ongK' inflii-
altei-ripciied seed, and there is some indieatioii enced In conditions during ripening (Cresswell
that it can reduce le\els of priinan'donnane\' as and (rrinie 1981, Gutternian 1982) and may
wc^II. Tlie warm-indnced reduction in iigitt xan consitlerabK' among the seeds of a single
recjuirement was less pronounced tor fresh plant (Silvertown 1984). The f! /w/z/icn seeds in
conij)are(l to after-ripened collections. these experiments demonstrated three lexels of
The res})()nse of F. pahiicri seeds to moist response to light, suggesting \ariable levels of
chilling and warm incnhation parallels those total or active phvtochrome in the seeds. Some
obsencd lor tall germinators (winter annuals) seeds germinated in the dark while others
(Baskin and l^askin 1985). This is supported bv required light, and a few remained dormant
the lack of primaiy dormancy in fresliK- har- e\en with light. The proportion of seeds that
vested seeds. Nevertheless, a significant portion could germinate in the dark was increased bv
of the seeds was not induced into secondarx incubation at 30 C> (Table 3).
donnanc\(lnringchilling. This suggests tliat late Light sensitixitv can be altered b\ tempera-
winter/eady spring germination of some seeds ture shifts during seed imbibition (Toole 1973,
is likely. It is of littU^ surprise^ that recentK Franklin and Ta\lorson 1983). This ma\- be due
emerged .seedlings wt>re foimd in /' jxibiwri to temperature effects on the production,
populations in both spring and fall. Such destruction, or dark rexersion of pin tochrome.
biuiodal germination patterns are txpical of tac- Temperature shifts mav al.so alter other factors
ultatixe winter annuals (Ba.skin and Baskiu associated with plntochrome action, thus
1985) and would be selected for in uupredict- resulting in an increase or decrease in light
able habitats where the best season for seedling seusitixitv. Hendricks and Tavlorson (1978) sug-
.sur\i\al maxdiffer from year t()year(Sil\-ertown gested that temperature effects on plnto-
1984). Such germination patterns xxould also be chrome action in s(>eds max be due to changes
adaptix-e for .species that colonize different kinds in membrane llniditx. It is iikelx that the effects
of habitats xxith xaning degrees of threat from of tcMuperatiuc on light .sensitixitx in seeds are a
fro.st and drought. Both .situations occm- xxithin r(\sult of mor(> than one process acting in concert,
the range of /^ /W///t'n. A light recinirenient max Iielp (k'tennine
(Tixen its small .seed .size (Pluminer et al. sc^ison of germination for buricnl /' jxilntcri
1968), alight requirement for germination of F. seeds. I labitatsxvith adecjiiatewintcr snowspn
pahiicri is not .surprising (Fenner 1985). Th(> xide enough moi.stiu-e for .spring germination of
lexel of actixe phx tochrome in dn- .seeds and, suriace seed. Long periods (8-16 xx'eeks) of
1992] Fi:\sti:m()\ rMMF.iu Sv.KD Ckhmiwtiox 57
T\ni I o. The cITcct of chilling (1 C; lor 2<S days), wanii iiiculnition (oO (.' for 7 ila\s>, and diilling followed In- warm
iiniiliation on the ligiit r('(jnir(>nient of nine after-ripened collections of P. palnieri. Tlie germination temneratnre was 15 (].
Ciermination percentage''
Light Dark
(loilection Control C^oiitrol IC
30 C
1 C/30 C
751.
17e
6S1)
13d
551)
24c-
77]i
34f
7()a
33b
S7a
65ab
S3a
38b
76a
46ab
fila
35b
721.
34(1
Browse
yoa
Leeds
89a
'/ions
72a
Kololi H(ud
95a
Utah Mill
89a
Monnt.iiii 1 l(
ime
88a
MiTcnr ( 'an\
on
86a
SaltCrei-kC.
ui\on
58a
N'el.o l^ooj)
75a
Means
.S2a
5(-;c
oZd
45c Ux\
37c 35e
49c 31c
41b 231)
54b 591)
42b (iv
26b .541)
12c He
4()c 27e
'Williin ..collrctio.i. inr.iiis l.ill.mfil In till- s.uuv U-ttc-r.iiv not siniiiliciilK dillrn-iit ..t tl.ry, lir, 1,a, I ,SNk
Tahi.I-; 4. Friman tlormancv, light re<|nirement. and the effect of warm incnl.ation i 14 da\s at 30 (.') on tlu' germination
I nine tresh collections of P. palnieri seed. The germination period was 28 da\s at 15 C. Light treatments reeeixcd a 12-hr
ihotoperiod. (ierminatnon in GA3 (2.50 mg L' ) was nsed as a measnre of xial.ilitA for each collection.
(termination percentage''
Control 30 C pretreatment
Collection Light Dark Light Dark C.\.i
Snow's CaiiNon 94a .')lli S5a 34l. 97a
Browse 86a 25c SOa 53b 93a
l.ce<ls 92a 35b 91a 511) 92a
/.ions 70a 38b 72a 24c 74a
Kolol. Boad 83a 30b SSa 171. 87a
\iounlam Ih.me 96a 56b S7a fifih 94a
MercnrCaTiyon 87a 58b S7a 76a 94a
Salt Creek CainoM 77bc 45d S6h 67c 98a
Nel.oi,oop 55b 16c 71a 401) Sla
.Means S2h .'57(1 S.51. fSc 9()a
'Willim .ndllcctuHi, iM.-.iiis hillcurcl In tin- samr l.ltcr .i.c rinl sii;inlK .uilK <l)ll, r,-iil ..I llic/i <^ .11.5 Ic-M-l .S\ki.
moist eliilliiio; rcdiicc tlic time lU'cdcd tor o;cr- liiiricd sccd.s witli a \'\\l\\l rcMjuinMiicnt arc liiiif-
miiuition to occur, thus incrca.siuo; the cliancc.s tioiialK dormant and would contriI)utc to the
oi .spring-germination and sectHing cstahhsh- seed hank. ,\i)[)arcntly, chilhng docs not reduce
ment from .seeds not inchiced into secondarN the hght re(iuirenu^nt in F. /jr////u'n seeds, while
donnancv (Kitchen and Me\er, unpuhlished warm incuhation ehminates it in a .significant
data on file at the Shnib Sciences Lahoraton, fraction of tlie. seeds (Table 3). This .suggests that
Provo, Utah). Rapid dning of the .soil surface huried seeds nia\' be more Hkely to germinate in
would make the gerniination of surface seeds tlie fall after (experiencing sufficient warm incu-
tollowing summer or autumn rains less likeJw bation to eliminate their light re(juirement.
58
Great Basin Naturalist
[Volume 52
Whether current-vearF. palmeri seeds germi-
nate in tlie fall or spring may depend as much
on time of seed dispersal as temperature and
moisture eonditious that follow. Tlu^ collection
dates for each population (Table 1 ) and field
obsenations regarding the timing of fruit dehis-
cence suggest that populations from areas with
milder winters (lower elexations) tend to ripen
and disperse seed during late summer. At higher
elexations where cold weather would occur ear-
lit^-, seed ripening and dispersal are delayed.
Habitats with mild winters and unpredictable
spring moisture sei^n to favor early dispersal
and fall eerinination. Such sites select for the
maintenance of a seed bank because extended
periods of drought are t\pical and conditions for
successful establishment may not be met for
many years. Cold-induced secondaw dormancy
and burial of light-requiring seeds should facil-
itate the buildup of this soil seed resene. In
habitats with more se\'ere winter conditions dis-
persal is retarded and spring germination of a
portion of tlu^ seeds is both probable and less
riskA'. The presenation of a seed reser\e through
cold-induced dormanc\' may also be important
in these more mesic habitats.
Fcustcinon pdhiich appears to be adapted for
(^stal)lishment in a variety of habitats. Two phe-
nomena are important in this success. First,
individual seeds seem to be capable of respond-
ing appropriat(^l\ to different environmental
stinmli. S(^cond. variability in germination
respon.se among .seeds within a population is
indicative of a bet-hedging strateg)' increasing
the chances for successful establishment across
a range of variabe and unpredictable environ-
ments. 1 labilat-related between-population
variation in germination timing mechanisms
appears to be n^lativelv unimportant.
ACKXOW LKDCMKNTS
This research was funded in part bv grants
from the Pittman-Hobertson Federal Aid to
Wikllife Project \\<S2-R and the Utah Depart-
ment of Atiriculture.
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n-qiiirnieiits lor seed germination of three Pcnstcmon
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Fewer, M. 1985. Seed ecolog\-. Chapman and Hall,
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Received 25 October 1991
Accepted 23 Xoveinher 1991
Crcat Basin Xaturdist 52( 1 ). 1992. pp. 59-67
LATE QUATERNARY ARTHROPODS FROM
THE COLORADO PLATEAU, ARIZONA AND UTAH
Scott \. Elias . |iiii 1. .Mead", and Lam D. A'^ciihroad"
AbsTIUCT — Late (^)iiatfniaiA-aL:;c arthropods wvre recowred from tin cavi^ deposits and pai'kiat middens located in the
Grand Camon. CaiiNonliuids. and Cden C^anNon region ol the (Colorado Phiteau. This QnaternaiA data re.source has not
been anaKzed before from the Colorado Plateau national parks. Radiocarbon dates on tlie xarions deposits containing
arthropotls range from 1510 to 30,660 \t B.P. The fossil assemblages \ielded 57 identified t;L\a of insects, arachnids, and
HiilHpedes. including 15 ta\a taken to the specie.s level. The information from tlic fossil insect record of the (>olorado Plateau
is not \et sulficieut]\' detailed to permit precise paleoeu\ironmental reconstructions. However, preliminan' conclusions
suggest a cooler, moister climatic regime during the late Wisconsin glacial and a mosaic of vegetation tvpes, such as grassland
and shnibln conunnnities. unlike the present vegetation at tiie localities.
Ki'ii uonl.s: Qudtcnuirij. Citlonulo PUiicau. iiiilirojuxls. \\ iscoiisin ijjdc'uil. CrautI ('(uii/oii. races.
This paper discusses the results of a prehiiii-
uan- stucK' of late Quateman" arthropod fossils
from ca\e deposits and packrat unddens from
southern Utah and northern Arizona. This Qua-
teman data source has not been anal\"zed
before from the Colorado Plateau, although the
arid Southwest has been the focus of pale-
oen\iroinuental studies for appro.ximateK* half a
centuiA' (Antevs 1939). Arid climate, coupled
with episodic fluctuating water tables, has
[)ro\en detrimental to the preser\'ation of most
exposed fossil remains. However, the same xeric
conditions, when coupled with a stable rock
shelter, pnnide a tmique situation — dn' preser-
vation. Such xeric locations provide the preser-
V ation of not ouK' pollen and plant niacrofossils,
but also soft tissues and other usualK' degrad-
able remains of animals (such as skin, hair, kera-
tinous tissues, and dung; Wilson 1942). The
studx of packrat middens in the Southwest has
provick'd a reconstruction of the Wisconsin gla-
cial biological conuuunities never before
obsenablc in such detail (see various chapters
in Hetancouil ct af 1990). Thus, an entirelvnew
held of research has been opened, and it should
[)rove valuable in understanding tlie latest
Pleistocene.
On cave deposits were (juickK discovcMcd to
])(' a warehouse of late Pleistocene information.
C\psum Cave (near Las \egas, Nevada) and
Rampart Cave (western (Trand Can\on. .Ari-
zona) were the .scenes of the first paleoecologi-
cal studies utilizing drv-preserved dung ol an
extinct animal. Landermilk and Munz ( 19.34.
1938) found a wealth of information presened
in the dung of extinct Shasta ground sloth
[Nothrotlichops shastciisis). Later studies con-
cerned witli dietaiT recon.stnictions expoimded
on the data axailable from dung of extinct her-
bivores, including Shasta ground sloth, mam-
moth [Manuntitluis). Harringtons mountain
goat {OrecDHiios liarhn^totii), and bison
(Bison), among others (.\hutin et al. 1961.
Hansen 1980. I3avis c-t al. 1984, Mead,
O'Rourke, and Foppe 1986, .Mead, Agenbroad
et al. 1986, Mead et al. 1987, Mead and
Agenbroad 1989).
Packrats iXccHoiiui: Hodentia; (dicetidae)
build nests surrounded bv construction materi-
als collected from within 30 to 100 m of the
house. The construction components are pre-
dominantK plant materials, but the packrat also
collects small stones, skeletal remains, and
dung. .Adding to the mattMnals procured by the
packrat are various vertebrates and inverte-
brates who live in the nest and waste pile as
cornmen.sals. Periodic hou.se cleaning produces
a vv aste pile of debris. Urination on the waste
pile (a nudden) ultimately may cement the
remains into a rock-hard deposit, encapsulating
, Institute of Alpine Researcli. Box 4.50. University ofColorado. Boulder. (:olora<Io S().309-()4.5().
"Quateman.- Studies Program and Ue|)artinent oiC;<-o!oi,'\\ Bov .56-t4, Nortlieni Ari/ona University. FlagstafT. Arizona S6()l 1-5644.
59
60
Cheat Basin Naturalist
[N'oluine 52
105
..Albuquerque
B-K = Bida a Kaetan caves
E = Escalante River localities
eek Canyon
-40
-35
Fig. 1. .Map ol'tlic Coloradi) i'latcan with sites disfiissrd in text.
the coiilcnt.sot tliut tiiiic. W'licii tlicsc iiKliiratcd
(cemented) inicklens arc located in a dn alcox c,
rock .shelter, or caxc, tlic contents nia\ he pre-
served lor as lon<j;as tlie slielter exists, i^adiocar-
bon dalint^ol indurated midden layers proxides
a chronoIoij;icaI framework (or the associated
plant and animal remains. Micklens, then, pro-
vide a imicjnc examination ol local past hiotic
connnnnities.
The investigation of insect fossils from ancient
packrat middens and cave (k'posits is a new
approach that is jnst !)e<i;innino; to.show snl)stan-
tial resnits. One of the anthers (SAE) recently
performed mon^ extensive res(>arch on a seri(^s
of insect fossil asseml)la<ji;es from packral mid-
dens in the (>hihnahnan desert regions of west-
em Texas and sonth central New Mexico (Elias
I9.S7, Elias and \an Devender 1990, 1991).
Elias (1990) also recently pnhlished the resnits
of a taj^honomic stnd\ designed to reveal the
sonrces and possible biases of insect exoskele-
tons in packrat middens.
Mkthoi:)S
1 .ocalities
.Matrices Irom packrat micklens and cave sed-
iments were washed or hand picked for arthro-
pod and other animal and plant remains.
Packrat midden and ca\e deposits from two
caxc sites were analyzed from (irand Canyon
National Park (GRCA), Coconino Conntv; Ari-
zona; three packrat middens from Salt Creek,
Canyonlands National Park (CANY), San jnan
19921
QUATKHWm Al'.TIIHOl'ODS, Coi.Oim^X) Pl.ATKM"
61
(]ounh; Utali; and three paekrat middens and
one ca\'e de[)()sit tioiii tlie Kscalante Hi\er
region ol Cdeii (.'aiixoii National Hecreation
Area (GLCA), Kane County, Utah (Fig. 1 ).
Bida Ca\e is a large limestone eaxc located in
])in\()n-jnniper woodland at 1430 ni ele\ati()n in
CHCA. Cole (1990) reported on the paekrat
niid(k'ns recovered from the ca\'e. Test pit e\ca-
\ati()ns produced a multitude ol faunal and
lloral remains (Mead 1983, OUourkeand Mead
1985, Mead, O'Rourke, and Foppe 1986,
Mc\'iekar and Mead ms). Radiocarhon dat(\s
(spanning from 2960 to 24,190 \t Bd'. ) on \ari-
ous remains are presented in Mead (1983) and
Mead, Martin et al. (1986); those ages from
units containing arthropod remains are listed in
Table 1.
Kaetan Ca\e is a medium-sized limesttjue
cawat 1430 m cdexation in GRCA. Mead ( 1983)
e\ca\ated portions oi the deposit in tlie
entrance room for the remains ol extinct moun-
tain goat (Orcainnos Jiarhiif^^toiii) (O'Rourke
and Mead 1985, Mead. O'Rourke, and Foppe
1986). Paleoenxironmental I'econstrnction
l)as(^(l on the macrohotanical remains reco\(^red
honi paekrat micklens and stratilied sediments
is in manuscript (McV^ickarand Mead). Radio-
carhon ages span the period from 14,220 to
30,600 vrB.R (Table 1).
ThrcH^ paekrat luiddens selected from a series
collected from Salt Creek Canyon, CANY (1505
to 1755 m elevation), have radiocarbon ages
spanning 3830 to 27,660 yr B.R; toda)- the
region is piuNon-juniper woodland with sage-
brush Hats. Hie analysis of the maciobotanical
remains and [)aleoen\iromueiital reconstruc-
tions ol the middens is in man nsciipt (Mead and
Agenbroad).
Bechan C.dw contains copious remains ol
extinct lied)i\()re dung ( Daxiset al. 1985, Mead,
.\genbroad et al. 1986, Mead and Agenbroad
1989) recovered from floor .sediments dating
I 1 .600 to 1 3.505 yr B.R Arthropods were recox -
cred from tlu^ dung kucr and from an isolated
ilolocene-age paekrat midden in the ca\e
liable 1). Other nearl)\ [)ackrat middens con-
tained additional arthropod remains dating
Irom 1510 to 8640 vr B.R
Insects
Fossil insect sclerities were sorted from
washed paekrat middens and ca\e sediment
matrices. Robust specimens were mounted on
modilied luicropaleontological cards with gum
'i"\ lii I I 1 „itc (,)u;itcTnai-\ deposits and ladicK-arhoii dates
1)111 sites on tlie (loiorado l^latean eontaininij artliropods.
l.oealit\
'Cane
l>al) nnniher
Ciiand Claiuon National Park, .Vri/.ona
HidaCaw
l.iver2
29(S() ' 200
.\-2836
L.a\vr 4
Hi, 150 r 600
HL- 11.35
l.a\cr .â– )
none
—
r>averS
24,190 + 4.3(X)
2800
.A-2.373
Kaetan ( -avc
I,a\er i
1 1.220 - .â– 520
.•\-28.'3.5
Laser,".
IT,.!!)!) + .'jOO
.'\-272.3
I^a\ci" 3
none
—
l,a\er fi
.■30.600 ± 1800
.\-2722
I,a\erS +
none
—
I'aekrat niid(
len 11) 17.100 - .500
.\-2719
Owl Hoost
1^2
21.430 i 1.5(X)
A-;3082
none
—
Canyonlaiuls National Park, I tali
Salt (Ireek (.'an\on i paekrat miildens)
Head ( )\\l 1 A 38:30 ± 70 lieta- 18267
Woodenslioe 1 6980 ± 120 Bcta-27214
Hoodoo 1 27,660 ± .•340 Beta-27213
Glen Can>«)n National Hecreation .Vrea, I tali
Ksealante Ki\cT region i paekrat nnddens)
13eehan ( :a\c 3 1510 ± 60 Beta-2.-3706
C;o\v-Perfeet 1 1820 ± 100 Beta-2;371 1
Bow lis 1 8640 ± 140 Beta-2.3704
Beelian Caw 15S 1 1.600-13..505
»Mshnilu,\:,lM
.■I A..,nl,i„a,l M
• a...iK/i-cl on MatHinuthiis (TiiaiiinKilli i ami cf. EuccratUcr-
«■<• i)a\iM-t,il, 19S.5. Mead. .\<;ciilm)ail .-I .il. 19S(i, Me.ul
tragacauth. a water-soluble glue. Fragile sp(>ci-
meus and dnplicates wvvv stored in \ials of
alcoliol. Fossils wcrv identified chiefl\- through
comparisons with modern identified specimens
in the U.S. National Museinu of Natural Iliston
(Siuithsonian institution). Washington, D.C>.
Some sjK'cimens were referred to taxoiiomic
specialists, as noted in the acknowl(Hlgments.
Mod(Mn ecological re(|uirements and distribu-
tions for species identified in the fossil assem-
blag(\s were comj)iled from the literature and
from s])ecimen labels in the U.S. National
Museum. All s|)ecinients will be curated in the
National I'ark Service Repositorx, Laboratoiyof
(,)naternar\ Paleontolog\-, Quatemaiy Studies
l^rogram. Norihern .Arizona Unixersih.
Results
The fossil assemblages \ielded 57 identified
taxa of insects, arachnids, and millipedes,
including 15 taxa taken to the .species level.
Table 2 shows the taxa identified from the
62 Great Basin Natuhai,ist [Volume 52
Tahi.F. 2. Fossil arthropods klciitificd from Rida and k'aetmi caves. GRCA. Arizona, in miiiinniin number of indi\idu;Js
per sample.
Rida Ca\e Kaetan Cave
Taxon 2" 4 5 S l'' 5 S ()RR2' ()R2'' 11."
colkoi'tkka
Cakabidai-:
Cahmwui cf. scnttator Fal). 1 — — — — — — — — —
Aoonuni (Hlui(liiie) pcrlciis (.'sy. 2 — — — — — — — — —
Afi^oiiiiiii {Rh(i<liii(') sp. — 11 — — — — — — —
SCAHAHAIIDAK
Ai)h(>cliiis nr. nijicldrus Fail — — — 1 — — — — — —
Aplioiliiis sp. — — — 1 — — — — — —
OntliopJiOfius sp. — — — 1 — — — — — —
Serial sp. 1 — — — — 1 — 2 1 —
Phi/ll(>j)li(i^a sp. — — 1 — — — — 1 — —
Diplotdxis sp. 1 — — — — — — 1 — —
(^enus indeterminate 1 — — — — 1 — 1 — —
Sii.l'iiii:)AF.
Thdiuttopliilus tntn(tiiu\ Sav 1 — — — — — — — — —
PriMDAK
Ptinis ap. — — — — — — — 1 — —
Nipttt-s cf, ventrirulns LeC, — — — — 10 1 — 9 — 4
NlTIDUl.lOAE
Genus indeterminate — — — — — 1 — — — —
Dk.kmkstidak
Genus indeterminate — — 1 — 1 — — 1 — —
HiSTKKIDAi:
Ck^nus indeterminate — 1 — — — — — — — —
El.,\TERID AK
Genus indeterminate — — — — 1 — — — — —
Tf-nkbriomdaF':
Eleocles cf. ni^rina LeC, — — — — 1 — — 4 — —
Eleodcs spp. 1 1 1 — 14 2 4 11
Coniontis sp, — — — — — 1 — — — —
Mkloidai.
Genus indeterminate — — — 1 — — — — — —
Mki.andhyidak
Auaspis nifd Sa\ — — — 1 — — — — — —
ClIHYSOMEI.IOAK
Ia'hui trilined White — — — — — 1 — — — —
Chdetocncmd sp, 1 — — — — — — — — —
Genus indeterminate — — — — 1 — — — — —
Clf.ridak
Acantlioscelidcs sp. — — — — — — †” 1 — —
CURCULIOMDAK
Sapotcs sp, — — 1 — — — — — — —
Oplin/dstcs sp, — 2 — 1 — — — — — —
Scijphophonts dcupunctatus C,\]\. 211 — — — — — — —
Orinuxlciiw protrartd Horn 1 — — — — — — — — —
Clcoiiklius triiittdttis or
C (jiiddriliiu'dtits — 1 1 — — — — — — —
Apleums an<:,ul(iri.'i (IjL'C) — 1 — — — — — — — —
Genus indeterminate — 111 — — — —
Sc:OLYTIDAF,
Genus indeterminate — — — 1 —
Nkukoptf.ha
MVRMFl.ON-riDAF
Genus indeterminate — — — — 1
HOMOI'TFRA
ClCADIDAE
Genus indeterminate — — — 1
Hf.miftf.ha
Genus indeterminate — — — 1
19921
Quaternary Arthhofods, Coioi^mx) Pi.atkmj
63
Tahi.k 2 covriMED.
T;l\()ii
Bida Cave
Kaetaii Cave
4 5 S
S ORR2' ()H2''
Okthoptkra
ackididae
Germs indeterniinate
Lkpidoptf.ra
(»enu.s indeti'rniiiiatc
I I'l MF.NOI'TKHA
Apoidea
Genus indeterminate
DlPTKHA
Geims indeterminate
Abac ii\ II) \
ACAHI
IXOUIDAK
Dcnnaccutor mulcrsoiii Stiles
Dcrmaccntor sp.
scohpiomda
Bv:tiiidae
Centtiroides sp.
DiPLOPODA
Genus indeterminate
'Niimliers refer to laver numbers at Bida Cave-
NiiinlxTS refer to la\er numbers at Kaetan Cave.
'Owl R<x)st R2
â– 'Owl Roost 2.
' Paekrat midden lb.
Grand C>an\on region, and Table 3 lists taxa
identified from Glen Canyon. The assemblages
are dominated hv taxa still foimd todax in the
American Southwest, but many of the
Pleistocene assemblages contain species that
Ii\e toda\- at elevations higher than the fossil
localities. As in other packrat midden and ca\e
assemblages from the American Southwest, the
fossil faunas are dominated b\' a few families of
insects and arachnids. The beetle (Coleoptera)
families (;aral)idae (ground beetles), Curculi-
onidae (wee\ils), Ptinidae (spider beetles),
Scarabaeidae (dung beetles and chafers), and
Tenebrionidae (darkling beetles) were repre-
sented in most assemblages. A few packrat and
other mammalian parasites were found, includ-
ing a tick (Ixodidae) and a blood-sucking bug
(Rediniidae) that are knowni to parasitize
packrats in their nests. A number of the identi-
fied species merit indixidual discussion.
Discussion of Selected Species
The ground beetles from the fossil assem-
blages include both ca\e dwellers and open-
ground species. Th(^ cateipiHar hunter,
CalosoDia scndaton was found in a late
Holocene assemblage from the Grand Canvon
(Table 2). This beetle is widespread in the
United States, southern Canada, and northeni
Mexico (Gidaspow 1959). It has been collected
from the floor of Havasu (^ainon, GRCA (Ehas,
unpublished data). The ca\e beetle. A^omni
perlcvis (Fig. 2A), pre\'s on other arthropods. It
is relatively coimiion in caws and near the
mouths of mammal burrows. It is found toda\'
from the state of Chihuahua, Mexico, northwest
to southcni .Arizona (Barr 19S2). This species,
found in Iat(^ Holocene asseml)lages in both tlie
GLCA and (tHCA regions, was identilicd from
Holocene packrat middens from sites in th(^
(>hihuahuan desert region of Mexico (Elias and
\'au Devender. unpublished data). Another
groimd beetle from the kite Holocene record at
CtLC'A is Disrodcrus inipolrus. which Hxcs in
open countiA'. It is common throughout the
American Southwest and is found in the
Chihuahuan, Sonoran, and Mojave deserts.
The checkered beetle (Cleridae), Cynmioclcra
pallida (Fig. 2E), is a predator of bark beetles in
coniferous forests in the ('hiricaiiua, Rincon, and
Huachuca mountains of .Arizona, as well as in
mountainous regions of (Chihuahua. .Mexico
(Wiurie 1952). C. pallida was found in a late
Pleistocene sample from tlu^ (irand (]an\on.
The dung beetle (Scarabaeidae), Aphodius
nificlanis. was found in a late Pleistocene
64
Great Basin Natuhalist
[\<
olunie oz
Tablk 3. Fossil arthropods idcntiUcd from the Cainoiilaiids and Clcn Caiixon region, Utah, in miniinnni ninnh(>r of
indixiduals per sample.
Taxon
CANY'
DOl.A' WSl HDl
COLKOl'TKUA
C.\KAI5ID.\K
A}i,onum (Rltadiiic) pcrlevis (Isy. —
Aiiwra sp. —
Dlsaxlcnis inipotciis LeC. —
Ciemis et sp. indeterminate —
S(.ak.\b.^kii).m:
Apliodius spp. —
Atdcnius sp. —
Scrira sp. —
Mcloloiillia sp —
Diplotdxis sp. —
Genus et sp. indctciininatc —
Ptinioak
Niptus sp. 10
Ptiiiiis spp. —
El.vikkioai:
Genus et sp. indeterminate —
BVKHIIIUAK
C^enus et sp. indeterminate —
TF.NKBKIOMDAK
Eleodcs spp. —
Couiontis sp. —
Genus et sp. jniletcnninate 1
Di:hmi:stii)ak
(k^mis et sp. intieteiniiiiatc' 1
ClIKVSOMKLIDAF.
Altica sp. —
PachtjhnicJiis sp. —
(n^nus et sp. indeterminate —
Cl.KKIDAK
Ctjinatodcrd pdUuld Sehlir —
IIOMOPTF.HA
Rh.ni VIIDAK
Tridtomd sp. —
Lki'idoptf.ka
Geinis et sp. indeterminate —
MVMKNOI'TKH \
FOKMICIDAK
' Forinicd sii. I
glc:a''
HC.r' C-IM Bl BC;i.5S
9
â– "CANY = Cany<)i)l;imls National Park.
''GLCA = C;leii C.'anvon National Uecrcalion Area
'Sites in Caiivoiilamls are: DOl A. Dead ()«1 1 A; W .SI . WikkUh
''sites in Clen Canyon are: B(:.3. Beclian Cave .^: C PI Cm-Pii
»■1: HDl, lie;
(I 1. HI Hour
1: H( 1")S. Beeli.mCave 1,5S.
asscml)laL!;(' from (;IX>.\. This hectic lix'cs lodax
throughout much ol western North .America
from Saskatchew au iu the north to New Mexico,
Arizona, and Clahiornia in the south. At the
southern limit of its range, it liws in iiionntain-
ous regions.
The carrion beetle (Sil[)hidae), Tliaiialophilii.s
tntitcaftis (Fig. 2B), lives in die southwestern
U.S. and northern Mexico in habitats spanning
altitudinal gradients from grasslands and arid
scmb desert through oak-piinon-juniper wood-
lands, pine forests, and montane meadows
(Peck and Kaulbars 19S7). T. truiiaitus was
loimd onK in a late Ilolocene assemblage from
the (irand (lauNon.
The spider beetle (Ptinidae), Niptus ventric-
iiliis. is a scaxcnger that ranges from Texas west-
ward to C'alilornia and south through Mexico to
C»natemala. it probabK breeds in rodent nests.
Modern specimens lia\t' been collected from
packrat nests and from the fur of kangaroo rats,
Di))()(l()i>u/s spp. ( Brown 1939, Papp 1962). This
beetU^ speeic^s was common in sexeral assem-
blaties from GLCJA.
19921
Qr ATKKNARY AUTI IH()I'()i:)S. COLORADO Pl.ATKAU
65
Fig. 2. SciUining electron iiiicrographs of fossil beetles from sites discussed in text: A, liead capsule, prouotuni, and eKtra
of A<i(»min jH'rh'vis from the i^owns packrat midden, C^len C'an\ou; B, pronotuni of Tliaiuitophiliis tniiiciilits from Bida
(!a\(', (Jrand Can\on; (J. prouotum of Elcodes ui'^rina from Kaetan Ca\(', (Jraud Cau\on: D, exoskeletou of Aiuispis nifa
from Bida ('a\e, Craud Canyon; E. left eKtron of Ctjmatoclcni pallida from Hoodoo packrat midtlen. Caii\()nlands. Scale
l>ar e(|uals I nun.
The ilarkliiiij; beetle (Teiiebrionidae). Elcodes
ni^^riiui (Fig. 2C), was fountl in a late
Pleistoc-ene a.sseiiiblage (roni tlu^ (tL(>.\. Tliis
-scaxenger i,s known todax from tlie Pacilie
Northwest sontli t(j the nionntains oi Aiizona. It
is a eold-harcK species, foinicl at eknations iij) to
3050 HI in the Colorado Rockies (Blaisdell
1909).
The false darklin'j; beetle (M(^landi-\idaei,
Anaspis nija (Fig. 21)), is \\ides[)read toda\.
Beetles in this faniik are fonnd nnck-r bark, in
fun<j;i. and in decaxing logs (Liljeblad 1945).
The leal beetle iChrwsoinelidae), Lcma
Irilinca. feeds on Datura (jinison weed) antl
other [)Iants in the .southern hallOf the United
States. It was identified from a late Pleistocene
as.semblage in the GRCA. Other jilanl-feeding
beetles identified from the fossil assemblages
inclnde the weexils (Cnrcnlionidae) Sci/j)h<>-
plionis acnpiincfatits. Oninodcina pfoiracla.
Aplcnni.s (iii^^iddhs. and Clconidiiis triiattalus
orC. cjiiadriliiicattts. all Irom the ( irand (.'anxon
assemblage. Of the.se, O. protracfa was lound
onK in the late Ilolocene, A. au<^idaris and C
Irivilfaliis or (.'. (pi(idnli)icaliis were found onl\
in the late Pleistocene, and S. acu))Uiirfaius was
i(l(Mitified (rom both periods. O. protracta li\es
at elevations from 2250 to 2700 m in the moun-
tains of .\ri/.ona. It is a soil dwellcM- that feeds on
loots (K. S. Anderson. National .\Insenm ot
Natural Sei(Mices, Ottawa, written comimmica-
tion. |nl\ 1990). A. aii^idaiis. C. tiiviHaliis. and
C. (piadriliiicatiis are all widespread toda\
throughout western North America, while S.
(iciipttiiclatii.s has been collected from Arizona
and Mexico, where it feeds on A<i^ave, Dasijlihoii
isotol), and Lopliopfxom (pexote) (R. S. Ander-
son. National Museum of Natural Sciences,
Ottawa, written communication. July 1990).
FinalK. the tick (Ixodidae), Dcnnacentor
(indcrsoiii. is found todax in the western United
States as far east as Montana. Immature
66
GiiEAT Basin Naturalist
[Volume 52
D. ondersoni parasitize small mammals, while
the adult stage parasitizes large nuuumals. This
tick is a x'ector for Rock)' Mountain spotted Fever
and Colorado tick fever (|. Keirans, National
Institutes of Health, BetlK\sda, Maiyland, writ-
ten cominuuication, |uiie 1990).
Paleoenn'ihonmkntal
intehphetations
The infonnation from the fossil insect record of
the Colorado Plateau region, is not yet sufficiently
detailed to allow precise paleoenvironmental
reconstnictions. Ilowexer, for both the C^raiid
CcUiNon and CAvn CJanvon regions, the axailahle
in,sect data suggest a cooler, moister '•liniatic
regime during the late Pleistocene. Montane-
adapted species lived at lower elevations. The
in.sects document the presence of conifers at the
sites but also suggest that a mosaic of ve<ietation
t)pes was locally represented, including grtissland
and shnibln terrain. The shift to postglacial cli-
mates occurred sonietime after 14,()()()\TB.P.,and
the most ain( 1 c( )i iditions appeal" to have developed
within the last 15()() vears. Additional studies of
regional insect iissemblages will unck)ubtedl\clar-
if}- the nature and timing of environmental
changers.
Altliougli prcliiiiiuaiA and incomplete in
nature, the arthropod data presented here are
in agreement widi the detailed plant recon-
struction proxided b\ the macrobotanical
remains bom the packrat middens. C'ole (1990)
concludes that a compari.son of modern and
full-glacial ass(MnbIag(\s from th(> eastern Cl^C'A
packrat mickleus (kMuoustrat(\s tliat individual
plant taxaaiid comparable couiiiiiiiiities shifted
upward appro\imat(4v 800 m at the close of the
Wisconsin glacial (ca 11, 000 yr B.R). Cole
(1990) concludes that the climate at the eleva-
tions of Bida and Kaetan caves was nion^ conti-
nental during the late glacial. This result is in
contradiction to the equable climates that may
have occurred in western and low(M--ele\ ation
regions of the CRCA and to (he south of the
Colorado i^lateau (Mead and PhiJlip.s 1981,
VanDexender 1990). Our arthropod data pre-
.sented here do little to clarify the continental \ s.
equable climatic reconstruction contradiction.
Our "cooler, moister climatic regime" recon-
struction could be interpreted as a continental
climate; however, it couklalso represent a n^ginu'
with slightly cooler winters and cool sunnners.
and therefore more available moisture.
ACKNOWLEDCMENTS
The scarab beetle, Aphodius ruficlanis, was
identified by Robert Gordon, U.S. Department
of Agriculture and U.S. National Museum,
Washington, D.C. The weevils, Sct/pJioplwnis
aciipiiiwtatiis. Oriinodcnui protracta, Cleo-
nidiiis trivittatiis or C. cjuadrilineatus. and
Apleiinis aiifi^idaris. were identified bv Robert
Anderson, National Museum of Natural Sci-
ence, Ottawa. The tick, Dernuicenforandersoni,
was identified bv James Keirans, National Insti-
tutes of Health, Bethesda, Mankind. We appre-
ciate the help of Emilee Mead, Paul Martin,
Bob Euler, and Bill Peachy. Scanning electron
micrographs of insect fossils were taken with the
assistance of James Nishi and Paul Carrara, U.S.
Geological Sunev, Denver. Emilee Mead
drafted the figures. Financial support for this
studv was provided bv National Science Foun-
dation grants EAR 8708287 and 8845217 to
Mead and Agenbroad, and National Park Ser-
vice contract CX-12()0-4-A062 to Agenbroad.
Thanks are also extended to the staff at Ralph
M. Bilby Research Center, Northern Arizona
Universitx', for their support.
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19921
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Rvci'ivcd 20 jiiiw imi
Accepted 14 Idiiiian/ 1992
Great Basin Naturalist 52( 1 ). 1992. pp. (iS-74
MICROIIABITAT SELECTION BY THE JOHNNY DARTER,
ETHEOSTOMA NIGRUM RAFINESQUE, IN A \WOMING STREAM
Hohcrt A. Lcidy'
Absth.u:!". — .Vlicroliahitat sek'ction b\- the johniiN darter (Ethcostotim uignun) w'-dv, examined in die North Laramie Ri\er,
Platte (xnmtw WXoming. where it does not oeenr with odier darter speeies in die same stream reaeh. Eleetixity indices
based on microhahitat ohsenations iniheate diat K. iii<s,ni>n avoids riffles and selects certain mierohahitats characterized by
intermediate water depths in [lools and slow-m()\insi; nnis with a snbstrate composed piimaiilv ot silt and sand. Niche
lireadth and electi\it\ \alnes for total deptli. bottom water \(locit\. and snbstrate measnrements from this shidv indicate
tliat E. nif^niin is a habitat generahst. except at the extreme endsol the liabi tat gradient. Habitat use here is generajlv similar
to other studies where E. nii^niin occurred with one or more otiier darter species. This stnd\ found little e\idence for
competitive release in the absence ot other dartirs.
Ki'ij words: microli/ihitat use. I'crcidiic. tiichc hrcadth. coinpditirc release, electicities. inoqiliohxj^iedl sju-euilizafions,
Etlieostonia iiisinnH.
Tlir joliiiiiN darter c.xliiliit.s the lafgest geo-
graphic distnhution among the Noith Aineiicaii
darters (Etheostomatini: Percidae), with the
possible exception of Pcrchui capnxh's. It
occurs farther west than an\ other darter except
Ethcosfoiiia exile ( l^age 1 983). Tlie ecologv^ of E.
nigniDi has ix^ceived consideral)Ie study, often
in conjunction with other darter species (e.g.,
Winn 1958, Smart and Gee 1979, Paine et al.
1982, p:nglert and Seghers 1983, Mimdahl and
Ingersol]'l983, Martin 1984). Tiie aliiiit)- of E.
nigni)n to colonize such a large geographic area
may he explained in part I)v its tolerance of a
varietx' of emironmental conditions (Scott and
Grossman 1973, Trantman 1981, Becker 1983).
Throughout most of its range, E. /H'gn///i coex-
ists with one or more darter species in streams
(McCJormick and .Aspinwall 1983, Schlosserand
Toth 1984, Todd and Stewart 1985). E. iii<iniiii
is also conunonly found in lakes with weedx or
sand)' shorelines (Page 1983). (^ot^xisting dait-
ers txpicalK- show resource^ partitioning along
food and habitat ax(^s (Smart and (iee 1979,
Paine et al. 1982, Matthews et al. 1982, White
and Aspinwall 1984, Todd and Stewart 1985). In
addition to E. iu<iniiiL the low a darter (E. exile)
and tlie orangethroat darter {Etlieostonui
speetihile) occur \u the upper Platte Ki\(M-drain-
age of eastern Wyoming. Both E. iii<iniin and E.
exile occur in a tributaiA of [\\v North l^latte
Ri\er, the Laramie Ri\er, and se\eral of its trib-
utar\' streams, but ha\e not been recorded as
co-occurring there (Baxter and Simon 1970,
Page 1983).'
The ptu'pose of tliis paper is to examine the
microhahitat use of E. ni^nini at the western
extreme of its range where it does not coexist
with other darter species in the same reach of
stream. Two basic (juestions are addressed: (1)
Are the microhahitat recjuirements significantly
different for E. ni<inini in the stud\' stream
compared to other streams in North America
where it is found? (2) Does E. iii^ntm show
signs of competitive relea.se in the absence of
other darters?
Study Area
The North Laramie Riwr, Platte (>oimt)',
Wyoiuing, drains the central .Medicine Bow
Mountains and is a tributan t)f the Laramie
Rixer, which in turn joins the North Platte Rix'er
near the town of Wheatland. The stud\- was
confined to a lOO-m reach of ii\er approximately
10 km upstream from Interstate^ Highwa\ 25 (ele-
\ation 1420 m). .At this location the ri\er tra\erses
a broad floodplain a\eraging().75-1.0kiu in widtli.
Dominant oxenstoiA' ripaiian \egetation includes
Cottonwood (Pojniliis dehoides) and \arious tree
and shrub willows (SV/Z/.v spp.). The stucK area is
U.S. Kiiuronmenlal Pr(>tci.tii>ii .Ay.-iicv. WVllands S.clion (\\-7-2). 75 I hiwllionic Slnct, Sail Kiaiici.scu, Caliloinia 94105.
68
19921
ETHF.OSTOMAMCIH M H AKINKSOI K in a WYOMIXC STIUvWI
69
s])ai"S('l\ populated \\ itli lai"<i;c' rattle laiielies and
allalla (anus hordeiiug the lower to middle
icaelies. Hie most noticeable iwsult ol tliese
land-us(^ practices has been renio\al ol ri])ai"iaii
\ ('fetation and consequent associated sedimen-
tation; h()\\'e\"er, fencing has ellecti\el\'
exchiiled cattle from tlu^ Xoitli Laiamie Hi\er
alou'j; the stnd\ reacli.
T\\c stucK reach, chosen as representati\e of
the lower portions of the North Laramie Ri\er,
is gtMKMalK cliaracterized b\' large, relativeh'
uniform, shallow pools connected hv short rif-
lles and nms of xaning water \elocities. W'ettetl
stic^uu channel width within the study reach
a\ crages 6.5 m with a gradient of 4.7 ni/km. This
contrasts with gradients within the middle
reaches of the North Laramie Ri\er of 15.1
m/kni. Stream discharge at the stud\' site a\er-
ages 0. 1 7 nV Vs, although short-term fluctuations
in flow ma\' occur from summer thunderstorms
and irrigation dixersions. The substrate ranges
from a dominance of small graxel and sand, silt,
and detritus in pools to medium to large graxel
and cobble in riffles and runs. Diel water tem-
peratures in sunnner t\picall\ range from 13.5
to 21 C Minimum undeiwater \isibilitA in the
rixcr was 2.5 m or greater during the stud\.
liooted acjuatic vegetation within the stud\
reach includes waterweed {Elodcti rc///c/Jr//.s/.s),
perfoliate penmcress {TJiIaspi pci-folidtiiin),
and Ranunculus lonf^irostris.
MKTII()1:).S
Microhabitat obsenations of E. ni<inint wcvv
made 7-12 September 1988. Undisturbed fish
were located In a single obsener snork(^ling in
an upstream direction. Because of the high
water claritA', relati\el\- close spacing of indixid-
ual fish, and their obsened habit of remaining
ill direct contact with the substrate, marking the
location ol lish was not a [)roblem. Txpicallv the
locations ol 4-7 indixiduals w(M"e noted and
marked l)\ placing a wliite golf ball on the sub-
strate. This ap[)roacli allowed the siioikler to
ina\iiiii/e the nimiberol undisturbed indi\ idiial
observations and niiiiimize disturbance to
upstream fish.
For each indi\ idual obsen ation the lollow ing
microhabitat data were recordetl: ( 1 i total depth
of the wattM- column, (2) focal point elexation
(\ertical distance of the fish from the bottom),
(3) focal point \elocit\- (water velocit) at the
fish's snout), (4) mean water cohnnn xelocitv.
(5) surfac-e \elocit)-, (6) substrate composition,
and (7) co\ (M hpe. \ elocit\- measurements were
mad(^ w itli a mini flow meter (Scientific Instru-
ments, Inc., .Mock'l 1205). .Mean water column
\elocit\- was measured as the \-el()c itA at 0.6 of
the total depth when the total deptii was less
than 0.75 m, or the mean \elocities at 0.2 and
0.8 of the total (k^ptli wlu^n greater than 0.75 m
(Bo\ee and Milhouse 1978). Helati\e depth, a
measurement ol the location of the hsh in the
water colunm, was calculated b\ subtracting
focal-point (dexation from total deptli and divid-
ing by total (k^pth. All obsened indixidnals were
greater than 25 nnn standard length; howexer,
no effort was mack' to distinguish between ju\e-
nile and achilt fish.
Nine codes were used to characterize sub-
strate composition (percentage) in an area 0.15
m on a side measured from beneath each fish:
1. tines (sand and smaller); 2. small gra\el (4—25
mm); 3. medium graxel (>25-5() nun); 4, large
graxel (>5()-75 nnn); 5, small cobble (>75-150
mm); 6, medium cobble (> 150-225 mm): 7.
large cobble (>225-300 mm); 8. small boulder
0300-900 mm); and 9. large boulder/bedrock
(>9()() nnn). A cover rating (0-2) as measured
b\ the relatixe degree of protection offish from
stream \ elocit\', \isual isolation, and light reduc-
tion (i.e.. shading) was assigned to each obser-
vation. A rating of denoted no protection; 1.
moderate protectic^n; and 2, major protection.
The general ty|3e and location of co\ cr in rela-
tion to fish also wcm'c noted.
Habitat a\"ailal)ilit\ was ck'terniined randoiiiK
each dav innnecliat(d\ following the collection
of microliabitat-u.se data (Mcnie and Baltz
1985). The lollowingavailabilitN' measurements
were made along 10 ranck)ml\' selected tran-
sects within the stuck reach: total depth;
bottom, mean w ater cohnnn. and snriace \eloc-
ities; substrate compcxsition: and co\er t\pe.
Between 15 and 30 ecjualK' .spaced measure-
ments were made along each transect. To ade-
(|uatel\- characterize habitat a\ailal)ilit)- within
tlie c()iiiparati\cl\ short stucK" reach, an effort
was made to collect a[)[)r()>dmately t\\ice as
iiiaii\ measurements of habitat axailabilitA' as
microhabitat obseivations.
.\n electi\it\ index was used to determine
selectiv it\ In E. ni<irunt for total depth, bottom
water \c'l()citA, and substrate composition. Elec-
ti\ities were calculated from the fonnula
D=r-p/(r+p)-2ip, where r is the proportion of
the resource used and p is the propoition axiiilable
70
(;hkat Basin Naturalist
[N
olunic oz
0.5
â– Habitat Use
n Habitat Availability
^j3^
0-10 >10-20 >20-30 >30-40 >40-50 >50-60 >60-70 >70-80
/^ Total Depth (cm)
Fig. 1 A. Hclatiw t'recjucncv distributions of microhahitat nsv ami a\;ulal)ilit\- for total water roliiimi tk^pths lor E ni'^niin
in the Xortii Laramie River. Eleeti\ities are indicated ++ (>().5(). strong preference), + (>0.25 lint <().5(). moderate
preference). {) ( +0.25. no preference), - (>-0.()5 hut < -0.25,. moderate a\-oidance), and = (<-0.()5, strong avoidance).
u
c
0)
3
a>
B
0.8 -
•i 0.2-
â– Habitat Use
m Habitat Availability
.^
i/ ^
i^
0-5 >5-10 >10-1S
Bottom Water Velocity (cm/sec)
>15-20
Fig. IB. Relative frequency distrihntions of microhahitat use and a\ailal)ilit\ for bottom water velocities for K.
tlie Nortli I^iramie Rixcr. Klectivities are indicated ++ (>0.5(), strong pn-ierencel. + (>0.25 hut <().50.
preference), ( +0.25. no |)reference), - (> -0.05 but <-- 0.25, moderate avoidancii. and = (<-(). 05, strong aM
in;^nini in
motlerate
in the .stream eiiNiroiuuent. Tlii.s iiide.x i.s based test for goodness of fit wa.s applied to freqnencv
on the fonnula by Jacobs (1974), as modified b\- di.stributions lor habitat use and a\ailabilit\ to
Moxle and Bait/. (1985) for detc>rminino; determine whether ma.ximnm differences
niicrohabitat .selectivity- from variables .similar to between the obsent-d and expected distribn-
thosensedin thisstndv .A KolmotioroN-.Smirnov tions were simiiheant (Sokal and !\ohll' 1981).
19921
ErUEOSTOM.WlClUM HAI'IM'.SgUE IN A WVOMINC; STIUvWI
71
0.8 n
â– Habitat Use
m Habitat Availability
( (
.
r
y
'
/ /
}
C Substrate Codes
P'ig. KJ. Ht'lathc frecjueiicA clistrihiitioiis of niicrohahitat use ami a\ailal)ilit\ loi' substrate codes for ¥,. iii^niin in tlie
Noitli Laramie Ri\er. ElectKities are indicated ++ {>0.50, strong preference), + (>0.25 but <0.5(), moderate preference),
(+0.25, no preference), - (>-0.05bnt <-().25, moderate avoidtmce), and = (<-0.05, strong avoichuice).
An additional measure of microliahitat utiliza-
tion, niche breadth, wa.s cakulated for E.
ni<i;runi. Two niea,sures of niclu^ breadth were
calculated to adequately characterize the effect
that the selectixitv of rare and common
resources might have on niche-breadth \alues.
Hurlbert's measure of niche breadth ( B' ), which
is sensitive to the selection of rare resources, was
calculated as follows: B' = l/S(pj""j/aj). Smith's
measure of niche breadth (FT), which is less
sensitive to the selecti\it\ of rare resources, was
calculatcnl as follows:
FT = 2( Vpjaj)
where pj ecjuals the projiortion of indixiduals
found in resource /(ipj= 1.0), and a| is the pro-
portion of total axailable resources cousisliiiij; ol
resource 7(Xaj= 1.0) (Krebs 1989). B' \ahies
were standardized to a scale ofO-l. using the
efjuatiou B'.\ = B' -amin/l -ainm. where B'
ecjuals liulbert's niche breadth, and amm e(juals
the smallest obsened proportion of all
resources (minimum aj). The larger the B' and
FT values, the less individuals discriminate
between resoinx-e states (mininumi specializa-
tion); the smaller the B' and FT \alues, the
greater the resource discrimination (iiiaximuni
specialization).
Results
Eight species of fish were obserxed with E.
nignnu at the stud\' site. These were sand shiner
{Hybo(^iuithii.s lumkiii.so)ii), suckemiouth minnow
(Phenacohiiis iiiirdhilis). creek chub (Scniofihis
atromocuhiius). common sliiner (Notropi.s cor-
niitiis), red shiner (A^. Itifrciisi.s), bigmouth .shiner
{N. clorsali.s), white sucker {Catostomus coinmcr-
â– soni), and rainbow trout {OiicorJii/iicluts nujkiss).
Microliabilat ObseiAations and
Habitat A\ailabilif\
Microhabitat-use data indicated that E.
iii^ntm alwavs occurred in continuous contact
w ith the substrate where water \elocities were
low (Table IV Eflico.sfoina /i/gn///i was almost
(â– \clnsi\cl\ found ()\(>r a substrate of sand or
small graxt'I, usualK in pools and slow-mo\ing
nms of intermediate de])th (Table 1. F'igs. lA-C).
In contrast, surface xclocitic^s often were rela-
ti\el\- high.
In tills stiuK. obsen ations Indlcatetl that indi-
vidual fish wt>re positioned ( 1 ) on the surface of
the exposed substrate with no apparent co\er,
(2) immediateK below the front edge of a slight
depression in the sand that .sened to protect fish
from the current, or (3) rarely on the dowii-
streani slope of a small cobble also protected
from the current. In all cases, E. nignim
(;he.'\t Basin Naturalist
[Volume 52
T.Mii.K 1. Means (± S.D.) from iiiicroliahitat use and
aviiilahilitv measurements lor E. nifinini in tlie North Lara-
mie Ri\er, Wyoming.
Habitat use
Habitat
\'arial)le
obsenations
availability
Total depth (cm)
40.5 i 8.S
27.1 = 16,8
Focal point e\alnati(jn (cm)
0.1 i 0.01
—
Relative depth (cm)
0.9 ± 0.02
—
Mean water cohunn velocity
(cm/s)
2.6 ± 4.5
3.7 ± 6.4
Focal point/ix)ttom \elocit\
(cm/s)
0.2 ± 0.7
1.8 ± 3.1
Surface velocit) (cm/s)
5.2 ± 7.3
5.4 ± 8.2
Substrate t\pes (%)
(1) fines
62.1 T 35.8
34.1 ± 36.3
(2) sniiill gra\-el
16.5 i 19.6
21.6 ± 25.6
(3) medium gravel
7.6 ± 14.7
6.4 ± 13.3
(4) large gra\el
4.7 ± 13.5
5.8 ± 14.9
(5) small cohhle
6.3 ± 15.7
9.7 ± 21.5
(6) medium cot)l)Ie
2.1 ± 11.3
15.5 ± 28.7
(7) large cohhle
0.7 ± 0.20
6.5 ± 21.5
(8) small houlder
—
—
(9) large houlder
—
—
Cover code (()-2)
Stream \elocit\-
] .5 ± 0.6
—
\'isu;il isolation
0.5 ± 0.6
—
Light reduction
0.1 - 0.3
—
Sample size
9!
1(>S
'HfrerloMelliods
T.\BI,k2. Niclic breadth values (/^',\aiid FTi lor E. iiip-iiin
for total depth, bottom water velocitv. and substrate in the
Nc)rth Laramie River, W'voming (approximate 95% conli-
dcTice interval shown in parentheses).
Bottom
Total (l(
â– j)th
velocitv
Substrate
Hurlbert's B' \
,45(,n.
,49)
,76 (,72, ,80)
,70 (,66, ,74)
Sniilh's /• r
.72 1,65,
,7S
,89 1,84, ,93)
,9.) (,89, ,96)
positioned itself in close proxiniit)' with other
t\pes of instream cover (e.g., stones, cobbles,
branches, or small depressions in the sand). The
average distance to such cover was less than 6
cm for 89% of the observations.
Measurements of microhabital a\ailal)ilit\
indicatc^d that average water depths a\ ailable to
E. ni<:^nini \v(M-e shallo\\'(>r than the depths at
which it was topically observed (Kohnogorov-
Smirnov te.st, .23, p < .01), and available mean
bottom water velocities were greater than
where fish were ol)seived(K-S t(\st, .25,/; < .01;
Figs. ] A, B). In addition, available sul)strate was
dominated by fines and small gravel (55%), but
this was disproportionatelv low when compared
with microhabitat use obsenations for these
same substrate t\pes (79%; K-S test, .28,
/; < .01; Fig. IC). '
Habitat Selection and Niche Breadth
Electivitv indices indicate that E. nigrum was
selecting certain microhabitats while avoiding
others. E. nigami selected intermediate water
depths and avoided high mean water column
velocities (Figs. lA, B). There w^as a strong
selectivity for a substrate composed of sand, and
an avoidance of medium to large cobbles (Fig.
IC). Fish generally avoided areas that ( 1 ) exhib-
ited high surface water velocities, (2) were iso-
lated visually, or (3) were well shaded by
physical cover (Table 1). Rather, fish utilized
relatively barren substrates exposed to full sun-
light but close to cover. Microhabitat niche
breadths (6'.\ and FT values) for depth, v elocit\',
and substrate indicate little resource specializa-
tion b)- E. nignnu (Table 2).
Discussion
The results of the electivitv indices and the
K-S test indicate that E. iu<iruin is highly selec-
tive in the microhabitats it occupies. Hovv^ever,
niche breadth values suggest that E. ni^iniin
does not discriminate between available
microhabitats (i.e., minimal habitat specializa-
tion). Tlie apparent inconsistencv between
niche-breadth values and electivitv indices may
be explained bv two factors: (1) the relative
scarcitv in the studv area of gravel/cobble riffle
habitats and their avoidance bv darters, and (2)
the preference bv darters for lovv-velocitv pool
habitats characterized bv sand and small gravel,
a habitat that was abundant in the studv area.
Values for Hurlberts measure of iiiche brc\idth
(B',\) were consistently lower than values ior
Smiths measure (FT) for depth, velocitv, and
substrate. This is expected because B' a is sen.si-
tivc to the selection of rare resoiu'ces that are
more lieavilv weighted in the calculation of
niche breadth, while FT is less sensitive to the
selection of rare resources (Krebs 1989).
nart(M' species tvpicallv are restricted to a
narrow range of microliabitats. This is especiallv
evident in their use of certain substrates (Page
1983). E. ni<^niin has an imusuallv broad toler-
ance among darters lor variable env iionmental
conditions and has been obseiAcnl over widely
vaning vcloc-ities, de[)ths. and substrates
between drainages and within a [)articular
stream reach (Smart and Gee 1979, Angenneier
19921
ErHF.OST()\f.\ Mcni M H \i-|\i:soi'K i\ \ W'vomixc Sthf.am
rs
I9S7). This stiulvand others (e.g., Becker 1959.
I'aiiic ct al. 19S2, Englert and Seghers 19S3)
geiieralK show that E. nig^nim occurs most hc-
(jiieiitlx in pools and sluggish reaches ol stream
oNcr sand or silt substrates, although this darter
also regulark occurs in riffles (Lachner et al.
1950. Smart and Gee 1979. Trautman 1981). In
other streams, pool and riffle habitats are often
coinhahiled l)\ one or more daiter species. II
competition with other darter sp(X'i(^s restricts
E. )ii<j^niin to microliahitat t\])es in which the\
arc conunonK' foiuid, then in the absence of
other daiter .species one might expect E. nipiiin
to experience competiti\e release. Efheostonui
iiii^niiit wlien alone should occupy a wider rang(^
ol habitat in a particular stream reach, without
as much specialization for a particular range or
resource t\pe. Obseixed [)atterus of
iiiicrohabitat use from this stud\ found little
c\ idencc^ of conipetiti\e release, suggesting that
other darters are probabK- not restricting
/'". iii<inini to a particular habitat txp(^ in streams
where the\ coexist.
Electi\it\ and niche-breadth \ alues lordepth.
\elocitx, and substrate measurements from this
stud\' sup])ort the conclusion of Coon (19(S2)
and Others (Winn 1958, Karr 1963) diat E.
iti<^riun is a habitiit generalist, except at the
extreme ends of the habitat gradient (i.e.. shal-
low cobble riffle and \en shallow pool liabitats).
Howcxer, in contrast to tlie studies of (^oon
( 1 982 ) and Smart and Gve ( 1 979 ), that rec< mlcd
I'., iiiiiniin in riffle and run/pool habitats with
one or mon^ darter .species, in this stud\ E.
iiiilfiniL w liile it was connnon in pools, did not
occur in riffles e\(^n in the absence of otiier
darters.
Schlos.ser andToth (1984) suggested that dif-
lerences in niicroliabitat use in two sxinpatric
darters ap[)ear to be constrained b\ mor])h()l()g-
ical s])eciali/,ations ol eacli .species rather than
by interspecific competition. As with most small
darters, E. ni^nini is characteri/cnl In morpho-
logical sj:)eciali/ations best suited to the beuthic
stratum of pools and othei' sluggisli stream hab-
itats, often with a sand or silt substrate ( I'age
1 983, Page and Swofford 1984). Support lor the
role of moipliologx in drixing habitat utilization
\i\ E. iii^niin in the stucK area conies from data
on co\-er utilization. Protection Ironi stream
M'locities in the absence of am a[)pareut i)h\si-
cal instream co\er ma\- be explained In this
species' small size and benthic habits. X'elocities
immediatek- abo\e the substrate wlu-re fish
w(>re obseiAcd were negligible when compared
t()\(4(R ities at the same location a few centime-
ters higher in the water column or at the surface.
.Mso, subtle (Kpressions in the sand sub.strate
olteii were occupied In indi\idual fish presum-
ably for protection from stream \elocit\. One
might expect that the small size and ob.sened
patterns of habitat utilization b\ E. iu<iniin
would increa.se its risks to predation. llcmcxer,
small size, drab coloration, speckling, \\'-marks,
and partial traiisluceiice, combined with expo-
sure to full sunlight, made detection of indi\id-
iial fish on the speckled sand substrate difliciilt.
The increased risks of exposure to predation
from small size alone would appear to be com-
pensated l)\ the combination of \arious mor-
phological features. The same moiphological
features tliat act as camouflage in (|iiiet pools
likeK ina\ not senc the same function in rillle
habitats (Page and Swofford 1984).
A(:K\(.)\\\.KDC,\[ESTS
1 am especialK indebtetl to Barbara l-^iedler
and Rand Fanclier for assistance in the field,
and to the owners of the IIR Ranch for gener-
ously proNiding access to the stud\ site. 1 am
sincereK' grateful to P(»ter B. Mo\le, Pegg\ Lee
Fiedler, and two anoiix iiioiis nniewers for crit-
ical comments (ju the manuscript. Thanks also
to George R. IxmcK- of BKAK Gonsnltants. Sac-
ramento, ( 'alilornia. tor lending the flow meter
Liti;h ATUHi". GrrKD
Ax(a;i;\n;iKH V. I.. 19S7. .Spatiotcinponil xariation in luil)-
itat .si'icctioii In lislics in small Illinois stiranis. lit: W. j.
Matthews and 1). (,'. ileins. eds.. ('()nninniit\ and
('\()lnti()nar\ ccolog) of North American stream fishes.
Uni\ersit\ <)( Oklidioma Press. Norman.
lUxii'.H (;. T, and |. R. SiMOX 1970. WVominsj fishes.
\\\()min<^(;aineand Fish Department. (>he\enne. IfiS
pp.
I5i;< kii; (;. (,'. 19.59. Distribution ol central W'iseonsin
fishes. Wisconsin Acadenn ol Science, .Arts, and Let-
ters 4S: 6.5-102.
. 19S.3. Fishes olW'isconsin. Uni\ersit\ ol Wis-
consin Press. Maiiison.
HoM.i; K, D.. and H. T. .Mll.lioi SK 197S. Hydranlic simu-
lation in instream How studies: theor\ and techni(|ne.
U.S. Fish and Wildlile .Seivice Biolosjical .Serxitvs Pro-
gram FWS/()ISS-7.S/:5;3.
(;()()X T. (;. 19S2. Coexistence in a "jnild oflK-nthic stream
fishes: the effects of'tiistnrhance. Unpublished doctoral
dissertation. University of C;alilbniia. Da\is. 191 pp.
FxcLKirr J..aud B. II. Si:(aiRi{S. 198.3. Habitat segregation
1)\ stream darters (Pisces: Percidae) in the Thames
River watershed ol southwestern Ontario. (Canadian
Field Naturalist 97: 1 77-180.
74
Ghi:at Basin Naturalist
[\ blume 52
Jac:obs, J. 1974. Quantitative meiusurenient of food selec-
tion: a niodifkation of the forage ratio and Ivlev's
eleeti\itv index. Oeeologia 14: 413—417.
K.\HH. J. R. 1963. .\ge. growth, and food hahit.s ol johnny,
slenderhead. and l)Iack.si(le darters oi Boone (lounts,
Iowa. Proceedings of the Iowa AcadcniN <>( Science 70:
228-236.
KkkBS. C. J. 1989. Ecological melhodolog). Ilatpcr and
Row, Publishers, New York. 6.54 pp.
L\(:il\KH, E. A., E. F. Westlake, and R S. Handwerk. 1950.
Studies on the I)iolog\ of some percid fishes from
western PennsxKania. .\inerican Nlidland Naturalist
43:92-111.
M.MrriN. D. J. 1984. Diets of four sympatric species of
Etheostoma (Pi.sces: Percidae) from southern Indituia:
interspecific and intraspecific nniltiple comparisons.
Environmental Biolog\- of Fi.shes 11: 11.3-120.
M.ATTllFWS, W I., J. R. Bkk, and E. SUR.vr 1982. Compar-
ative ecology- of the darters Etheostoma poclosteinoiic,
E. flahcllarc and Pcrcina nmnoka in the upper Roanoke
f\i\er drainage, N'irginia. (Jopeia 4: 80.5-814.
McCoKMKk K II., and N. A.si'in\\'all 1983. Habitat
selection in three species of darters. Environmental
Biologv of Fishes 8: 279-282.
MoYi.K, R B., and D. M. B.altz 1985. Microhabitat use bv
an assemblage of California stream fishes: developing
criteria for instream flow determinations. Transactions
of the Aiiiencan Fisheries Societv 114: 69.5—704.
.\Ii \i)\iii. \. D.. and C. G. iNGF.HSOi.L. 1983. Earlv
autumn movements iuid densities of johnnv
(Etiu'ostoiiui lu^ntin) and fantail (E. flahcllarc) tlarters
in a southwestern Ohio stream, [onnial of Science 8.'3:
10.3^1 OS.
Pack L. M. 1983. The handbook of darters. T F II.
Publications, Neptune City, New Jersey. 271 pp.
Pack E. M., and D. L. Swokfohd 1984. Morphological
correlates of ecological specialization in darters. Envi-
romnental Bif)log\()f Fishes 11: 1.39-1.59.
Pain'k .\I. D.. |. j. DousoN. iuid C. Power. 1982. Habitat
and food resource partitioning among four species of
tlarters (Percidae: Ethco.stoimi) in a .southern Ontario
stream. Canadian Journal of Zoolog)' 60: 163.5-1641.
Sciii.ossKH I. J., and L. A. ToTll 1984. Niche relationships
ami population ecologv of rainbow {Etheostoiria
cacntlcnm) and fantail (E.flabcllare) diuters in a tem-
poralK variable environment. Oikos 42: 229-2.38.
SctriT. \V. B., luid E. J. Cr()S,sman 197.3. Freshwater fishes
of Canada. Bulletin of the Fisheries Research Board of
Canada 1984.966 pp.
Smart H. J.,andJ. H.Cek 1979. Coexistence and resource
partitioning in two species of darters (Percidae),
EthcostoDw nigrum and Pcrcina maculata. Canadian
Journal of Zoology .57: 2061-2071.
SoKAL, R. R., and F.' J. Roiilf 1981. Biometiy W". H.
Freemiui, San Francisco.
Todd, S. C, iuid K. W. Stewart 1985. Food habits and
diet;uA overlap of nongame insectiv orous fishes in Flint
Creek, Okkdioma, a western Oziu'k foothills stream.
Great Basin Naturalist 45: 721-733.
Traitman, M. B. 1981. The fishes of Ohio. Rev ed. Ohio
State University Press, Columbus. 782 pp.
White. M. M.,andN. Aspinwall. 1984. Habitat partition-
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Received 1 October 1990
Revised 1 May 1991
Accepted 1 October 1991
Creat Basin Natmalist 52( 1 ), 1992, pp. 75-77
NOMENCLATURAL INNOVATIONS IN INTERMOUNTMX llOSIDAE
Arthur Croiuiuist
1,2
\hs IH \c:'l'.-New ta\a include Ijniiuliuin juiikurdidc (j'oikj. (Apiat-cai'). Crotoit tcxciisls (Klotzscli ' Mucll. Ar". \ar
utiilicitsis (joncj. I KupliorbiafiMf' Other noinenclatnral innox ations inelnde: Cyntoptcnts longipcs v;ir. ibapensis (M. E.
Jones) (aonij.. I.Diiiatiinn nisraniini (.'i()n(j. (Apiaceae); ('(iiiiissoiiia hootltii (Douglas) Haven vm: dccorticans (Hook. &
Am.) Croncj., C/iinis.snniti hootltii (Douglas) Ra\en \m: (Iciri-tonnit iMunz) Croiiq., Caini.ssonid chivaefonni.s (Torr. &
Frem.) Raven \ar. aurantiaca (Munz) Cronq., Cdinissoiiia cliiKicfoniiis (Ton: & Freni.) Ha\en \ar cnicifoniii.s (Kellogg)
Cronq., Cami.s.soitia chivaefonni.s (Torr. & Frem.) Ra\cn \ar fniicrcd i Raven" (joiki . ('ainissonid clavaeformis (Torr &
P^rem.) Raven var lancifolia (A. A. Heller) Cronq., Ctiinissonid lictcrocliroiiKi \S. WatsJ Raxcn \ar inoiioeiisus (Munz)
(.'ronij., CamLssonia kcnicnsis (Munz) Ra\en viu. gilmanii (Munz) Croncj.. C.(i]iiissoiii(i sciqioidfn (Torr & Cray) Raven \'ar
macrocai-jui (Rawn) Cronq., Oenothera Inennis L. var strigfisa (Rvdl). ) Cronq., Oeiiolheid pallida I.indi. \ar nnieinata
(Engelm.) Cron(]. (Onagraeeae).
Kci/ irords: nciiicnclatnrc. Rosida(\ taxoiiouui.
M\ iiianiisc'ri[)t on a nunihcr ot tamilifs ol
Hosidae for Iiitermountain P'lora has been com-
pleted and awtiiting pul)lieation for .sexeral
\ (nirs. These famihes should constitute a large
part of \olunie 3A (Rosidae except Fabales).
Since I cannot now anticipate when \olunie 3A
\\ ill be published, the followino; nonienclatural
inno\ations are liere \alidated.
Apiaceae
Ctjmopteriis longipes S. Wats. var. ibapen-
siH (M. E. Jones) Cronq., conil). nov. [based
on: Cijmoptcnis ihapci}sis \l. E. Jones, Zoe 3:
302. 1893].
Lotruitium packardiae Cronq., sp. now
(Fig. 1). Ilerba ptM'ennia caespitosa radice
crasse et caudice nianifeste ranioso, omnino
sulnelutina, foliis omnibus Ixisalibus. teniato
(\el quinato)-pinnatifidaet dcuuo plus-niinus\e
pinnatifidis, .segmentis ultimis augustis, 1-2 nun
latis. iiiiparibus, eis majoribus 1-3 cm longis;
scapi maturi 1.5-4 dm alta, umbella ])rr
anthesin compacta, pana, ca 2 cm lata, ladiis
imparibus, demum aperta radiis longioribus 4-fi
cm longis, bracteis inxolucelli panels, lineari-
attenuatis \el nullis; flores flaxi, lobis caKcis
minutis \('l obsoletis; pedicelli fructiferi 3-7
nun longi: nuMicaipia glabra \el interdum
patenti-hirtella, S-9 X ,'3-3.5 nmi. maiiilcste
alata, alis uscjue ad 1 mm latis.
HOLXrrvrE. — Packard 74-46. in ash (hat has
not disintegrated into clax. along Old Succor
Creek Rcjad, near Sheaxille, \ev\- close to the
Idaho border, T27S, H46K, Malheur Co.,
Oregon, 19 Ma\ 1974; NV! I.sot\pe at ClC
Habitat and distrihutiox. — bi volcanic
ash and rhyolite on rock\ cla\' soil in the sage-
brush zone. Malheur and Lake cos.. Oregon, S
to \\'ashoe and Humboldt cos., Nexada. Flow-
ering from April to )un(>.
COMMENTAR')'. — Lo null ill m packardiae has
.sometimes passed in the herbarium as L.
tritcniattiiii (Pursch) Coulter & H().s(\ which
howcNcr has solitan or few stems or .scapes on
tlie sinij)l(' or occasionalK' few-l)ranched crown
or short caudex atop the taproot. The ultimate
segments of the leaxes of/,, packardiae are also
shorter than is tvpical lor L. triteniaiiim. the
larger ones ouK 1-3 cm long, so that the lea\es
haxc a dillercnt aspect.
Lomatium roHeanum Cronq., noni. nox.
Lepiotaenia leiher^ii (>()ulter 6c Hose, Contrib.
U.S. Natl. Herb. 7: 202. 1900. Not Lomatium
liihen'ii(.\m\[vybc Ho.se, 1900.
,The New York Botanical Clarde
"Deceiised March 22. 1992.
Bronx, New York 1(M.5S-.5126.
76
Ghka'i" Basin Naturalist
[Volume 52
Fig. ]. I .ouKil'nnn juickind'u,
Euimi{)HI5iakc:eae
Croton texensis (Klotzsch) Muell. Arg.
var. utahensis Cronq., \ar. lun-. A var. texeiisis
loliis supra glahris diffcit.
HOLOTVPK. — Cwntjuist 6 K. Thonic 11839.
sand dunes ca 1<S km airline N of L\nnd\l, [uab
Co., Utah, T13S, R5W, ca 1500 m ele\.,'2.s"jul\
1983, at NY! Isot>pes at BRY!, UTC:!
Co\IMl-:\TAKV.— Crofo/j tcxciisis is \ariahle
in densit\()t ])ul)escence, hut tlir()u>i;houl most
of its ran^e the upper surface ol the lea\es has
at least a few stellate hairs (though these- ma\
eventnalK- fall off). An ahuudant population on
the sand dunes nc^u- lAnnd\l in |ual) and Mil-
lard COS., Utah, n-pre.sents the least pubescent
extreme. In these plants the upp(>r surface of th(>
Iea\es is wliolly glabrous or proxided willi ouK
a lew (|uickly (k'ciduous stellate scales. The
L\nindyl plants and some .similar ones from
Kane and San Juan cos., Utah, and from northern
Coconino Co. in Arizona, are here considered
to form the \ ar. titahciisis Cronq. The othen\i.se
fairly widespread var. texensis, with the upper
surface of the leaves evidently (and more or less
persistentlv) stellate-hain', is largely allopatric
with \'ar. ufdhcnsis, bareK' entering Utah in San
Juan Co.
Ona(;raceae
Camissonia boothii (Douglas) Raven var.
decorticans (Hook. & Ai-n.) Cronq., comb.
no\. [based on: Gaurd dccoi'ticans Hook. &Arn.
Bot. Beechevs Vo\age343. 1S39].
CamisHonia boothii (Douglas) Raven var.
desertorum (Munz) Cronq., stat. nox. [based
on: Oenothera dccoiiicans \ar. (h'sciit)niin
Munz, Bot. Gaz. 85: 246. 192S|.
Camissonia clavaeformis (Toit. & Frem.)
Raven var. aurantiaca (Munz) Cronq., stat.
no\-. [basetl on: Ocnothcni scdpoidca \ar.
aunintiaca S. Wats. Proc. Amer. Acad. Arts 8:
595, 613. 1873; an illegitimate name which as
defined by Watson included the t\pe of the
earlier O. scapoidea xar. clavaeformis S. W^its.
1871. Oeiiotliera clavaeformis \'ar. aurantiaca
Munz, Amer. J. Bot. 15:237. 1928].
CflmissomV/ clavaeformis (Ton*. & Frem.)
Raven var. crucifonnis (Kellogg) Cronq.,
stat. nov. [based on: Oenothera cniciformis Kel-
logg, Proc. Calif. Acad. Sci. 2: 227. 1863].
Camissonia clavaeformis (Torr. & Frem.)
Raven var. fmierea (Raven) Cronq., stat.
no\. [based on: Oenothera clavaejormis subsp.
fu)H'rea flaxen. Uni\. Calif Pub." Bot. 34: 106.
1962].
Camissonia clavaeformis (Toit. & Frem.)
Raven var. lancifolia (A. A. Heller) Cronq.,
stat. nov. [ba.sed on: Clu/lismia lancifolia \. A.
Heller. Muhlenbergia 2:"226. 1906].'
Camissonia heterochroma (S. Wats.)
Raven var. monoensis (Munz) Cronq., stat.
now [based on: Oenotlwra heterochroma \ar.
)iionoeiisis Mnn/, Aliso 2: 84. 1949].
Ckimissonia kernensis (Munz) Raven var.
^ilmanii (Munz) Cronq., stat. now [based on:
Oenodicra dentata \ar. <j^ilmanii .Munz, l^eatl.
W. Bot. 2: 87. 1938|.
Camissonia scapoidea (Torr. & Gray)
Raven v ar. macrocarpa (Rav en) Cronq., stat
noN. Iba.sed on: Oenothera scapoidea subsp.
macrocarpa iiaxcn, Uni\. Calif. I^nb. Bot. 34:
95. 19621.
1992 NOME\(:i..\TllHAI. I\\()\\TI()\SI\ HOSIDAE 77
Oenothera biennis L. var. strigosa (Rydb.) ACKNOW i,i;i)(;mi:\ts
Cronq., coinl). iua'. [based on: Ociiothci'd
.slri<^o\(i H\(ll). Mem. \. V. I^ot. (iard. I: 27S, The work here reported was suhsidi/ed oxer
19()()|. a period of years In sueeessixe grants from tlie
Oenothera pallida Lincll. \ar. runcinata National Seienee Fonndation to tlie New York
(Engelni.) Cronq., stat. now |l)ased on: Botanical (lank-n in snpport oltluMnternionn-
Ociiothcrd (lU)ic(ndis wir. niucinata Engclni. tain l^dora project. The drawing ol LoiiuiHidii
Anicr. J. Sci. Arts 84: 334. 1.S621. jxickardiac was done b\- Bobhi Angell.
Received 30 Aii'^tisi 199]
Accepted 26 November 1991
Great Basin Nat malist 52(1). 1992, pji.TS-S.S
NOMENCLATURAL CHANGES AND NEW SPECIES IN PLATYPODIDAE
AND SC:OL\TIDAE (COLEOPTERA), PART II
Stephen L. Wood
Ai?sthac;t. — In PlatNpoclidac the new name Gcni/occni.s stroliincycri replaced the jnnior homonvni G dlhipennis
Strolunever, 1942, luid the new nanii- Pliili/pii.s apphinatulus replaced the junior homonvin Platypus applanatus .Schedl,
1976. New names are presented in Scolvtidae as replacements for junior homonyms as follows: Cn/pluihi.s hmicnei for
CnjpJialus ai-t(>caif)iis Schedl. 195S; Ci/clorhipidion diJiinisiniin kn Xtjlchorm diJungensis Schedl, 1951; HijpotJicnemus
(itcrriimilus for Lcpiccroi/lcs (now Hi/pothcucniu.s) (itcrhiuiis Schedl, 1957; Hypofliciiciiiit.s khinliitskiiyac for
Hypotluncinus iiisnlnri'^ Kn\()lutska\a: Piti/ophthoni.s nfricdiiiilits {'or NaHlnjococics (now Pityoplithonis) (ifricaiiiis Schedl,
1962; ScohjtogeiKs /)(//)(/(//,s;,s for \iil()cn/i)tii\ (now ScDlijto^enes) papiKinus Sclicnll, 1975; Scolytogcncs panuloxiis for
Scolyt()<:,cii('s paptiauiis SL\\ri]\. \'>n't>:\iililHiniui\\pi)iipi>slinis (or Eidopliclus (now Xylel)(»iiiti.s) spiuipciinis Schedl, 1979;
Xi/lebonis fonno.sac for Xi/lchonis foniuisdtiiis Browne, 19.S1. New combinations for fossil Scolvtidae include Dnjocoetes
diliaidlis for Pifi/oplitlwmidcd diliniiilis Wickliam, 1916. and Hi/lcsiniis liydropicus for Apidnccp1i(dus hydmpictis
W'ickham, 1916, Phlocotiihtis ziiiniuTintmui Wickliam, 1916. is transferred to the famiK C'nrculionidae. In Scolvtidae,
Crypludiipliilu.\ Schedl. 1970. is a junior generic sviionvm oi Sail ijt a ^c lies Eichhoff; Mdcrocn/phidiis Nohuchi. 19S1. is a
junior generic s\non\ni o( tli/pnthciicimis Westwood, 1836; Ni})poiiopolt/<^raphiis Nohuchi, 19S1, is a junior generic
s\nonvm o'i Pohi'^niphiis Erichson, 1S36; Pseiidocosinodercs Nobuchi, 1981, is a junior generic .svnonym of Cosiiiodere.s
Eichhoff, 1878; 'I'dpiiwcocfcs Pfeffer, 1987, is a junior generic synonym of Tc//;/(/v)/-)/r/i!/.s- Eichhoff; Tnjpdnophellofi Bright,
1 982, is a jiinior generic synomvm of Lipdiilirnin Wollaston. New .specific .sviionymv in Scolvtidae includes: BrdcJiyspaiius
moiitzi Ferrari (=C()i-tlii/liis ohtnsiis Schedl), Cdrpliolionis iniiiiiims (Fabricius) (=Cai'i>lwhonis hdlj^ciisis .Mnrayama),
Cocc()tn/))('s dddiilipcrdd (Fabricius) (=Cocc(>fn/])cs tnipiciis Eichhoff), Cn/pludits sctdiricollis Eichhoff (=Cn/plidlus
hrevicollis Schedl), Ficicis dcspccts (Walker) (-Hi/lr\iiiii.s stiinodiuis Schedl), Hijld.stcs pluinhciis Blantltord [=Hijlun^ops
fusliiincnsis Muravama), Hi/liir^op.s intcrsfititdis ((^hapuis) (=Hyliirgi)p.s nipoiiiciis Muravama), Hi/litri^ops spcssivtscvi
Eggers (=Hi/liii'<s,op.s modest us Mura\amai. //).s stehhin<^i Strohmever (=Ipsseliiimtzenhoferi Holzschnh), Pldoeosinus nidis
Blandiord (=Plil()ei)sinus sliDtneiisis .\Iura\ama. PoJif'^rdphus kdimochi (Nobuchi) (=Pi>li/<^rdphus qtierci Wood), Poly-
•n'dplius pnixiinus iilandford {=P(ih/^rdjilius iiu/i^iius Mura\ama), Sei>Ii/t(><yne.s brdderi Browne ( = Seoliit(>^enes orientdlis
Scliedl), Seoli/tiipldli/pus pdniis Sampson (=Sa>li/topldti/pus rnfifiiudd Eggers), Sphdciolnipes ipierci Stebbing
{ = Chr<iinesus 'jjoliulus Stebbing, Sjiluierotiypi's teetiis Beeson). Siiens niisiinai (Eggers) ( = Spli(ier(itn/pes eoiitrorersae
.Muravama). Tainiens hrei ipilosus (Eggers) ( = Bldsli)plidiius klidsiaiiiis Muravama. Bhistophdiius iiiultisetosus Mura\ama).
The European lli/ldstes updeiis Erichson is reportetl as an establishi'd breetling population in New York ( US.-K). Pliloeosiiius
annatus Heitter of Asia Minor is rcpoiteil as causing economic tlamagi' as a new introduction to Los .-Kngeles County,
California. The following species arc named ;is new to scit-nce: Cijcloiiiipididii siihdiiiidtiiiii (Pliilippine Islands),
Dendwtmpes zcdhiudleiis (New '/e;ilaudl, Pohiiiidjilms lliitsi d^urma). rrinteiiiiuis pilieoiiiis (buli;i). and Xi/lehonis
ina<inifirus (Peru'.
Key uords: iKiiiiciicldttirc. Phili/jiodiddc Srali/tidae. Idxoiioiini. hark hectics. Colcopteni.
Durin<r the conipilatioii ol' a vvorlcl catalog of (e) two new in.trodiictioii.s of a European and an
Flatvpodidae and ScoKtidae, a nuiiilxM- of A.sian .scoKtid into North Ameinca, and (0 five
nonienclatnral iteiii.s vvcn^ ionnd that i('(|uire
vaHdation and/or [)nhHcation prior to relea.se of
the catalog. The.se items inchide: (a) two new
rej)Iacenient names for jnnior homonyms in
Idatvpochdae and nine in ScoKtidae, (b) three
spc^cies named as new to science
New Names in Pe.\tvp()didae
new combinations in fossil Scolvtidae, (c^ si.\
cases ol new generic SNiionv ni\ in ScoKtidae, (d)
17 cases of new specific .s\ iioin ni\ in ScoKtidae,
Gciii/occriis stnilintci/cri. n. n.
Didpus (dhipeiiiiis Strohme\er. 1942, .\r!)eiten uber
Moiphologische iiiul laxonomisclie Eutomogie 9:284
(SvntA'pes; Insul Simaloer, westlich Sumatra; Strohmever
Collection), preoccupied In .Motschulsk)-, 1858
.332 Lilf Scifiiti- Miisciiiii. Brii;liam Voiiiij; b'nivcrsitw I'n
78
19921
N()MEi\(:LATri{Ai. C:nA\c;Ks i\ PiAriTontim: wi:) Scoi.^TinM
79
Tlic naiiic Clcm/occnis alhipcimis Motscliiil-
sk\', 1S5S. was c()ii.si(l(M'(Hl lost for moro than a
centun (Wood 1969: US). In an attempt to
assiiiin a species to this iianic, Stn)hn)(>\'er
named Diapus alhipennis. cited ahoxc. When
the Motschulslcv' hpe was r(nlisco\(M-ed (Wood
1969:118), it was recognized that two distinct
hut congeneric species were representetl.
Because the Strohme\er name is the juuioi-
homouNin in this case, the new name stroli-
nict/cri is [proposed as a replacement name lor
(ilhipctDiis Strohme\er as indicated ai)o\e.
Pl(iti/})iis applanatulus, n. n.
rliiti/pns tijiplintdtiis ScIr-iH, 197(i, .\l)liaiKlluii<ieii
Stiuitliches Museum fur Tierkkunde IDresden 41(3):S5
(Ilolotvpe. male; Manaus, Amazonas; Naturhistorisches
Museum W'ieuK preoccupii'd In Wootl, 1972
rUitijpus applanatus Schedl, 1976, cited
al)o\e, was named fi\e \ears after the same
name had been used b\ Wood (1972:244). In
\i(n\ ol this homonxniiv, the new name
(ippltniatiihis is here proposed as a replacement
lor the junior name (ipphnuitiis Schedl, as intli-
cated al)o\e.
New Names in Scolytidae
Cn/f)luiliis hnnviwi. n. n.
Cn/plialitsai-toc(ii-f)u.s Schedl. 1958, Sarawak Museum Jour-
ual 8(11):498 (Holotxpe; Sarawak. Seuien2;oli: British
\Iuseuiu [Natural Ilistorxli. preoeeupietl h\ Schedl.
1 9:39
T\\r name Crijpiuilus aiiocaipus Schedl,
195S, cited ahoxe, was established even though
its author had previously named Eiicn/pltaliis
(iiiordrpiis Schedl, 1939, and had considertnl
Cnjpluiliis and EricnjpJuilus .s\nion\nious. This
generic s\non\-m\ was confirmed (Wood
1986:91). In view of this oversight, Schedls 1958
name is a junior liomonym of the 1939 name and
must be replaced. The new name hrowiici is
pioposed as a replacement, as indicated aboxe,
in recognition of the late F. G. Browne who
contributed significantK to our knowledge of
t]ies(^ insects.
(â– i/clorliipidioii (lOiiixincuni. n. n.
Xijichonis (liliiii^fiisis Scliedl, 1951. Tijdschrilt \oor
Entomoloi^e 93:71 (S\nt\pes, 2 f'euiales, 1 uiale: Ja\a:
Batoerraden. G. Slauiet: Naturliistorisches Museum
W'ien), preoccupied In Eiiijers 1930
The name Xylehonis dihinfj^ensis Schedl, cited
above, was proposed at a time when it was
preoccupied l)\ lvj;gcrs, 1930. .\ltliou'j;h both
names were reccMitK transferred to other
genera, the [)riman' homoimnv remains. The
new name (liltiii<^icuni is proposed as a replace-
ment lor the Scliedl name as indicated abo\e.
Hi/})c>lliciicimis (itcrriniitlus. n. n.
lA})kcr()khs (ilcrhiims Schetil. 1957, .\miales du .Miisee
H()\aK(lu ( 'oiiiro Ik'Ige, ser 8. Zoologie 56:59 (HoloUpe;
i-iuaiida: lliruil)e: Belgian Congo Museum. Ter\iiren),
preocciijiicd In Schedl. 1951
The generic name LrpUrwUk's ScIuhII was
placed in synon\ui\ under Hijj)(>theiu'miis
(Wood 1986:92). This act transferred its t\j)e-
species, atcrrhnns Schedl, 1957, cited abo\e. to
HypotJieuciiuis where it became a junior hom-
omm of//, (itcrhmus (Schedl, 1951). The new-
name <7f<;'rn//(/////.s' is here proposed as a rej^lace-
ment name for (ilcniiinis ScIumII, 1957. as indi-
cated aboxe.
Hijpothcncinns krii oliitskai/ac. n. n.
Ui/j)()tliciuiiiu\ iiiMilanini Krixolutskava, 1968. ;/( Kureu/.cn
& Konoralova, The insect iannaof the So\iet Ear East ami
its ecologv', p. 56 (Ilolorspi-; Kiiriie Islands; presumahK
at \1adi\()st()ki. pre()ccuj)ied l)\ Perkins. 1900
Hijpotheneitiu.s iiisulanim Kri\()lutska\a.
cited above, was gi\en a neuter specific name in
a masculine genus. When the gender is cor-
rected, as re(|uire(l under tlu^ C^ode, this name
becomes a junior honioii\m ol Hi/pothcucmus
insuloris Perkins, 1900, and must be replaced.
The new name khrolutskat/dc is proposed as a
replacement name, as indicated al)o\e.
Fiti/oplilltiinis (ilricdiiiihis. n. n.
Meocln/ococtis iifiicdiiii.s Schedl. 1962. Re\ista de
Entomologia de Mocamhique 5(2);1079 (Holot\pe;
("ongo; Ma\uml)e; Belgian ('ongo Museum. Tennren),
preoccupied l)\ Eggers, 1927
Schedl naiiK'd Xcodn/ococtcs (ifricaiuis. cited
aboxe, from fi\e specimens that did not e\hii)it
sexual (hflerences. Because the neotropical
genus. A/Y//;/f/.v ( -Xcodn/ococtcs) does not occur
in .Africa and tiiese specimens belong to the
related gcMius Piti/ophfJionis. Schedls name,
afriatnus. iinist l)c transh'iicd to that genus
where it becomes a junior homonxin and must
be replaced. The new wMwe ofriconuUis is pro-
posed as a replacement for the 1962 Schedl
name as indicated aboxc.
Scoh/fD^cncs papucnsis, n. n.
Xijlcciifptiis p/ipitiniiis Schedl, 1975, Naturhistorisches
Museum W ieu. .Annales 79:352 (Holotxpe; Upper Manki
80
(;i{KAT Basin Natuhaijst
[N'olunie 52
logging area, Biilolo, MoioIh^ District. New Ciiiiu-a: jt must he replaced. The new name, formosae,
Naturl,i.st()risd.e.s Mu.seuin Wicni. pre.Kcnpu.l Ia ■p,-„po.secl a.s a reiilacement as indicated ahoxe.
Schedl. 1974 ^ ^ ^
The genus Xijl<)cn/j)tiis Schedl, 1975, was
estahhshed with X. papuduns Schedl as the tyj)e-
species. When Xi/l<)cn/})fus became a junior s\ii-
omm of Sc()lylc)<i,('i}cs (\V''o()d 1986:90), the
transfer of papuanus to that genus caused
papuanus Schedl, 1975, to become a junior
homonvm o\' Scoh/to^otes (originally Cnjphalo-
inoiyhus) papiKnuis (Schedl, 1974). In order to
correct this duplication of names, the new name
papiicnsis is here proposed as a replacement for
ptiptumus Scluxll, 1975, as indicated alxne.
Sci>lij((><s,('nes jjaradoxiis, n. n.
Sa)lijh><s,cn('.s papuanus Sciiedl, 1979, Fauiiistisflit'
Ahhandlungen 7:97 (Hoiotxpe; I'apua, New Cruiiiea;
Naturiii.stori.sclie.s Museum Wien), preoccupied In
Schedl, 1974
When Sc(>h/f()<iciics papuaiius Schedl, 1979,
was named, Schedl ve^^ardedCnjphdloinoqjJtus
as a distinct genus. The placement of CnjpiidJo-
nioiyhus in sviiomniv under the senior name
Sc(>h/t()<^('ncs (Wood 1986:90) and the conse-
quent transfer of C. pnpuanns Schedl, 1974, to
Scolijto<^enes caused the name S. papuanus
Schedl, 1979, to becouie a junior homouN in. For
this reason, the new name paradoxus is pro-
po.sed as a replacement for papuatnis Schedl,
1979, as iudicated above.
Xiflchoriiuis spi)iip()sticus, n. n.
EidophcUis .spinipcnnis Schedl, 1979, New Zealand Ento-
mologist 7:106 (Holotxpe, leniale?; Fiji: Schedl C^ollee-
tion ill Natiirhistorisches MuseuiiiW'ieii), preoccupied In
loggers, 19:30
Bea\-er (1990:94) transferred Eklophflus
spU\ip(')u\is Schedl, 1979, to Xi/lchoriiuis where
it is preoccupied hy sj)inij)cii)iis (Eggers, 1930).
Inordertorenunetheduplicatiouofnames, the
new name spiniposticus is heie proposed as a
replacement kn spiniju-iniis (Schedl, 1979) as
indicatcnl abo\e.
Xijlehonis jonnosac, n. n.
Xijichonis foniio.sanits Browne, 19S1, koiitsu 49(1):1:)1
(llolot\pe. female: Ilualien (Formosa) tf) Yat.su.shiro
(Japan), imported: British Mu.seuin [Natural IlistotA]),
preoccupied In Fggers. 19.30
When Browne named Xijlehonis forniosauus.
cited aboxe, he (nerlooked pre\ious usage oi"
this species-group name in the combination Xi/le-
bonis nuniciis foniwsanus Eggers, 1930:186.
Because the Browne name is a junior homonxm,
Generic Ti^ANSFERS of Fossil
SC;OLYTIDAE
Drijococtcs (liluvialis (Wickham)
l'lli/(iplillii>ri(lc(i (liluiidlis \\ ickliam, 1916, State Unixersity
of Iowa. Eahoraton- of Natural IIistor\; Bulletin 7: IS
(IIolot\pe: fossil in Miocene, Florissant, Colorado: not
located)
The photograph of the holot)pe that w-as pub-
lished with the original description of Piti/oph-
thoridca diluvialis Wickham ( 1916:18) suggests
that tins species is a member of the genus
Dn/ococtcs. Because there appears to be no
justification whate\er for recognizing a separate
genus, the name Pitijoplifhoroidcs is placed in
synonymy under the senior name Dnjocoefcs,
and diluvialis is transferred to that genus, as
indicated aboxe.
Hi/lcsiiuis hijdntpicus (Wickham)
Apidoccpliiihis }u/(lri)})inis Wickham, 1916, State Universitv
III lo\\:i. Laboraton of Natural Iliston; Bulletin 7:18
(Holotspe: fossil in Miocene, Florissant. Colorado: not
located)
The photograph of tlie holotxpe that was pub-
lished with the original description of Apido-
ccphahis lu/dropicus Wickham indicates that
this species is a member of the genus Hi/lesinus.
The generic name Apidoccphahis is here placed
in .synonymy imder Hijlcsiuus and the fossil spe-
cies hijdropicus is transferred to that genus, as
indicated above.
Plilocotrihus ziiunicniumiii Wickham, to
C>urculionidae
Pliliicdlrihu.s ziiiiincniianiii Wickham, 1916. State Uni\er-
sil\ ()l low:i. Lalioratonof Natural Histon-, Bulletin 7:19
( I lolohpe: fossil in .Miocene. Florissant, ('oloratlo: not
located)
The photograph of the holotxpe o\ Phhwofri-
hus zinintcrnunnii Wickham (1916:19) that was
[)ublishedwith the original description indicates
that this species is not a member of this family
and nmst Ix^ transfernxl from ScoKtidae to the
famil\- (Jurculiouidae.
New Synonymy in Scolytidae
(Uisiuodcrcs Eichhoff
CoMnodcrcs Eicliln)!!, 1S7S. Societe Entoniolo^iijiR' de
Liege, Memoires (2)<S:495 (Tvpe-species: (".osinodcrcs
monilirollis Eichhoff, monobasic)
19921
NOMENCLATl'HM, C:iIA\CE.S IN PLATVrODlI) \I-: WD SCOI.^TIDAK
81
Fscu(l(>C()siiu>elcrcs Nobuchi. 1981. Kont\ii 49(1 ):16 (T\pc-
sprcii's: I'sciKlocosiiuHlcrcs atictiiiatiis Nohuchi =CV).s-
I node res inoiiilhcllis I'iclilioll, original (Icsii^natioii). ,\V(c
siiiiiinijxui
TIk' ^('iius FscikIocosiikxIci'cs Xohuflii. citctl
al)<)\'(\ was named lor Pscu(l(>c()s))U)(lcrcs
atlciiuatus Nobuchi, 19S1. The photojiiiaph ol
iho hpe material that accompanied the oriij;inal
description is an ilhistration of ('.osnuxicrcs
iiK'nilicollis Eichhofi, 1878. The Nobuclii genus
is an ohxions .sviionvui of Cosinodcrcs. The
sj)ecilic s\ iionymy requires confirmation, l)nt is
almost certainlx' correct.
Dnjocoetcs Eichlioff
l)n/()cc)iii:s Kic-liliotf, 1S64, in Sthiciik, Hii'st-ii unci
Forsclmngeii in .\niur-Landf 2:155 (T\pf-,specirs:
Biisfnchiis tiut()<s,r(ij>lius Ratzel)uit^, snlisequent designa-
tion InWood 1974)
I'id/oplithoridca W'ickliaiii, 191fS. .State Uni\ei".sit\' ot Iowa,
Lalioraton' of Natural Histon. Bulletin 7:18. figs. 27-28
(T\pe-speeies: Piti/oplithoruica dilurialis Wickliani. orig-
inal designation). Xcic si/ndinfiuii
Tile figtu-es of the liolotxpe of Pifijopli-
tlioridcd that were publislied with the original
d(\scri[)tion indicate that the tspe-species, P.
(liliiiialis, is a meml)er of the genus Dn/ococtcs.
(,'()nse(|uentl\, Wickhanis name Pifi/oplifhor-
ulcii is [ilaced in s\iion\ni\ under the senior
name, as indicated aboxe.
Hijpothenemus Westwood
lUijH'ihdicuins W'esbivood. 1836. Entoniologieal Soeiet\ ol
London, Transactions 1:34 (Tvpe-species: Htjpotliciicnius
cniditus Westwood. monobasic)
Macrornjphaht.s Nobuchi, 19S1. Kontvu 49(1 ):14 (Tvpe-
speeies: Mdcrocnjpludns ohlougna Nobuchi, original des-
ignation). Frohaljje s\non\in\'
The g(^nus Macrocn/plialiis Nobuchi, cited
abo\e, was named InrMacrocn/phalus olAoii'^us
Nobuchi. .'\ close examination of the photo-
gra])hs of t\pe material pul)lislied with the orig-
inal descriptions clearK indicates that the
species ohlonous is composite. Tlie "male"
illustrated is a female of Ht/potlwncnnis
Jiiscicollis Eichhoff a sj^ecies ra])idl\ b(^c-oming
[)antropical in distribution through commerc(\
rlie â– female' is a female of another
//7/)e//H'(/r///?/\ speci(^s that cannot be identified
with certaint\ from the illustrations. It repre-
sents an ob\ious introduction from another
area. The name Macrocn/pluilus is lu^-e placc^d
in sNuonxniN until tlie name ()l)l(»i<^iis can be
clarified.
Lipai-tltnim Wbllaston
Lipaiiltnnii Wbllaston. 1854. In.secta Maderensia. p. 294
(T\pe-s|X'cies: Lipaiihniiii hUiihcrnilatuiii Wbllaston.
original designation)
'I'njpuiioplicUos Bright. 1982. Studies on Neotropical Fauna
and Kn\ironnient 17:166 (T\pe-species: TnipauophcUos
iicc(>])iitus Bright). Newstpioinpni/
Tii/paiioplK'Hos iiccopimis j-iright was based
on a unicjue female collected bv Schwarz at
Cayamas, (^uba. I examined this specimen in
1976 at the U.S. National Museum and recog-
nized it as a (listincti\e, undescribed species of
Lipaii]iruiii.T\\(.' holot\pe was recentk' reexam-
ined and compared to otluM- Lipartlii-uni spe-
cies. Because I am unable to see an\ generic
characters that might possil)l\ distinguish
Tnjpan()j)licll()s front Liparfhnou, Bright's
generic nanu^ is placed in s\ iionxinx- under the
senior name as indicated abox e. The species, L.
necopinus, is uni(jue among .\merican Lipar-
thniin species in liaxing a double row of scales
on the decli\ ital interstriae.
P()li/<irapluis Erichson
Pch/'^rapliiis Erichson, 1836, .Arclii\ ffir Naturgeschichte
2(1):57 (T\pe-species: Ili/lcshiiis puhescem Fabricius
= Dcnnestcs polif^rapliiis Linneaus, monobasic)
Xipponopoli/griipliiis Nobuchi, 1981, Kontxu 49:12 (Tvpe-
species: SippoudpoliinrtipJius: kaiinochi Nobuchi, origi-
nal designation). \ctr sijiioiiipiu/
The holotxpe and two paratxpes of
Nipp()ii()p()li/<ii'(ipliiis kaiiiuxhi Nobuchi were
examined and found to be normal specimens of
Polijgraplms Erichson in w Inch the eye is deepK"
emarginat(\ but not dixided. Approximatelx'
one-fifth of the species in this genus haxe the
halves of the eye connected. The Nobuchi
genus xvas based on this one unusable character-
and must be placed in sxnonxnix as indicated
aboxe.
Scohjto^ieiw.s Eichhoff
Sci>li/t()gc)ics l'',ichhoff". 1878, preprint of.StKiete Roxaledes
Sciences de Liege, Memoires (2)8:475. 479 (T\pe-spe-
cies: S(()h/I()<ji'iics danciiii Eichhoff, monolia.sic)
('njpluilopliilus Scliedl, 1970. Kontxii 38:358 {Tvpe-s[X^cies:
('n/phalophihis afer Schedl. monobasic). Correction of
sifnoiiipitii
Due to a clerical error in Wood (1984:228),
the name Cni])Jialop1ulus Schedl xx'as incor-
rectlx placed in .s)nionyinx under the name
Scohjtodcs, a neotropical genus. CnjpJial-
ophiliis is actuallx a .sxiionxin of Scohjtoocncs. a
circumtropical genus. The holot^pe of the t\pe-
species, C. afer, was examined.
82
Gheat Basin Naturalist
[\'()li
Tapli ronjclms Eichhoff
Taplironjchits Eiclihoff, 1<S78, prcpiiTit ol Socic'ti'" lloxalf
des Scieiitc's de Eiege, Memoires (2).S:49, 204 (Tspe-spe-
cies: BostricliuM hicolor Ilerhst, .siil).sc(jucnl clesigimtion
bv Hopkins 1914)
Taphrococtes Pfeffer. 1987. Acta Entoiiiologica
BoluMiioslovaca 82:22 (T\pe-specie.s; Taphronjcliiis
Itiiicllits Eichlioff, oritiiiial designation). \'cw sipioiujiiuj
The name Taphrococtes Pfeffer, cited above,
was proposed as a means to subdivide the genus
Taphron/cJtiis using the size and distribution of
asperities on the anterior slope of tlie pronotum.
Because Taphrorijcluis is much more wide-
spread and diverse (\Vood 1986:74) tlian was
known to Pfeffer, a division of the genus using
the pronotal characters lie proposed is not
possible or meaningful. Several examples of all
European and most Asiatic species of this genus
were examined in my review of this problem. As
indicated above, Taphrococtes is placed in svii-
oiniuN' under the senior name.
Brachijspartus inoritzi Ferrari
Biachijspartns inoiitzi Ferrari, 1867, Die Forst- uiid
Hanni/nelitseliadlichen Borkenkafer, p. 68 (Holotvpe,
tenure; \'ene/.neki; Naturhistorisclies Museum Wien)
Cotihijhis ohtiisiis Schedk 1966, Entomologsehe Arbeiten
ans der Museum Frev 17:122 (Hok)t\pe, female: Wne-
/.uela; Naturliistorisches Museum \\ ien). Ncic sipioiuiini/
The female holotyj^ies of Brachi/spartus
nioritzi Ferrari and Co)~tJtt/his obfiisus Schedl
were compared directK to one another by me
and were found to be identical in all respects.
Thev obviouslv represent one species in which
Ferraris name has prioritv, as indicated abo\e.
Carphohonis ntiiiiinus (Fabricius)
Hijlesinu.s iitininttis I'abrieius, 1801, S\stema Ele-
utlieratoruni 1:395 (Syiitypes, 4; Saxoniae: (Copenhagen
Museum)
Ciiq)li(>l)(>nis /w/g('»i.v/.s .Muravama, 1943, .Annotationes
Zoologicae Japonenses 22:99 {Lect()t\pe, male: District
of Halga, Manclioukuo, China; U.S. National Museum.
present designation). Xcic sipiniupin/
Caqyhohonis IxiU^cnsis Muravama was
named from one male and one female syntvpes
mounted on separate microcards on one pin.
The male is in recognizable condition and is
here designated as the lectot>pe for this Mura-
vama name. The "female" has been damaged
and only the head remains; its face is entirc>l\
iuunersed in glue. This lectotype was compared
to males of my .series of C. Diininiiis (Fabricius)
from Europe and northern Asia. While no two
males of this species are ever exactly the same,
tlie halgen.sis lectotvpe is of the same size and
proportions as C niininiiis and falls well within
the limits of variabilit)- and geographical range
for this species. Because only one species is
represented by this material, the name balgcnsis
is placed in .synonymy as indicated above.
Coccotnjpcs dacttjlipcrdd (Fabricius)
Bnstrichus dactijlipenla Fabricius, 1801, Systema Ele-
utheratoruni 2:387 (S\ait\pes, female; date pits inter-
cepted in Europe; Copenhagen Museum)
Coccotnjpes tropicus Eichhoff, 1878, preprint of Societe
Royale des Sciences de Liege, Memoires (2)8:312 (Holo-
tvpe, female; .America Meridionalis (Peru); Hamburg
Museum, lost). New .siptoiii/iiii/
Eichhoff states in the original description,
cited above, that his Coccotnjpcs tropicus is
near C. dactijlipcrda. Because the description
fits the pantropical dactijlipcrda. because there
are no knowii endemic Coccotnjpcs in South
America, and because the unicjue holotvpe and
only known specimen of tropicus was lost in the
destniction of the Hamburg Museum, C. tropi-
cus is here placed in synonymy under the senioi
name, as indicated abov^e, as a means of dealing
with this unidentifiable species.
Cnjphalus scabricollis Eichhoff
Cnjphalus scaljiicollis Eichhoff, 1878, preprint of Societe
Rovale des Sciences de Liege, Memoires (2)8:36 (Holo-
tvpe; Hindustan Asiae; Hamburg Museum, lost)
Cnjphalus hreiicolli.s Schedl, 1943, Entomologische Blatter
39(l-2):36 (Leetotvpe, female; Bagnio, Luzon,
Philippineu; Naturhistorisclies Museum Wien, desig-
nated b\ Schedl 1979:47). \'cw sipidiiiinu/
The holotvpe of CrijphaJiis scabricollis
Eichhoff was lost in the 1944 destiiiction of the
Hamburg Museum. My concept of this species
is based on a series of specimens in the Forest
Research Institute, I>=>hra Dim, that was com-
pared 1)\- Beeson and Eggers to the hoK^tvpe
before it was lost. Mv series was compared
directly by me to this series; then these speci-
mens w t're later compared to the holot)pe of C.
brcvisctosus Scliedl. All represent the same
coimnon, widely distributed species that infe.sts
various species oi Ficus from bidia to the Phil-
ippine Islands. For this reason, Schedls name
C. brcvisctosus is here placed in svnionvmy
unck'r the senior name, as indicated above.
Ficicis dcspcctus (\\'alker)
llylcsiiius cicspcdus Walker. 1859. Annals and Magazine of
Natural lliston (3)3:261 i llolotNpt'; Cevlon: British
Mu.seum [Natural Histon])
Hylcsiiius siniiiKniiis Schedl, 1951, Bishoji .Museum Occa-
sional Papers 20(10): 142 (Sviitvpes, male; Upolu,
1992]
NOMENCL.\TUHAl. Cll A\(;KS IN PLATYI'ODH) \i; AND S( iOLVniMK
83
Tapatapao; British Miisriiiii | Natural llistorvj and
.NaturliistorisflK's Muscuiii Wiciii. Wu \i/iu>iiijiiii/
Tli(^ Schc'dl sMihpes of Hylesiinis saDioanus
Scliedl in the W'ien Museum were examined 1)\
me and were c()m[)ared dii'eetK to m\ liomo-
t\pes ol H. (Icspcciits Walker. C)nl\ one speeies
was reeoifnized. On {\\v hasisof tliis c'<)ni[)ai"i,s()n.
Scliedls name is plaeed in s\non\in\. as indi-
cated abo\e.
Hi/lasics pliiiiihciis Hlandloixl
Ih/liislcs j)liiiiiliiii\ 15landford, 1894, Entomological Socich
oi London, Transactions 1894:57 (S\'nhpcs; Nagasaki ct
a Ilioga, Japan: Brnssels Museum)
//(//» /"ijo/n fttslimiciisis MuraNama, 1940, Annotationcs
Zoologicac japoncnsis 19:235 (Lectohpe, feniide:
Fuslicn. .Mancinuna: U.S. National Museum, present des-
ignation). Scic si/noin/ini/
Hijliir^Dps fiisliiiitoisis Mnraxama was hased
on one male and one iemale s\iit\pes that are
mounted on one pin. The callow female is
mounted upright; the callow male is moimted
upsitlc> down with the dorsal surface imbedded
ill glue. The female is here designated as the
lectot\"pe for //. ftishiniciisis Mura\ama. This
lectot\pe was compared directK t(i ni)' Ussuri
specimens of Hylastes pbimhens Blandford that
were identified b\" Kurenzow These specimens
clearlv represent one species. For this reason,
fuslunwnsis is transferred to Hi/lastes and is
placed in s\non\-my under the senior name, as
indicated aboxe.
I li/liir<j_ops iittcrsiiiialis (C'hapuis)
Hijldstcs interstitiiilis (lliapuis, 187.5, Societe Entoinolo-
liique Belgifjuc. Aiinalcs 18:196 (S\iit\pes; Nagasaki and
Kiuslui, Japan; Bnissels Museum i
lliilitn^oj)s nipdincns Mura\ama, Ui.'id Tcntlin-di) i:12.).
149 (Ilolotxpe, mule: Kamikoclii, Nagano prelect mc:
IS. National Museum). Nnc si/noni/intf
The uiii(|ue male holot\pe ot lhjluriH>ps
niponiais Muraxama was examined and com-
pared directK to m\ long series ol //. ntlcr-
stifiali.s (C^hapnis) from |apan (detcMiiiiiicd 1)\
Nobuchi) and Siberia ({l(4(M-iiiiii('d b\ Kiiicii-
y.ov). The Miiraxaiiia holotxpe is an axciage
Japanese specimen ot this species. The name
nipoiiicus is here placed in sxtioumux under the
senior name as indicated aboxe.
Hifltir^ops spcssivtsevi Eggers
Htjlnrgops spessivtsevi Eggers, 1914, Entomologisclie
Blatter 10:187 (Lectot\pe, male; Ostsiberien, USSR; U.S.
National .Museum, designated bv Anderson & Ander.son
1971:;30)
Htjlur'^ops niodcstus .Muraxama, 19.37. Tentbredo l:.3fi7
(Syutxpes; Pic Biro du Kongosan. Korea; .\Iura\ama C^ol-
lectiou in U.S. N;ition;il .Museum). Ncic sijnont/nit/
Txxo Iemale six'cimens in the .\hnaxama (Col-
lection are labeled as "paratxpes" (.){ Hijlur'^ops
ni()(l('siiis .Muraxama. Their label indicates that
thex xxere taken at "Yalelomia. Mancliiiria, 25-
MII-f94() bx \. Takagi"; a second label gixes
"Manchoukuo, (,'ollected 1940, J. Miuaxama,
Hylurgops nuxlcstus Muraxama, parat)pe."
Because this Muraxama species xx'as named in
1937, it is presumed that these "paratxpes" are
actuallx metatxpes that xxere compared bx-
Mtnaxama to his t\pe series. Murax ama told me
in 1955 that xirtuallx' all of his Manchurian col-
lections had been destroxed during World War
II. Con.seqnentlx, the aboxe "paratxpes" are
probablx the onlx knoxxii existing .specimens of
nuxlcstus that are reasonablx autlientic. These
"paratxpes" xx'ere compared directlx to m\'
homotxpes of H. spessivtsevi Eggers and xxere
found to be normal, axerage specimens ol this
Eggers species. For this rea.son, the name iiuxl-
estiis is placed in .sxnonxinx under the scMiior
name, as indicated aboxe.
Ips stchhiiigi Strohmexer
lp\\trhhiii<^i StroinncNcr, 1908, Entomologi.scben Wbclien-
hlatt 25:69 (Sxnhpes. male. lemiJe: Kula. Himalava
occidentalis: Strolunevi'r (Collection. Eberswald. Forest
Research Institute. Dehra Dun, etc.)
Ijis sclmiutzeiiliofcn llolzschuh, 1988, Entomol()gic;i
Basilieusia 12:481-485 (Ilolotxpe, male; W'e.st-Bluitan,
Cham^ang, 3000 m: Naturhistoriscbes Museum Wien).
.V(7r siiii(})iijiiit/
1 examined txxo sxiitxp(\s ol Ips stehhin^i
.Strohmexer in the Forest Research institute
(.'ollection, Dehra Dun, as xxell as approxi-
matelx 2. ()()() other specimens of this species
from l^ikistan, Nepal, Bhutan, and India
(Kashmir, Punjab, Uttar Pradesh) from species
of.\/>/'r.s. C.idnis. Picra. and riniis <s^ri[fitliii. I am
unable to distinguish inx specimens that xxere
compared to the Strohmexer sxiitxpes from t\\-o
paratxpes of /. scJinuitzenhofer Holzschuh or
from a series taken in 19(S0in Bhutan '(xomPicea
spiinilosa bx P. Singh. It is apparent from the
description of /. sehmutzenhoferi that .speci-
mens cited as /. stehhiii<ii xxere actuallx of 7.
longifolia, a distinct, but related, species. In
xiexx' of the aboxe, /. sehmutzenhoferi is here
placed in sxnonxnu', as indicated aboxe.
84
Ghkat Basin Naturalist
[\-
online o2
Plilocosiiuis nulls Blandford
Plilocosiinis nidis BlaiKifbrcl, 1894, Entoinolo^iciil Society
ol LdikIoii. Transactions 1894:73 (Sxntvpes; Kaslii\\'aij;('
and K()II)e, Japan: Britisli Mnseuni |\atnrai Ilistonj)
Plilocosiniis shotociisis Muravatna, 1955, Yaniagnti Uniwr-
sitA Facnlh of Aijricnitnrc. Bulletin 6:88 ( Holotspe, male:
Japan: Onnde, SluHlojinia. Kapma pref.: U.S. National
Mnsenin). New si/iioiiyini/
The tN'pe .series of Plilocosiiuis sliolocnsis
Murayama consisted of one male and six
females from the t\pe localitv and seven females
from other named localities. Murayama clearly
states that the male is the t)pe. All 13 specimens
in the tvpe series were compared to my homo-
t)pes of P. nulis Blandford. The Murayama
sjiecimens fall well within the range of varial)il-
it\' of nidis. Because it is ohxious that only one
species is represented by these specimens, the
name sliofociisis is placed in SMion\-m\' as indi-
cated aho\ e.
Poli/^r(ij)liiis kaintorlii (Nohuchi)
Nippoiu>p(>h/^raj)hiis kaiiuo<-lii Nohuelii, 1981, KontMi
49:1.3 (Il()lot\pe, female; Sliionomisaaka, \\'aka\ama:
Nobnchi Collection, Ibaraki)
Pohj<ir(ipluis qticrci Wood, 1988, (ireat Basin Naturalist
48:195 (nolot\ix>. female: Melialkhali [Bnrma?]: Forest
Research Institute, Dehra Dun). Xcu: si/noiiiiiuij
The female holotspe and two parat)pes of
Ki])])onopohj<^ropluis kaiinorhi Nobuchi were
compared directly to one another and to the
t\pe series of Poli/<^rapliiis cfncrci Wood bv me
and were foimd to represent onK' one species.
The junior name, qiicrci, is placed in s\iionvm\'
as indicated above.
Pohj<^raj)liiis f)ro.\ii)iii.s l^landford
Pohj^raphus proxiinus Blantllord, 1894, Entomological
Society of l^)nd()n. Transactions 1894:75 (Sviit\pes, 2;
Sapporo, Japan; British Museum [Natural Ilistonj^
P<>ltl<ir(i})liii.'i m(i<iiiits Mnra\ama, 1956, Yamaguti Uni\ersit\
Faculty- of Agriculture, Bull(>tin 7:279. 282 (IlolotApe.
Icniale: Nishiniata, Aki C^onntA, Kochi pref., Japan; U.S.
National Museum). Sen: si/uoiti/iiii/
The unique female holotApe oi' Poh/<irapluis
nia<inus Muravama was examined and com-
pared to my series of /^. proxiiniis Blandford that
had been identified b\ Kureuzox, Nobuchi. and
Pfeffer. A .series of this species receixxnl from
Mura\ama had been id(^ntified as P. oblon^^ns
Blandford and is presumed to be incorrectK
placed by him. The ina^^mis holotvpe is 3.2 mm
in length (exclusive of the head), which is sub-
stantially smaller than stated in the original
description. The jironotum ol this specimen is
contaminated In host resin, thereb\ gi\"ing both
tlie stout biistles and scales the false impression
that they are all scalelike. In realit\', these setae
are precisely as in normal specimens of prox-
iiniis. In addition, the size falls well within the
upper limits of size for /;r<u"//////.s'. The nui^iiiis
holotvpe obviously is a normal, large female of
proxiniiis. For this reason, the Murayama name
is j)laced in s\nonvm\' as indicated abo\e.
Scoli/to<s,ciics hradcri (Browne)
C'n/pliahiiiHirpluis hradch Browne. 1965, Zoologische
Mededelingen 40:191 (Holot\pe; I\on C'oast:
Adiopodoume; Leiden Mu.seum)
Cn/])luih>uu>rphns oriciifalis Sclietll, 1971. Opu.scula
Entomologica 119:11 (Holot\pe; Clliana, BekAvai;
Naturliistorisches Museum W'ieni. \civ si/ni>iu/uu/
The holotvpe of Crijplialoinoiylius orientalis
Schedl, cited above, was compared directly bv
Schedl to the holotvpe of C n/phaloinorphus
bracleri Bro\\aie, cited abo\e, and (as indicated
in a note in his collection) he concluded that
only one species was represented. I examined
the Schedl holotvpe and compared it to speci-
mens identified b\' Schedl as hradch Brcmaie
and reached the same conclusion. In view of
this, the name orientalis is here placed in svii-
on\in\' as indicated aboxe.
Scoh/toplati/pns pairus Sampson
Snihjtopldti/jni.s parvus Sampson, 1921, .Annals and Maga-
zine of Natural I Ii,stoi-v (9)7:36 (Ilolotspe, male; Sarawak,
Mt. .Matang; British .Mu.semn [Natural Histon])
Scolt/foj)l(ifi/})us nifianula Eggers. 1939, .\yV\\ for Zoologi
31.'\(4):.36 (llolot\pe, female; Kamhaiti, .Nordost-Birma,
7()()() ft.; Stockholm Museum). Nnr sipioiu/Dii/
Four specimens of Scolt/toplati/piis parvus
Sampson that were compared to the holotvpe by
Brownie were compared directh" b\' me to nine
specimens in the Forest Research Institute,
Df^hra Dun, that had been identified bv Eggers
as his S. nificaiida. The\- all represent the same
species. Assuming that Eggers correctlv identi-
fied his species, tlu^ name s. nificaiida nnust be
placed in sviionx ni\ under the senior name S.
pan lis. as indicated abo\ e.
Spltacrotri/pcs cjiicrci Stebbing
Sphiicrotn/pv.s (jurivi Stehhing. 1908. hulian Forest Mem-
oirs, .series 5, 1(1):5 (Sviitvpes, sex?; India. N-\V Hima-
la\;i, Kunuimi: Forest Research Institute, Dehni Dun,
lost)
('ludincsiis 0(>hiiUis Stehhing. 1909, Indiiui Forest Mem-
oirs, Forest ZoologN- .series 1(2):21 {Hok)t\pe. Kathian.
(Ihakrata. U.I'., India; Forest Research Institute, Dehra
Dun). Preoccupied
19921
NOMKXCLATUHAI, CMl WCI'.S IN Pi, ATYl^ODIl) AI! WD SCOI.^TIDAP:
85
SjihdcwtnijH'S tectus Beesoii. 1921. Intliaii P'orestiT 47:514
I ll()I()t^pc^ sex?; Katliiaii, ('Iiakrata, V.\\. India; I'orcst
Hcscarcli Institute. ndiiM \1\\\\. ant i\lk-^.\cif'siiii(nii/iiii/
The .series of SpJiaerotn/pes cfucrci Stehhintj;
in llie Forest Research Institute, D(^lira Diui,
collected h\ Stebbing and otht^^s, does not
include oripnal specimens. H()we\(>r. Steh-
!)inii;'s identification, description, and notes
cleaiK indicate that this name was correctlx
applied to his .series. This material was examined
and compared directK to the holotxpe of
C'lti7inicsiis globulus Stebliing In' me. Both sets
olspeci uKMis clearK represent tfie same species.
Beeson recognized that the name S. g^lobosiis
was preoccupied hv Blandford and proposed
the re{)Iacement name S. tectus for St(^b!)ing's
species. The senior svnon\ni, .S. (jucrci Steb-
bing, lias priority" and is used to designate tliis
species, as indicated aboxe.
Sui'iis niisiituii (Eggers)
Ihliirrlii/iiclius iiiisiiiuii Eij;ijers, 1926, Kiit()in()l()u;i.sclu'
Blattrr22:133 lHolot:\pe. temair: |apan: Urakawa 1 1 loko-
ilate]: U..S. National Museum)
SjiliacrDtnjpcs rinitroveisae Mura\aiiia. ]95(), Iiisrcta
.Matsuiniiraiia 17:fi2 ( Lectotxpt'. tenialc; Daidoniinaini-
\aina. Kotlii pref.. Sliikokiii. |apan; l^S. National
Mnsciini. present designation). Xcw .\iiiu>iii/mii
xMura\ama named Sphacrotnjpcs con-
frovci'sae from six female .specimens mounted
on two pins. Although he refers to a t\pe, a
holotxpe was not marked or labeled In Mura-
\ ama. The^ two specimens mounted on separate
points on one pin are coxered by glue and are
recogni/ed with difficult\. On the other pin, the
third specimen from the top (or the second one
up Irom the bottom) is in the best condition and
is here designated as the lectot)pe of coii-
troiersdc. These specimens were compared
directK to m\ homotApes and other .series of
Siiciis niisi))t(ii in m\ collection and are identical
in all respects. Because oiiK one species is rep-
resented, the name coiitrover.me is placed in
.s\iionym\ under the senior name as indicated
.il)()\'e.
Toitiiciis i)rci i})il()siis (Eggers)
Blnslopliapis hrciipilosiis Eggers, 1929, Entoniologisclii'
Hlatter25:103 (Svnhpcs, 2; [Fnkien] China: Kggers (Col-
lection)
Bl(isiopli(i'j^\i\ khds'uiHHs .\Inra\ania 1959. HrookKn taito-
niologieal Societs. Bulletin 54:75 illolotxpe: .Shillon<j;.
Assam. India: U.S. National Museum). Scust/iioitijiiii/
Blastopha^iis imilti.sctosus Mura\aiiia. 1963, Studies in the
seoKtid fauna of the northern h;ilf oi the Far East.
Shukosh Press. Fukuoka, p. 37 ( Holot\pe. Ceinale: .Vlt.
.Man/a, CJununa prel., |apan: L'.S. .National .Museum).
Xcic stjuotupHii
The female holotype of Bla.stoplui<i^ns inulti-
sctosus .Murayama, m\ topot>pic homotvpes of
B. klidsiainis Muraxama. and mv homotxpes of
B. l)rcvipil()sus Eggers were all compared
directly to one another. Althougli the As.sam
specimens are st)me\vhat larger, all share the
\en short interstrial setae and are here placed
in the same species. This .species is ver\' closel\-
allied to pUiipcrda (Linnaeus) and is distin-
guished with some difhcnlt\' from that species
b)- the .setal characters. It is cmrentK' placed in
the genus Tomicits imd(M- the senior name
hrciipiliisiis as indicated abo\e.
New iN'i'KoDi ctions
Hijhistcs opacus Erichson
Hijlastcs oparua Erich.son. 1S36, .Arehix fiir Natnrgcschichte
2(1):51 (Syntxpes; presumabK' Germaiiv; Berlin
.Museum)
A series of Hi/lasfcs opaciis Erichson was col-
lected near tlie eastern tip of Long Island on
Fishers Island, Suffolk Co., New York, USA, 23
Ma\' 19S9, from an Ips plieromone trap, b\' T
W. Phillips, (circumstances of the collection sug-
gest that this species has established a breeding
population at that site. This species is conunon
throughout the pine belts of Europe and north-
ern Asia and it has become established in pine
plantations in Soutli Africa. While it breeds pri-
mariK in the roots and stumps of pin(^ (Piitiis
spp.) and spruce {Ficcd .spp.), it is known as an
economic p(\st of small .seedlings of these trees.
Plilocosiiiiis (initaliis Heitter
I'hlncDshiiis (innatu.s Heitter, i8S7, Wiener Entomologisehe
Zeitung 6: 1 92 ( I lolotxpe, male; Syrien; Naturhistori.sches
.\Inseinn \\ ien)
Tliis species was recentK foimd to be estab-
lished in Los .Angek's Co.. Califoniia, USA, in a
broad area in sufficient numbers to cause eco-
nomic losses in Cn})ri'ssus spp. It was prexiously
kucmn from (nprus, S\ria, and Israel, where it
is an impoilant pest of (jiprcssiis spp.
New Species
C'l/cloiiiipidioii siiha<iiiatiiiii. n. sj5.
Schedl (1957:100) cited Xylchonts stih-
a^natiis Eggers, nomen nudum. He later
(Schedl 1961:94) expressed the opinion that
86
Great Basin Naturalist
[\'()luine 52
X. suha<^n(dus Eggers, from tlie Philippine
Islands, was actuall\ X. parvus Lea (ol Aus-
tralia), and he published a complete description
of the Philippine series in that article under the
name of X. paiijiis. Later, he (Schedl 1964:314)
saw the t\pe ofX. p(imts\ recognized the differ-
ences in the two taxa, and presented the new
name S. siiha^iiatns Schedl for the Philippine
series. He then (Schedl 1979:239) designated a
"lectot\pe" forX. â– sul)a<j^iuifus Schedl.
Because X. .sitba<^natti.s Eggers was never val-
idated, Schedl s presentation of a new name for
it did not meet the recjuirements of the Code of
Nomenclatin-e e\en though a description exists
for the taxon. This taxon has l)een transferred to
the genus Cijclorhipidion, where it is treated
here.
Ct/clorliipidioii sulxipiatuni is presented here
as a species new to science. The validating
description is published in Schedl (196L94-95)
under the misidentified name Xylebonis parvus
Lea. The female holotype is the specimen
labeled as the "lectot}pe" of Xi/leborus suh-
apiatus Schedl in the Naturhistori.sches
Mu.seum Wien. The tNpe localitv is Mt. Irid,
Luzon, Philippine Islands. Other specimens in
this Schedl series from this localitv in the Wien
Museum are paratxpes.
Dcudrotrupes zcdhnulicus, n. sp.
Tliis s[)ecies is distinguished from cosficeps
Broun, the ouK' other named species in this
genus, by the smaller body size, by the less
strongly impressed male frons that lacks a
median epistomal denticle, and b\ the more
evenlv romidetl el\ tral (k^cli\it\.
MalK. — Length 1.5-1.7 mm, 2.7 times as
long as wide; color brown, eKtra mostK liglit
brown.
Frons broadK, uioderateK- concaxe from
epistoma to slightK' above eyes, deepest at its
center, upper area subrugulo.se and punctured,
lower third more nearh' shining and snbacicu-
late; lateral margins subacute ouK- near antennal
in.sertious, ronndcHl ab()\c>; a finc^ median carina
from center ol conca\it\ to (>pistonial margin,
usually higher on lower third, without a denticle
near epistoma (as seen in co.sticcps). Xestitiu-e
hairlike, ratlier sparse and inconspicuous; not
conspicuousl) longer and more alnmdaut on
margins as in costiceps.
Pronotum 0.9 times as long as wide; similar to
co.sticcps except punctures more shaiply, more
stronglv impres.sed, hairlike setae shorter, less
con.spicuous.
EKtra 1.7 times as long as wide, outline similar
to costiccps: striae 1 slightl), others not
impressed, punctures rather small, round, deep;
interstriae as wide as striae, smooth, shining,
punctures minute, confused, moderately abun-
dant. Declivdt)' gradual, not steep, evenly, rather
narrowlv convex; sculpture as on disc except
interstriae 1-3 each with a row of about six
minute granules; \estiture much less abundant
than in cosficeps . interstrial rows of erect setae
rather slender, each about as long as distance
between rows, groimd cover recumbent, each
seta about half as long as erect setae.
Fe.MALE. — Similar to male except frons
convex, carina less conspicuous.
T^TE MATERIAL. — The male holot)pe, female
allotxpe, and two male paratxpes are from
Rot()nia, New Zealand, Hopk. US 3726-U, C. L.
Masse\. The holotxpe, allot\pe, and parat)pes
are in m\ collection.
Poh/j^raphus fliifsi. n. sp.
The name Spoiif^occnis tliitsi Beeson
( 1941 :387), nomen nudimi, was used b\' Beeson
without a description or designation of t\pe
material, either in the original publication or on
specimens in his collection. Browne (1970:550)
recognized this deficienc\' and attempted to
correct the problem b\- designating a Beeson
specimen as "lectot)pe" and presenting a
description of it. Howe\'er, in order for a lecto-
t\pe to become a primaiy t\pe it must be validly
designated (Code of Nomenclature, 1985, Arti-
cle 74a). In the present case, because
Spo)i<^occrus fliitsi Beeson was a nomen nudum,
a t)pe series did not exist; and because there
were no sviitxpes, a lectotvpe could be not be
\alidl\- designated. Therefore, regardless of the
action h\ Browne (1970:550), Beesons nomen
nudum remained inxalid. The name
S})on<^otarsus is currentK" a s\nomni of Poh/-
^r(ij)lnis\ consequentK; the .species cited as
ihilsi is here transferred to tliat genus (^^bod
19Sfi:56).
I'^or the [)uipose of \alidating this name, Poh/-
oraphus tliitsi is presentetl here as new to sci-
ence. It is allied to P. kainiocliii Nobuchi, from
Bunii'i, but it is distinguished In the much
larger size (4.7-5.S iinn). In the completely
dixided e\e, bv the laigcr pronotal punctures,
b\ the more slcMider eKtral scales, and In the
host.
19921
NOMEXCLATUIiM, CllWClvS l\ Pl.ATVI'ODIl ) M', AND SCOLVPIDAE
87
Browne (1970:550) presents a lull (Icscriptioii
oi P. fJiifsi. Browne's inxalid '"l('et()t\]H'" is lierc
(k'si^natccl as the female liolotxpc ol /' lliitsi.
Except that the tApe loealitN. Xamina Kesene
(Burma) is IneorrectK spelled. Browne's data
are correct; it is in the British Museum (Natural
Histon K The male allotvpe has the lower hall
of the Irons shallowK. almost concaxt'K
impressed on the median third; it hears data
identical to the holotvpe and is in m\ colK^ction.
One female paratspe in m\ collection and 47
parat\pes of both sexes in the Forest Research
Institute bear data identical with that of the
holot\]')e.
TriotcDiiuis pilicon}is. n. sp.
This species is distinguished from zei/ldniciis
Wood, below, h\ the slightK larger size, b\ the
lighter color, bv the coarser pronotal punctures,
l)\ the \er\' large, median horn on the male
\ertex, and bv tlie \en' small mandibular spines
in the male.
Male. — Length 1.5-2.2 nun (female slightK
smaller); 2.5 times as long as wide; color brown.
Frons strongk; trans\'erselv excaxated, feebh'
if at all concaxe between eyes; a veiy large,
dorsoxentralK flattened, median spine on xertex
(this spine often more than twice as long as
scape); surface smooth, shining, glabrous, dorsal
surface of spine strongK' pubescent, the.se setae
ver\' long.
Pronotum ver\' slightly longer than wide, snb-
(|nadrate; surface smooth, shining, punctures
coanse, deep. Vestiture sparse, rather short, \en
long and conspicuous on lateral and antcMior
iiiargins.
Ektra similar to zci/laiiicus exce[)t punctures
slightK' smaller; setae more slender, decli\it\'
more broadlv com ex.
Fe.MALE. — Similar to male except: Irons
weakK-, transversely impressed (stronge-r than
f(Mnale zei/lanicus), moderateK punctuicd:
w ithout spines on vertex or mandibles.
Type M vrEHIAE. — The male holotxpe, female
all()t\]H'. and six jiaratxpes were taken at
Chikalda, Malgahat, C.P.. India, 16-X-193fi
R.R.D. 106, R.C.R. 181, Cage 660. Iroui
EiipJiorhid sp. b\- N. C. Chatterjee; all are
mounted on hvo pins. The holotxpe is the
n[)permost specimen and the allot\pe is the
third specimen downi on the same pin. The
holot\pe, allotxpe, and parat\pes are in ni\ col-
lection. More than 480 non-t\pe specimens
were examined at the Forest Research Institute,
Dehra Dim. Ironi th(^ states of Karnataka,
Madliya Pradesh, and Maliarashtra from
Eiiphorhid spp.
Xi/I chorus iiia^nificiis. n. sp.
This species is distinguished from X spdthi-
peiinis Eichhoff b\ its larger bod\' si/e. In the
much mon^ broadK, less steepiv comex elxtral
declivit); In the nmch less strongK' impressed
eKtral striae, and In other details described
below. It is a unich stouter species than X.
princcp.s Blandlord. In a series of spatliipciinis
from the same localit\ and date, the strial punc-
tures on the disc are mostlv confluent; in iiui'^-
nificiis the\' are mostlv separate.
Female. — Length 5.6 nun (paratspes 5..5-
5.7 nun). 2.3 times as long as wide; color xeiA'
dark browni.
Frons about as in spafliifx-nnis.
Pronotum similar to spathipennis except:
anterior margin less stronglv produced
(.straighter), serrations less well dex'cloped:
discal area smoother, punctures smaller.
Elvtra similar to spathipennis except: form
slightlv stouter, posterior margin more broadK"
rounded; profile ol upper decli\it\' more
strongK', less exeuK' arched; striae nnich less
strongK impres.sed on di.sc, not at all impn^ssc^d
on declixitx ; interstria(^ much more broadK con-
\ex on disc, flat on declix i(\. punctures smaller,
more numerous, more ob.scure and almost
ne\er replaced In* minute granules on declix itv;
declivital interstriae 2 and 4 ne\er with setae (a
few short .setae present in spatJiipennis).
T^TE MATERIAL. — The female holot\pe and
five female paratopes are labeled: lunin [pre-
.sumabK Peru], ()'l-IX-79, S. Poncor, EESC. 5-
80. The holot)pe and paratypes are in m\
collection.
Lite HATE RE Cited
I5l \\ i:n. R. A. U)91. \r\\ s\-nonvmv and taNoiinmic
ciiangc-s in Pacific .ScoKtidac (Coleoptera). Natnrlii.s-
torisclie.s Mn.scnni W'ien, .Annalcs, serie B, 92:87-97.
Bkkson (". E. C. 194L Tlie ecologx' and control of the
forc.st in.sect.s of India and the neigliborino; coniitries.
Pnhlislied 1)\- the anthor, L>ina Dim. 5 + ii + 1007 pp..
20;3fig.s. 36pis.
BuowNK. F. C. 1970. .Some .Scolvtidae and Platvpochcke
(C'oleoptera) in the collection of the British Museum.
journal of Natmal I liston 4:539-583.
SciiKi)!.. K. E. 1957. ScoKtoidea nouveaiix du Congo
Beige. II: Mi.ssion R. 'Ma\iie-K. E. Schedl 1952.
.Annales du .VI usee Royale dn Congo Beige. TerMuen,
serie 8, Sciences Z(X)logiques 56:1-162.
Great Basin Naturalist [Volume 52
. 196]. Fauna of the Fliilippiius, IX. I'liilippiiu- . 1972. New records and species of American
Journal of" Science 9()(l):87-96. Plat\poclidae (Coleoptera). (ireat Ba.sin Natur;ilist
1964. Zur Sviionvniie der Borkeiikaler, W. 31:243-253.
Reichen!)acliia 3(29):30.3^3I 7. . 1984. New generic .sMionvmv and new genera of
. 1979. Die Tvpen der Saninilung Schedl, Faiiiilic Scolytidae (Coleoptera). Great Basin NaturiJist
Scolvtidae (Coleoptera). Kataloge der wissenschait- 44:223-230.
lichen Sannnlungen des Natin-historischen Museums . 1986. A reclassification of tlie genera of
in Wien, Entomologie 3(2). 286 p. Scolvtidae (Coleoptera). Cweat Basin Naturalist Mem-
WOOD. S. L. 1969. New .svnonvnn and records of oirs 10. 126 pp.
Platspodidae and Scolytidae (Coleoptera). Great Basin
Naturalist 29: 1 13-128. Received 6 januanj 1 992
Accepted 24 januanj 1 992
(ircat Basin Naturalist 52i 1 i. 1992. pp. S9-92
NOMENCLATURAL CHANGES IN SCOIATIDAE
AND PLATYPODIDAE (COLEOPTEUA)
StcpllCH I .. \\ 0()(1
.VliSI'KACr. — New s\ii()ii\iii\ in ScoI\ tidac includes C.ii/pluiliis picfdc i Hat/churi;, 1S37) {=Cn/pluilH.s siih(lcj)r(:ssus
Kijgers, 1940), Gnathotnipes lon'^iusculus (Scliedl, 1951 ) {^C^iuilliolrupcs ciliiitus Schedl. 1975). Hiipoilwuvmus cniditus
Wc'Stwood ( = Steph(inodercs coiitniiinis Schanfnss, 1891). In ^lat^p()(lidae tlic new name Plfiti/jiiis ahniptifcr i.s proposed
as a replacement for the jnnior liomonNin Plati/pii.s ahntptits Browne. 1986: t\pe-species designations are proposed for tlio
genns-gronp names Scittopi/'^its Nnnberg, 1966. Pi/<^(Hl(>liiis Nunherg, 1966, Mix<)})i/<ius Nunherg. 1966. Mcs(>i)i/<iiis
Xnnherg, 1966. Asctiis Nunherg, 1958, Stciioplati/piis Strolnne\er. 1914, Pidtiipiiiiis Schedl, 1939, Pltiti/scapiis Scliedl.
1939, Tix'f)tiiplatypit.s Sehedl, 1939, Tcsscroplati/jm.s Seliedl, 1935; pri'\ionsK nnpuhlislied .specific svnonvmv is pre.sented
lor Cmssotarsii.s cxtcnwdentatus (Fairmaire, 1849) [=Dui])iis tahirae Stebbing. 1906), Crossotarsu.s tcnniiuitus (>"liapnis,
IS65 (=Crossot(ir.sus nicohariais Beeson, 1937), Phiti/pits ahditus Schetll, 1936 ( =Phiti/ptis transHus Scliedl, 1978). Phili/piis
nifftsifrons ,Scliedl, 1933 ( =Pl(iti/pits pretio.sn.s .Schedl, 1961 ), Platypus tirio.seii'iis Reicliardt. 1965 i =l'l(ih/pjis silicdli Wood,
1966), Trcpti>phiti/j)us midtipoms Schedl, 1968 (=Platiipus fastiiosus Schedl, 1969).
Kct/ words: Scolijliddc. I'liili/pailitliii-. ('olcoptcni. iioiiicncldtinv.
The following page.s record iteni.s affecting
lion ienclati.n-e in Scolvtidae and Platvpodidae tliat
are pre.senttxl here in order to make tlie changes
a\ iiilahle for the world catalog now in preparation
for these families. Included are three ca,ses of new-
specific sviionvmy in Scol\tidae. In Platypodidae
are (a) one new replacement name for a junior
hoinornm, (b) 10 t\pe-.species designations for
genus-group names, and (c) six new ca.ses of spe-
cific s\iioimn\.
Nkw Synonymy in Scolytidae
C.i'ijjilialtis piccdc (,Kat/el)in-g)
Boslrichus picc<ic Hat/.eburg, 1837, Die Forst-insekten.
Killer 1:163 (S\iit\pes; Oberschlesien nn B;uern: Institut
flir Pflanzen.schntz, Eberswalde)
Cnjplmlus stihdcprcssus Kggers, 1940, Centralblatt liir
(Tcsamte Forstwescn 66:37 (HoIot\pe; Kleinasien
[Ayancik in northern Turkey]; Eggers C-ollection, in
Naturhistorisclu's Mnsemn Wien). New stiitoni/mi/
A Schedl note in his collection indicates that
Cnjj)luilns MilMlcprcs.siis Eggers (from northern
Turke\ ), cited al)o\c. is s\"non\nious with C'.
kiircnzoii Stark (=C. puiictiildtus Eggers) from
the Far East of USSR, and with C. picctic as
identified b)' Reitter. In die absence of known
specimens of kiirciizoii west of Ussuri and of
the occurrence of pircac Ratzel)urg, cited
abcne, throughout Enro])e and northern .\sia, it
appears prudent to follow Reitter and recognize
the Turkish population as piccac. For this
reason, the uixn\t^ sulxleprcsstis is placed in s\n-
onvm\ as indicated abo\'(\
Gitalliotnipcs l()ii<iiiisciiliis (Schedl)
C.iialltofrichiis loii<^iiiscidiis Schedl, 1951, Dnsenia 2: 121
(Ilolots'pe, male; Tierra del Fnego, Via .Monte; Eggers
Collection, Naturhistorisches Mu.seuin Wien)
GiKitliotnipcs rilidtiis Schedl, 1975, Studies on tin-
Neotropical Fauna 10:4 (IIolot\pe, female; Argentin;i,
Nahnel Iluapi National Park: Natin'historisches Miisemu
Wien '. Sctv si/nonipni/
The male holotspe of GiuiHiotricluis
l()ii<j^iusni}iis Scliedl, cited abo\e, and the
female holotxpe of Cudihotnipes ciliatns
Schedl, cited abo\c. were compared directK' to
one another and to other males and females of
this species in the Schedl (Collection and in my
collection, l^ecause distinguishing characters
that iiiight ])e used to .separate species are
absent, it is apparent that onK" one species is
re[)resented b\ this material, 'flic name ciliatns
is placed in sNuoininx in tlie genus
Giiatliotnipes as indicated abo\ e.
.«2I,it'eSci<MUrMi
Bri<4li.im VoMiii; IJniversih. Vm\a. Ll.ili 846(12.
89
90
Great Basin Naturalist
[N'olume 52
Hijpolhcmnnus cnuJifus Weshvood
Hi/potlu-nciniis enidUtis Weshsood, 1S36. Entoinoiogical
Socieh- of" London, Transactions 1:34 (S\iit\pes, female:
England: some in British Museum [Natural Iliston].
London)
StepJuntoderes coiiununis Selianfuss, 1891, Tijdschrift \()or
Entomologie 34:11 (Holotvpe, female; Madagascar;
Scliedl (Collection in Naturhistorisches Mnsenm W'ien).
.Wic sijuoiiijinii
The female holohpe ot StcplunHxIcrcs coin-
miiiiis Schaufuss, cited aboxe, has the head
missincf and most of the body scales have been
nibbed off, bnt there is no doubt whatexer that
it represents a normal female of Hi/pothcncnuis
cniditiis Westw'ood. The holotxpe of coDiniiinis
was examined b)' me and compared directh to
my homot\pes of cniditus. This is the most
common species of ScoHtidae in the world,
although it is often recognized with difficult\; as
in this case. The new synoimtn is indicated
above.
New Name in Platypodidae
Flati/pus ahiiipfifci: n. n.
Vlatiiyiiisdhniptus Browne, UJSfi, Kont\()54:337 ( Ilolotspe,
male; New C^ninea: Adi Island to Nagoxa [[apan],
imported; British Mnsenm [Natural Iliston], London),
preoccupied h\ Sampson 1923
The name Plat i/ pus ahniptiis Browaie, cited
abo\e, is a junior homonym and must be
replaced. The new name, ahniptifcn is pro-
posed as a replacement as indicated aboxe.
Generic Chances in Platypodidae
DoJiopij^us Schedl
D(>li(>i)i/ffts Schedl, 1939, International Congress of Ento-
molog); Procei'dings 7:402-403, t\pe-species: Cr<«.s-
otarsus hohcinani Chapuis, designated by Schedl 1972
Scut(>i)t/ffis Nunherg, 1966, Re\iie de Zoologie et de
Botaniqne Airiciiines 74:1S7-1S8, t\pe-species; C/v«.s-
otarsit.s nipax Sampson, present designation. Nnvsipwiii/im/
Pijgodolim Nunberg, 1966, Revne de Zoologie et de
Botanicjne Africaines 74:1S(S-189, tvpe-species: C'/o.s.s-
otaisus vc<ira)i(lis .SampsoTi, present designation. \cic
sijnonijini/
Mixopt/fitis Ntniberg, 1966, Re\ue de Zoologii- et de
Botani(jue Africaines 74:188, tvpe-species: Crossotarsiis
conmdti Strohniever, present designation. New sipionipiu/
Mvsopiji^ii.s Nnnberg, 1966, Revne de Zoologie et de
Botaniqne Africaines 74:187-188, t\pe-species: Cross-
oiarsu.s ukcrcicccnsis Schedl, present designation. Wir
stjnomjmij
V\)r the genus Doiiopij<ius Schedl, Nunberg
named the four subgenera cited above, without
designating a t)pe-species for them. To remove
this ambiguitv' from these names, a t\pe-species
is designated abo\e for each of them. Because
Doliopi/ffis contains only 142 species and the
di\ersit\' within the genus is only moderate, it is
felt that subgenera in this genus are not needed
at the present time. These Nunberg names are
regarded as .s\iion\nis of D()Uopi/<iiis, as indi-
cated above.
PcrioDinuitiis Chapuis
PcriDiniiKitii.s Chapuis, 1865, Monographic des Platvpides,
p. 42, 316, t\pe-species: Perinmtnntus lon'^icoUis Chapnis,
monobasic
Ascius Nnnberg, 19.58, Acta Zoologica Craco\iensia 2:10,
tvpe-species: Periomnwtus sevcriiii Strohniever, present
designation, ,svmonvm\' bv Schedl 1972
The name Asctiis Nunberg, cited above, was
established and then placed in s\nionvniy under
Perionunaftis as indicated. E\en though it is an
essentialK unused name, in order to remove
ambiguity from citations of it, a t\pe-species
must be designated. This designation is gixen
above.
Plati/piis Herbst
Phitifpus Herbst, 1793, Natnrswstem aller bekannten . . .
Insekten, Der Kiiler .5:128, t\pe-species: Bitstrichus cijl-
iiulnt.s Fabricins, monobasic
Stcnopliitypits Strohmeyer, 1914, Cenera Insectonnn, Fasc.
163:35, tvpe-species: Crossofcirsits sphuilosiis Stroh-
niever. present designation, .sviionvmv bv Schedl 1939
Platyp'mufi Schedl, 1939, International Congress of Ento-
mologv'. Proceedings 7:397, tvpe-species: Platypus ciwtns
("hapnis, present designation, sviionvmv by Wood 1979
Pldti/scaptis Schedl, 1939, International Congress of Ento-
nKilogv, Proceedings 7:397, tvpe-species: Platypus car-
hiulatus Chapuis, present designation, preoccupied by
Hnistache 1921. renamed PJatyscapuJus Schedl 19.57,
s\iion\u)\ b\ Browne 1962
The genus-group names Stenophiti/piis Stroh-
me\'er, Plati/piniis Schedl, and Plati/scapits
Schedl (-PJdtijscdpulus Schedl), cited abo\e,
were named without the designation of a t\pe-
species. To remove this deficienc\" and the con-
secjuent ambiguity" associated with them,
tvpe-species are designated as indicated abo\"e.
All three names are junior s)iion)ms oi Platypus
Uerhst.
Tcsscrorcnis Saunders
Tcsscroccnis Sauntlers, 18.36, Entomologiciil Societv oi
Loudon. Traus;ictions 1:1.55, tvpe-species: Platypus (Tcs-
scroccnts) iuscguis Saunders, monobasic
TcssiToplali/pus Schedl, 19.35, Entomologische Nachricli-
teu 9:149, tvpe-species: Tesseroplatypus ursus Schedl
= Tcsscroccnis iusionis Saunders, present designation.
s\iionvm\' bv Schedl 1972
1992]
NOMENCL.'\Tl'H \l, ClI ANCKS IN SCOLVPIDAK AM) PLVHTODIDAK
91
The o|enus-<i;i"()U[) name Tesscropldiijpus
SeliecU. cited aboxe, was proposed without the
tlesignation of a t\pe-speeies. To reino\'e this
deficiency a t\pe-species is designated as indi-
cated al)o\e. The name was plactxl in s\iion\ ni\
se\eral \ears ago, as indicatcnl.
Trcploplati/pii.s Schedl
Tirplopldli/juis Sc'iiecll. 1939, Inteniatioiiiil Congress of
l'",nt()in()l()'j>. Pmceediiigs 7:401. t\pe-species: Cross-
(il(irsu\ trcjxiiKiliis Chapuis, prcst^nt (k'siiiuatioii
The generic name Trcptoplati/pus Schedf
cited aho\e, was named witliout the designation
of a t\pe-species. To renioxe this deficiency, a
t\pe-species is designated al)o\ e. as indicated.
New Syxoxymy in Platypoimdae
Cro.ss(4(irstis cxtcnicdcutatus (Fainnaire)
Pldli/jiiis cxtcnu'dnitdtiis Eairniaire, 1S49. Rexuc et .\Iag-
asin de Zoologie Pure et Appli(juee, ser 2. 2:78 (Molo-
tApe. male: Taiti: Mu.seiim Nationd d'Histoire Naturelle.
Paris)
l^iapits tdlnrac Stel)bing. 1906. Departiuental notes on
insects tliat affect forestiT (Calcutta), No. 3, p. 418 (Smi-
tvpes; India: Madras Presidencw N. Coimbatore Forests:
Forest Hi'searcli Institute. Delira Dun. Xcic si/iioiit/ini/
The species Diopiis tahirae Stebliing, cited
ah()\e, was described as occurring tliroughout
India in economicafl\' significant numbers.
Reports from 1906 tlirougli 1908 repeat the
original report. It was last mentioned in original
literature in Stebbing 1914 (Indian Forest
Insects, p. 626), where it was transferred to tlie
genus Platijpii.s. There are no specimens under
this name or host {Shorca tdhira) in the Forest
Research Institute, Dehra Dun, nor is the t\pe
localits' represented on an Indian platspodid.
The Stebbing 1914 illustration is of a Cross-
otarsus species, probably cxfcntcdoitdfus
(=saiin(lcrsi). Becau.se so main' of Stebbing's
Platxpodidae in the PT-II (Collection are misiden-
tifications of this species, ialunic is placed in
s\non\niy under cxlcntcdoitatiis, as indicated
aboye, based on the Stebbing illustration in the
absence of other e\idence. The fact that it was
said to be a common, economic species supports
this placement.
Cr()ss()t(irsu>i IcnniiKitiis (Chapuis
Crossotami.s tci-miiuitiis Cliapuis, 1865, Monographie ties
Plat\pides. p. S3 (Holot\pe, male; Singapour; British
Museum [Natural Histor\], London I
Cros.mtaisus nicoharicus Beeson, 1937, Indian Poorest
Records, Entomolog\' 3:86 (S\nt\pes; Nicohars: i'.ixr
Nieohar; fairest Ixestarcli Institute. Delna Dun). XiCW
Sl/IIOIltlllll/
The male hol()t^pe and .se\en parat\pes of
CU'ossotarstis nicoharicus Beeson. cited abo\(\
were compared In- me directly to the Beeson
series of C. vciuistiis (.'hapiiis (=C. terminatiis
Chapuis), cited abo\(\ and tAyo of these to m\
series of C. feniiiiiafits. In the absence ofdistin-
guishing characters, all were considered to rep-
resent the same species. For this reason the
name )iicohariciis is placed in s)non)m); as indi-
cated ab()\(".
Fifiti/pus (ilxlitus Schedl
rlaitjpus (ihflilii.s Scliedl. 1936, Hexiie Fran^iiise
trEutoinologie 2:246 (Holot\pe. male: Naturliistorisches
.Museum W'ien)
Pidtijpiis tmmittis Schedl. 197S. Entomologische
Abhandlimgen Staatliches .Museum tiir Tierkunde in
Dresden 41:309 (Holotvpe. male: Brasilien. Linliares. E.
Santo; Naturliistorisches Musemn W'ien '. \rusiiiutiu/iiu/
Tlie male lioIotApes, cited aboxe, of Platypus
abditus Schedl lukI of F. transitus Schedl were
compared by me directly to one another and to
other representatiyes of this species. Because
distinguishing characters could not be found,
the junior name, transitus, is placed in smioii-
> ni\, as indicated aboxe.
Plat 1/ pus ru<i(isifroiis Scliedl
Platiipiis ni<:^(>sifnuis Sc-hedl. 1933. Ke\istade Entomologia.
Sao Paulo 3: i73 ( I lolotxpe. male; Brazil, S. Paulo. .Ylto da
Serra; Naturliistorisches .Museum W'ien)
Platijpus pniiosHs Scliedl. 1961, Pan-Paciiic Entomologist
37:233 (Holohpe, m;ile; Venezuela, Mt. IDuitla; Califor-
nia .Acadenn of Science. San Francisco). Scic sijnon\jm[i
The male holot\pe of Platypus nt<i^osifro)is
Schedl, cited aboxe, and the male paratope of P.
prctiosus Schedl in the ScIkhII (x)llection were
compared directlx to one anotlKM" and to my
homotspes of this sjiecies. Because ouK one
species appears to be represented In this mate-
rial, the junior name, prctiosus, is placed in
s\ nonx ni\ as indicated aboxc.
Plati/pus tirioscnsis lieicliardt
Pidti/pits tirioscnsis Heichardt. 1965. Papeis .Axiilsos do
i^iepartamento de Zoologia. Secretaria de .Agricultma.
Sao Paulo 17:53 (Ilolotxpe. nnile; Brasil, Estado de Para.
Tirios (.Alto rio Paru d'Ocste: Departamento de Zoologia.
Secretaria da .Agricnltnra. Sao Paulo)
Platt/ptis sclicdii Wood, 1966. Creat Basin Naturalist 26:51
(ilolotxpe, male; .Mauaka. Briti.sh Cniana; British
Museum [Natural Ilistorxj. 1 x)n(lon). .Vcif .s;//i()/i(/;/i(/
Although direct comparisons of h()lof\p(\s
were not made, it is appan^it from published
92
Great Basin Natuhaust
[\'oluine 52
illustrations and from niy examination of the
Seliedl male oi Flafi/pns fihosciisis R(Melianlt,
cited alx)\ (', and of the F. schcdli t\ pe series, that
these names are s\iionvms. Both Reiehardt and
I sent specimens of this species to Schedl in
1964 for comparison to related species. We both
received enconrai^ement from him to name the
species, althon^h snbseqnent events clearly
indicated tliat he was fnIK aware we were both
working witli the same species. The name
schcdli is placed in sviionvniy as indicated
al)o\e.
Trcf)f(>j)hilijj)iis ntulfiporns Schedl
Treptoplatijpns initltiponis Schedl, 196S, Pacilic lust'cts
10:270 (Ilolotvpe. f'rmale; Okapa (kasa), E. lliglilands
District [XewGuincal: CSIHO. Canberra)
Plali/ptis fdstiumis ScliecU. 1969. Linneiui Society of New
South Wales. I'roceedings 94:226 (Holot\pe, male; New
Cuinea: Maral'unpi, 2S00 in: CSIRO, Canherra). \\-w
SljIIOIII/lltll
Schedl named Tn'ptoplati/piis nuilfiponis,
cited above, from the female and Plati/piis
jastiiosns, cited above, from the male. Subse-
quent collecting has demonstrated that these
names represent the opposite sexes cjf the same
species. A note in his collection indicates that
Schedl was aware of this problem. Both holo-
t\pes, as well as additional material, were exam-
ined. The jnnior name, fastiiosiis, is placed in
.s\-nom ni\ as indicated above.
Received 3 March 1992
Accepted 13 March 1992
Cicat Basin Xatnialist 52(1 ). 1992. p. 9o
BOOK REVIEW
Plant biolog\ of the Basin and Range. C. B.
Osmond, L. F. Fitelka, and (;. M. IIid\.
Springer-\erlag. BtM-lin, 1990. 375 p]).
$69.50.
This iiitriiLj;uiii<j; xolnint' will Ix' ol intcre.st to
man\ people for a \ari('h ol rea.son.s. It was
w litten to honor W. Dwight Billings, who began
his (hstingnishetl career in what is now called
ph\ siological ecolog\ at the UnixersitA of
Ne\'ada at Reno. Althongh he nuned to Dnke
Uni\ersit\' in 1952, liis heart, and considerable
research, remained in Ne\ada. Twent\-se\tMi
anthors contrihnted the nine chapters of the
hook. While tliat is generalK' enongh to make
one mow on to something else, in this case it
would he a mistake. Althongh the hook was not
w hat I ("xpected, I was pleasantK" sniprised. The
chapters are \en nnexen and range from the
hroad and general to the narrow and higliK
technical. The contributors are first rate and the
chapters well written. I suggest tliat the reader
browse, first reading whate\er appeals and then
perhaps returning to some of the other areas.
The strangest chapter in the book is the first
one. It is a nice introduction but in spite of its
title is neither about anthropologv or botauN.
The cKriamics of climate in the Basin is the
subject of tlieuext chaptt^r. Bri(4 but inttM-esting,
it is clearK written for the nonclimatolo'^ist. The
heart of the book is the 4()-page chapter b\
Billings himselt on mountain forests of North
.\nierica. It clearK extends beyond the Great
Basin but should be required reading of e\ er\
stnck^it of plant ecologw Here is the master
'jjixinsj; us the distilled wisdom of decades ol
research and thinkiufj;. We then moxc on to
hi<j;h-ele\ation forests in an excellent cha[)ter on
the difficult problems imposed on li\ing thin*j;s
In the harsh conditions associated with altitude.
There are high mountains not only surromiding
but running tlnough the Basin in a north-south
direction. Edaphic fac-tors and their influence
on water and nutri(^nt a\ailabilit\ and sub.se-
(juent [)]ant distribution aic next considered.
Tliere are islands of xcrx disjunct .soils thr()n<j;h-
ont the Basin.
Chapter 6 examines wiiat uiost of us think ol
in the (Ticat liasin — the lowland plants. The
emphasis is on ecopln siologw and broad pat-
terns are the theme. Maitxn Caldwell and his
co-workers ha\e spent man\ \ears stuck inti; tlu^
root ,s\ stems of desert plants. This sunuuan of
their work is well worth careful stuck. 1 lowexer.
I was sniprised to find onk a on-son- mention
of the role of mvcorrhi/ae. (^ha])ter 8 deals with
isotopes and \egetation changes. That sounds
narrcjw and well focused but the chapter was
not. It is an oxeniew of the potential use of
carbon isotopes in pli\ siological ecologx. The
last chapter deals briefK with climatic change in
the (.reat Basin. The ])ast has been \\'\^^^
cKnamic and exciting. WTiat ma\ wc expcx't in
the future?
Whilc^ I was disappointed l)\ some ol the
things the title seemed to promise and did not
deliver, I did like the book and recommend it
liighlv. As in man\' boc:)ks with contributes! cha[)-
ters, thc^ lack of continnitx or transition between
chapters left an oxerall im])ression ol a dis-
jointed and uncncn apjiroach. in spite of this,
we can be grateful forwiiat was ck-lixcred: wcll-
w ritten text that was fa.scinating and stimulating
in placets, nicc^ illustrations. <j;ood index. Pin sio-
logical ecologists interested in the (.reat Basin
should spend some time with this xolume.
Bruce \. Smith
Department olliotaiix and Hange Science
Brigham Young l'ni\c'rsit\
Prcno, Utah <S46()2
93
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agement 44: 695-700.
Sousa, W. P. 1985. Disturbance and patch dynam-
ics on rocky intertidal shores. Pages 101-124
in S. T. A. Pickett and P. S. White, eds.. The
ecologx of natural disturbance and patch dy-
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(ISSN 0017-3614)
GREAT BASIN NATURALIST voi 52 no i March 1992
CONTENTS
Articles
In memoriam— A Perry Plummer (1911-1991): teacher, naturalist, range
scientist E. Durant McArthur
Secondary production estimates of benthic insects in three cold desert streams
W. L. Gaines, C. E. Gushing, and S. D. Smith
Effect of rearing method on chukar survival Bartel T. Slaugh,
Jerran T. Flinders, Jay A. Roberson, and N. Paul Johnston
DNA extraction from preserved trout tissues D. K. Shiozawa,
J. Kudo, R. P Evans, S. R. Woodward, and R. N. Williams
Relating soil chemistry and plant relationships in wooded draws of the north-
ern Great Plains Marguerite E. Voorhees and Daniel W. Uresk
The genus Aristida (Gramineae) in California Kelly W. Allred
Temperature-mediated changes in seed dormancy and light requirement for
Penstemon palmeri (Scrophulariaceae)
Stanley G. Kitchen and Susan E. Meyer
Late Quaternary arthropods from the Colorado Plateau, Arizona and Utah
Scott A. Elias, Jim I. Mead, and Larry D. Agenbroad
Microhabitat selection by the johnny darter, Etheostoma nigrum Rafinesque, in
a Wyoming stream Robert A. Leidy
Nomenclatural innovations in Intermountain Rosidae Arthur Gronquist
Nomenclatural changes and new species in Platypodidae and Scolytidae
(Goleoptera), part II Stephen L. Wood
Nomenclatural changes in Scolytidae and Platypodidae (Goleoptera)
Stephen L. Wood
Book Review
Plant biology of the Basin and Range C. B. Osmond, L. F. Pitelka, and G. M. Hidy
Bruce N. Smith
H E
MCZ
LfSRARY
OCi 1 4 1992
HA-^VARD
Ll^JiVL.F^;oli^Y
GREAT BASIN
NMRALIST
VOLUME 52 NO 2 - JUNE 1992
BRIGHAM YOUNG UNIVERSITY
GREAT BASIN NATURALIST
Editor
James R. Barnes
290 MLBM
Brigham Young University
Provo, Utah 84602
Associate Editors
MiciiAi-'-LA. Bowers
Blandy Experimental Farm, University of
Virginia, Box 175, Boyce, Virginia 22620
J. R. Caliahan
Museum of Southwestern Biolog)', University of
New Mexico, Albuquerque, New Mexico
Mailing address: Box 3140, Hemet, California
92.546
Jeanne C. Chambers
USDA Forest Service Research, 860 North 12th
East, Logan, Utah 84322-8000
Jeffrey R. Johansen
Depiirtment of Biology, John Carroll University,
University Heights, Ohio 44118
Paul C. Marsh
Center for Environmental Studies, Arizona State
University, Tempe, Arizona 85287
Brian A. Maurer
Depiirtment of Zoology, Brigham Young University,
Provo, Utah 84602
JiMMIE R. PaRRISH
BIO-WEST, Inc., 1063 West 1400 North, Logan,
Utah 84321
Paul T. Tueller
Department of Range, Wildlife, and Forestry,
University of Nevada-Reno, 1000 Valley Road,
Reno, Nevada 89512
Editorial Board. Richard W. Baumann, Chairman, Zoology; H. Diiane Smith, Zoology; Clayton M.
White, Zoology; Jerran T. Flinders, Botany and Range Science; William Hess, Botany and Range
Science. All are at Brigham Young University. Ex Officio Editorial Board members include Clayton S.
Huber, Dean, College of Biologiciil and Agricultural Sciences; Norman A. Daniis, Director, University
Publications; James R. Barnes, Editor, Great Basin Naturalist.
The Great Basin Naturalist, founded in 1939, is published quarterly by Brigham Young University.
Unpublished manuscripts that further our biological understanding of the Great Basin and surrounding
areas in western North America are accepted for pubhcation.
Subscriptions. Annual subscriptions to the Great Basin Naturalist for 1992 are $25 for individual
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directed to the Editor, Great Basin Naturalist, 290 MLBM, Brigham Young University, Provo, UT
84602. ^ ^ ^
Scholarly Exchanges. Libraries or other organizations interested in obtaining the Great Basin
Naturalist through a continuing exchange of scliolarly publications should contact die Exchange
Librarian, Harold B. Lee Library, Brigham Young Universitv, Provo, UT 84602.
Editorial Production Staff
JoAnne Abel Technical Editor
Carolyn Backman Assistant to the Editor
Natalie Miles Production Assistant
Copyright © 1992 by Brigham Young University
Official publication date: 22 September 1992
ISSN 0017-3614
9-92 7501162
The Great Basin Naturalist
PUBLISIIKD ATPKOX'O, UTAII, HV
Brigham Younx; Um\'krsit\
ISSN 0017-3614
Volume 52
June 1992
No. 2
Great Basin Natunilist 52(2), pp. 9.5-121
RED BUTTE CANYON RESEARCH NATURAL AREA:
HISTORY, FLORA, CEOLOCY, CLIMATE, AND ECOLOCY
James R. Ehleriiii^er , Lois A. Amovv , Ted Aniow",
Ining B. McNulh , and Norman C. Negus
AliSTlUCr — Red Butte Canyon is a protected, near pristine ctmyon entering S;ilt Lake Valley, Utdi. It contains a
\\ell-de\'eloped riparian zone iuid a perennial stream; hillside vegetation r;uiges from grasslands on the lower limits to
Douglas-fir and aspen stands at the upper ele\ations. In this paper we describe the histon,' of human impact, natural histon
aspects of climate, geologv', and ecolog\', and faun;il and floral information for kev species in the canvon. The role and
importance of Research Natural Areas is di.scus.sed, particularly with respect to the need to protect Reel Butte Can\()n — one
of the few remaining undisturbed riparian ecosystems in the Intermountain West.
Ki'tj words: <ir(i.ssl(in(l. Iiitcnnoinitaiii West, onk-tiuiplc. plant (uhiptiition. Red Butte Caiiijoii. Researeh yatund Area,
rijxniiin eenlof^i/.
Red Butte Canyon, one of many canyons in
the Wasatch Range of Utah, opens westward
into Salt Lake Valle\, immediately east of the
Uni\ersit)' of Utah (Fig. 1). Like most canyons
along the Wasatch Front, it is a grassland at the
lowest elevations, is forested at its upper end,
and has a perennial stream. What makes this
canyon unusual is its history. The canyon was the
watershed for Fort Douglas, the U.S. Arnnpost
built in 1862 that oyerlooked Salt Lake Cit\'. As
a protected watershed, these lands were, for the
mo.st part, kept free from grazing, hirming, and
other human-impact actixities. When the U.S.
Army declared the.se lands suq:)lus in 1969, the
U.S. Forest Serxice assumed responsibilit)' for
the canyon. Since that time, Red Butte Canyon
has been kept in its protected state and desig-
nated a Research Natural Area (RNA).
The Research Natural Area designation
denotes an area that has been set aside because
it contains unusual or unique features of sub-
stantial yalue to society. These might include
unique geological features, endangered plant
and animal species, or areas of particular \alue
for scientific research as ba.seline bench marks
of ecosystems that haye been largely destnned
by human impact. In the case of Red Butte
Canyon, the RNA designation was given
because this canyon is one of the few reniiiining
(if not the last) undisturbed watersheds in the
Great Basin. The U.S. Forest Service report
proposing that Red Butte C>anyon be declared a
Research Natural Area described the can\-on as
". . . a hviu"; nuiseum and biological libraiv of a
size that exists nowhere else in the Great Basin
... an invaluable bench mark in ecological
time." The Red Butte Canyon RNA is unique
becau.se it is a relativeK' undisturbed watershed
adjacent to a major metropolitan area (Salt Lake
\ alley). To protect this \aluable re.source, access
to the Red Butte Canyon RNA has been largely
restricted to .scientific investigators. One of the
,Depiirtnieiit iit Binlcirx . Uiiiveisitv of L't.ili. S.ill l„ike- Cil\. Ut.ili S41 12.
"CoiiMilting irt-nloyisl, 1064 E. HilNneu Dnve , Salt L.ifce (iin. Ut.ili S4124.
95
96
Great Basin Naturalist
[Volume 52
Salt Lake City ^^^^^^^^^^'
Intl. Airport y////////.
lie f//////////////////////////////y///////y//////y/yy/y////
Pinecrest
CO
o
CD
CD
i.:Ia».^ A»Ar^ f//y/y///////y/yy//yyyyyyyyy/yyyyyyyyyyyyyyyyyyyyyyyyyyy k Aill PrPGK ^' '/
— I Kilometers ///,/, ///fy/y/,y,yy/yyy/yy//y///yy////y/y/////''/y//////' Mil' ^"
Fig. L Ijocation of Red Butte Ciinvon and other sites referred to in text.
goals of the RNA Program is to protect and
preserve a representative array of all significant
natural ecosystems and their inherent processes
as baseline areas. A second goal is to conduct
research on ecological processes in these areas
to learn more about the functioning of natural
versus manipulated or disturbed ecosystems.
Research activities in the Red Butte Canyon
RNA are directed at both of these goals: under-
standing basic ecological processes (physiologi-
cal adaptation, drought adaptation, nutrient
c\'cling, etc.) and also the impact of humans on
our canyons through both airborne (air pollu-
tion, acid rain, etc.) and land-related (grazing,
human traffic, etc.) activities. The latter are
conducted through comparison of Red Butte
with other canyons along the Wasatch Range.
In size. Red Butte Canyon is relatix elv small
compared with other drainages along the
Wasatch Front. The drainage basin covers an
area of approximately 20.8 km" (5140 acres)
(Fig. 2). The drainage arises on the east from a
minor divide betvveen City Creek and Emigra-
tion canyons and drains to the west. The canyon
has two main forks (Knowltons and Parleys) and
many side canyons. Near the canvon base, a
resen-oir was constructed earlier this century to
prcAide a more stable water supply to Fort
Douglas. The diversity of slope and aspect com-
binations of the terrain contributes to a variet)'
of biotic commimities along an elevation gradi-
ent from about 1530 m (5020 ft) on the west end
to more than 2510 m (8235 ft) at the crest.
The puipose of this paper is to provide a brief
description of the histoiy, flora, geology, cli-
mate, and ecology of this unusual and valuable
resource. There is increasing interest in Red
Butte Canyon, in part by scientific investigators
because of its utility as a protected, undisturbed
watershed, and in part by curious citizens from
the nearby Siilt Lake Valley. Yet, there has not
been an overall reference available for those
interested in general features of the canyon or
past ecological studies within the canyon. Most
of the information on Red Butte Canyon is
scattered. With the closure of Fort Douglas in
1 99 1 , many of the historical records will become
more difficult to access. It is hoped that the
synthesis presented in this paper will provide
the necessary background for those interested
in the histoiy and ecologv of the Red Butte
Canyon RNA. Irving McNulty first summarizes
the history of the canyon, followed by Ted
Amow's description of geologv' and soils. James
Ehleringer contributed the h)'drology, climate,
and plant ecology sections. The section on vas-
cular flora was prepared by Lois Amow, and
Norman Negus wrote the mammalian and avian
fauna sections.
19921
Rkd Butte Canyon Heskahch Natural Area
97
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\0
Q
i^
<$>.
/O
.x^P^
'Vo-
^0'
1^^
^&-
.^^
\^-®
a^
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Q
,^
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9.e'
se^
^o^^
^vgS
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,Q Q
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mile
kilometers
Fig. 2 Major drainages and weather and bench mark stations within the Red Butte Canyon Research Natinal Area. B
represents the location of the USGS Bench Mark station; circles numbers 2, 4, and fi represent the locations ot weather
stations known as Red Butte #2, Red Butte #4, and Red Butte #6, respecti\el\ .
History
The historv' oi Red Butte Can>oii comes as
bits and pieces from many sources, including
Arrington and Alexander (1965), Hibbard ( 1980),
and the Fort Douglas Army Engineers Office
(1954), records of the Fort Douglas Museum,
and discussions with C. G. Hibbard (Fort Doug-
las historian) and Harold Shore (Fort Douglas
water master oxerseeing Red Butte Canvon). It
is primariK' a hist(nA' of human iiupact on the
utilization of natural resources provided bv the
canyon. Major resources were water from the
stream and sandstone quarried for use in con-
struction. Of minor importance were grazing
and timber In 1848, just one year after the
arrival of the first pioneers in Salt Lake Vallev,
red sandstone was first quarried in the canyon
to be used in construction in tlie building ot Salt
Lake Git)'. It was the closest source of construc-
tion-quaHt)' sandstone and was quarried for
almost 100 years. This mining had considerable
impact on the plant and animal life in the lower
portion of the canyon. The major use of Red
Butte Greek water was by the U.S. Army at Fort
Douglas, which was establish(nl at the mouth of
the canvon in 1862. This utilization of water
outside the canxon had little effect on the canyon
itself, as U.S. Army administrators worked over
many years to protect the watershed and water
qualit)'. In fact, protection has grown steadily
since Fort Douglas was first established, and
particularly since the canyon was acquired by
the U.S. Forest Service in 1969 and declared to
be a Research Natural Area.
98
Great Basin Naturalist
[Volume 52
A
M(jb
Mdo
Mdo
Township IN. Range IE
23
see I ion
22
) kilometers
Fig. 3 Geologic map of Red Butte Canyon Researcli Natural Area. See Table 1 for a de.sc ription of abbreviations. Solid
lines represent contacts l)ehveen torniations, dashed lines represent norniiil faults, and T-ditsJied lines represent the Black
Mountain thnist fault. The transect A-A' is shown in cross section in Figure 4. Adapted from Marsell and Threet {I960)
and Van Honi and CMttenden (19S7).
Red Butte saudstoue (Nuggett Sandstone)
was the first resource utilized from the canyon.
Most sandstone was obtained from Quarr>-
Canyon on the south side of the canyon, 4.4 km
(2.9 mi) from the mouth of the canyon. Because
of the proximit\- of Quarr)' Canyon to Salt Lake
C]it\', sandstone was (juarried there from 1848 to
the end of the centur\- by private companies and
intermittently by the Army until 1940. This
required a road in the bottom of the canyon and
housing for workers. In 1889, 66 men and 38
oxen and horses lived at the canyon bottom,
contributing considerable downstream pollu-
tion to Red Butte Creek. In 1887 the U.S. Con-
gress provided a railroad right-of-way to be built
to the rock (|uarr\' to increase the amount of
sandstone removed. Stream pollution caused by
quarrying activity' brought many complaints
from Fort Douglas and ultimately a court action
in 1889, which required the Salt Lake Rock
Company to control stream pollution and cease
housing men and animals in the canyon.
Red Butte Creek was used for irrigation by
a few pioneers east of Salt Lake City in the early
185()s. When Fort Douglas was established in
1862, Armv personnel initially depended mostly
on water from nearb\' springs. However, by 1875
Armv personnel constnicted two reseivoirs east
of Fort Douglas and diverted water from Red
Butte Creek to fill them. In response to the
recurrent stream pollution problems caused by
quarrying activities, the Territorv' District Court,
in 1890, declared that the waters of Red Butte
Creek were the sole propeiiy of the U.S. Army
and under the jurisdiction of Fort Douglas. Also
in 1890, the U.S. Congress passed a law to
1992]
Red Butte Canyon Research Natural Area
99
Tablk 1. Description of geological formations in Red
Butte Canvon.
Cenozoic era, Quatemarv .s\steni. Holocene scries
fa Fldod-jilriin iillin iiiiiL Sand. eohhK to silt\, dark gra\' at
top: grading ilownuaril to medium to liglit gra\, sand\' to
cohbK' gra\el; kxalK bouldeiA .
fc En^iiwered fill. Selected earth material that has been
eniplaced and compacted.
Cenozoic era, Quateman^ and Tertiary systems,
Holocene and Pleistocene series
/g Allurial-fdii deposits. Boulder\' to claye\' silt, tlark gra\' to
brown; rocks angular to subrounded.
Id Landslide deposits. Composition similar to material
npslope.
Mesozoic era, Jurassic system
Jtc Tain Creek Liiiwstoite. BrtnMiish gra\' ;uid pale gra\ to
pale yellowish grav silt\- limestone, intercalated with
greenish gray shale.
Mesozoic era, Jurassic? and Triassic? systems
JTii Su^et Saitdstone. Pale pinkish buff, fine- to medium-
grained, well-sorted Siuidstone that weathers or;uige-
brown. Massive outcrops form the ridge c;illed Red Butte.
Mesozoic era, Triassic system
Tail Ankareh Formation, upper member Reddish brown,
reddish puqole, grayish red, or bright red shale, siltstone,
and sandstone.
Ta<^Ankareli Formation. Gatira Grit Member White to pale
purple, thick-bedded, crossbedded, pebbK' quartzite.
Forms a prominent wjiite ledge for long distances.
Tatn Ankareh Format i(n}. Mahogany Meml)er Reddish
brovyn, reddish purple, gravish red, or bright red sh;ile,
siltstone, ;ind sandstone.
Tt Thai/nes Formation. Medium to light gray, fossiliferous,
locall) nodular limestone, limy siltstone, and sandstone.
Tw Wood.side Shale. Cravish red, grayish purple, or liright
red shale and siltstone.
Paleozoic era, Permian system
!'))(â– Park City Formation and related strata. Fossiliferous
sandy limestone, calcareous sandstone, iuid a medial
phosphatic shale tongue.
Paleozoic era, Pennsylvanian system
Ptv Weber Quartzite. PiJe tan to nearly white, fine- to
uiedium-gr;uned, crossbedded (juartzite and medium
gray to pale gray limestone.
Pn Round Valley Limestone. Pale gra\- limestone with pale
gray siltstone partings. Contiiins pale pinkish chert that
forms irregular nodules.
Paleozoic era, Mississippian system
Mdo Doughnut Formation. Medium gray thin-bedded
limestone with pods of dark gra\ to black chert and
abundant brachiopods and brvozoa.
A/g/; Great Blue Formation. Thick-bedded. localK clilT-
forming, pale gray, fine-grained limestone.
A//( Hnmbuo Formation. Alternating, tan-weathering. lim\
sandstone and limestone or dolomite.
Md Deseret Limestone. Thick ledges of dolomite and lime-
stone v\ith moderately abundant lenses and pods of dark
chert.
Paleozoic era
P Paleozoic rocks, undifferentiated.
protect the water .suppK oi P^ort Dougla.s. This
hiw prevented any .sale of land in the eanxon or
fnrther watershed development. In 1906 the
U.S. Army built a dam on Red Butte Creek to
-suppK- additional water for Fort Dougla.s. The
present dam was constructed between 192S and
1930, and the reservoir provided water ibr Fort
Douglas until its closure in 1991.
There are no grazing records available lor
Red Butte Canyon prior to 1909, by which time
the United States had acquired title to most of
the land in the canyon. Cottam and Evans
(1945) reported evidence of some gulK' erosion
occurring in the canyon prior to 1909 and
assumed it was due to overgrazing. Although we
lack quantitative data, there are a few isolated
incidents indicating the occurrence of grazing,
including an 1 854 report of a young man drowii-
ing in a flash flood in Red Butte Canyon while
herding animals. Over forty head of oxen used
to haul sandstone from the quarrv in the late
1800s reiuained in the canyon during that time.
In 1869 the War Department appointed a
herder to control loose cattle gnizing on Fort
Douglas and in the canyon. In 1890 three squat-
ters had settled into the canyon, and their forty-
head of cattle were grazing in the Parleys Fork
area before being evicted. B\' 1909 the Armv
had built a gate at the mouth of the canvon to
control access, thus further protecting the
watershed. Although this did not prevent occa-
sional animals from wandering into the canvon
from adjacent canyons, it did reduce both their
numbers and their length of stav. Consequentlv,
most of the canyon has not been grazed b\ cattle
or sheep through most of this centur\.
Portions of the upper reaches of the can\on
were timbered. In 1848, when a road was built
along the canyon bottom, it was reported that
there was an abundance of timber suitabk^ for
fence poles. Later The CJhmch of Jesus C'htist
of Latter-day Saints built a bowen on Temple
S(|uare in downtown Salt Lake ('its' in the ISoOs
with wood obtaiiu^l from Table .Moinui
(between Knowltons Fork and Beaver C>an\()n).
In 1863 the Arm\ constructed 34 buildings at
Fort Douglas from "timber hauled from the
canyons," but there is no indication as to how
much timber came from Red Butte Canyon.
However, apparently not manv timber-size trees
were available in the lower canyon as indicated
by a pioneer who built a log cabin in the canyon.
He stated he had to tra\el five miles up the
100
Ghkat Basin Naturalist
[Volume 52
canvon to obtain enough logs iov the cabin in the
early 1860s.
There are no available records of fires that
niav liave occurred in the canvon. In 1988 a fire
from Emigration Canyon spread into the upper
headwaters of Red Butte Creek before it was
contained. The land was subsequently reseeded
with native species bvthe U.S. Forest Service.
Land ownership within the canyon changed
several times during the late 1800s and early
1900s. Land occupied by Fort Douglas in 1862
was officialK' given to the U.S. Army in 1867
when President Johnson withdrew four square
miles from public domain for the use of the
Anny. However, this included only a small por-
tion of the mouth of Red Butte Canyon. The Salt
Lake Rock Company, which quarried most of
the sandstone in the canyon, owned part of the
canyon, and the Union Pacific Railroad Co.
acquired four sections in the lower portions of
the canyon in the 1860s. Smaller portions of the
canyon were claimed by private indi\iduals
under the Homestead Act of 1862. Such ckiims
could be acquired easilv under this act, which
was veiT liberal and required onl\' a small claim
fee. Graduall)', between 1884 and 1909, through
a combination of acts of Congress, exchanges of
property, and outright purchases, Fort Douglas
obtained title to most of the canyon b-\' 1896 and
almost the entire canyon by 1909. Only three
small parcels of a total of less than 90 hectares
(—200 acres) are still privately o\Aiied today, and
these are close to the margins of the canyon. In
1969 the U.S. Department of Defense relin-
(juished ownership of Red Butte Canyon. The
U.S. Forest Service is now responsible for these
lands. The Forest Service recognized the natu-
ral state of the area had been preseived through
many years of closure to the public and desig-
nated Red Butte Canyon a Research Natural
Area in 1970. By definition such areas are tracts
of land that liave not been stronglv impacted b\'
human-related activities such as logging or graz-
ing by domestic livestock. Tl un are permanently
protected from devastation by humans so they
may serve as reference areas for research and
education.
Red Butte Can\'on has sened as a research
site for biologists for over fifty years and w ill
continue to do so in the future. Public education
about conservation and the need for the public
to better understand the importance of
Research Natural Areas are major concerns.
Recently the Forest Service briefly opened the
canyon to the general public. In 1987 the canyon
was opened to the public in late spring for
several days; this weekend opening attracted
over 5000 visitors and led to a trampling on
vegetation along the main road in the canyon.
This opening was repeated in 1988 and
attracted 1100 people. Currently the State
Arboretum at the University of Uttili conducts
natural history education classes (—10 individu-
als per group) in the lower portions of the
canyon. Limited deer hunting has been permit-
ted by the Forest Service each fall, but the
impact of the hunts is unknown. A Red Butte
Steering Committee, consisting of representa-
tives from the Forest Service, the University of
Utali, and other government agencies con-
cerned with preservation of natural areas, is
involved in making decisions pertinent to the
jurisdiction and management of the Red Butte
Canvon Research Natural Area.
The histoid of Red Butte Canyon, with the
exception of the quari-)ing acti\it\' and some
grazing in the past century, is largely a histon" of
preservation. The U.S. Army at Fort Douglas
was concerned with the protection of the water-
shed and gradually acquired sufficient control
to protect it. The U.S. Forest Service declared
the entire canyon a Research Natural Area and
thus insured its protection for the future as a
bench mark of riparian and shrub ecosystems in
the Intermountain West.
Geology
The rocks underl)ing Red Butte Canyon
range in age from recent Holocene deposits of
our time to Mississippian rocks that are about
360 million years old. Holocene and Pleistocene
deposits are unconsolidated, consisting mostly
of landslides or alluvium deposited by existing
streams. Their aerial distribution is shovvai in
Figiu'e 3, and a description of the deposits is
given in Table 1.
The older rocks range in age from
Mississippian to )urassic, a span of about 220
million vears. The)' are all consolidated now, but
originallv they were formed as deposits in
oceans or inland seas or as sand dunes in an arid
environment. No rocks representing the
approximatelv 140 million vears between the
end of Jurassic time and the Holocene are pres-
ent in Red Butte CJanyon. Either they were
never deposited or they have been eroded.
The consolidated rocks in most parts of the
lower walls of the canyon consist chiefly of shale,
1992]
Red Butte Canyon Researchi Natural Area
2500 -
2000 -
1500 -
1000 -
meters
Fig. 4. Geologic cross section of Red Butte Canvon. Explanation as in Figure 3. Adapted from Van Horn and Crittendei
(1987).
with some gritt)' (juartzite and sandstone. The
upper southeast-facing slopes consist mostly of
limestone with some sandstone and limy shale.
Tlie upper northwest-facing slopes are made up
mostK' of sandstone with limestone and limy
shale near the southeast divide. Figure 3 shows
the distribution of the rocks in the canyon, and
they are described in Table 1.
The older consolidated rocks in the canyon
generally dip toward the southeast (Fig. 4), and
they form the northern flank of a large s\iicline
whose axis trends toward the northeast and
whose southern flank is in Mill Creek Canyon,
about 6.5 km to the south. The rocks are cut by
numerous normal faults that are part of the
\\asatch fault zone, a lengthy fault zone that
bounds the west face of the Wasatch Range for
ahnost its entire length. Movement along these
normal faults has resulted in horizontal dis-
placement of the rock formations, whereas
nio\'ement along the Black Mountain thrust
fault in the northwestern part of the canyon has
raised older rocks to a position o\erl\ing yovm-
ger rocks. The faults and their effects on the
consolidated rocks are shown in Figures 3 and 4.
Soils
bedrock. The distribution of the soils in the
canyon is shown in Figure 5. The relationship of
the soils to the bedrock is apparent by compar-
ing Figure 5 with Figure 3, a geologic map of
the canyon. The soils map (Fig. 5) was adapted
from Woodward et al. (1974). Soils in Red Butte
Canyon have been characterized as dominantly
strongly sloping to ver)' steep and well drained.
According to Bond ( 1979), most soils are neutnil
to sliglitK basic, xarv' in color from brick red to
dark browni, with textures generalK- ranging
from sandy to loamy clays. Depth of the soil is
irregular, with depth to bedrock varying from
nearly 2.4 m (94 in) at the canyon floor near the
mouth to as little as 60 cm (24 in) or less on the
slopes. Soil tvpes include loams, silt loams, and
dry loams. There is little profile development,
but a pronoimced litter layer and appreciable
incorporated humus exist in places. CJeneralh'
the soils are approximately 1 m (39 in) deep,
especially those adjacent to streams. However,
the steep, rocky upper slopes have shallow and
cobbl\- soils. Table 2 includes a description of
each of the soils shown in Figure 5. The descrip-
tions were ackpted from Woodwiuxl et al. ( 1974).
Hydrology and Nutrient Flow
Soils in Red Butte Canyon are derived from Red Butte Creek is a perennial third-order
the weathering and erosion of the underKing stream without upstream regulation or dixersion
102
Great Basin Naturalist
[Volume 52
Township IN. Range IE
23
suction
22
kilometers
Fig. 5. Soils map of Red Butte Canvon. See Table 2 for a description of abbre\iations. Adapted from Woodward et ;J.
(1974).
vintil flow is collected in the reservoir located
near the base of the canyon. The stream has
creatcnl a narrow-based canvon with sides rising
abniptly at an average slope of about 35 degrees
to the north and about 40 degrees to the south.
Immediately upstream of the reserxoir is a U.S.
Geological Survey Hvdrologic Bench Mark Sta-
tion. This gaging station has been maintained b\
the U.S. Geological Survey since October 1963.
Priortothat, the Corps of Engineers, U.S. Armv,
recorded monthly discharge at this location
beginning in Januarv 1942.
The average monthly discharge (1964-88) is
0.133 mVsec (~4.7 ftVsec) as it enters the res-
en'oir at 1646 m (5400 ft) elevation (U.S. Geo-
logical Suivcy records). The stream flow
exliibits a straightforward annual pattern, char-
acteristic of this geographic region — high spring
flows driven by snowmelt followed by very
much reduced flows derived from groundwater
throughout the remainder of the vear (Fig. 6).
Spring melt flow, which is t\pically an order of
magnitude greater than other periods of the
year, peaks in Ma\- and persists for 6-8 weeks.
The average monthlv stream flow rate during
May is 0.416 mVsec (14.7 ftVsec). By Septem-
ber, the lowest average monthly flow rate,
stream discharge has decreased to 0.058 mVsec
(2.0 ftVsec). Mean stream flow rates do not
increase durino; the summer months, althouo:h
nearly one-fourth of the annual precipitation
falls during this period.
Average monthly stream flow \alues, how-
ever, hide much of the stream dynamics and
resultant impact on riparian vegetation. On a
daily basis, stream flows can vary tremendously
1992]
Red Butte Canyon Research Natiral Area
103
Tablk 2. Description ol units on the soils map ol Red
Butte C]an\on.
AGG Agassiz association, ver\ steep. 40-7U percent
slopes; nioderateK permeable, well drained. Agassiz — .35
percent, verv col)bl\ silt loam on ridges and convex areas
of upper slopes. Picaviine — 55 percent, nonc;ilc;ireous
variant, gravelly loam in concave areas tuid in draws.
Other soils — 10 percent.
BCG Brad ver\' rocIv\' loamy sand, 40 to SO percent
slopes. \('i"\ [X'rmeahle, extremelv well drained. \en
rocla, cohhlv. loamv sand; dark retklisli-hrowii; shallow.
BEG Bradshaw-Agassiz association, steep. 40-70 per-
cent slopes; moderatelv permeable, well drained.
Bradshaw — .55 percent, very cobblv silt-loam in slightlv
concave areas. Agiissiz — 3.5 percent, v erv cobblv silt-loam
in convex areas and ridgetops where soil is shallow. Other
soils — 10 percent.
DGG Deer Creek-Picayoine association, steep. 30-60
percent slopes; nioderateK permeable, well tlrained.
Deer Creek — .55 percent; loam; verv dark brown; deep
on very steep, north- and northeast-facing mountain
slopes. PicavTine — 35 percent; gravelly clav loam; verv
dark brown, deep, calcareous on west-facing slopes.
Other soils — 10 percent.
EMG Emigration very cobbly loam, 40 to 70 percent
slopes. Moderatelv permeable, well drained. Cobblv
loam; facing south; dark, gravish brown; shtJlow; patches
ot bedrock.
HGG Harkers-VV'allsburg association, steep. .Moder-
ately permeable, well drained. Harkers — .55 percent,
loam, 6—40 percent slopes, ver\' dark browTi, deep in
drainageways and concave areas of slope faces. Walls-
burg — 35 percent, very cobbly loam, .30-70 percent
slopes, on ridges luid convex areas of slopes where bed-
rock is near the surface, verv dark gravish browii, shallow.
Other soils — 10 percent.
HHF Harkers soils, 6 to 40 percent slopes. .Vlotleratelv
permeable, well drained. Loam and cobbly loam, on
sloping old alhiviiil ftuis and steep mountain slopes.
LSG Lucky Star gravelly loam, 40 to 60 percent
slopes. Moderately permeable, well diiiinetl. Wrv dark
gravish brown, deep on northerly slpes.
Mu Mixed allu\ial land. PoorK drained, highly stratified
mi.xed alhiviiini on undulating, gently sloping, and nearly
level flood iihiiiis.
during snowinelt, depending on air tempera-
tures and sncmpack depth (priuiaril\- tliat of"
upper Red Butte Canyon and Knowltons Fork).
The 1982-(S.3 winter was one of unusually high
precipitation along the Wasatch Front. Heavy
snows in mid- May 1983 were followed b\-
equall)- unusual wann temperatures at the end
of the month. As a consequence, stream flow
rates peaked at record \'alues. On 28 May 1983,
Red Butte Creek crested at a discharge rate
exceeding 2.97 mVsec (104.9 ftVsec) (stream
flow was above the maximum gage height), and
(nerland flow was substantial. This was !)\ far
the greatest discharge rate in recent times,
having eclipsed the previous maximum single
day rate of 1.70 m^/sec (60.0 ftVsec) measured
on 18 May 197.5 (U.S. Ceological Survey
Records).
The unusually high stream discharge rate in
May 1983 is of particular significance because
of its impact on stream geonioq)holog\- and
adjacent vegetation. The high flows (juickly
scoured the streambed, taking out beaver dams,
eroding stream banks, knocking down riparian
trees, and causing massive erosion. Gullies .5-10
m (16-33 ft) deep were cut into permanent
streambeds in Knowltons Fork and throughout
Red Butte Creek. Sediment flow associated
with this record stream discharge was in excess
of 269 metric tons (~.593.(){)0 lbs) per day in
mid-Mav (compared to tvpical spring melt con-
centrations of 1 metric ton [—2200 lbs] per day)
(U.S. Geological Survey Records); this resulted
in a delta formation at the mouth of Red Butte
Resenoir Prior to the 1982-83 winter, no delta
had existed. The delta was soon ~30 m (-100
ft) long. By 1990 the delta had fanned out more
than 60 m into the reservoir The heaw winter
rains of 1982-83 saturated soils all along the
Wasatch Front, and landslides were common.
Red Butte Canyon was no exception. Slope
sloughing, which killed the overlying perennial
vegetation, was common throughout the canvon.
No doubt this compounded the stream sedi-
ment load during the spring of 1983 and tor
several years thereafter. In 1990 signs of the
1982-83 slope sloughing were still clearlv obvi-
ous in Knowltons Fork as well as in the upper
and lower portions of the main canyon. Natunil
revegetation of both riparian and slope vegeta-
tion t)pes has occurred since these floods. In
particular, Acer neffimlo (boxelder) and Salix
cxiffia (willow) have increa.sed in frecjuencv in
the nevvlv deposited alluvium along the stream-
sides (Donovan and Ehleringer 1991). Recov-
erv of the sloughed slopes, which were for the
most part covered bv/\.<^m/i<'//V/<7jfr/ff///i (bigtooth
maple) and Qticrats ^amhclii ((»ambel oak), has
proceeded at a slower rate, with those slopes still
dominated by herbaceous species.
As part of the bench mark analysis, the U.S.
Geological Sunev monitors .several major iLSj^ects
of stream qualitv in addition to stream discharge,
including water temperature, suspended sedi-
ment, and chemical qualit)'. Included with
chemical rjualitv are specific conductance. pH.
104 Great Basin Naturalist
"1.50 I I ' I ' 1
[Volume 52
C/5
CO
1.25
E
(D
1.00
C5^
\-
CO
JZ
0.75
o
C/5
"a
E
0.50
03
C/D
0.25
Fig. 6 Mean monthly discharge rates of Red Butte Creek just before it enters Red Butte Reser\'oir. Large and small
tick marks indicate end-of-year and mid-year points, respectively. Data are from U.S. Geological Survey records.
di.s.soK'ed oxygen concentration, coliform bacte-
ria, and ionic and dissolved elemental concen-
trations (ammonium, arsenic, beryllium, cadmium,
calcium, carbonate, chloride, chromium, cobalt,
copper, fluoride, iron, lead, lithium, magnesium,
manganese, mercury, molybdenum, nickel,
nitrate, nitrite, phosphate, potassium, selenium,
silver, sodium, sulfate, strontium, vanadium,
and zinc). The stream itself is strongly alkaline
(pH 8.0-8.6), and travertine is deposited at sev-
enil points along the stream channel (Bond 1979).
Summertime stream flow represents
groundwater discharge, while the spring flows
result primarily from snowmelt at higher eleva-
tions. Not all of the grovmdwater originatine;
from upper-elevation sources enters the stream
before it leaves the canyon. Tracing the possible
sources of water into stream, and therefore that
water which is a\ailal)le to plants, is possible bv
analyzing the isotonic composition of that water.
The deuterium ("H or D) to hydrogen (^H)
ratios of stream waters have been measured
since June 1988 at the USGS Bench Mark sta-
tion and at the mouth of Parievs Fork by the
Stable Isotope Ratio Facility for Environmental
Research at the University of Utiili (Dawson and
Ehleringer 1991). These naturally occurring
stable isotopes of hydrogen provide long-term
data that are usehil in addressiu"; both Ions-
term regional climatic patterns and the .specific
water sources used by plants for growth (see
discussion below). Hydrogen isotope ratios
(ratio of D/H of a sample to that of a standard)
are measured relative to an ocean water stan-
dard; samples lighter than ocean water have less
deuterium and are therefore negative in their
values. Over the four-year measurement period
(1988-91), hydrogen isotope ratios of stream
waters have averaged near -122%o, with the
only seasonal changes being more negative
viilues occurring during spring snowmelt. Typi-
cally the hydrogen isotope ratio of winter stonn
events (snow) is more negative than that of
summer storms. The hydrogen isotope ratios of
wells and springs near Pinecrest (immediatelv
east of Red Butte Canyon) are - 132%p, slightly
more negative than Red Butte Creek (Dawson
and Ehleringer 1991), and suggest that a frac-
tion of the groundwater originating from the
upper portions of the canyon may persist as
underflow and does not enter the creek before
leaving the watershed. Hely et al. (1971) indi-
cated that substantial fracturing occurs in the
bedrock of Red Butte Canyon, which would
have the effect of increasing groundwater loss
from the canyon through these layers and not
\'ia stream discharge.
Bond (1977, 1979) investigated nutrient-
concentration patterns of stream flow in Red
Butte Creek. In particular, his studies focused
19921
Red Butte Canyon Research Natuiul Area
105
Tablf. 3. Locations of wcatlicr stations of Red Butte C^iuivon. All stations were operattd 1>\ tlie U.S. Arniv between
1942 and 1964, and onI\- precipitation was recorded. The U.S. Geoloijical Siir\e\ has maintained a storage gage at Red
Bntte #2 since 1964. The BioIog\ Department at the Universit)' of Ut;ili has maintained daik temperature, humidity, and
wind speed records at Red Butte #2, Red Butte #4, iuid Red Butte #6 since 1982. Red Butte #1 . while technicall\ outside
the canyon, forms an integrated part of the weather station complex.
Station
Location
Latitude
Longitude
Elevation
Period
Red Butte #1 Fort Douglas 40° 46'
Relocated to Biolog)' 40° 46'
Experimental Garden
Red Butte #2 Head of Red Butte 40° 47'
Resenoir
Retl Butte #3 Along Red Butte Creek 40° 48'
at Brtish B;isin
Red Butte #4 Along Red Butte Creek 40° 48'
100 m west of Bea\'er
Canvon
Red Butti- #5 Parleys Fork 100 m above 40° 47'
inlet to Red Butte Creek
Red Butte #6 Upper end Knowltons Fork; 40° 49'
relocated to top of Elk Fork 40° 49'
110°
'50'
110°
.50'
IIP
48'
111°
47'
iir
46'
111° 48'
111° 45'
111° 46'
1497 111
1515 in
1653 111
lS65m
lS90ni
17.53 111
2195 m
2195 m
1942-1964
1991-pre.sent
1942-19fS4
1982-present
1942-1952
1942-1971
1982-preseiit
1942-1956
1946-1971
1982-present
on relationships between ntitiient transport out
of the watershed and stream diseharge rates.
Sokite concentration was not necessarilv pro-
portional to stream discharge. Instead, for many
ions, such as magnesium, sulfate, and chloride,
the relationship was logarithmic. The slopes of
these relationships depend on whether stream
flow is increasing (i.e., spring snowmelt) or
decreasing. Over the course of the year, a loop
or directioucil trajectory was formed by having
two different slopes. For most of the major ions,
the trajectorv' was clockwise; that is, ionic con-
c(Mitration was greater in winter when flow rates
were low than during summer. Plant growth of
the dominant riparian species commences near
the end of the snowmelt period, and it is ques-
tionable whether riparian species are able to
utilize the greater nutrient aviiilabilitv durino;
the snowmelt period. After snowmelt, stream
discharge is based primarily on groundwater
input. Nitrate, ammonium, and phosphate con-
centrations in Red Butte Creek during ground-
water discharge are low (Bond 1979). In
contrast, overall concentrations of calcium,
magnesium, sodium, chloride, and sulfate are
much greater because of parent bedrock char-
acteristics.
Climate
Climate within Red Butte Can\on is charac-
terized by hot, dry summers and long, cold
winters. Most precipitation occurs in winter and
spring, with the summer rains less predictable
and dependent on the extent to which mon-
soonal systems penetrate into northern Utah.
Mean annual precipitation ranges from about
500 mm (20 in) at the lower ele\ation to appro.x-
imatelv 900 mm (35 in) at the higher elexations
(Hely et al. 1971, Bond 1977; Table 3).
Precipitation stations have been monitored
in Red Butte Canvon by several groups. The
U.S. Army had six rain gages in operation
between 1942 and 1964 (Table 3). Bond (1977)
collected data at several of these stations
between 1972 and 1974. In addition, the U.S.
Geological Sune\' maintained storage gages at
Red Butte #2, Red Butte #4, and Red Butte #6
between 1964 and 1974. Since that time, they
have maintained a storage gage at Red Butte #2.
Within the watershed, diiiK' precipitation as
rainfall is collected at eacli of the weather sta-
tions; snowfall is not adequately measured by
the sensors in place. However, these data are
currently collected at Hogle Zoo in Salt Lake
City (same elexation as pre\ious Red Butte #1,
but 4 km south).
Variation in annual precipitation w ithin Red
Butte CJanxon is strongly dependent on eleva-
tion (Fig. 7). The slope of this relationship is
similar to that obser\ed for other mountainous
areas within the Great Basin (Houghton 1969),
and precipitation at the Salt Lake Cit\' reporting
station (Salt Lake Citv International Airport)
falls on this relationship. Thus, while lacking
continuous precipitatif)n records for the canyon
proper, precipitation records a\ailable for Salt
Lake City can be used as a preliminar\- basis for
estimating mean annual precipitation at differ-
ent locations within the canxon.
106
Great Basin Naturalist
[Volume 52
o
400
1200 1400 1600 1800 2000 2200
Elevation, m
Fig. 7. Relationship between mean annual precipitation
and elevation for Red Butte Canyon storage gages Red
Butte #l-#6. Shown also is the mean annn;il precipitation
for the primarv station of Salt Lake City (Salt L;iJ<e City
International Airport) as the open symbol.
Fig. 9. Mean monthly maximum and minimum air tem-
perature at Red Butte #2 (165.3 m elevation). Red Butte #4
(1890 m elevation), and Red Butte #6 (2195 m elevation)
during the growing season between 1982 and 1990.
Air teinperatiire.s have been collected from
automated weather .stations at Red Butte #2,
Red Butte #4, and Red Butte #6 since 1982.
Mean monthly air temperatures at Red Butte #2
were below freezing in December and fanuaiy
and above 20 C in June, July, and August (Fig.
8). In contrast, mean monthly temperatures at
Red Butte #6 were below freezing only slightK
longer, from November through February, and
abo\'e 20 ( ] in July and August. During the main
growing period (May through September), day-
time maximum temperatures ranged between
30
20
10
-10
8
6
4
2
H 1 1 h
H 1 1 h
-H \ 1 1 1 h
M A M J
A S N D
Fig. 8. Mean monthlv ;ur temperature, vapor pressure,
and photosvntheticallv active solar radiation (400-700 nm)
measured at Red Butte #2 between 1982 and 1990.
18.7 and 31.8 C (66-89 F) at Red Butte #2, while
nighttime minimum temperatures ranged
between 5.2 and 16.4 C (41-62 F) (Fig. 9). At
the higher-elevation stations, davtime maximum
air temperatures were lower. The difference in
maximum temperatures was negatively related
to elevation (maximum temperature [°C] = 34.3
- 0.00494 • elevation [m], r = .91) at approxi-
mately half the diy adiabatic lapse rate. On the
other hand, nighttime minimum temperatures
were not related to elevation, because of cool-
air drainage effects (Fig. 9). Red Butte #4 is
located streamside within the canyon, whereas
the other two stations are above the channel of
cold iiir that develops at higher elevations and
pours down the canx'on at night. As seen in
Figure 9, this cold-air drainage effect at Red
Butte #4 (1890 m [6180 ft] elevation) depressed
nighttime mininuim air temperatures bv 4-8 C
(7-14 F) below that obsened at Red Butte #6
(2230 m [7292 ft] elevation).
Photosynthetically active solar radiation
(PAR, 400-700 nm), atmospheric vapor pressure.
1992]
Red Butte Canyon Research Naturae Aiu<:a
107
and wind speed are also recorded at each of
these stations. Between 1982 and 1990, mean
daiK' total PAR \iilues have exceeded 40 niol
m "' d~ ' ( Fig. 8), which is t>pical for mid-latitude
sites ha\ing onK' moderate cloud cover and little
sunuiier precipitation. This number is quite
useful not only in estimating the available
photon flux for photos)Tithesis, but iilso in pro-
\iding an estimate of the extent of solar heating
of the surface, which ultimatelv affects air tem-
peratures. Elevation has a limited impact on the
PAR values within Red Butte Canyon, since the
difference in elevation is relatively small. How-
ever, we suspect there may be relatively large
differences in PAR betv\'een Red Butte Can)'on
and Salt Lake Cit\' because of increased mv
pollutants within the city that tend to reflect the
sunlight before it strikes the earth's surface.
Most notablv we would see this as haze or smog
within the \alle\' that is lacking once in the
canyon.
Average monthly atmospheric vapor pres-
sure at site #2 showed little annual variation,
ranging onlv about 3 nibar throughout the year
(Fig. 8). Other sites exhibited a similar pattern.
This parameter is largel)- affected by large air
mass movements; and since subtropical air
masses do not move into this region during the
summer, the monthly changes in atmospheric
\'apor pressure change little during the course
of the year. However, because of the large
annual change in air temperature and the non-
linear dependence of the evaporative gradient
on temperature, relative humidit\' levels are
substantially lower and evaporative gradients
are substantially higher during the summer
months.
Vascular Flora
From the mouth of Red Butte Canyon at
about 1530 m (5020 ft), its walls rise to their
highest point— 2510 m (8235 ft)— at the head
ofKnowltons Fork in the northeast corner of the
canyon. Within this modest rise of 980 m (3215
ft) occur four distinct plant communities: ripar-
ian, grass-forb, oak-maple, and coniferous.
Piiion-juniper and ponderosa pine communi-
ties, which often occur in this ele\ational range
in Utah (Daubenmire 1943), are not present in
Red Butte Canyon. Billings (1951, 1990), in
discussions of vegetationtil zonation in the Great
Basin, cites a greater incidence of winter
cyclonic storms and slightly more moist sum-
mers as factors producing the xariatioii in the
vegetative zones of the eastern boundary' of the
Great Basin. Juniper is present in the central
Wasatch Range, i)ut onlv three Utah juniper
ijunipenis osteospenmi) are known to exist in
Red Butte Canyon: a mature tree with a 0.5 m
(1.6 ft) diameter trunk, located on the south
slope of Parleys Fork and nearly obscured by the
more mesoph\tic vegetation, and two shniblike
plants 1-1 .3 m (3-4 ft) tall growing on the south-
west divide.
With few exceptions, notably the naturalized
grasses Agrostis stolonifera (redtop bentgrass),
Bromits tectonim (cheatgrass), and Poa praten-
sis (Kentucky bluegrass), onK the most common
indigenous plants that occur in the \arious plant
communities are listed below, primarily because
the presence of introduced plants is usually
dependent on disturbance and tends to fluctu-
ate accordingly. Some of the more frequently
occurring introduced plants are listed in a sep-
arate section.
Riparian community— From the point at
which Red Butte Creek emerges from the
canyon and throughout the floor of the cam on
the streamside vegetation (plants residing in soil
kept moist to wet by the stream) consists chiefly
of western water birch (Bettila occidcntalis) and
mountain alder {Aliuts incana), accompanied at
intervals by usuiilly dense stands of red osier
dogwood {Corrms sericea) and willow {Salix spp.).
Adjoining the stream along the floor of the
canyon below and above the reservoir is an often
densely wooded strip consisting chiefly of
Gambel oak {Quercus gambelii), boxelder {Acer
ncgiindo), and bigtooth maple {Acer grancli-
dentatinn), many of these trees ranging from 9
to 18 m (30 to 60 ft) or more tall. Also included
in this plant connnunit} are wideK scattered
individuals or small populations of cottonwoods
{Populns frenwntii, P. angustifoUn, and P. x
acuminata), chokecherry {Pniniis virginiana).
Woods rose {Rosa woodsii), bearbern,- honey-
suckle {Lonicera invulucrata), thimbleberry
{Rubus parvifloms), serviceberry {Amelanchier
ainifolia), western black currant {Rihes htid-
soniamini), and golden currant [Ribes aurenin).
Relatively few species of grass and forbs are
found here, among them:
Ehjitms i>l(innis
Loiiuitiitin (iLsscctiim
Mahouia refjens
( B c rb c n.s ref)e ns)
Osmorhiza chilemis
Poa comprcssa
blue wildrv'e
y;iant lomatium
Oregon grape
sweet cicelv
Canada bluegrass
108
Great Basin Naturalist
[Volume 52
P. pratensis Kentucky hliiegiiiss
Smilacina stellata wild lily-of-the-valley
S. raccinosd false Solomon-seal
Solidago canadensis goldenrod
Bcaven once native, were reintroduced into
Red Butte Canyon in 1928 (Bates 1963) and
were active along Red Butte Creek and some of
its tributaries for 54 years thereafter. Numerous
marshy areas between elevations of 1645 m
(5400 ft) and 2133 m (7000 ft) were created by
the impoundment of water due to their dam-
building activities. To prevent the beaver popu-
lations from becoming undesirably large, the
Utiili Dixision of Wildlife Resources in 1971
undertook management of the populations. In
December 1981 a recommendation was made,
based on an analysis of the water supply to Fort
Douglas from Red Butte Canyon, that all beaver
be eliminated from the canyon because their
feces could contaminate the water with the par-
asite Giardia Jamhlia. Accordingly, in 1982 the
colonel in command of Fort Douglas applied for
and received from the Utah Division of Wildlife
Resources a permit to remove the beaver from
the canyon. Subsequently, all beaver were "har-
vested."
Bates (1963) studied the impact of beaver on
stream flow in Red Butte Canyon. The vegeta-
tive cover was affected for approximately 91 m
(298 ft) on either side of the portion of the
stream in which the beaver were active, and
sediment deposited behind the beaver dams in
the canyon varied from 0.6 to 2.4 m (2 to 8 ft) in
depth. He also noted that the small alluxial
plains formed by the sediment made it apparent
that during periods of high rimoff, and perhaps
during normal flow, the dams allowed the reten-
tion of quantities of suspended materials. Schef-
fer (1938), in a report on beaver as upstream
engineers, ascertained that two beaver dams
retained 4468 m' (157,786 ft^) of silt. It is not
known whether an actual count of the number
of beaver dams in Red Butte Canyon was ever
made; but the environmental change effected
by their ultimate displacement during the 1983
flooding of what had to have been enormous
quantities of sediment has been significant. The
removal of all inactive beaver dams has inevita-
bly led to the elimination of or significant reduc-
tion in the densitv' of some 55 species of t^'^iicalK
wetland plants from once marshy areas wdthin
Red Butte Canyon. For example, in 1990 it was
noted that in an area which once supported a
nearly pure stand of closely spaced cattails
{Typha Idtifolia) covering approximately 0.25
hectare (0.62 acre), only a few scattered clumps
remained. According to Forest Service person-
nel, these losses would not have been as severe
had the beaver dams been active during flood-
ing. Species in the following genera are among
those undoubtedly affected: Eleocharis, Scir-
pus,Junnis, A<i^rostis, Catahrosa, Deschampsia,
Ghjceria, Poa, Polijpogon, Eqnisetum, Angelica,
Betula, Ciatta, Heracleum, Rudheckia, Soli-
dago, Barbarea, Cardamine, Nasturtium,
Rorippa, Lonicera, Corniis, Trifoliiim, Mentha,
Nepeta, Lenina, Epilohinni, Hahenaria, Pole-
nioniiim, Polygonum, Rumex, Aconitum,
Ranunculus, Geum, Rihes, Salix, Mimulus,
Veronica, and Urtica.
The U.S. Forest Service, Salt Lake Ranger
District, requested the Utah Dixision of Wild-
life Resources to reintroduce the beaver during
the summer of 1991. At the time of this publi-
cation, bea\'er had not vet been reintroduced. It
is hoped that with time the plant diversit}' typi-
cally associated with beaver dams will be rees-
tablished.
GRASS-FORB community. — According to
Stoddart (1941), the grasslands of northern
Utah form the southernmost extension of the
Piilouse prairie. Of the two communities into
which the Palouse prairie is divided, onlv that
dominated by bluebunch wheatgrass {Ehjmus
spicatus, originally known as Agropyron
spicatum) occurs in Red Butte Canyon. Rela-
tively large open areas inhabited by grasses and
forbs, wath an occasional big sagebnish {Artemi-
sia tridentata), squawbush {Rhus trilohata), and
bitterbmsh {Purshia tridentata), are found
chiefly below the 1829 m (6000 ft) contour
(Kleiner and Harper 1966), although smaller
grass-forb associations also occur in forest clear-
ings at higher elevations. Some of the more
commonly occurring species wdthin the grass-
forb communitv' at lower elevations are:
Achillea inillifolinin
Allium acianinatuin
Ambrosia psilostaclnja
Arahis hollniellii
Aiistida piiijiurea
(A. l()n<i^isefa)
Artemisia huloviciana
Astra<iahis utahcn.sis
Aster adscenden.s
Balsanu>rhiza macrophijlla
Bal.samorhiza sagittata
Bromns teetoniin
Cirsium undulatiim
CoUomid linearis
Comandra innhellata
milfoil Narrow
tapertip onion
western ragweed
Holhoell rockcress
pnr][ile threeawn
Louisiana wormwood
Utah milkvetch
everywhere aster
cutleaf balsamroot
arrowleaf biilsamroot
cheatgrass
gray thistle
narrowleaf collomia
bastiird toadflax
1992]
Red Buttk Canyon Research Natural Area
109
niomitain li auks heard
loiiji-stalk spriiig-parslev
sleiulcr wlu'at grass
aiituinii willowherh
spreading ckisN'
broom siiiikeweed
northern sweetvetch
showy goldeneye
temate lomatiuin
silveiT Kipine
little polecat
threadleat scorpionweed
longle;if phlox
Sandberg bhiegrass
needle-and-thread
mnlesears
Crepls (icuininatd
Cynioptents lon^ipes
Ely mils traclii/caiiliis
{Ai^ropyron caiiinuin)
Epih >l>i uin h rack ycarjnim
(E. panicuhtum)
Erigeron diveraens
GuticiTczUi sarothrae
Hcclysani in horcale
Helionwris mitltiflora
( V'(g(»V'ra niiiltifliira )
Lonuitium tritenuituin
Lupinus argenteiis
Microsti'ri.s gracilis
Phacelia linearis
Phlox longifolia
Poa scninda [P. sandhcrgii)
Stipa conuita
Wt/ctliia ainplcxicaidis
Oak-MAPLE communit\'. — Gambel oak
{Querciis gamhelii) is the dominant type of veg-
etation tliroughoiit the altitudiniil range of the
canvon. It forms what appear to be randomly
spaced clones throughout much of the area. In
accordance with the moisture regimen, the
clones may range from thickets 0.3 m (1 ft) or
less in height in dr\' upland sites to stands of
stately, well-spaced trees in lowland areas. Both
walls of the canyon support often nearly
impenetrable oak in association with bigtooth
maple {Acer grand identatiun) , the latter grow-
ing chiefly in drainageways. Few species thrive
as understor\' with dense oak cover. The most
common are Galium aparine (catchweed bed-
straw) and Mahonia repens (Oregon grape).
Others appearing seasonally under oak are
Enjthroniiim grandiflonim (dogtooth violet),
Claijtonia lanceolata (lanceleaf spring beauty),
Hydroplujllum capitatum (ballhead waterleaf),
and H. occidentale (western waterleaf). Among
plants commonly fringing oak clones are:
Agoseris glaura mountain dandelion
Apocyniun androsacinifolinin spreading tlogb;xne
Arabis glabra tower mustard
Bromus carinatus mountain bronie
Comaiidra itmbellata bastiird toadflitx
Delphiniinn niittallianinn Nelson larkspur
Descurainia pinnata blue tansv nuistard
Eriogunum heracleoides whorled buck-wheat
E. racenwsiim redroot buckwheat
Geranium viscosissimum sticky geriuiinm
Hcliandiella unijlora one-headed sunflower
Heliomeris multiflora
(Vigiiiera multiflora) hairv' goldeneye
Hydrophyllum spp. waterleaf
Koeleria iiuierantha
(K cristata) Junegrass
Leucopoa kingii
(Hesperochloa kingii) spike fescue
Lomatium dissectiim giant lomatium
Machacrantlicra canescens hoar\' ;ister
Meiiensia brei isti/la
Microseri.s nutans
Pha celia heterophylla
Poa fendleriana
P. pratensis
Senecio integcrrimiis
Wasatch bluebell
nodding scor/onella
varileaf scoq:)ionweed
muttongriiss
Kentucky bluegrass
Columbia groundsel
Mountain mahoganv {Cercocarpus ledifo-
litis) occurs as individuals and as scattered,
mostly small populations, often in association
with oak, sagebrusli, or other mountain shrubs,
generally on northwest-facing, sparsely vege-
tated slopes. It can be seen from the main road
through the canyon as small trees against the sk\'
along the exposed, rock-v, south rim of the
canyon, especially toward its western end. As
low shmbs it occurs sporadicalK; chiefl\' on
exposed diy sites above 1980 m (6500 ft).
Big sagebrush {Ariemisia trident ata) occurs
sporadically in drier sites throughout the
canyon's altitudinal ran^e. Low sao;ebrush
(Artemisia arbnscula) occurs as relatixely pure
stands at about 2133 m (7000 ft) along the
southeast rim of the canyon.
Coniferous community. — Douglas-fir
{Pseudvtsuga menziesii), white fir (Abies con-
color), and aspen (Popnlus trenmloides) domi-
nate this community, either in pure or in mixed
stands, growing chiefly on north- to northeast-
and northwest-facing slopes; the aspen reach as
low as 1706 m (5600 ft) and the firs occur mostly
above 1828 m (6000 ft). Achlorophyllous
CorallorJiiza spp. (coralroot orchid) are ainong
the few plants able to flourish in the shade of
dense stands of mixed conifers. Many small
trees, shrubs, forbs, and grasses thrive in less
dense stands or in openings between stands of
trees in this commimit)'. Among them are:
Aeerglabntm
Anwianehier ainifolia
Acjiiilegia eoendea
Aniiea spp.
Castilleja spp.
Ccanothiis vcliitiniis
Elymus glaueiis
Erigeron speciosus
Galium spp.
Hordeum braeliyantlicntin
Lathy nis paiieiflonts
Physoca rjnis nuilvaceus
Poa nervosa
Pninus virginiana
Rihes viscosissimum
Riibus paniflora
Sambncus spp.
Sorbiis seopuliua
Symphoricaiyos oreophilus
Thalictniin fendlcri
Rocky Mountain maple
Saskatoon seniceberry
Colorado columbine
arnica
Indian paint brush
mountain lilac
blue wildr\e
shouy fleabane
bedstraw
meadow barley
Utah .sweetpea
mallow ninebark
Wheeler bhiegrass
chokecherrv'
sticky currant
thinibleberr\
elderberr)
American mountain ash
mountain snowberr\'
FendJer meadownie
110
Great Basin Naturalist
[Volume 52
Plants endemic to Utah. — Only two spe-
cies occurring in Red Butte Canyon are said to
be endemic to Utah: Ang^elica wheeleri Wats.
(Mathias and Constance 1944-45) (Wheeler
angelica) and Erifieron arcnarioidcs (D. C.
Eat.) Gray (rock fleahane). Angelica icJieeleri
has, however, been collected close to both the
Idaho and the Nevada boinidaries with Utah
(Albee et al. 1988). Ehgeron arciiaiiokles is
kn(nvn from Salt Lake, Utah, Tooele, Weber,
and Box Elder counties (Albee et al. 1988,
Cronquist 1947).
Plants introduced to Utah. — In Red
Butte Canvon, plants introduced to Utali, either
from other portions of the United States or from
another country, are largely restricted to road-
side and trailside sites and to open grassy or
rocky slopes below 1829 m (6000 ft). Some of
the more commonh' occurring plants in this
categorv are:
Ali/ssu in ahjssoulcs
Artibiclopsis thaliana
B ramus hriziformis
(B. hrizacfomiis)
B.Japonicits
B. tctiontm
Capsclla hu rsa-pastoris
Ct/n()<^l().s.sum officinale
Dactijlis t^loinvrata
Draha vcrna
Erodiuni cicutarium
Grin deli a sqiia rrosa
Holostcum iiinhcllafnin
Isatis tinctoria
Ladiica scrrioUi
Lcpidiuin jx'iidliiitunt
Linaria dahnatica
Lithospcnnti nx ancnac
Mdlva nc'^lcctd
Mdilotus alha
M. officinalis
Poa Indhosd
Ranunadiis tcsticiilatiis
Sisijmhrinni altissiiinun
Tanixdciim officiudle
Thlaspi dncnsc
Trdff)po<ion dnhius
Veroiiicd dnagallis-dtjudticd
alyssum
mouse-ear cress
rattlesnake chess
Japanese or meadow cliess
cheatgrass
shepherd's purse
hound's tongue
orch;u'd grass
spring draha
storkshill or ;ilfileria
curhcup gumweed
jagged chiek'weed
dvers woad
pricklv lettuce
peppergrass
Dahnation toadflax
com gromwell
cheeses
white sweetdover
yellow .sweetdover
bulbous bluegrass
bur buttercup
|iui Hill uuistard
conuiiou dandelion
pemivcress
goatsbeard
water speedwell
The incidence oflsatis tinctoria and Linaria
dahnatica increased greatlv between 1970 and
1990.
Floristic DIXERsrn.— The following .spe-
cies were reported from Red Butte Canyon b\
Cottam and Evans (1945) and by Bates (1963).
Not only is the presence of these plants unveri-
fied by herbarium specimens (see Albee et al.
1988, which is based on specimens in the herba-
ria of Brigham Young Universit); Utiili State
University, and the University of Utah), but at
least SLX of them wot
within the elevational 1
A<^rostis scniivciiicilldtd
Anisinckid tessclldtd
Angclicd pinndtd
"Bhckcllid ^rdndijlora
Cdstillcjd dnffistifolid
Cirsiiim flodnwnii
Cryptdnthd fldvoctddtd
Dcsclunnpsia cacspitosd
"Erifieron ^Idhelhis
°Eriog(miiin ovalifoliitin
Gdt/oplii/tu m rdmosissi inu ni
Geraniuin bickncllii
Ghjccrid ^rdndis
Jtincns uicricnsidnits
"Ldthi/nis hrdclnjcaliix
Mentzclid dlhicdulis
Scirjnts inaritimiis
"Stcllarid lon^ipes
Vdlcridnd edulis
lid not ordinarilv occur
limits of the canyon:
water polypogon
rough fiddleneck
small-lea\ed angelica
tasselflower
Indian paintbnish
Flodnian thistle
yellow-eve crvptanth
tufted hairgrass
smooth tleabiuie
cushion buck"A\'heat
branchy groiuidsmoke
Bicknell cr;uiesbill
American mannagrass
Merten's rush
Rvdberg sweetpea
whitestem blazing star
alkali bulnish
long-stalked starwort
edible valerian
The following species were reported by
Amow ( 1971 ), but, for the reasons stated below,
can no longer be considered part of the flora of
the canyon:
Arahis pnbenila Nutt.
(pubenilent rockcress)
Calypso hulhosd (L. ) Oakes
(fair)' slipper orchid)
Collection identified by
R. C. Rollins as an anom-
alous A. lenwwnii Wats.,
the correction too late for
the 1971 publication.
1971 report based on a
basal leaf, no subsequent
evidence of its presence
available.
A misidentification.
Carcx muricata L. (as C.
ani^ustior Mack)
Species names now submerged with those of
other species present in the canvon (also
included in section on nomenclatin-al changes):
Arabis divaricaijja A. Nels
= A. holbocllii Horneni.
Bromits coniinutatus Schrad.
= B. japonicus Thunb.
Gli/ccria data ( Nash )
M. E. Jones = G. striata
(Lam.) Hitchc.
jiincus traci/i Rvdb.
= J. cnsifoliiis Wikst.
Taraxacum laeii^atum
(\Villd.)DC. = T officinale
W'iggers
Thus, the 511 species representing 73 fami-
lies reported from Red Butte Canyon by Arnow
(1971) can now be placed at 484 species (390
indieenous and 94 introduced) known to have
Holboell rockcress
Japanese or meadow chess
fowl mannagrass
swordleaf nish
common dandelion
°\\'itli tlie iis.si.staiice of Kave Thome and Leila Shiiltz, curators of the herbaria
at Brigliam Yoiinj; and Utah State universities. respecti\ely. a herbarium check
u'iLs made to l)e certain tliat no Hed Butte Canvon s[)ecimens exist for those
s])ecies marked with an asterisk tliat. .iccordiny to .\Miee et al. ( 19.S8), are not in
Ked Butte Canyon or its vicinitv.
1992]
Red BrrrK CIwyon Rkskahcii N'vii uai. Akea
111
2200
2000
1800
1600
Fi<j. 10 Distribution, b\' elcnation, of the major ]ilaiit
C'oniniunitics in Red Butte Cainon.
been present in tlie ean\()n at one time or
another. Onl\' two populations present in 1971
are definitely knowni to have been eliminated:
Lactuca biennis (biennial \v\\d lettuce), which
w as introduced into Utali from the nortli about
1967 but did not survi\'e; and SoJid(i(H)
occidental is (western ii;oldem"od), a single
streamside population at the mouth of the
canvon taken out by the 1983-84 flooding.
According to Albee et al. (1988), the 390
indigenous species reported from Red Butte
Canx'on (Arnow 1971) also occvu" in at least one
other canvon to the south. Arnow et al. (1980)
and Albee et al. ( 1988) indicate that roughly 1 30
native plants not found in Red Butte C>an\'on
ha\e been collected between an ele\ation of
1S2S and 2438 m (fiOOO and 8000 ft) in can\ons
liaxing a greater altitudinal range in southern
Salt Lake Countw This figure indicates tliat the
Holistic di\ersit\' in Red Butte Cainon, while
greater than that in hea\ih" disturbed Emigra-
tion ('aiiNon (Cottani and E\ans 1945), is less
than that in camons farther south.
Nomenclatural changes since Arnow (1971)
are listed in the Appendix.
Plant E(;ol()(;y
Vegetation distribution. — A number of
studies ha\e focused on describing the \egeta-
tion distribution within Red Butte Can)'on
(Kleiner and Harper 1966, Swanson, Kleiner,
and Haiper 1966. Kleiner 1967). There is a
strong xeric to mesic elexation gradient, with
lower portions of the canxon dominated b\- a
spiing-actixe grassland communitx and the
upper portions ol tlu^ cainon txpicaJK consisting
oi suinmer-actix'e scrub oak, aspen, and conifer-
ous forest cominunities (F'ig. 10). CJomposition
within each of these communities is not con-
stant, but instead species \an' in their impor-
tance within a communitv t)pe as orientation
and ele\ ation change. These elevation gradients
n^present a continuum of moisture axailabilitx;
with high temperatures and low precipitation
amounts at lower elevations making conditions
more xeric, while slope orientations less south-
vv\\' in exposure become progressivelv more
mesic within an elevation band. Soil txpe (Fig.
5) and depth also play a major role in afflicting
plant distribution by providing variation in the
water-holding capacity of the substrate. The dis-
tribution of the sciTib-oak communitx- to the
highest elevations within tlie canxon is most
likelv related to soil conditions, sinc(^ at liigh
elexations scrul) oak persists on south-, east-,
and west-facing slopes that would normallv be
expected to be dominated b\ aspen if it were not
for the \en' shallow, rock^' soils that txpif\ these
elex ations witliin Red Butte Ciinvon.
Red Butte Canvon has been largeK pro-
tected fr(jm grazing since its ac(juisition by the
U.S. Army almost a centuiy ago. The conse-
(juence of this lack of grazing pressure at lower
elexations is a recoxerx' to near pristine levels,
and this is clearly reflected in the earl\- commu-
nitx- anaKses of Exans (1936) and Cottam and
Exans (1945). \\'ithin the .scrub oak and grass-
land communities of Red Butti^ Camoii and
adjacent Emigration Can\-on, a canyon annually
expo.sed to sheep griizing, there are large differ-
ences in plant densitx' (Fig. 11). Emigration
Canvon was originally described by early pio-
neers as haxing a dense vegetation at lower
elevations. However, grazing not onlv reduced
that coxcr but also increa.sed the fraction of the
plant cover occupied In- mderal, weedv .species
(Cottani and Exans 1945). While plant densit)'
in Red Butte Canyon mav be greater and weedy
species composition loxx'er as a result of reduced
disturbance and grazing, the canvon is not free
of these vxeedx components and historical
effects (as noted in earlv- sections). Dam con-
struction during thi> 1 920s and other U.S. Army
actixities vxithin the lower portions of Red Butte
C^anxon have resulted in sufficient disturbance
that main mderal, weedy species, such as
Crindelia sijuarrosa (curlv gumvx'eed), Lactuca
serriola (pricklv lettuce), and Polygonum avi-
culare (knotxveed), are novx- common.
112
Great Basin Naturalist
[Volume 52
Saniuelson (1950) conducted an analysis
similar to that of Cottam and Evans (1945) on
the algal components of the streams in Red
Butte and Emigration canyons. He observed
that as a result of livestock grcizing and human
settlement, sediment load and turbidity were
much greater in Emigration than in Red Butte
Creek. The consef juence of this stream-qualitv
difference was the dominance by algal genera in
Emigration Creek that are turbidity tolerant,
such as Oscillatoria and Phonnidium. Con-
versely, in the clear waters of Red Butte Creek
filamentous algae, primarily Nostoc, were most
common. Overall algal densities were three
times greater in Red Butte Creek, owing to the
greater light penetration into that stream. At the
same time, Whitney (1951) compared the dis-
tributions of aquatic insects in the two streams.
He found that densities of aquatic insects were
greater in Red Butte Creek. Of those insects
persisting in Emigration Creek, there was a
preponderance of species characterized by gills
protected from silt, which would better allow
them to tolerate the more turbid conditions in
Emigration Creek.
Phenology, — Plant activity is governed by
t^vo parameters: temperature and soil moisture
availability. Cold winter temperatures limit
growth activity between November and March
(Caldwell 1985, Comstock and Ehleringer
1992). While a limited number of species, such
as the early spring ephemeral Ranunculus tes-
ticulatus (bur buttercup), may begin activity
during warm periods in Eebmary, most annuals
do not begin growth until the warm periods
between snowstorms in early March. At lower
elevations, a number of herbaceous perennials
such as BalsainoHiiza macroplujUa (cutleaf
balsamroot) may begin to leaf out during March,
but most woody perennials do not leaf out until
mid- to late April. The annvials and most herba-
ceous species at lower elevations have com-
pleted growth and reproduction by mid-June
and then remain dormant until the following
autumn or .spring (Smedley et al. 1991). In con-
trast, woody species at lower elexations remain
active from April through October, although the
vast majority of the growth will occur during the
spring (Donovan and Ehleringer 1991). At
higher elevations, vegetative and reproductive
growth are delayed imtil late May or June by
cold temperatures. Plants at the higher eleva-
tions vdll remain active throughout the summer,
30 r
20'
^ 10
m Red Butte
n Emigration
*i>.^
â– A
1515 1625 1700
Transect elevation, m
2060
Fig. IL A comparison of the plant cover in open grass-
Ituid communitie.s of different elevations in Red Butte and
Emigration ciinyons. Adapted from Cottam aiid Evans
(1945).
even though there may be httle summer precip-
itation (Dina 1970, Dina and Khkoff 1973).
Adaptation. — In the nonforested portions
of the Intermountain West, plant growth is
largely restricted to spring and early summer
periods by cold temperatures during winter and
limited water availabilitv during the summer
(Caldwell 1985, Dobrowolski, Ciildwell, and
Richards 1990, Comstock and Ehleringer 1992).
A number of recent reviews have addressed
adaptation characteristics ot plants growing in
these environments (Caldwell 1985, DeLucia
and Schlesinger 1990, Smith and Knapp 1990,
Smith and Nowak 1990). For the most part,
plants within Red Butte Can von are exposed to
a hot, diy environment, with little relief from
developing water stress during the summer
months. The onlv clear exception to this pattern
is the series of plants within the riparian com-
munities cilong the canyon bottom. To giiin a
better imderstanding of this occurrence, many
of the recent ecological researchers within the
Red Butte Canyon RNAhave focused on mech-
anisms by which plant species have adapted to
limited water availabilitv.
Among the first ecophysiological studies was
that b)' Dina ( 1970), who examined water stress
levels of the dominant tree species in the lower
portions of the canyon: Acer firandidcntatum
(bigtooth maple), Acer negundo (boxelder),
Artemisia tridentata (big sagebrush), Purshia
tridentafa (bitterbrush), and Quercus ganibelii
(Cambel oak). Dina (1970) observed that
1992]
Red Butte Canyon Research Naturae Area
13
o
E
o
E
E
>.
o
c
o
o
CD
en
ZD
I
CO
grasses
forbs
April
May
June
Fig. 12. The mean water-use efficiency viilues for
grasses and forbs within the grassland community of Red
Bvitte Canyon during main period of the growing season.
Water-use efficiencies were calculated from ctirbon isotope
discrimination values from Smedlev et al. (1991) ;uid the
\apor pressure data in Figure S.
middav leaf water potentials of -30 to -65 bans
develop in perennials occupying slope sites
during late sunniier, whereas water potentials of
adjacent riparian tree species are maintained
between -20 and -30 bars during the same
periods. Water potentials in the range of — 10 to
-15 bars cause many crop species to wilt and
close their stomata, reducing transpirational
water loss. Tolerance of water stress le\els as low
as -40 to -60 bars is thought to occur in only
the most drought-adapted aridland species.
These late-summer water potential \alues on
slope species are sufficientK' low to close sto-
mata and reduce photos) nthesis to near zero
values. In Dina's (1970) study photosynthetic
rates of riparian species decreased bv 50-80%
from nonstress \alues, l)ut riparian trees were
able to maintiiin positive net photosynthetic
rates throughout the summer. More recentK;
Dawson and Ehleringer (1992) and Donovan
and Ehleringer (1991 ) conducted related stud-
ies and again obsened that photos\iithetic
carbon gain of slope species is largely limited to
spring and early summer, whereas riparian spe-
cies are able to maintain photosNuthetic rates
throughout the \ear, albeit that photosxiithetic
rates are lower in summer than in spring.
Two common responses to limited water
a\ailabilit> are axoidance and tolerance. Axoid-
ance of water stress is accomplished by comple-
tion of growth and reproductixt* activities before
theon.set of thesunimer drought, whereas toler-
ance is associated with the e\olution of features
that allow plants to persist through the drought
period.
Several interesting studies ha\e been con-
ducted in Red Butte Canyon that shed liglit onto
the nature of a plants ability to tolerate water
stress and persist through time. Treshow and
Harper (1974) examined longevity of herba-
ceous perennials in grass, mountain bmsh,
aspen, and conifer communities throughout the
canyon. They observed that life expectancies of
dominant herbaceous perennial species, such as
A.sf/7/gc////.s utahcnsis (Utah milk\etch), Balsa-
niorliiza inacwpJu/lIa (cutleaf balsamroot),
Hech/sanini horcale (northern sweetvetch), and
WyctJiia ainplexicaulis (mulesears), are rela-
tiveK' short (3-20 vears) when compared to the
longer-li\ed (>65 years) grass species, such as
Ag^ropyron spicatum (bluebunch wheatgrass)
and Stipa comoto (needle-and-thread). The
inabilitA- to persist through successive drought
years ma\' be one of the reasons that dic()t\Ie-
donous species have shorter life expectancies
than monocotyledonous species. Related to this,
Smedlev et al. (1991) examined the water-use
efficiency of these and other herbaceous grass-
land species. Water-use efficiency, the ratio of
photosynthesis to transpiration, serves as a mea-
sure of how much photosynthetic carbon gain
occurs per unit water loss from the leaf. Dicot
herbaceous perennials had consistently lower
water-use efficiencies than their monocot coun-
teq^arts (Fig. 12). The differences in intrinsic
water-use ef^ficiencv within this life form maybe
a major contributing factor to the shorter life
expectanc) in dicot herlxiceous species. Consis-
tent with this pattern, Smedley et al. (1991)
observed that wat(^r-use efficienc\- of annual
species is significantK' lower than that of peren-
nial species in grasslands along the lower por-
tions of the canyon. The\' also obsened that
perennials which persist longer into the summer
drought period have higher water-use efficien-
cies than those species that became dormant in
late spring. During 1988-90, precipitation was
unusualK- low. The effects of the three-year
drought are now seen in Canibel oak and
bigtooth maple at their lower distribution limits,
especialK- on shallow soils, where stem dieback
has become pre\alent.
114
Great Basin Natuhalist
[Volume 52
10 cm
March
April
Fig. 13. Heiglit of Ci/iiuijiti'ni.s lunfiipcs ahoM^ tlic- u;ii)uik1 siiriace at differt- nt
Afler'wVrketal. (19.S6)'.
on til
May
urm\i till- .spriii<j; tjrowinfj .season.
Ehleringer (1988) examined leaf-lex'el
adaptations of plants along the entire elevational
transect within Red Butte Canyon. This stud\'
focused on determining patterns of leaf angle
and leaf absoiptance variation among species
within communities exposed to different degrees
of drought stress. Increased leaf angle and
decreased leaf absoq^tance reduce the solar
energ)' incident on lea\es and are \'iewed as
mechanisms for both reducing leaf energ\' loads
(reducing leaf temperature) and increasing
water-use efficienc\'. Along a transect from
grassland through coniferous forest, \'ery few
plant species exhibit any significant changes in
leaf absoiptance. However, leaf angles among
species become progressively steeper in drier
habitats. This pattern is consistent with the
notion that as plants are exposed to progres-
sivelv drier en\iroiunents, the general adaptixe
response of species within the communit>- is to
incnnise leaf angle, thereby rechicing incident
solar radiatioji levels.
In the grasslands on the lower portions of
Red Butte Canyon is a most unusual plant spe-
cies, Cijmopfcnis lon^ipes (long-stalk spring-
parsley). Sometinu^s knowm as the "elevator
plant," C. I()i}<i^ij)cs is a prostratt^ lu>rbac(n)us
perennial with an elongating pscudosca[)e (a
scape is a leafless flowering stalk arising froiu
ground level; the pseudoscape is an elongation
of the leaf-bearing stem in the retnon between
the roots and existing leaves). Other
(-ijmoptcnis species also have a pseudoscap(\
but in none of the other species is it as well
dexcloped as in C. loii^ijx's. In spring, solar
heating of the ground surface increases soil and
leal temperatures and can n^sult in moderateK'
warm knif temperatures (3()-.35 (]). These tem-
peratures are substantialK' higher than the opti-
mimi photosvnthetic temperature for the eleva-
tor plant and result in both a decreased
photo,s\nthetic rate and a decreased water-use
efficiencN' (Werk et al. 1986). To increase both
the rate of photosvnthetic carbon gain and
water-use efficiency, the pseudoscape elongates
as spring temperatures progressiv^ely increase
(Fig. 13). The result is that what was once a
prostrate canopv is elevated abo\e the warm soil
surface and now exposed to cooler air tempera-
tures abo\e the ground surface. Werk et al.
(1986) showed that the rate at which the
psuedoscape elongates is dependent on the rate
of soil-surface heating. Plants from protected or
north-facing sites elongate less than those from
exposed, southerly sites.
Donovan and Ehleringer (1991) examined
relationships between water use and the likeli-
hood of establishment b\' common shnib and
tree species in the lower portions of Red Butte
Canyon. They obsen^ed that photosvnthesis is
greater in seedlings than in adults throughout
most of the growing season, but that water stress
and water-use efficiencv' are lower in seedlings.
Seedling mortalit\ in several of the species is
associated with highei- water-u.se efficiencies,
suggesting that mortalitv' seU^ction occurs with
greater fr(H|uencv in seedlings that are conser-
vative in their water use before tlun ha\ e estab-
lished sufficiently deep roots to suni\ c the long
stunmer drought period.
Few studies have addressed ecophvsiologi-
cal as])ects of riparian ecosvstems in the Inter-
mouutain West. This is somewhat surprising
since riparian ecos\ stems are most often among
the first to be damaged bv human-related activ-
ities, Irom outdoor recreation to water
1992]
Ri<:n BuTTK Canyon Heseaiu:ii Natural Area
115
g
c5
L_
(D
Q.
O
O
CO
c
o
â– D
>^
X
o
CD
03
X
-50
-70
-90
O -110
-130
â– 150
-| r
A
^'-E^'
â– D Acer grandidentatum
• o Acer negundo
A Quercus gambelii
.a
o
precipitation
stream water
ground water
.%",S^*^„%o^ ^
o oo
J L
J L
12.5
25
37.5
50
DBH of main tree trunk, cm
Fig. 14. Hydrogen i.sotope ratio of stem waters ot tliree eoninion streainside luul adjacent nonstreaniside tree species
in Parle\s Fork oi Red Butte Canvon as a function of the diameter at breast height ol the main tnuik. Plotted as gray bars
are also the h\(b-ogen isotope ratios of the tluee possible water sources for these plants: local precipitation, stream water,
and groundwater. Open symbols represent streamside phuits and closed symbols represent nonstreaniside plants. .Adapted
Irom Dawson and EhlenniTer (1991).
iiiH)()tin(lnient to grazing. Red Butte Canyon, a.s
one of the few remaining riparian systems in the
Intermountain West not severely impacted h\
hiiuuin actixities, is ideal for studies of the adap-
tations of riparian plants and for comparatixe
.studies of .species .sensitixities to human-related
actixities.
in a recent studx Daxxson and l^lileringer
1 1 992) examined xvater sources used by riparian
plants species. In their study, plants xx'ere segre-
gated according to microhabitat antl size:
streamside xersus nonstreaniside and juxenile
xersus adult (based on diameter at breast
height). Their results xvere ratluM- startHng and
suggest that a uexx' per.spectixe is necessan'
xxhen exaluating riparian communities, their
establishment potentials, and their sensitixitA' to
disturbance. Dawson and Ehleringer (1991)
used hydrogen isotope anah'ses of stem xxaters
to determine the extent to xx'hich different cat-
egories of riparian trees utilize stream xx'aler,
recent precipitation, or groundxxater. I lydrogen
isotopes are not fractionated b\' roots during
xxater uptake; therefore, the hydrogen isotope
ratios of stem xxater xxill reflect the xxater
sources currently used by that plant. Rain,
groundxxaters, and stream xvaters differ in their
hxdrogeu isotope ratios, proxiding a signal dif-
ference that could be detected bx' stem-xx'ater
analxses. Daxx'son and Ehleringer (1991)
obsei-xed tliat among matui(> tree species none
xxere directlx using stream xx'ater (Fig. 14). All
xx'(M-e using waters from a nuich greater depth,
x\ Iiich had a hxdrogen isotope ratio more nega-
tixc than either stream xxater or precipitation.
Young streamside trees utilized stream xxater,
but onlx when small. Young trees at nonstream-
side locations utilized precipitation, haxing
access to neither stream xxater nor deeper
groundxxater. One possible reason that stream-
side trees max not depend on stream xx'ater is
that this surface xx-ater source ma\" occasionallx'
drx up during extreme drought years and
become unaxailablc^ to these trees; another is
that stream chaimels occasionally change their
course, and dependence on sinface moisture
xx'ould then result in iiu-reased drought stress
and likely increased uiortalitx" rates. The long-
term stream dischariie rates suggest that stream
116
Great Basin Naturalist
[Volume 52
water ma\' be less dependable than deeper
groundwater sources (Fig. 6).
Man\' plants do not contain both male and
female reproductive structures in their flowers,
but are present as either male or female plants
(dioecy). Freeman et al. (1976, 1980) noted that
dioecy is a common feature of plants in the
Intermountain West. Furthermore, they
obsened that the two sexes are usually not ran-
domly distributed across the landscape. Rather
there is a spatial segregation of the two sexes
such that females tend to predominate in less
stressful microsites (wetter, shadier, etc.),
whereas males occur wdth greater frequencies
on more stressful sites (drier, sunnier, saltier,
etc.). In Red Butte Canyon, Freeman et al.
(1976) investigated spatial distributions of Acer
lu'f^iindo (boxelder, a riparian tree) and Thalic-
tniDifeiulh'ti (Fendler meadowixie, a perennial
herb). In both species, there was a strong spatial
segregation of the two sexes.
Dawson and Ehleringer (1992) have fol-
lowed up on the initial obseivations of spatial
segregationin Acer negimdo (boxelder), seeking
to determine whether intrinsic physiological
differences among the sexes may contribute to
plant mortalit)' in different microsites. They
observed that female trees have significantly
lower water-use efficiencies than male trees on
both streamside (where female predominate)
and nonstreamside locations (where males pre-
dominate). Male trees exhibit a higher water-
use efficiency in drv sites than in streamside
locations, but female trees exliibit no such
response across microhabitats. The lack of a
change in water-use efficiency b\' female trees
on dr\', nonstreamside locations ma)- contribute
to an increased mortality rate, which then
ultimately results in a male-biased sex ratio at
these .sites.
Mammalian Fauna
The mammalian fauna of R(^d Butte Canyon
is remarkably diverse, due in part to the altitu-
dinal gradient and mmierous small patches of
various plant conununities indigenous to the
area. A particularly rich small mammal fauna is
associated with the patches of riparian habitat
along Red Butte Creek and its tributaries. Prior
to the iim-off of 1983, riparian habitats were
much more extensivek dexeloped than at pres-
ent. Numerous marshy meadows existed in
association with large, actixe l)ea\er dams prior
to 1982. The loss of acti\e beaxer dams in the
early 1980s has doubtless greatly reduced the
populations of small mammals that are
restricted to the mesic-marshy habitats of the
canyon.
Nonetheless, based on the altitudinal gradi-
ent and vegetational diversity of Red Butte
Canyon, a total of 51 species of mammals should
hyj^othetically occiu" there. Below is a list of the
39 species of mammals knowni to occur in Red
Butte Canyon.
I NSKCTIX'OKA — SOHICIDAE
So rex palustris
water shrew
Sorex vagmns
wandering shrew
So rex cinereus
masked shrew
CHIROPTEKA — VESPERTILIONADAK
Eptesiciis fuseiis
Lagomorpha — Leporidae
Lepiis townsendi
StjJvilagus mittallii
big brown bat
white-tailed jaekrabbit
Nuttall cottontail
RODENTIA — SC1URID.\E
Tainiascinnis liudsonicus
red sfjuirrel
Mannota flaviventer
yellow-bellied marmot
Speniiophihis annatus
Uinta ground squirrel
Spermophihis variegoftis
rock squirrel
Eutamias ininiinus
least chipmunk
Glaticomijs sabriniis
northern living squirrel
RODENTIA — GeOMVIDAE
Tfioinoini/.s t(dpoidcs
northern pocket gopher
Tlioinotnijs hottac
RODENTlA — CaSTORIDAE
botta pocket gopher
Castor canadensis
beaver
RODENTIA — MURIDAE
Reithrodontoinij.s megaloti.s
western hanest mouse
Peronnjsctis maniculatu.s
deer mouse
Peroiui/sciis hoijUi
Clcthrionomys gapperi
Ondatra zihetlucns
bnish mouse
red-backed \'ole
muskrat
Phenacomtjs intenncdht.s
heather \ole
Microtiis niontantis
montane vole
Microtus longicandiis
long-tailed \ole
Arv'icola ricliard.soni
water \ ole
RoDENTiA — Zapodidai:
Zapu.s princeps
Rodentia — Eretuizontida}-:
Erethizon dorsatuni
western jinnping mouse
porcupine '!
Carni\ora — Canidae
Canis latrans
coyote
Ca RN I\'0 lU — P ROCYON ID AE
Bassarisciis astutiis
ring-tailed cat
Procyon lotor
racoon
CaRNI\OR.A — MUSTELIDAE
Mtistela frenata
long-tailed weasel
Mii.stela cnniiwa
ermine
Mustela vison
mink
Taxidea taxiis
badger
Mephitis mephitis
striped skunk
Carninoiu — Fei.idae
Lynx nifiis
bobcat
Fells concolor
mountain lion
ARTlODACmLA — CER\aD.\E
CeiTus canadensis
Ochcoileus hem ion us
elk
mule deer
Alces anwrieanus
moose
1992]
Red Butte Canyon Research Natural Area
117
Some of the larger species ha\e been
observed only occasionally, such as the bobcat,
mountain bon, and moose. But others such as
the mule deer, elk, and coyote are obsen'ed with
high fre(juenc\' at some seasons. A rather rich
rodent fauna inhabits the canyon, with many of
the species preferentially occupying the moist
riparian communities of grasses, forbs, and
shrubs. Thus, the red-backed vole, heather vole,
montane vole, long-tailed xole, water vole, and
jumping mouse are \irtuall\' restricted to the
small mesic meadows along Red Butte Creek
and its tributaries. Similarlv, the three species of
shrews in the canvon are distributed almost
exclusively in the riparian habitats.
In some larger meadows, such as along Par-
leys Fork and at Porcupine Gulch, the microtine
rodents are distributed in a strongK' zonal pat-
tern. Long-tiiiled voles are found in the driest
parts of the meadows, montane \ oles in the
more mesic areas where grasses, sedges, and
forbs comprise a diverse community, and water
voles in the immediate streamside area, their
burrows often entering the bank at the waters
edge. Red-backed voles and heather voles are
t\picalK' found around the bases of willows in
the meadows, as well as around the edges of
conifers at higher elevations.
A few species are found onl) at higher eleva-
tions in association with Pseudotsuga menziesii
(Douglas-fir) and Popiihis trcmuloides (aspen).
These include the red squirrel, Uinta ground
squirrel, yellow-bellied marmot, and least chip-
munk. The oak-mountain mahogany zone
seems to be the preferred habitat of the rock
squirrel and perhaps the ring-tailed cat as well.
Sexeral dissertations dealing with the ecolotA"
and plnsiologiciil adaptations of shrews, microtine
rodents, and jumping mice have utilized studv
sites in Red Butte Canyon (Forslund 1972,
Cranford 1977).
A\'iAN Fauna
In his studv of the birds of Red Butte
Canyon, Perr\- (1973) found that 106 species
occurred in the area during his studv. Of these,
32 species are penuanent residents and 44 are
summer residents. The remainder (30) are
migrants or winter residents. The permanent
resident birds include:
F.\LCONIFOKMES — ACCIPITRIDAE
Accipiter gentilis Goshawk
Accipiter striatus Sharp-shmned Ha\\k
Accipiter cooperi Cooper's Hawk
Gai.i.ifohmks — Tithaonidaf:
Dciulragapus ohscu nts
Boiuisd mnhclltis
GaLLIFOKMKS — PllASIAMDAK
Lopliortijx califoniiciis
Ph as ian u .s colcli i ctis
Alcctoris graced
Stricifokmks — Stri(;idae
Otiisflatniiwoltis
Btiho virginianiis
Asia otus
Coa\CIIFORMES — Au.edinidaf.
Mcgaccnjlc ah yon
PiCIFOKMES — PiClDAE
Colaptes cafer
Sphyrapicus varius
Dcndrocoptis villosus
Denclrncopus puhescens
PaSSERIFORMES — COR\ID\E
Cyanocitta stclleri
Apheloconui coenilescens
Pica pica
PaSSERIFORMES — PaRIDAE
Pants atricapilliis
Panis aanJ)eli
Psa It rip a nis m i niu ms
PASSERIFORMES — SlTTIDAE
Sitta canadensis
PaSSERIFORMES — CeRTIIIIDAE
Ccrthia familiark
PaSSERIFORMES — CiNCLIDAE
Cinclus mexicanus
PaSSERIFORMES — TURDIDAE
Myadestes townsendi
PaSSERIFORMES — SYL\IID.\E
Regiihis satrapa
PaSSERIFORMES — STURMDAE
Sturnns vulgaris
PASSERIFORME.S — ICTEHIDAE
Stiimella neglecta
Passeriforme,s — Fhincillidae
Ca qwdaciis mexica nus
Spinas pinus
jiinco orcganns
Blue Grouse
Ruffed Carouse
C'aliforuia ^uail
Riug-neeked Pheasaut
Chukar
Flammulateil Owl
Great Homed Owl
Long-eared Owl
Belted Kingfisher
Red-shafter Flicker
Yellow-bellied Sapsucker
Hair\' Woodpecker
Downv Woodpecker
Steller's Ja\
Scnib JaN'
Magpie
Black-capped Chickadee
Mountain Chickadee
Common Bushtit
Red-breasted Nuthatch
Brown Creeper
Dipper
Towiisend's Solitaire
Golden-crowned Kinglet
Stiirling
Western Meadowlark
House Finch
Pine Siskin
Oregon Junc(j
In addition to the species that are permanent
residents in Red Butte Canvon, the following
list of summer residents represents .species thiit
probably also nest in the camon:
Anseriformes — Anatidae
Anas platyrhtpiclios
Falconiformes — .'\(x:ii'itridak
Biiteo jainaiccnsis
Acjuila chn/saetos
F AI ,C:ON IF( )R M ES — FaLC:ON I DAE
Falco sj)arcerius
ClIARADHIIFOR.MES — ScOU)I'ACIDAJ-:
Aciitis nuictdaria Spotted Sandpiper
COLUMBIKOHMES — COLL.MHIDAK
Zi'naidnra macraura
Apodiformes — Tr(k:iiii.idae
A rch ill )clt us alcxandri
.Mallard Duck
Red-tailed Hawk
(Golden Eagle
Sparrow I lawk
Mourning Do\e
Sclasf)lu>nis platyccrcus
P.VSSERIFOHMES — TlR^NNIDAE
Empidonax ohcrholseri
Black-chinned
Hummingbird
Broad-tailed
Hummingbird
Dusk-x Flycatcher
118
Great Basin Naturalist
[Volume 52
Empidonax diffirilis Western Flyeatclier
Coittopiis surdidulus Western Wood Peewee
PaSSEHIKORMES — HiKUNDIMDAK
Tacliijcincia tluilassina N'iolet-green Swallow
Iridoprocnc hicolor Tiet> Swallow
Rifxiiia riparia Bank Sw;i!low
Stel^idof)tcn/x nificollh- Rough-\\ino;ecl Swallow
Iliniiidc nisticti Bam Swallow
PeiroclichdoH piirrlumotii (."lift Swallow
Passf.hifohmfs — TK(x;i.t)i)rrii)AK
Tn)<ilodif1es acdon Honse Wren
Salpimics ohsulctus Rock VWen
PaSSEKIFOKMES — TUHDIDAF
Ttirdtis iiii^ratoriiis Robin
Hi/lorirhia ^iitlala Hermit Thnish
Ilijlocicida nstidatti Swainson's Thnisli
Sinlia ciirnicoidcs Monntain Bluebird
PaSSEKIFOHMES — SVIMIDAF.
Polioptilci cacndca Blue-gra\ Cinateatcher
PaSSEKIFOHMES — VlHEONlDAE
Virco ^dvtis Warbling Vireo
PaSSERIFORMES — P.\i^ULIDAE
Vermivura celata Orange-crowmed Warbler
Vennivora virginiae Virginia's Warbler
Dc'iidwica pftcchia Yellow Warbler
Deiidroica andtd)oiii Audubon's Warbler
Opomrrm tohnici VlacCIillivrav's Warl^ler
Wilsonia pusilla Wilson's Warbler
PaSSERIFORMES — ICTERIDAE
Ictcnis hullickii l^ulloek's Oriole
Molothnis alcr Brown-headed Cowbii'd
PASSERIFORMES — TllRAUPIDAE
Pirani^fi hidoviciana Western Tanager
PaSSERIFORMES — FRINCnLElOAE
Pliciiticiis iiicldiKH cplKiliis Black-headed Cirosl)e;ik
Ptisseriiia innociui La/.uli Bunting
Caiynddcus cassinii (>'assin's Finch
Spiniis tristis American Croldtinch
Cdilonira cldoruni (ireen-tailed Towhee
Pipilo crytlirotlxihiuis Rufous-sided Towhee
Pooecetes '^rainiiifiis Vesper Sparrow
Jtinco caniccps (irav-headed Jmico
Spizella pdsserina (Shipping Sparrow
Melospiza inelodia ^"'igi Sparrow
Role of Research Natural Areas
Research Natural Areas proxide several spe-
cific acKautages to the natiou's scientific
comniunit)', which are tvpically not othenvise
available. These include potential use of an area
that has had minimal human interference and
has a reascjnable assurance of long-term exis-
tence, and the potential association and interac-
tion of scientists from different disciplines
leading to discoveries unlikely to occur without
such an association. Conducting research at
common locations is kev to developing these
interactions. Research Natural Areas not onlv
assist in the progress of basic science, but also
provide federal and state agencies with informa-
tion upon which to base management decisions.
The melding of ecosvstem presenation and
research on basic ecological processes at
Research Natural Areas provides numerous
valuable options to societv. The Red Butte
C'anvon RNA serves this puipose well. Although
initially affected bv human activities during the
early settlement of the Salt Lake Valley, the
canyon was soon set aside bv the federal govern-
ment and has now had nearlv a centuiy to
recover (tliough the loss of beaver represents a
significant impact to the ecologv of the riparian
ecosystem). Other canyons in the \Vasatch
Range have not received equivalent protection.
As we move into the twenty-first centuiy,
there will he increasing pressure to understand
the dynamics of ecological systems and man s
impact on ecological processes. Maintained as a
protected watershed, the Red Butte Canyon
RNA provides a unique oppoitunitv' for
addressing these important issues to human
societ)' and to the presenation of our environ-
ment. Unprotected, it is an invaluable resource
lost forever.
Federal laud-management agencies have
been developing a national system of Research
Natural Areas since 1927. More than 4{)() areas
have received this designation nationally. Since
inception of the RNA Program, there have becMi
two priman puqx).ses for Research Natural
Areas:
1. to presene a representative arrav of all
significant natural ecosystems and thtii-
inherent processes as baseline areas; and
2. to obtain, through scientific echication and
research, information about natural svstem
components, inherent processes, and com-
parisons with representative manipulated
svstems.
Literature Cited
An asterisk (°) refers to studies conducted in
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this manuscript.
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Rkd Butte Canyon Reskarch Natuhai. Area
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Appendix
Nomenclatural Changes in the Flora,
1971-1990
The following is a list of nomenclatural and
orthographic changes made since pubUcation of
the Vascular Flora of Red Butte Canyon, Salt
Lake County, Utiili (Amow 1971). Family
names of flowering plants are changed to accord
with those used by Cronquist (1981). All other
name changes are contained in Welsh et al.
(1987) unless otherwise specified.
Amaranthace.\e
Anuiranthus graecizans of Americiui authors, not L. = A.
hlitoides Wats.
AMARYLLIDACEAE = LiLIACEAE
Brodiaea douglasii Wats. = Triteleia grandiflora Lindl.
Anacardiaceae
Blius radicans L. = Toxicodendron rijdhergii (Small)
Greene
Berberidaceae
Berheris repens Lindl. = Mahonia repeiis (Lindl.) G. Don
Boraginaceae
Cnjptantha nana (Eastw.) Pays. = Cnjptantha humilis
(Gray) Pays.
Hackelia jessicae (McGregor) Brand = H. micrantha
(Eastw.) J. L. Gently
Lappula echinata Gilib. = L. squarrosa (Retz.) Duniort.
(Weber 1987)
Cactac'eae
Opuntia aitrea Baxter, misapplied to O. macrorhiza
Engelm,
Caryophyix.'^ceae
Cerastium vulgatum L. = C.fontanuin Baumg.
Stellaria janwsiana Torr. = Pseudostellaiia jamesiana
(Torr.) Weber & Hartman (Weber and Hartinan 1979)
Cel,\straceae
Pachistinui = Paxistima
ChEN()POL5IACEAE
Sal.sola kali L. = Sal-sola iherica Sennen &; Pau
CX)MP0SITAE = ASTEIUCEAE
A-iter chilensis Nees = A. ascendens Lindl.
Haplopappus n/dhergii Blake = H. watsonii Gray
Lactuca pulchella (Pursh) DC. = L. tatarica (L.) C. A.
Mey
Matricaria matricarioides (Less.) Porter = Chamomilla
suaveolens (Pursh) Rydb.
Solidago nemoralis Ait. = S. sparsiflora A. Gray
S, occidentalis (Nutt.) T. 6c G. = Eutluimia occidentalis
Nutt. (Sieren 1981)
Taraxacum laevigatum (Willd.) DC. = T. officinale
Wiggers (Weber 1987)
1992]
Red Butte Canyon Research Natural Area
121
Vit^uicra inultiflora (Nutt.) Blake = HcUomrris tnullifliu-a
Niitt.
CORNACEAE
Comtts stolonifcra Michx. = Coniiis scrirca L.
Ckuc:ikkiuk = Bkassicaceae
Arahi.s divaiicar-jui A. Nels. = A. Iiolhorllii Homem.
Rorippa islandica (Oed.) Borb. = R. palnstris (L.) Besser
R. tninaita (Jeps.l Stuckev = R tciicrriitKi (»reene
CUSCUTACEAE
Cusctita campcstiis Yiinck. = C. pciifdfiona Engelm.
Cypehaceae
Carex utriculato Boott = C. rastmta Stokes
Gramineae = PoACEAE (Amow 1987)
Agroprjron caninum (L.) Beaiiv. = Eh/iim.s tiiiclit/caulns
(Link) Shinners
A. dasijstdcJujum (Hook.) Scribn. = Eh/iims lanceolatus
(Scribn. & Sin.) Gould
A. ititcniirdiuin (Host) Beaux'. = Eh/uins hi.spiilits (Opiz)
Meld.
A. siuithii R\dl). = Ehpiius sntithii (R\db.) (iould
A. spicatum (Pursh) Scrilin. = Eh/mtis spiciittis (Pursli)
Gould
Agrostis alba L. = A. stolonifcra L.
A. semiverticillata (Forsk.) C. Christ. = Poli/pofioit scmi-
verticillatHS (Forsk.) Hylander
Arktida loiigi.scta Steud. = A. purpurea Nutt.
Bromus hrizacfonnis Fiseh. & Mev- = B. hhzifonnis
B. commutatiis Schrad. = B. japonicnsThymh.
Gltjceria data (Nash) M. E. Jones = G. striata (Lain.)
Hitehc.
Hesperochloa kiiigii (Wats.) Rvdb. = Leucopoa kingii
(Wats.)W. A. Weber
Kiieleria cristata Pers. = K macrantha (Ledeb.) Schiilt.
Onjzopsis hijmenoides (R. & S.) Ricker = Stipa
lupncnoidcs R. & S.
Poa saiidbergii \'ase\- = P. secunda PresI (Amow 1981)
Sitanioii juhatum ]. G. Smith, misapplied to Eltpnus
ehjinoidcs (Raf.) Swezev
Stipa occidi'ittalis Thurb. = S. ueisouii Scribn.
JUNCACEAE
Junais bait ic US W'iWd. = J. ar(tki/.v Willd.
J. traciji R\-db. =/. cnsifolius Wikst.
LaHLYPAI-; = L'WIIACKAE
Moldavica parviflora (Nutt.) Britt. = Dracocrpluiluin
paniflonun Nutt.
Lkcuminosae = Faba(;E;VE
Mor.\{:eae = Cannabaceae
Hnmidus lupulus L. = H. amcricanus Nutt.
Ona(;iu(:e.\e
EpilohiuDi pauiculatu)n T. & G. = £. braclniraqyuin Presl
E. ivatsoiiii Baibev = E. ciliatuin Raf.
Oenothera hookeri T & G. = O. data H.B.K.
Zaudincria iiaiTcttii A. Nels. = Z. latifolia (Hook.)
Greene
Orobanchaceae
Orobandw califoritica Cham. & Schleclit. = O.
conpnbosa ( Rydb. ) Ferris
Polemoniaceae
Iponiopsis aoare<iata (Pursh) \. Gnuit = Gilia a<i^regata
(Pursh) Spreng.
PoLYPODi.At;EAE. as it occurs in Red Butte Canvon, is now
dixdded into the following families (Trvon and Tr\c)n
1982):
DennstaEDTWCEAE, of which the genus Pteridiuin is a
member
Dryopteridaceae, which includes the genera Ci/stoptcris
and Woods ia
Ctjstopte'ris fragilis (L.) Benih. is now known to include
two taxa (Leilinger 1985), of which only C. tenuis
(Michx.) Desv. occurs in Red Butte Canvon.
Ranunculaceae
Ranuncuhis longirostris Godron = R. aquatilis L.
R. tcsticulatus Crantz = CeratocepJudus oiihorenis DC.
(Weber 1987)
S.\lk;aceae
Salix rigida Muhl. = S. lutea Nutt.
Saxifragace.\e:
Lithophragnia bulbifera R\db. = L. ^ahra Nutt.
Scrophuiv\ria(:eae
Castilleja leonardii Rvdl). = C'. rhexifolia R\db.
Tamaricac;e.\e
Tamarix pentandra Pall. = T. raniosissiina Leck'b.
Umbellifer.'^e = Api.\(;eae
Cictita dou<!jasii (DC.) Coult. & Rose = C nuiculata L.
Lonwtium nuttallii (Cnw) Macbr. = L. /cZ/ig// (Wats.)
C'roiui.
Creat Basin NatiirtJist 52(2), pp. 122-130
INFLUENCES OF SEX AND WEATHER ON MIGRATION OF
MULE DEER IN CALIFORNIA
Thomas E. Kiiccia
Abstiuct. — I examined differentes In sex and influences of weather on timing; and patterns of migration of RockA'
Mountain mule deer (Oclocoilcus h. Iwinionus) in the eastern Sierra Nevada, (>alifoniia, during 1984-87. Deer initiated
.spring migration from the v\anter range at about tlie same time in all )ears and made extensive use of holding areas at
intermediate ele\ations. Radio-telemetered deer showed strong fidelitv^ to summer riuiges o\er as manv as four years. Fall
weather produced different patterns of fall migration. Storms during October produced a pulsed migration, in which most
animals migrated to the winter range during or soon after the storm; in a year without a storm, fall migration was gradual.
Despite the influence of storms on the pattern of ftdl migration, the median date of fall migration bv females did not var\-
over vears; howe\'er, among males it was later in a year without fall storms.
Kcij words: mi^ratioiK mule deer. Otlocoileus hemionus, sex differences, icetitlier radio teleinctn/. C'alifoniia.
Seasonal migration is common amongawdde
variety of vertebrates (Baker 1978), including
large terrestrial mammals (McCullough 1985,
Fn'xell and Sinclair 1988). Migration ultimately
contributes to individual reproductive success
(Baker 1978). Proximally, however, migration is
related to the seasonal availabilitv' of resources
(Sinclair 1983, Garrott et al. 1987). Migration is
a common phenomenon among mule deer
{Odocoileus lieiniontis) in the mountainous
western United States, and various studies have
described aspects of nuile deer migration (Rus-
sell 1932, Leopold et al. 1951, Gniell and Papez
1963, McCullough 1964, Bertram and Rempel
1977. Garrott et al. 1987, Loft et al. 1989).
Ilowexer, questions remain as to the influence
of proximate factors, especially weather, on the
timing of migration. In addition, because .stud-
ies of mule deer involving radio-telemetn' rarely
have inchuk'd males (e.g., Garrott et al. 1987,
Loft et al. 1989), little is known of differences
between the sexes in migration patterns.
My objectives were (J) to describe the
timing and pattern of seasonal migration of
mule deer in the ea.stern Sierra Nevada, C'alifor-
nia; (2) to test the hvpotheses that there were no
differences b)- sex or year in the timing and
pattern of luigration and degree of summer-
range site fidelity-; and (3) to relate ob.sc'ncd
migration patterns to other aspects of tlie (X'ol-
ogy- of these animals.
Study Are.\
The Sierra Nevada is a massive granite block
tilted toward the west, extending for 600 km in a
generally northwest-southeast direction (Storer
and Usinger 1968). The west side of the moun-
tain range slopes gradually for 75-100 km, from
the foothills near sea level to the crest at 3000-
4500 m. The eastern Sierra Nevada is more
narrow and steep than the west side, with fre-
quent elevational changes of 3000 m in <10km.
A population of 3000-6000 Rocky Mountmn
mule deer (Odocoileus Ji. Jieniioiuis) wanters at
the base of the eastern escaipment of the Sierra
Nevada in Round Willev. Invo and Mono coun-
ties, California, about 15 km west of the town of
Bishop (Fig. 1). An area of about 90 knr of
Roinid \^alley is used bv' mule deer as winter
range, at elevations from about 1450 to 2100 m.
Pine Creek forms the dividing line between
what is termed the Shetwin Grade (SG) deer
herd to the north and the Buttermilk (BM) herd
to tht" south. These deer are hunted under
bucks-onlv regulations, and posthunt adult sex
ratios of 7-12 males: 100 females occm"red
dvning this studv" (California Department of
Fish and (rame. Bishop, California).
As winter storms h'oni the Pacific Ocean rise
up the western slope of the Sierra Nevada, thev
ck^posit rnoistiu'e, leaving a mucli more arid riiin
sliadow on the t>ast side. Precipitation in the
nepartincnt oC For.sin .nul Kcsourcc VIaiiui;i-iii.-nt. .iiul Vli
)t\rrt,-l)r.i(.-'/.(K)l(>i,r\. Iniu-rsilN <if CalilDmia, Brrk.'l.'v. Calilomia 94720.
122
1992]
Mk;iutiunof Muli<: Dkkk
123
_OlVENs
CROWLEY LAKE
Fig. 1. Map of the stuil\ aiva sliow ing tlic dcc-r winter range as the shaded area ni Konnd \alley; the crest of the Sierra
Nevada is from nortliwest to southeast, witli elevations (m) of" selected peaks and major passes.
area ranges from an animal mean of 14.5 cm at with ai)ont 757c of the annnal total oc'cnrring
the Bishop aiqx)rt at 1240 m to 40.6 cm at between November and March. Summers are
2860 m in Pine Creek Canvon (Vaughn 1983, hot, witli davtime temperatures in Jul\ often
National Oceanic and Atmospheric Administra- >37 C. Jannarv is the coldest month, with
tion 1987). Precipitation is strongly seasonal an a\erage temperature of 4 C and frequent
124
Great Basin Naturalist
[Volume 52
nighttime lows of <-15 C. Potential evapo-
transpiration is 66.8 cm, or more than four times
the mean precipitation.
Vegetation on the winter range is t\|:)ical of
the Great Basin Desert and conforms to the
sagebrush belt of Storer and Usinger (1968).
Shnibs are dominant, and blackbmsh (Coleoayne
ramosissiina), rabbitbnish (Clin/sotJunnnus
spp.), big sagebnish {Artemisia trident at a), and
antelope bitterbrush (Purshia trident at a) are
most common. Deer summer ranges are on
both sides of the Sierra crest, at elevations from
about 2200 to >3600 m (Kucera 1988), and
include the sagebrush, Jeffrey pine {Piniis
jeffretji). lodgepole pine {P. murraijana)-red fir
{Abies ma^nifica) , subalpine, and alpine belts
(Storer and Usinger 1968).
Livestock use of deer winter range was light,
consisting of 129 animal-unit-months of use by
cattle, restricted to part of the SG range from
1 April to 15 October (U.S. Department of the
Interior 1990). Use of deer summer areas by
livestock (including horses, cattle, and sheep)
varied from ver\' heavy in more accessible loca-
tions on the east side of the mountain range to
none at higher elevations and more remote
areas.
Methods
Fieldwork was conducted from Januar)' 1984
through Mav 1987. Deer were captured on the
winter range Januar)' through March 1984 and
January and February 1985 with a variet\' of
methods including Clover traps (Clover 1956)
baited with alfalfa, drive nets using a helicopter,
and remotelv triggered drop-nets; net guns fired
from a helicopter and tranquilizer darts also
were used to capture selected males. Deer cap-
tured in 1984 in Clover traps were chemicalK
immobilized with Rompon (xylazine hvdrochlo-
ride), the effects of which were reversed with
yohimbine after handling (Jessup et al. 1985).
Deer were captured also during May 1984 and
1985 witli tran(|uilizer darts on a spring migra-
tion "holding area ' (Bertram and Rempel 1977)
about 50 km north of the winter range. This is
an area where deer congregate for 2-6 weeks
before continuing to areas occupied during the
summer.
I fitted 8 males and 9 females from the BM
winter range, 7 males and 10 females from the
SG winter range, and 10 females captured on
the spring holding area with radio collars
(Telonics Inc., Mesa, Arizona). All deer were
<2.5 years of age. I attempted to distribute cap-
ture efforts throughout accessible areas to min-
imize biases in the marked sample. I selected
females for telemetry to include all age classes
of adults; however, I selected males to receive
radio collars on the basis of large size and rela-
tivel)' old age. I excluded smaller, younger males
because of concerns arising from body growth;
males do not approach maximal neck circumfer-
ence until about 4 years of age (Anderson 1981),
and this, combined with seasonal neck swelling
during rut, could result in injury caused by
radio-telemetry collars. Older males have
achieved nearly maximum body growth; I
allowed for seasonal neck swelling bv attaching
the nonexpandable collars with a circumference
20-25% larger than the animal's neck circum-
ference after rut, measured midway between
head and shoulders. I noticed no serious prob-
lems resulting from the use of radio collars on
male deer in this study, although after a )ear or
two, some fur appeared to be rubbed off the
backs of the necks; a similar situation occurred
with telemetered females. Collars on the males
moved toward the head when the necks swelled
during rut and hung loosely at other times.
While animals were on the winter range, I
determined at least once per week, and usually
more often, whether each radio-marked animal
was on the BM or SG winter range bv observing
the direction of transmitter signals received
from standard locations. These data were sup-
plemented bv additional radio locations and
visual locations as observers moved through the
winter ranges. During spring and fall migra-
tions, and during summer, locations of teleme-
tered deer were determined from a fixed-wing
aircraft, from a vehicle, and from the ground.
During the spring, locations were determined
several times per week until the aniniiils crossed
the crest of the Sierra. Due to the remoteness
of most summer ranges in roadless wilderness
areas, frequency of locations of animals, deter-
mined from the air and the ground, on the west
side of the Sierra Nevada was approximately
twice per month. Of 42 deer that reached
summer ranges, I located 38 from the ground.
Twenty-two deer were followed for more
than one sunmier. Of these, 10 (45%; 1 male, 9
females) were located in two consecutive sum-
mers, 9 (41%; 3 males, 6 females) in three con-
secutive summers, and 3 (14%; 1 male, 2
females) in four consecutive summers. For
1992]
Migration of Mule Deer
125
these animals I expressed ficlelih' to summer
range as the greatest linear map distance
between mean locations in consecutive sinii-
mers (1 July-7 September). During the fall,
locations of animals were monitored from the
east side of the Sierra crest at least several times
per week, and frequently daily. I could thus
determine, within several davs and often within
one dav, when telemetered deer from the west
side of the crest crossed to the east side.
I dixided annual migration into three peri-
ods: ( 1 ) leaving winter range, defined as ascend-
ing to an elexation >2100 ni; (2) crossing the
Sierra Nevada crest in spring; and (3) crossing
the crest in fall. The last two applv only to those
animals (n - 34) that summered west of the
crest. Because of logistic difficulties in locating
animals on the west side of the crest, I did not
attempt to determine precisely when animals
crossing the crest reached their summer ranges.
The steep eastern slope of the Sierra Nevada
provided the opportunity to determine the pres-
ence or absence of a radio-marked animal on the
east side with little error. In situations in which
I could not deteninine an exact date of crossing,
I estimated the date as the midpoint of the
interval in which I did and did not receive a
signal.
For analysis I determined frequencies of
movement by week during an 8-week period of
leaving the winter range beginning 1 April, a
7-week period of crossing the crest in spring
beginning 15 May, and an 11-week period of
crossing the crest in fall beginning 1 1 Septem-
ber. I used the Kolmogorov-Smimov test with
chi-square approximation (Siegel 1956) to test
for sex differences in the timing of these com-
ponents of migration. Steep mountains on the
west side of Round Valley constrained move-
ment off the winter range to northerlv or south-
erly routes; I tested for sex differences in the
direction (north or south) of migration from the
winter range with the binomial test (Zar
1984:591 ). I expressed temponil patterns of fall
migration as the percentage of radio-marked
deer in an annual sample crossing the crest
during any week. I tested for differences among
years in the largest weekly percentage crossing
the crest in any year with the Z-test (Zar
1984:396).
From April through June of 1985, 1986, and
1987, commencing as soon iis snow conditions
permitted, deer were counted from a vehicle
along a standardized route of 1 1 km that passed
through a major spring holding ari'a located 1-8
km south of the town of Mammoth Lakes,
approximately 50 km north of the winter range.
These weekly surveys began 30 minutes before
sunrise, and direction of travel was alternated
on consecutive survevs.
Daily precipitation in the fall was measured
at the U.S. Forest Service (USFS) weather sta-
tion at the Mammoth Lakes Ranger Station,
Inyo National Forest, Mammoth Lakes, Califor-
nia, at an elevation of about 2400 m. Winter
snowfall totals were from the USFS weather
station on Mammoth Mountain, at about 2940 m.
Results
Spring Migration
From 1984 to 1986 the first radio-marked
deer left the winter range during the first or
second week of April in anv vear; in the same
years the last radio-marked deer left during the
second, third, and fourth weeks of May. For
femiJes the median departure date from the
winter range was during the third, second, and
third weeks of April 1984-86, respectivelv'; for
males, the median was during the second week
of May and second and third weeks of April,
respectively. The frequency differences by sex
in vveeklv migration approached statistical sig-
nificance (X- '= 5.94, df = 2, .05 <P< . 10).
Of the 17 telemetered deer from the BM
range, 10 (3 of 8 males, 7 of 9 females) migrated
north, through the SG range, to reach their
summer range; 5 males and 2 females moved
south. Of the 17 deer telemetered on the SG
range, 15 (5 of 7 males, 10 of 10 feinales)
migrated to the north; 2 males went south.
Overall, more (P = .0003) females migrated
north (n = 17) than south (n - 2). Analysis by
herd showed a significant difference (F = .0001)
in migration direction among SG females {n - 10);
the difference among BM females (n = 9)
approached statistical significance (F = .07).
There were no significant differences among
niiiles in migration direction, either with all
males combined {n = 15, F = .196), or bv herd
(BM: n = 8, F = .22; SG: /i = 7, F = .16). Of the
10 females captured on the spring range, 4
wintered on the BM range, 5 wintered on the
SG range, and 1 died before the fall migration.
Holding Areas
After leaving the winter range, telemetered
deer moved to higher-elevation holding areas at
126
Grkat Basin Naturalist
[Volume 52
22()()-24()() 111 on the east side of tlie Sierra
Nevada. Hundreds of deer already were present
on the first road suneys of the spring, and
patterns of oecurrence were similar in all years
(Fig. 2). Largest numbers were counted in late
April and early Ma}'; numbers then decreased
through mid-Jime as deer moved to summer
rang(\s. During early spring a portion of the
winterino; animals also foraged in irrigated
meadows immediately adjacent to the winter
range in Round Valley.
Diminution of deer counted on the holding
area \vas reflected by an increase in deer cross-
ing the crest to summer ranges. Of the radio-
marked deer that summered west of the crest,
the first crossed the crest during the third or
fourth week of May in any year, and the last
crossed during the third or fourth week of June.
There were no sex differences in timing of
spring crossing (X" = 3.50, df = 2, F > .10). The
median for both sexes in all vears was the first
week of June.
The temporal uniformit)' over years in leax-
ing the spring holding area for simimer ranges
occurred despite greatly different snow condi-
tions. In the winters of'l983-S4, 1984-85, and
1985-86, the USFS recorded total snowfalls of
671, 767, and 1021 cm, respectively, on Maiu-
moth Mountain, geographically close and at an
elevation similar to the passes that migrating
deer crossed to reach summer ranges on the
western slope. Despite these differences in
snowfall and consequent snowpack at higher
{4evations, no differences in the timing of spring
migration were evident. The snowfall of winter
1 986-87 was only 246 cm, or less than one-{|uar-
ter of that of the previous year. Although the
.sample si/.(> is small, the median week that three
radio-marked males and tu^o radio-marked
females crossed the crest in the spring of 1987
was the same as the prexdons year, the first week
of June. Thus, the amount of snow on the
ground did not appear to inlliience the timing
of migration o\-er the Sierra crest in the spring.
SunmuM" Range
()1 the 32 deer captiuvd on the winter range
that reached summer ranges, 28 (87.5%)
crossed the Sierra crest and snnunered on the
west side. Sununer range locations of these
deer, plus thosc^ of deer captured on the spring
rangi\ extended from the headwaters of the
Middle Fork of the San Joacjuin Ri\-er south
throughout the upper San Joaquin Ri\(M- drain-
700
600
^
500
a>
0)
Ti
»*-
400
o
k.
300
E
3
200
100
^ I I I
1 3 Apr 3 May 23 May 1 2 Jun
Figf. 2. Nuniher of inuk' dciT fountcd Iroiii a \ L-Iiicle on
standardized weekly sin"\evs at dawn through a spring hold-
ing area near the town of Mamniotli Lakes, Mono Countv,
Cahfoniia, 1985-87. Suivevs begiui in the .spring when snow
conditions made the roads passable.
age above about 2134 m into the North and
Middle forks of the Kings River (Kucera 1988).
Two males and 4 females sunnuered on the east
side of the Sierra, from Manuuoth Pass on the
north to the North Fork of Bishop Creek on the
south. Thus, an area nearly 100 x 25 km seived
as sunuuer range for deer from the BM and SG
herds.
Sunuuer Range Fidelits'
Distances between smumer ranges of 22
tle(M' located in consecuti\e \ears averaged
0.7 km (range - 0.2-4 km) for both males (/i = 5)
and females (n - 17). Onl\ 1 deer, a female, was
>1 km from a prexions location in successive
summers; she spent her second sununer about
2.5 km from her first, and her third and fourth
about 1.5 km farther awax'.
Fall Migration
In 1984, 1985, and 1986 die first radio-
marked deer crossed to the east side dining the
first week of (October and second and fourth
weeks of September, respectively; all were
females. The last crossed during the fourth
week of October and second and fointh weeks
19921
Mk;kati()N()f Mule Dker
127
80
60
40
O
0) 20
"D
0)
1984
Deer, n = 15
Precipitation
1985 Deer, n = 26
Precipitation
r~[
/ \
; \
/ \
/ \
I \
I \
/ \ ' ^ .^
\ ' ^
\ /
\ /
-i, — /
20
— Deer, n = 16
— Precipitation
4.0
2.0
0.0
4.0 £
O
c
o
^-»
a
"o
0.0 2
Q.
4.0
11 Sep 25 Sep 9 Oct 23 Oct 30 Oct 13 Nov
2.0
0.0
Fig. 3. Percentage of telemetered mule deer per week crossing the crest ot the Sierra Nevada, ln\o and Mono counties,
California, and weekly precipitation measured at the town of Mammoth Lakes, Mono Countv, in the fall of 19S4-86.
of Noxember; all were males. In 1984 and 1985
the median week of crossing the crest was the
same for both sexes, the third and second weeks
in October, respecti\elv. In 1986 the median for
females was the third week in October, but was
tvvo weeks later for males {X' = 18.72, df = 2,
P< .001).
Length of time during which fall migration
occurred also varied among years. In 1984, 11
of 15 (73%) and, in 1985, 14 of 26 (54%) tele-
metered deer, including both sexes, crossed the
crest in a one-week period. These proportions
were not different (Z = 1.2, F > .11). Howevei;
in 1986 no more than 4 of 16 (25%) radio-
marked deer crossed the Sierra crest in any
week. This proportion was smaller than those of
the previous two years (Z = 2.45, P < .007),
indicating that in 1986 there was no mass move-
ment of deer in a short time period.
Differences among years both in timing and
in pattern of fall migration were related to the
presence or absence of major fall storms (Fig.
3). In 1984, 1.8 cm of precipitation in the form
of about 20 cm of snow was recorded on 17
October at Mammoth Lakes; no doubt snow at
the passes (400-1500 m higher) used b\- migrat-
ing deer was much deeper This storm was
accompanied by a rapid moxement oi radio-
marked deer over the crest and to the winter
range within a few davs. Earlier storms, which
resulted in virtually no snow at the recording
station, did not trigger movement. In 1985,
shortK after a storm on 7 October, there was
another rapid movement of deer o\er the crest.
The remaining deer appeared gradually on the
east side of the crest through 13 November,
when the last radioed animal, a male, migrated
over the crest following a major winter storm.
In both 1984 and 1985 1 saw dozens to hundreds
of deer migrating simultaneouslv with the tele-
metered animals, and man\' tracks and deep
trails in the snow were evident. In 1986 there
were no major fall storms. Migration was grad-
ual and unpunctuated by am rapid, mass mo\e-
ments (Fie. 3). In all cases deer returned to the
128
Great Basin Naturalist
[Volume 52
winter range (BM or SG) occupied in previous
years.
Discussion
In this study the timing of mule deer migra-
tion from the winter range did not differ among
years. This occurred despite large differences in
animal condition and vegetation growth mea-
sured on the winter range (Kucera 1988). One
explanation mav be that these deer had well-
defined spring holding areas where they could
predictably obtain nutritious forage, avciilable
even in years of hea\/y snowfall such as 1986,
when hundreds of deer were on the holding area
when counts began (Fig. 2).
Adult males may leave the winter range
somewhat later than females, as reported from
western Colorado (Wright and Swift 1942).
Given the demands of pregnancy, females might
be under greater nutritional stress than males,
and if better forage conditions exist on spring
ranges, females may tend to leave the winter
range sooner to take adxantage of them. Garrott
et al. (1987) reported that spring migration of
female mule deer in northwest Colorado varied
between years by as much as one month, and
they attributed these differences to the severity
of winters and consequent energetic demands
on deer. Bertram and Rempel (1977) reported
that California mule deer (O. h. californiciis) on
the western slope of the Sierra Nevada varied
the timing of their spring migration by two
weeks, and attributed this to differences in plant
phenology both on the winter range and along
the migration route. Loft et al. (1989) also
reported a similar relationship between initia-
tion of spring migration and anioimt of snow and
stage of plant growth in the western Sierra
Nevada.
In my study most telemetered females
migrated from the winter range to the north;
males showed no significant selection for
direction. I contend that this sex difference is a
product of local geomoipliolog)' and land man-
agement patterns. Animals moving north had
access to an extensive area of the west slope of
the Sierra Nevada on national forest lands at
elevations of 22()0-28()() m. .'\nimals moving
south had access to sunmier range in King's
('anyon National Park at higher and steeper,
and thus more barren and less vegetated, eleva-
tions (Kucera 1988). The presence of more and
better summer range to the north expkiins why
most deer of both sexes would migrate to the
north. However, those animals migrating to the
north were in areas open to hunting both on
their summer ranges and along the migration
routes. That telemetered males showed no
apparent selection for migration direction,
whereas most females migrated to the north,
probably resulted from the higher hunting mor-
talit)-' of males summering to the north, and the
absence of hunting in the national park.
Although as many males as females would be
expected to migrate to the north, the higher
mortality of adult males moving north could
expUiin the apparent pattern of no directional
preference. Because older males are dis-
proportionately reproductively successful
(Kucera 1978, Geist 1981, Glutton-Brock et al.
1982), the national park may act as a refuge for
a large proportion of the most reproductively
successful males.
Deer in this studv made extensive use of
holding areas in the spring (Fig. 2), which may
be beneficial because of higher elevation,
greater precipitation, and absence of winter
f^eeding. Vegetation in these holding areas was
largely sagebrush scrub (Munz and Keck 1959),
a common vegetation type in the eastern Sierra
Nevada. These areas are among the last large
areas with vegetation suitable for deer present
in the spring before the deer cross the Sierra
crest. Large aggregations of deer on the holding
areas may result from animals simply collecting
in these areas for several weeks before ascend-
ing over the crest. Bertram and Rempel (1977)
and Loft et al. ( 1989) described a similar pattern
of use of spring ranges in the western Sierra
Nevada and emphasized the importance of
these holding areas in providing herbaceous
forage. Further, Bertram and Rempel (1977)
reported that spring holding areas typically
occurred at the base of an abnipt elevation
change, which was true in mv studv.
Timing of movement off the holding area
and over the crest in spring did not differ among
vears or between sexes, suggesting that animal
condition or vegetation did not greatly affect
this stage of migration. The passes had snow in
all years of study when deer crossed, but snow
depths differed greatly. However, by spring
snow was consolidated, enabling deer to walk
over the surface.
In 1951 Jones (1954) found that BM deer
began moving off the winter range about 1 April,
and began crossing a nearby pass about 15 May.
1992]
MiciuTioNOF Mule Deer
129
This agrees well with the present obsenations
made more than three decades later. In the
western Sierra Nexada, Rnssell (1932), Leopold
et al. (1951), Bertram and Rempel (1977), and
Loft et ill. (1989) described spring migration as
an "upward drift" of deer, controlled by the
receding snowline and spring plant growth. My
study showed a different pattern in the eastern
Sierra Ne\ada. The upward moxement of deer
w as blocked by the abiiipt elevation change of
the mountains. On the more gentlv sloping west
side, deer can follow spring gradualK' up slope.
On the abnipt east side, the need to cross high-
elexation passes prevents such a pattern.
The strong fidelity to specific summer home
ranges shown b\- individual deer in this stucK
is characteristic of mule deer (Ashcraft 1961,
Gmell and Papez 1963, Robinette 1966, Bertram
and Rempel 1977, Garrott et al. 1987, Loft et al.
1989). With few exceptions, both males and
females returned to the same summer home
ranges, and winter ranges, for as many as four
consecutix'e years.
The temporal pattern, pulsed or gradual, of
the fall migration in the eastern Sierra Nevada
is largeK- determined by weather, particularly
snowstorms. In both years with simificant
snowfall in October, radioed deer moved rapidly
and in a pulsed fashion from summer ranges to
the winter range (Fig. 3). In a year without
significant fall storms, movement was gradutil,
and males migrated significant!)' later than
females. Previous studies discussed the relation-
ship of snow.storms to fall migration (Russell
1932, Dixon 1934, Leopold etal. 1951, Richens
1967, Gilbert et al. 1970), although some cases
were based on anecdotal evidence. Bertram and
Rempel (1977) stated that deer on the west
slope of the Sierra Nevada moved in anticipa-
tion of fall storms, but I found no evidence of
this. Garrott et al. (1987) speculated that in
northwest Colorado deer moved not because of
snow, but to maximize the qualitv of their diets
prior to winter. Differences in details of deer
migration apparent between mv studv and stud-
ies in the western Sierra Nevada and in north-
west Colorado indicate that deer migration can
be influenced b\- local conditions.
Females may be constrained in their timing
of fall migration by the nutritional and energetic
demands of lactation and smaller body size, by
the inabilitx of fawns to cope with severe fall
conditions, or both. Males do not ha\e the same
energetic, nutritional, or parental constraints.
Additionall), as consequence of hunting regula-
tions, those males that do migrate early are likely
to be killed.
ACKN OWLEDG M E NTS
Financial support was provided bv Invo and
Mono counties, the Sacramento Safari ('lub,
National Rifle Association, Mzuri Wildlife
Foundation, Boone and Crockett Club, and
Theodore Roosevelt Memorial Fund of the
American Museum of Natural Historw I thank
the California Department of Fish and Game
and U.S. Bureau of Land Management for their
personnel, logistic, and administrative support.
T. Blankinship, X. Koontz, D. R. McCullough,
T Russi, T. Taylor, R. D. Thomas, and others
were instnnnental in various parts of this work.
I thank V. C. Bleich, R. T Bowyer, and D. R.
McCullough, and particularh- an anonviiious
reviewer for their thcnightful reviews of the
manuscript.
Literature Cited
Anderson. A. E. 1981. Morj^hological ;uid plivsical tluuac-
teristics. Pages 27-97 in O. C. Wallmo, etl.. Mult- and
black-tailed deer of North America. Uni\ersit\ of
Nebraska Press, Lincoln.
AsncHAFT, G. C, Jr. 1961. Deer movements of the
McCloud flats herd. CiJifornia Fish and Game 47:
145-152.
Baker. R. R. 1978. The exolutionarv ecok)g\- of animal
migration. Holmes tuid Meier Publishers, New York.
Bertram. R. G., ;md R. D. Rempel 1977. Migration of the
North Kings deer herd, (laliforiiia Fish and (^ame 63:
157-179. '
Clover. M. R. 19.56. Single-gate deer trap. Gaiifornia Fish
and Game 42: 199-201.
Glutton-Brock. T. II., F. E. Glinnes.s. and S. D.
A1.B0N. 1982. Red deer: ecologv' and behavior of two
sexes. Universitv of Chicago Press, Chicago.
Dixon, J. S. 1934. A .studv of the life history and food habits
of mule deer in C';ilifornia. Gaiifornia Fish and C^ame
20: 181-282.
FnvxELL, J. M., iuid A. R. E. Sinclair 1988. Gau.ses and
consequences of migration In' large herbi\'ores. Trends
in Ecologv' and Evolution 3: 237-241.
(iAHROTT, R. A., (;. C. White, R. M. Bartmann. L. H.
Carpenter. ;uid A. W. Alldreuc:e 1987. Move-
ments of female mule deer in northwest Colorado.
Journal of VVilcUife Management 51: 6.34-643.
Geist. \', 1981. Behaxior: adaptive strategies in mule deer.
Pages 157-223 in O. (-. Wallmo, ed.. Mule iuid black-
tailed deer of North .\uicrica. University of Nebraska
Press, Lincoln.
Gilbert. P R, O. G. Wallmo ant! R. B. Gill 1970.
Effect of snow depth on mule deer in Middle Park,
Colorado. Journal of Wildlife Management .34: 1.5-33.
130
Great Basin Naturalist
[Volume 52
Giu'KLi., G. E., luxl N.J. Papez. 1963. Movements of mule
deer in northeastern Nevada. Journal of Wildlife Man-
agement 27: 414-422.
JESSUP, D. A., K. JoNKs. R. MoiiK. and T. Kucera 1985.
Yoliimbine antagonism to .xylazine in free-ranging mule
deer and bighorn sheep. Journal of the Ameriean Vet-
erinary Medical Association 187: 1251-1253.
Jones, F. L. 1954. The Inyo-Sierra deer herds. California
Department of Fish and Game, Federal Aid Project
\V-41-R.
Kucera. T E. 1978. Soci;il behavior and breeding system
of the desert mule deer Journal of Manun;ilog\' 59:
463-476.
. 1988. Ecolog\- and population dynamics of mule
deer in the eastern Sierra Nevada, California. Unpub-
lished doctoral dissertation. University of California,
Berkeley. 207 pp.
Leopold. A. S., T Riney. R. McCain, and L. Te\is. Jr
1951. The Jawbone deer herd. California Department
of Fish and Game, Game Bulletin Number 4.
Loft, E. R., R. C. Bertra.m. and D. L. Bowman 1989.
Migration patterns of mule deer in the central Sierra
Nevada. California Fish and Game 75: 11-19.
McCuLLOUCH, D. R. 1964. Relation.ship of weather to
migratory movements of black-tailed deer. Ecolog\' 45:
249-256.'
. 1985. Long range movements of large terrestrial
mammals. Contributions in Marine Science 27: 444-
465.
MUNZ, R A., and D. D. Keck 1959. A California flora.
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National Oceanic and Atmospheric Administr.\tion.
1987. Local chmatological data, annual summar\' with
comparative data. Bishop, California. Nationtil Cli-
matic Data Center, Asheville, North Carolina.
RiciiENS, V. B. 1967. Characteristics of mule deer herds and
their riuige in northeastern Utah. Joiunal of Wildlife
Management 31: 551-666.
Robin ETTE, W. L. 1966. Mule deer home range and dis-
persal in Utah. Journal of Wildlife Management 30:
335-349.
Rus.sELL, C. P. 1932. Se;isonal migration of mule deer.
EcologictJ Monographs 2: 1-46.
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Kogakusha, Ltd., Tokyo, Japtui.
Sinclair. A. R. E. 1983. The function of distance move-
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and P. J. Greenwood, eds.. The ecologv' of animal
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Vaughn. D. E. 1983. Draft soil inventorv of the Benton-
Owens VtJley tirea. U.S. Department of the Interior,
Bureau of Land Management, Bakersfield District,
Bakersfield, California.
Wright. E., and L. W. Swift 1942. Migration census of
mule deer in the White Ri\'er region of northwestern
Colorado. Journal of Wildlife Management 6: 162-164.
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Hall, Englewood Cliffs, New Jersey.
Received 15 November 1991
Accepted 15 April 1992
Great Basin Naturalist 52(2), pp 131-138
DIATOM FLORA OF BEAVER DAM CREEK,
WASHINGTON COUNTY, UTAH, USA
Kiiitis II. Yt-arsk
â– , Sanmel R. Huslitortli . and Jeffre\' H. Joluuisei
Abstract — Tlie diatom flora of Beaver Dam Creek, Washington County, Utah, was studied. The study area is in a warm
Mojave Desert en\ironment at an elevation bet\veen 810 and 850 m. A total of 99 taxa were identified from composite
samples taken in the fall, winter, spring, and summer seasons. These taxa are all hroadlv distributed and no endemic species
were encoimtered. Three new records for the state of Utah were identified: Gomphoiwis cricnse Sk-v. & Maver, S'avicula
el sinensis \ar. lata ( M. Perag. ) Patr., and Nitzschia calkla Cnin. The most important taxa throughout the study as determined
hv multiplying percent presence by average relative density (Important Species Index) were Cijniljella ajfinis Kiitz.,
Epithemia sorex Kiitz., Naviaila vcneta Kiitz., Nitzschia palea (Kiitz.) W. Sm., and Nitzschia microcephala Grun.
Kcti liords: Beaver Dam Creek, diatoiris, desert streams.
The algal flora ot the Intermountaiii West of
North America is not well known despite the
fact that numerous studies dealing \\ith algal
systems of waters in this region have been com-
pleted in recent years. These studies have exam-
ined streams, fresh water lakes, saline lakes,
thermal springs, and terrestrial habitats
(Sommerfeld et al. 1975, Stewart and Blinn
1976, Czarnecki and Blinn 1977, 1978, Blinn et
al. 1980, Bush and Fisher 1981; for bibliogra-
phies see Rushforth and Merkley 1988, Metting
1991).
Algal floras of wanii desert systems are espe-
cially poorly known. The present study was ini-
tiated to provide additional information on the
diatom flora of a desert stream located in west-
em North America. We examined the diatom
communities of Beaver Dam Creek, a tributary
of the Virgin River in southwestern Utah. This
paper is intended as a baseline floristic and
communit\' study of the diatom communities
present in this Mojave Desert stream.
We had three objectives in this study: (1) to
identify all species of diatoms present in Beaver
Dam Creek, (2) to document seasonal variation
in the diatom communities of this stream, and
(3) to compare diatom populations according to
habitat t\pe. Our stud\- reports all diatom taxa
present in this stream across four seasons of
1987-88. We studied populations in (1) riffle
areas with erosional flow velocities, (2) deposi-
tional areas with slower flows, and (3) epipln tic
habitats on the stems and leaves of aquatic xas-
cular plant vegetation.
Site Description
Beaver Dam Creek at L\tle Ranch Preserve
is located 37°10' North latitude and 114° West
longitude in Washington Countv, Utah (Fig. 1).
The stream occurs in our study area at an eleva-
tion of about 850 m at L\i:le Ranch dropping to
810 m at Tenys Ranch. Our study sites are
located along the wash near the ranch house at
Lvtle Ranch Preserve and near a smaller out-
building at Tenys Ranch.
Beaver Dam Creek is a vigorous, braided
perennial desert stream. It is important to the
entire biota of the area since it is the main source
of perenniiil water. The stream through the
study area has formed a broad gravel flood plain
due to frequent flooding. The stream occurs in
bajada and alluxial fan materials derived from
the Bull N'alley, Pine Vallev; and Santa Clara
mountains (Welsh et al. 1987).
Beaver Dam Oeek is fed by seeps, springs,
and snowmelt primarily from the Pine Valley
Mountains. This area is also characterized by
flash floods caused by sevx^re periodic thunder-
storms in the summer and fall seasons. For
instance, prior to the April 1988 collection,
Beaver Dam Wash received 11 days of rtiin
J Department of Botany and Range Science, Bngliam Yonng lJnJ\ersit\ . Provo. Ltali 84602.
Departmentof Biology, John Carroll University, Universits Heiglils, bliio4411S.
131
132
Great Basin Naturalist
[Volume 52
Fig. 1. Map of Beaver Dam VVasli .sliowing tlie location of colletting lotalitit-s at Tern s Raiuli and Lxtle Ranch Preserve.
Due to the meandering and clianging nature of Beaver Dam Creek, the stream itself is not sliown on this map.
1992]
Diatoms of Bkankh Dam Chkkk
133
producing moderate to severe flooding along
the stream channel. This scoured the stream
channel, remoxing large amounts of ac^natic
\e2etati0n and causing; channel relocation in
some areas.
The gravel bar in Beaver Dam Creek is gen-
erallv higher in the center than at the margins,
causing the stream to meander over a wide area
\\1th frequent changes of channel during flood-
ing (Welsh et al. 1987). The fall in elevation
downstream is not constant. Gravel tends to pile
up in steps that vary' in length and height. This
uneven granular substrate causes the stream to
meander along the gravel bar and eventually to
sink underground approximatel) four miles
below the southernmost collection site (Welsh
et al. 1987). The perennial stream reappears
infrequenth' as seeps and springs lower in
Beaver Dam Wash until merging with the Virgin
Rixer.
Climate in the stud\' area varies consider-
ably, not only diunially and seasonally, but over
longer periods of time. Winters are generally
cool and drv; summers hot and dry. MiLximum
summertime temperatures have been recorded
at 45.6 C. Rainfall averages less than 15 cm a
year, although this is \ariable due to intense
storms (Welsh et al. 1987).
The biota of our study area is exceptionally
diverse. Mammals, birds, reptiles, amphibians,
invertebrates, and a great variety of plants occur
in Beaver Dam Wash (Welsh et al. 1987). The
stream supports a diverse riparian habitat con-
sisting of Fremont cottonwood (Populus
freinontii Wats.), Arizona ash (Fraxitms vehitina
Torr.), black willow (Salix oooddingii Ball), seep
wiWow {Baccharis emorxji Gray//; Torn), numer-
ous torbes, grasses, and grasslike species (Welsh
et al. 1987). Silty terraces occur immediately
adjacent to the wash and have been historically
used for cultivation. These areas are dominated
by catclaw acacia {Acacia greggii Gray), panicu-
late rabbitbrush {Chn/sotJiamnus panicidatus
[Gray] Greene), Ambrosia species, and numer-
ous others (Welsh et al. 1987). Adjacent uplands
support Joshua tree forests {Yucca hreiifolia
Engelm.), creosote bush {Larrea tridentata
[DC] Gov.), prickly pear cactus {Opuntia
en^ehnannii Engelm.), cholla cactus {Opuntia
hasilaris Engelm. and Bigel.), and numerous
other xerophvtic species (Welsh et al. 1987).
Methods
Water chemistn,' was sampled at the collec-
tion sites for Febmarv, April, and July 1988
using a portable Hach field water chemistry lab.
Air temperature and water temperature, dis-
solved oxygen, hardness, alkalinit\, and pH were
measured.
Diatom collections were taken on 21
November 1987, 20 February 1988, 30 April
1988, and 6 July 1988 to docvmient seasonal
\ariations in diatom populations. (Composite
samples were collected from three habitat
t)pes. First, riffle areas with erosional flow rates
were sampled by scraping algae from large
stones in the creek bed. Second, slow water
areas in the stream were sampled by obtiuning
sediments, rock scrapings, and visible attached
algae. Finally, submerged sedge stems and
leaves were scraped or collected at selected
localities to studv epiph\'tic assemblages.
Due to seasonal changes, it was not always
possible to sample all three substrate t\pes at
both locations. A total of 19 samples were ana-
ly7:ed during the course of the study. Samples
were stored at air temperature and retimied to
the laboratoiy at Brigham Young University- for
analysis.
Diatoms were cleared by boiling in nitric
acid and potassium dichromate (St. Clair and
Rushforth 1977). After rinsing, cleared fnistules
were suspended in distilled water and allowed
to air dry on cover slips. Strewn mounts were
prepared using Naphrax high-resolution resin.
Representative slides were examined with Zeiss
RA microscopes equipped with Nomarski
optics and bright field illumination. An Olym-
pus AD photomicrographic system was used to
record each taxon. Strewn mounts ha\e been
placed in the collections at Brigham Young Uni-
versity.
A minimum of 500 valves was counted for
each sample, and a percent relati\ e densit\- was
calculated for each taxon (Kaczmarska and
Rushforth 1983). An Important Species Index
(ISI) for tcLxa present was calculated by multi-
plving the percent frequency of occurrence of a
taxon in the samples 1)\- its oxerall average per-
cent relative densitv in all samples (Ross and
Rushforth 1980, Kaczmarska and Rushforth
1983). This method is useful since it considers
both abundance and seasonal distribution of a
taxon (Warner and Haqoer 1972). Species diver-
sity for each sample was calculated using the
134
Great Basin Naturalist
[Volume 52
T./VBU. 1. Mean values for air teinperatiue iuid water chemical paranieter.s taken from collecting loc;ilities in Beaver
Dam Creek, Washington Countv', Utah.
February
April
Ji
ily
L\tle Terry's
Lvtle
Terrv's
Lytle
Terry's
Air temp. (C)
16.3 17.3
20.5
20.5
33.0
26.0
Water temp. (C)
14.5 17.5
16.8
16.8
24.3
22.3
Di.ssolved O2 (mg/1)
9.5 10.0
9.0
9.0
( . 1
7.0
Hardness (mg/1)
247.3 276.1
707.5
707.5
281.9
362.4
Alkalinitv (mg/1
195.6 207.1
201.3
224.3
pH
7.3 7.1
6.9
7.0
8.1
7.7
T.\BLE 2. T;t\a present in samples collected from Beaver Dam Creek, 1987-88, Listed with Important Species Index
(ISI) values. When ISI is below 0.01, the species is listed ;is a trace (T).
Taxon
ISI
Lvtle
Terry
Achnanthes affinis Gnin.
Achnanthes exi^tia Cnm.
Achnanthes hnceolata (Breb.) Gmn.
Achnanthes miniitissima Kiitz.
Amphora libt/ca Ehr.
Amphora pedictihis (Kiitz. ) Gmn.
Amphora veneta Kiitz.
Cah>nei.s bacilhim (Cnm.) Cl.
Cah)neis siliciiht (Ehr.) Cleve
Cocconeis pedicuhis Ehr.
Cocconeis placentula viu". eti^hjpta (Ehr.) Cleve
Cocconeis placentula v;xr. lincata (Ehr.) VH.
Cyclosteplianos invisitattis (H. & H.) Ther., Stoerm. & Hak.
Cyclotella nu'nc<^hiniana Kiitz.
Ci/mbella affinis Kiitz.
Cifiuhclla niexicana (Ehr.) Cl.
Ct/inhella microccphala Gmn.
Ci/nihclla silcsiaca Bleisch
Ci/nihclla tiimida (Breb. ex Kiitz.) V.H.
Dcnticula dedans Grun.
Denticnia clc<ians f. valida Pedic.
Diatoma viil^arc Bott
Diatoma vnl^are var. breve CJrnn.
Epilhemia adnata var. proboscidea (Kiitz.) Hend.
Epitheinia sorex Kiitz.
Epithciiiia tur^ida (Ehr.) Kiitz.
Fra<iilaria constniens (Ehr.) Gmn.
Fraiiilaria constniens f. venter (Ehr.) Hust.
Fra^ilaria piwiata Ehr.
Fraiiilaria vaucheriae (Kiitz.) Peters.
Gomphoneis eriense (Grun.) Sk\'. & Meyer
Gon\phoneis olivacea (Home.) Dawson
Goniphonema acuminatttm Ehr.
Gomf)honcnui an<i^usttim Agardh
Gomphonenia clavatum (Ehr.)
Gomphonema p^ninotvii Patr.
Gomphonenia parvidnm (Kiitz.) Kiitz.
Gomphonema pseudoatiffir L.-Bert.
Gomphonema truncatiim Ehr.
Gtjrosi<ima nodulifemm (Gnm.) G. West
Hantzschia amj>hioxys (Ehr.) Grun.
Melosira variam Ag.
Meridion ciradare (CJrev.) Ag.
Navicida abiskoetisls Hust.
Navictda atomus var. permitis (Hust.) L.-Bert.
Navinila baeilhnn Ehr.
1.92
1.8
2.6
0.03
0.1
0.1
2.51
3.8
1.1
1.92
3.4
1.3
0.10
0.4
0.1
1.76
2.5
1.1
0.13
0.6
0.1
T
T
0.04
0.1
0.1
1.07
3.1
0.8
1.22
1.4
1.1
T
0.72
1.0
0.5
17.57
23.4
13.2
T
0.58
1.2
a.5
0.16
0.4
0.1
T
1.44
2.5
0.4
T
V
0.84
1.7
0.5
0.11
0.5
0.1
0.07
0.1
0.3
1.3.25
1.8
35.9
T
0.21
0.5
0.2
0.50
0.5
0.8
0.14
0.2
0.3
2.21
1.1
3.0
0.02
0.1
0.1
0.27
0.7
0.2
T
0.51
0.8
0.4
0.06
0.2
0.2
0.08
0.3
0.2
1.89
2.1
0.2
1.32
1.6
1.5
T
T
T
0.06
0.3
0.1
T
T
0.08
0.2
0.1
0.09
0.2
0.2
1992] Diatoms of Bea\'Er Dam Creek 135
Tablk 2. Coutiiiut'il.
Navicula capitatoradiata Germain
Navicula cincta (Ehr.) Ralfs
Ndviaila constans \ar. symmetrica Hust.
Navictihi aispidata Kiitz.
Naviada eli^ineusis var. lata (M. Perag.) Patr.
Naviada gregaria Donldn
Naviada menisctdus Schumann
Navicula minu.scida \ar. muralis (Gmn.) L.-Bert
Navicida jutptda Kiitz.
Navictila radiosa Kiitz.
Navicula tripunctata (OF. Miill.) Bor\'
Navicula tripunctata \ar. schiz-oneiiwidcs (V.H.) Patr.
Naviada trivialis L.-Bert.
Navicula vciwta Kiitz.
Nridium affinc (Ehr.) Pfitz.
Ncidium did>ium (Ehr) Cl.
Nitzschia acicularis (Kiitz.) W.Sm.
Nitzschia amphibia Gnm.
Nitz-schia calida Grun.
Nitz.schia communis Rabh.
Nitzschia constric-ta (Kiitz.) Ralfs
Nitzschia di.ssipata (Kiitz.) Gnm.
Nitzschia fonticola Gnm.
Nitzschia Jnistulum (Kiitz.) Grun.
Nitzschia hantzschiana Rabh.
Nitzschia inconspicua Gnm.
Nitzschia linearis (Ag.) W. Sm.
Nitzschia microce))hala Grun.
Nitzschia palea (Kiitz.) W. Sm.
Nitzschia si^moidca (Nitz.) W. Sm.
Nitz-scliia std)tilis Gnm.
Pinnularia appcndiculata (Ag.) CI.
Plcurosi<ima dclicatulum W. Sm.
Plcurosira lacvis (Ehr) Compere
Rcimeria sinuata (CJreg.) Kociolek & Stoermer
Rlioicosplwuia curvata (Kiitz.) Grun.
Rhopalodia hrchissonii Krammer
Rhopalodia oihba (Ehr) O. Miill.
Rhopalodia f^ihha var. vcntricosa (Kiitz.) Perag. & Perag.
Rhopalodia '^ihhcnda (Ehr) O. Miill.
Stauroncis smithii Gnm.
Stcnoptcrohia intemwdia (Lewis) V.H.
Stcphanochscus hantz-schii Grun.
Surirclla an^usta Kiitz.
Surirclla minuta Breb.
Surirella (nalis Breb.
Sipu'dra acus Kiitz.
Sijnedra fasciadata \-m: tnincata (Gre\-. ) Patr
Stjncdra radians Kiitz.
Syjicdra ntmpcns vnr. mcnc^hiniana (Irun.
Sijncdra ulna (Nitz.) Ehr
Sijnedra ulna viir. contractu Oestr.
Shannon-Wiener clixersiU' index (Shannon and averages. It is \videl\- nsed and has been found
Weaver 1949, Zar 1986). to introduce less distortion than other methods
Siniilarit)- indices were calculated for all (Kaesler and Cairns 1972).
pairs of samples following Ruzicka ( 1958). Clus-
ter analyses based on Ruzicka's indices using RESULTS AND DISCUSSION
unweighted pair-group technicjues (UPCMA)
were then performed (Sneath and Sokal 1973). Water chemistiv did not van- significantly
This method computes the average similarit\- of according to collection localit}' (Table 1). Stream
each site to e\'er\' other site using arithmetic tem^^eratnre increased somewhat during the
L99
2.7
2.0
0.17
0.5
0.1
T
T
0.06
0.2
0.1
0.16
0.5
0.1
T
0.06
0.3
0.1
0.16
0.4
0.2
0.19
0.3
0.3
0.12
0.5
0.1
0.10
0.4
0.1
0.28
0.6
0.2
8.78
9.0
8.6
T
0.1
T
0.02
0.1
0.1
1.51
2.4
0.4
0.02
2.0
0.30
0.9
5.9
0.19
0.4
0.3
L90
3.9
0.4
0.58
LI
0.4
0.01
0.1
0.20
0.2
0.1
0.65
1.4
0.2
T
5.44
4.7
6.3
5.76
8.7
2.4
0.01
0.1
0.1
T
T
0.1
0.02
0.2
T
T
0.1
2.73
1.4
4.9
T
T
0.01
0.1
0.03
0.2
0.1
T
0.1
T
0.02
0.1
0.06
0.2
0.1
T
T
T
T
0.01
0.1
0.1
0.11
0.1
0.3
0.40
0.7
0.5
0.20
0.3
0.3
136
Great Basin Naturalist
[Volume 52
summer uionths, hut it is uoteworthy that teui-
perature variations in the stream were relatively
small. The stream is circumneutral to slightly
alkaline.
A total oi 99 diatom taxa in 24 genera were
obserxed in our collections. Three new records
for the state of Utah were noted: Gomphoneis
eriense (Gnui.) Skv. & Meyer, Naviaila el^hien-
sis var. lata (M. Perag.) Patr., and Nitzsclua
calida Gnm. Taxa are illustrated and described
in Yearsley (1988). Nomenclature followed in
Yearsley (1988) was similar to that used histori-
calK' b\' researchers in our laboratoiy for com-
parative puiposes (Rush forth and Merkley
1988). Diatom taxonomy in this paper is based
primarilv on the recent texts of Krammer and
Lange-Bertalot (1986, 1988, 1991), although
other references were consulted and sometimes
followed. We did not follow the numerous
generic changes proposed in Round et al. (1990)
due to the controversy over many of their rec-
ommendations.
Eighteen taxa in Beaver Dam Creek had an
Important Species Index value greater than 1.0
(Table 2). The most important taxa in the overall
study with ISIs above 5.0 were CijmhcUa affinis
(ISI = 17.57), Epitlu'inia sorex (13.25), Navic-
iila verieta (8.78), Nitzschia palea (5.76), and
Nitz.schia microcephala (5.44). Taxa with ISIs
greater than 1.0 included Rlioicosphenia
curvata (2.73), Achnanthes lanceolata (2.51),
Frogilaria vaucheriae (2.21), Navicula capita-
toradiata (1.99), Achnanthes affinis (1.92),
AchnantJjcs niinittissinia (1.92), Nitzschia dis-
sipata (1.90), G()nif)lioneimi parviduni (1.89),
Nitzschia anipJiihia (1.51), Denticiila elegans
(1.44), Gomphoncnia psendoaiigtir (1.32),
Cocconeis placentida var. Uneata (1.22), and
Cocconeis placenfnia \ar. ew^Uipta (1.07). All of
these taxa are cosmopolitan and found in a vari-
ety of habitats.
In comparing the diatom assemblage from
Beaver Dam (Jreck with the floras of streams of
other arid regions, we noticed a striking similar-
it)'. The important taxa overlapped in all of the
studies even though the streams varied in terms
of their flow rate and climatic regime. Further-
more, each system was dominated by cosmopol-
itan species. Our preliminary data indicate that
a diatom flora unique to desert streams does not
exist. Further research to substantiate this con-
clusion is necessarx'; .some evidence is gi\en
below.
Blinn et al. (1980) considered substrate col-
onization in Oak Creek, Arizona. Thev reported
12 important taxa which, in order of decreasing
abundance, were: Nitzschia frustuhun, Epithe-
niia sorex, Cocconeis placentida var. euglifpta,
Achnanthes niintitissima, Navicida cnjpto-
cephala, Navicula veneta (as N. cnjptocephala
var. veneta), Nitzschia dissipata, Achnanthes
lanceolata, Ci/mbeUa affinis, Fragilaria con-
stniens, Navicida decussis, and Synedra idna.
These diatoms accoimted for 90% or more of
the total algal population on newly introduced
material in their study. Eight of these taxa were
also important in our stream, having ISI values
above 1.0.
Johnson et al. (1975) conducted further
study on the diatom flora of Oak Creek, Arizona.
They reported 41 diatom taxa, of which 25 are
common to our study area. Cijnihella affinis,
Epithemia sorex, and Nitz^schia palea were
reported as common or abundant. This com-
pares favorably with the results of our study
since these three were among the most common
diatoms in Beaver Dam Creek.
Rushforth et al. (1976) examined the algal
flora of Freshwater Wash, Arches NationiU
Park, in southeastern Utah. Their study docu-
mented 57 diatom taxa, 29 of which were tilso
observed in Beaver Dam Creek. Achnanthes
niinittissinia, Cijmhella affinis, Denticula ele-
gans, Goniphonenia acuminatum, Navicula
radiosa, Nitzschia linearis, Nitzschia palea,
Rhoicosphenia curvata, and five other species
not present in Beaver Dam Creek were the most
abundant taxa in Freshwater Wash.
In their analvsis of Sycamore Creek, Arizona,
Fisher et al. (1982) reported that diatoms made
up 77% of the total algal mass, \v\i\\ Achnanthes
exigua, Gomphonema parvidum, and Navicula
pupula being the most important taxa. These
taxa were present in Beaver Dam Creek but in
lower numbers. Gomphonema parvidum was
the most abundant of the three in our samples.
The flora of Beaver Dam Creek is also sim-
ilar to that of other streams of western North
America draining more mesic regions. Gushing
and Rushforth (1984) in a study of the Salmon
River, Ickilio, identified 145 diatom species, 48
of which were among the 99 taxa found in
Beaver Dam Creek. Half of their important
species (9 of 18) were also among the important
species in Beaver Dam Creek, several with sim-
ilar importance values.
Preliminar\' research also indicates that a
flora similar to that found in North American
1992]
Diatoms of Bean-er Dam Cheek
137
hardwater streams exists elsewhere. S(juires and
Saoud (1986) reported nine ta\a from the
Damour Ri\er, Lebanon, with Importance Spe-
cies Index xiilues above 1.0. Six of these also
were important in Beaxer Dam (>reek. In the
Damoin- Ri\er stiid\' Aclinantlics inlnutissiina
was the most important taxon with an ISI \alue
of 44.4, followed bv Nitzschia dissipata (5.12),
Cyniljella microcephala (3.63), and CifmheUa
affinis (2.62).
Shannon-Wiener diversit)' values for all 24
samples ranged between 1.95 and 4.59. Diver-
sit)' did not show any clear trends with regard to
season or substrate tyjoe. The overall mean for
the indices was 3.42, the median \alue being
3.57. These \alues are relati\el\' high and indic-
ative of unpolluted water.
Oiu" collections did not cluster well on the
basis of habitat t\pe or season. However, there
was a tendency for stands to cluster on the basis
of the Terpy's Ranch versus L)tle Ranch Pre-
serve collecting localities (Fig. 2). The upper-
most cluster consists of samples from Terrv's
Ranch, while the second cluster contains sam-
ples from the Lvtle Ranch Preserve. The third
cluster has a mix of all sites, substrates, and
seasons. The fall depositional sample from the
Lvtle Ranch Preserve is an outlier.
The reasons for the clustering b)' site seen in
the top half of the cluster are unclear. Water
chemistry' and temperature did not var\" greatlv
between the sites during the year (Table 1).
Likewise, insolation is approximatelv the same
for both sections of the creek. Stream velocities,
however, appear to be different. The creek at
Lytle Ranch Presene is generallv slower, shal-
lower (<15 cm), wider, and more meandering
than the stream at Terry's Ranch where pools
may reach depths of nearly one meter.
The cluster shows a number of samples that
paired b\- date of collection (Fig. 2). However,
seasonalit)- was ver)- weak. The absence of sea-
sonal changes is probably attributable to one or
two factors. First, temperatiu'e changes
tlu-oughout the year are minor, and changes in
photoperiod alone are not enough to drive suc-
cession. Second, storm events scour the creek
bed occasionally and may keep the diatom
assemblage in an early successional stage.
The habitat t\pes sampled did not cluster
separately, indicating they are fairly similar.
Because of scouring events, the depositional
areas initially sampled often had all sediments
remoN'ed at later sampling dates and so consist
PERCENT SIMILARITY
100 90
T M Nov.
T R July
T D/R July
L D/R Apr.
T D/R Apr.
L R Feb.
L R Apr.
L R July
T M Apr.
L M July
L D/R Feb.
L R Apr.
T R Apr.
Fig. 2. Cluster diagram of 19 samples collected from
Beaver Dam Creek. T = Terrv's Ranch, L = Lvtle 's Ranch
Preserxe, M = macrophvtic vegetation (sedges), R = riffle,
D/S = depositional area, .sediments, D/R = dejx)sitional
area, rock scrapings.
of rock scrapings, just as in the riffle areas. The
one sample that consisted of sediment only
(Lylile Ranch, November 1987, depositional
area) clustered separately from all other sam-
ples (see bottom line of cluster, Fig. 2).
In summary, the diatom assemblages
observed in Beaver Dam Creek consisted of
cosmopolitan species common to other hard-
water rixers. Seasonalit\' was minimal, as were
the effects of habitat t\pe.
LiTER.'\TURE Cited
Blinn. D. \V., a. Fhkdf.hickskn, and V. Koin k 19.S().
Colonization rates and community stnictnre of diatoms
on three different rock substrata in a lotic svstem.
British Phycologicd Jouniiil 15: .30.3-.31().
Bush. D. E., and .S. C. Fishkk 1981. Metabolism of a
desert stream. Freshwater Biologv' 11: .301-.3()7.
CusiUNC,. C. E., and S. R. Risiifortm 19<S4. Diatoms of
the middle fork of tlie Salmon River Dr;iinage, with
notes on their relative abundance and distribution.
Great Basin Naturalist 44: 421^27.
C/.'VRNKCKl. D. B., and D. \V. Blinn 1977. Diatoms of
Lower Lake Powell and vicinitv Bibliothcca Fh\-
cologica 28: 1-119.
. 1978. Diatoms of the Colorado River in Grand
Canyon National Park and vicinit\-. Bibliotheca Phy-
cologica 38: 1-181.
138
Great Basin Naturalist
[\blume 52
FisiiKH. S. C, L. J. GiuY, N. B. Ghimm. unci D. Busii 1982.
Teiinxiral succession in a desert stream ecosystem fol-
lowing flash Hooding. Ecological Monographs 52: 93-
110.
Johnson, R.,T. Ricii.^rd.s. and D. W. Bi.iw, 1975. Inves-
tigation of diatom populations in rhithron andpotamon
communities in Oak Creek, Arizona. Southwestern
Naturalist 20: 197-204.
Kaczmahska, I., and S. R. Rusiiforth 1983. The diatom
flora of Blue Liike, Tooele County, Ut;ili. Bibliotheca
Diatomologica 2: 1-123.
Kaesler, R. L., and J. Cairns. 1972. Cluster analysis of
data from limnological surveys of the upper Potomac
River. American Midland Naturalist 88: 56-67.
Krammkr. K., and H. Lange-Bertalot 1986. Bacil-
lariophyceae. 1. Teil: Naviculaceae. Volume 2/1 in
Pascher's Suesswasserflora von Mitteleuropa. Gustav
Fischer Verlag, Stuttgart. 876 pp.
. 1988. Bacillariophyceae. 2. Teil: Bacillariaceae,
Epithemiaceae, Surirellaceae. Volume 2/2 //( Pascher's
Suesswasserflora von Mitteleuropa. Gustav Fischer
Verlag, Stuttgart. 596 pp.
1991. Bacillariophvceae. 3. Teil: Centrales,
Fragilariaceae, Eunotiaceae. Volume 2/3 in Pascher's
Suesswasserflora von Mitteleuropa. Gustav Fischer
Verlag, Stuttgart. 576 pp.
Metting, B. 1991. Biological surface features of semiarid
lands iuid deserts. Pages 257-293 in J. Skujins, ed.,
Semiarid lands and deserts: soil resource and reclama-
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Ross, L., tmd S. R. Rushforth 1980. The effects of a new
reservoir on the attached diatom communities in Hunt-
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Round, F. E., R. M. Crawford, and D. G. Mann. 1990.
The diatoms, biologv' and moiphology of the genera.
Cambridge Universitv Press. Cambridge, United King-
dom. 747 pp.
Rushforth, S. R., and G. S. Merkley 1988. Com-
prehensive list by habitat of the algae of Utali, USA.
Great Basin Naturalist 48: 154-179"
Rushforth. S. R., L. L. St Clah^ T. A. Leslie, K. H.
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of two hanging gardens in soudieastern Utiili. Nova
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Methoden in der Geobotanik (Synthetische
Bearbeitung von Aufnahmen). Biologia, Brastisl. 13:
647-661.
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141.
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Received 16 September 1991
Accepted 31 March 1992
{ ;ivat Basin Naturalist 52(2), pp. 139-144
STRATIFICATION OF HABITATS FOR IDENTIFYING HABITAT SELECTION
BY MERRIAM'S TURKEYS
Mark A. Riinible and Stanley H. Anderson"
Abstract — Habitat selection patterns of Merriam's Turkevs were conipiired in hierarchical iuialvses of three levels of
habitat stratification. Habitat descriptions in first-le\el analyses were based on dominant species of vegetation. Habitat
descriptions in seconcl-le\el anaKses were biised on dominant species of vegetation and overstorv^ canopy cover. Habitat
descriptions in third-level anal\ses were based on dominant species of vegetation, o\erston' canopy cover, and stnictural
stages (dbh categories). First-level analyses showed turkeys selected for ponderosa pine and selected against meadow
habitats. No conclnsions could be drav\Ti regiucling forest management on habitat selection of turkevs at this le\el of habitat
stratification. Second-level analyses showed that selection of ponderosa pine and aspen/birch habitats \aried among seasons.
Implications for forest management actixities on turkevs at this level of habitat stratification could be made. Third-level
aiuil\ses added little to conclusions of habitat selection patterns drawn from second-le\el analyses and increased chances
for T\pe II errors. Habitat selection patterns of Merriam's Turkeys were best described when habitats were stratified by
dominant species of vegetation and overstorv Ciuiopy cover.
Kjcij words: Merriam's Wild Turkeys. Meleagris gallopa\o merriami, hiihitat descriptions, forest ntana^eiiwut, habitat
selection
Habitat use and management of Merriam's
Turkeys {Meleagris gallopavo merriami) in
northern latitudes have been studied in South
Dakota (Petersen and Richardson 1975) and
Montana (Rose 1956, Jonas 1966). These early
studies were limited to direct observation of
birds when assessing habitat use, and data con-
tained biases in the assessment of the birds'
habitat needs (e.g., Jonas 1966, Brvant and Nish
1975, Petersen and Richardson 1975, Shaw and
Smith 1977). Telemetrv' has allowed collection
of data on habitat use patterns in an unbiased
manner, but few studies have addressed the
detailed stratification habitats.
Studies of habitat use and selection patterns
by Merriam's Turkeys have delineated habitats
based primarilv on the dominant species of veg-
etation (DSV)' (Jonas 1966, Biyant and Nish
1975, Scott and Boeker 1975, ^Mackey 1982,
1986, Lutz and Crawford 1989). Because timber
management activities seldom result in conver-
sions of vegetation t)pes, understanding habitat
selection patterns at this level precludes under-
standing the effects of forest management activ-
ities such as logging or thinning on Merriam's
Turkeys. Increased value of ponderosa pine
timber resources, emphasis on old-growth
resource values, and improved technolog)' for
harvesting timber have potential to impact
Merriam's Turkey habitat (Shaw 1986). There-
fore, stratification beyond dominant species of
vegetation is necessary to elucidate the effects
of forest management on turkeys. Merriam's
Turkeys in southeastern Montana demonstrated
an apparent preference for pole-size (<23 cm
dbh) ponderosa pine habitats (Jonas 1966).
Merriam's Turkeys in Oregon avoided habitats
that had been logged by clear-cut or shelter-
wood methods (Lutz and Crawford 1989). To
our knowledge, no researchers have stratified
habitats in terms of size and densitv categories
of tree species. However, on lands managed by
the US DA Forest Service and other public
agencies, methods of habitat stratification that
include structural stages (SS) and overstory
canopy cover categories (OCC) have been
described (Thomas 1979) to further stratifv hab-
itats.
The objective^ of tliis stuck was to determine
the level of habitat stratification that best
described habitat use and selection patterns of
Merriam's Turkeys in the Black Hills.
, USD.-\ Forest Sen.ice, .501 E. St Joseph St.. South Dakota School of Mines. Rapid Cit\. South Dakota .57701 .
"USDI Cooperative Fisheries and Wildlife Research Unit. University of Wyoming. Laramie, \W)ming 82071.
139
140
Great Basin Naturalist
[Volume 52
Methods
Study Area
This study was conducted in the central
Black Hills of South Dakota, 16 km west of
Rapid Cit\'. Most of the land is vmder manage-
ment by the Black Hills National Forest, Pactola
Ranger District. Some private holdings associ-
ated with ranch operations are present in the
meadows, and several private homes and cabins
are located in the study area.
Vegetation of the study area is primarily pure
ponderosa pine forest (84%). Meadows and
aspen/birch (Popiihis tremtiloicles/Betida pa-
pyrifera) habitats occur in drainages.
This study was conducted over a three-year
period beginning March 1986 and ending Janu-
ary 1989. Because anahtical methods used to
make statistical tests were goodness-of-fit tests
and nonsignificance indicates lit by the pro-
posed model, hypotheses tested have been
stated appropriately. The hvpotheses tested rel-
ative to Merriam's Turkeys in the Black Hills of
South Dakota were that each of the following
habitats depict patterns of use and selection by
Merriam's Turkeys: (1) habitats stratified by
DSV (2) habitats' stratified by DSV and OCC,
(3) habitats stratified by DSV and SS, and (4)
habitats stratified by DSV, SS, and OCC.
Trapping and locations. — Turkeys were
trapped in late February or early March of each
year of the study with rocket nets and drop nets
over com bait. This study was primarih' con-
cerned with hens since they are the reproduc-
tive segment of the population. Forty-four (36
females and 8 males) of 82 turkevs trapped were
fitted with back|:)ack radio transmitters weigh-
ing approximate!)' 108 g.
Locating birds began after a one-week
period of adjustment to the radio transmitters
(Nenno and Healy 1979). Each bird in the study
area was located three times each week, once
during each of the following time periods: sun-
rise-1000 hr, 1001-1400 hr, and 1401 hr-
sunset. Birds that emigrated from the defined
study area were located at least monthly to mon-
itor their activities and determine if they had
moved back into the study area. Locations were
determined by plotting 2+ bearings (frequentlv
5+) from known locations on USGS 1:24,000
contour maps in the field using a luuul-held,
two-element yagi antenna. Bearings were usu-
ally taken from positions within 300 m of the
estimated location. Each location was assigned
to a habitat unit (see below) based on maps and
Universal Transverse Mercator coordinates
recorded to the nearest 100 m in the field. To
achieve independence of observations (All-
dredge and Ratti 1986), only one location was
recorded for each bird on any given day and
most were two days apart.
Habitat Descriptions
Habitats were numerically identified geo-
graphical units approximatelv 4-32 ha (10-80
acres) in size. Boundaries were usuallv defined
by watershed topography such as ridges and
drainages. Obvious changes in vegetation type
also were used to define boundaries of habitats.
In all, 513 habitat units were delineated.
Vegetative descriptions of habitats were
determined from five plots located within each
defined habitat unit. These plots were marked
on unit 1:24,000 contour maps in the lab and
distributed evenly across each habitat. Some
habitats were too small to effectively place five
plots, so fewer plots were used. Each plot was
then located in the field and sampled to deter-
mine tree basal area.
Habitat descriptions were made based on
DSV, SS, and OCC according to criteria devel-
oped by the US DA Forest Service, Region 2
(Buttety and Gillam 1983). DSV categories
were ponderosa pine, aspen/birch, oak, spruce,
and meadows. SS categories were pole timber
(trees 2.5-22.8 cm dbh) and sawtimber (trees
greater than 22.8 cm dbh). OCC categories
were 0-40%, 41-70%, and 71-100%. OCC was
estimated based on the following equation:
OCC(%) = 0.5 r BASAL AREA (FT'/AC) -
1.94 (Bennett 1984). Depending on the level of
stratification included in the analyses, .5-12 hab-
itats were delineated.
Analyses
Data pertaining to use of habitats described
above were stratified into seasons: December-
February (winter), March-May (spring), June-
August (summer), and September-No\ ember
(fall). Chi-square testof independence was used
to test the lupothesis that habitat use patterns
of Merriam's Turkeys were similar among sea-
sons. Because this test was significant (P < .001),
tests of habitat selection at different levels of
habitat stratification were made within seasons.
Chi-square goodness-of-fit tests with correc-
tion for continuit)' (Cochran 1963) were used to
test hypotheses regarding the level of habitat
19921
Turkey Habitat Stratification
141
stratification that best depicted habitat selection
patterns of Merriam's Turkeys in a hierarchical
structure. Bonferroni confidence intervals
around proportion of use (Neu et al. 1974, Byers
et al. 1984) were used to determine habitat
selection patterns that deviated from expected
use. We determined differences from expected
use of habitats for which utilization was by
examining chi-square residuals with G-stan-
dardization and Bonferroni correction to the
Z-statistic (Mosteller and Pamnak 1985). An
array of structural stages occurred only for
ponderosa pine habitats. Therefore, the test
for DSV X SS level of habitat stratification
was analyzed using data from ponderosa pine
habitats.'
Initial chi-square tests of use versus avail-
abilitv for DSV x SS, DSV x OCC, and DSV x
SS X OCC were made with oak, aspen, and
spruce habitats pooled to reduce as much as
possible the number of cells with fewer than five
expected observations. Selection of these hab-
itats by turkeys was evaluated individually with
Bonferroni confidence intervals for comparison
tests. The significance of confidence intervals
holds regardless of the overall chi-square test
(Neu et al. 1974).
Results
Habitats Determined by DSV
The hyjDothesis that habitats stratified by
DSV depict patterns of habitat use and selection
by Merriam's Turkeys was rejected (F = .06).
Meadows were selected less than expected
across all seasons (Table 1). Ponderosa pine
habitats were selected more than expected
during winter, spring, and fall; they were equal
to what was expected during summer. Aspen
habitats were selected more than expected
during summer. Oak habitats were selected less
than expected during spring, while spruce hab-
itats were selected less than expected during
winter and spring.
Habitats Determined bv DS\' and OCC
The hvpothesis that habitats stratified b\
DSV and OCC depict patterns of habitat use
and selection by Merriam's Turkeys was
rejected for all seasons (P = .04). Stratifying
habitats by DSV and OCC did not alter' the
results for meadow, oak, or spnice habitats
(Table 2). Oak and spruce were not represented
across all ov erston canopv cover categories on
this study area.
Aspeii/birch habitats with 41-70% OCC
were selected more than expected during spring
and sunuiier by turkeys in the Black Hills. Infre-
quent use of aspen/birch habitats with 7 1-100%
OCC was noted over all seasons. But statisti-
cally, this was less than expected onlv during
spring. Open ponderosa pine habitats (0-40%
OCC) were selected less than expected during
the winter and spring. Turkeys selected pon-
derosa pine habitats 41-70% OCC more than
expected during spring. Dense ponderosa pine
habitats (71-100% OCC) were selected more
than expected during fall and winter and less
than expected during summer.
Habitats Determined by DS\' and SS
The hypothesis that habitats stratified by
DSV and SS depicted patterns of habitat use and
selection by Merriam's Turkeys was not rejected
for winter, summer, and fall. During spring,
ponderosa pine habitats with stems greater than
23 cm dbh were selected more than expected.
Othenvise, no differences were apparent in the
habitat selection patterns of turkeys when pine
habitats were stratified based on dbh.
Aspen/birch, oak, and spruce habitats were not
adequately represented across structural stages
to make comparisons.
Habitats Determined by DS\' SS, and OCC
The hyjiothesis that habitats stratified by
DSV, SS, and OCC depict patterns of habitat use
and selection by turkev s v\'as rejected {P = .03)
during winter, spring, and summer (Table 3).
Data from fall indicated observed differences
from expected at F = .11. Since several habitat
categories were pooled to achieve minimum
sample .size in the overall chi s(juare test, F = .11
was considered sufficient indication of differ-
ence from expected to proceed with the
Bonferroni confidence intervals.
Use patt(M-ns of meadov\', oak, and spruce
habitats bv Merriam's Turkeys v\'ere unchanged
from previous levels of habitat stratification.
However, because more habitats were included
in the analyses, selection of spruce during
winter and aspen/birch habitats with 41-70%
overstory canopy cover during summer no
longer differed from expected.
Turkev s selected open ponderosa pine habi-
tats in both structural stages less than expected
durine winter, and the 2..5-22.8 cm dbh stnictnral
142
Great Basin Naturalist
[\blume 52
Table 1. Seasooiil utilization by Merriam's Turkeys of habitats described by dominant species of vegetation in the Black
HiUs of South Dakota/'-^'
Habitat
Proportional
area
Winter
(205)
Spring
(S78)"
Summer
(126)
FaU
(218)
Aspen
Meadow
Pine
Oak
Spnice
0.0516
4—
61
17+ +
14
0.1016
11—
9—
5 —
/ —
0.8371
186+ +
807+ +
100
195+ +
0.0044
4
0—
1
1
0.0056
0—
1—
3
1
"Sample sizes (teleinetn' fixes) are in parentheses. Expected use can he calculated from proportional use X sample size.
Differences [P < .101 among hahitats selected versus aviulahle are indicated hy — it used less than exjiected and ++ if used more than ex]«?cted.
Table 2. Seasonal utiUzation bv Merriam's Turkevs of habitats described bv dominant species and overstoiy c;xnopy
cover of vegetation in the Black Hills of South Dakota.' '
Habitat
Percent
Proportional
Winter
Spring
Summer
FaU
canopy cover
area
(205)
(878)
(126)
(218)
Aspen/birch
0-iO
0.0148
2
14
4
1
Aspen/birch
41-70
0.0191
46+ +
11 + +
12
Aspen/birch
71-100
0.0177
2
1—
2
1
Ponderosa pine
0-^0
0.1199
.3—
6.3—
26
29
Ponderosa pine
41-70
0.3760
65
430+ +
45
71
Ponderosa pine
71-100
0.3412
118+ +
314
29—
95+ +
Meadows
0.1016
11—
9—
.5—
7 —
Oak
0-100
0.0044
4
1
1
Spnice
0-100
0.0056
1—
3
1
''Sample sizes (telemetr\' fixes) are in parentheses. Expected use can be calculated from proportiona! use X sa
'Differences (P < .10) among habitats selected versus available are indicated by — if used less than expected ;
nple size
nd++if
used more than expected.
Table 3. Seiisonal utibzation by Merriam's Turkeys of habitats determined bv dominant .species, overstor\' canopv cover,
and structural stage in the Black Hills of South Dakota."'
Habitat
Structuriil
Percent
Proportional
Winter
Spring
Summer
Fall
stage
canopv cover
area
(205)
(878)
(126)
(218)
Aspen/birch
2.,5-22.8 cm
0-40
0.0148
2
14
4
1
Aspen/birch
2.5-22.8 cm
41-70
0.0191
46+ +
11
12
Aspen/birch
2..5-22.S cm
71-100
0.0177
2
1—
2
1
Ponderosa pine
2.,5-22.8 cm
0^0
0.0701
1—
9—
20+ +
18
Ponderosa pine
2.5-22.8 cm
41-70
0.1677
32
143
2.5
.'3.3
Ponderosa pine
2.5-22.8 cm
71-1(K)
0.2173
85+ +
222
16—
62
Ponderosa pine
>22.8 cm
0-40
0.0498
2
54
6
11
Ponderosa pine
>22.8 cm
41-70
0.2083
33
287+ +
20
38
Ponderosa pine
>22.8 cm
71-KK)
0.1239
33
92
13
33
Meadows
0.1016
11—
9—
5—
7
Oak
0-100
().(K)44
4
1
1
Spruce
0-100
0.0056
1—
3
1
J^Sample sizes (telemetry fi,\es) are in parentheses. Expected use can be calculated from proportional use X sample size.
Differences (P < .10) among habitats selected versus aviiilable are indicated l)y — if used less than expected and ++ if used more tli.u
stage was selected less than expected during
spring. No differences were noted for pon-
derosa pine with 41-70% overstory canopv
cover and 2.5-22.8 cm dbh across seasons.
However, the structural stage greater than 22.8
cm dbh and 41-70% overstory canopy cover was
selected more than expected during spring.
Dense ponderosa pine ( >71% overstorx' canopv
cover) 2.5-22.8 cm dbh was selected more than
expected during winter and less than expected
during summer. No differences were noted for
dense ponderosa pine >22.8 cm dbh.
1992]
TuHivt:Y Habitat Stratification
143
Discussion
The highest level of stratification of habitats
that added new information to use and selection
patterns of Merriam's Turkeys in this study area
was b\- DS\' and OCC. Despite statistical signif-
icance of differences when habitats were strati-
fied by DS\', SS, and OCC, trends in habitat
selection were similar to analyses for which data
were pooled across SS categories. Shaw and
Smith (1977) noted apparent habitat selection
b\- Merriam's Turkevs in Arizona when pon-
derosa pine habitats based on diameter classes
were ignored. However, pole-size ponderosa
pine habitats were used more than other size
classes b\' turkevs in Montana (Jonas 1966).
Within our studv area, 12 ot the 372 ponderosa
pine habitats had an average dbh of less than 15
cm (6 in); the lowest average dbh was 10.7 cm
(4.2 in). Thirt\-se\en of the ponderosa pine
habitats in the stud\ area had dbh greater than
30 cm (12 in), of which the majoritv" were in the
0-40% OCC category indicative of large over-
mature trees. Most of the study area had been
logged in the past one hundred \ears. Because
excellent germination conditions for ponderosa
pine in the Black Hills result in overstocked
stands with reduced growth rates (Boldt and
\'an Duesen 1974), ponderosa pine habitats
larger than 30 cm dbh were rare. Ponderosa
pine habitats in this study were representative
of a narrow range of the potential tree dbh
classes for ponderosa pine. However, they did
represent the size classes of ponderosa pine
throughout the Black Hills.
The tests of the model for DSV x SS sug-
gested good agreement between the model and
observed use bv turkevs from a statistical point
of \iew. These results suggest random selection
of habitats when stratified by DSV x SS. Non-
random selection of habitats had already been
demonstrated. We also beliexe that stratifica-
tion of habitats bv DSV x SS obscured biologi-
cal patterns alreacK' demonstrated In the test of
DSV X OCC. Many of the relationships of OCC
were contrasted between high and low OCC.
These results were pooled, resulting in the
apparentlv good fit of the DS\' X SS UKxlel.
Our approach to these anahses was hierar-
chical in nature; and since patterns of habitat
selection by turkeys had been demonstrated at
higher le\els, it would not be prudent to ignore
those biological patterns. Howexer, to ensure
that no oversights were made, we made tests of
hal)itat selection based on habitats stratified b\-
SS, OCC, and SS x OCC. The test of the model
for SS was not rejected. Tests of the model for
OCC and SS x OCJC were rejected, but were
influenced b\' the preponderance of the studv
occupied b\ ponderosa pine (84%) and the
range of dbh classes in the Black Hills. Interpre-
tations of results from these latter tests were
similar to tests of DSV X SS and DSV X OCC.
Stratification of habitats bcNond that neces-
sary' to depict the dispersion patterns of the
animal decreases the sensitivit)- of tests and
increases the probabilits of T\pe H error in the
anahses (Alldredge and Ratti 1986). The effect
of adding stratification factors is to dilute the
sample sizes in indixidual cells, thus increasing
the chance of Type H error. Apparent T\pe II
errors occurred in the determination of habitat
selection patterns when habitats were stratified
b\ DS\' X SS X OCC. At the highest level of
habitat stratification, apparent differences from
expected use for three habitat categories disap-
peared from the analyses.
Acknowledgments
This research was supported b\' the USDA
Forest Ser\ice, Rocky Mountain Forest and
Range Experiment Station; National Wild
Turkev Federation; Black Hills National Forest;
and South Dakota Came, Fish and Parks. We
extend special thanks for the support and
encouragement of Dr. A. J. Bjugstad
(deceased). Technical assistance of R. Hodorff,
T. Mills, C. Oswald, K. Thorstenson, K. Jacob-
son, and L. Harris was appreciated. M. Green
\'olunteered his time throughout this study, and
R. Taylor allowed access to his property- for
trapping and data collection. Dr. G. Hur.st,
Dr. R. fonas, and H. Shaw reviewed earlier
drafts of tliis manuscript.
LiTER.ATURE CiTED
Alldrf.dgf.. J. R.. and J. T Rvni 1986. Comparison of
some statistical techuiejiii's for iuialysis of resource
selection, jounial of Wildlife Management .50: 157-
16.5.
Bf.wktt D. L. I9S4. Criizinn potential of major soils
within the Black Hills of South Dakota. Unpublished
master's thesis. South Dakota State Uni\ersit\, Brook-
ings. 199 pp.
Boldt, C. E., and J. L. Van Duf.sf.n. 1974. Sikiculture of
ponderosa pine in the Black Hills: the status of our
knowledge. USDA Forest Ser\ice Research Paper
RM-124. Fort Collins. Colorado. 45 pp.
144
Great Basin Naturalist
[Volume 52
Bkvant. F. C]., and D. Nisii 1975. Habitat use by Merriains
Turkey in southwestern Utdi. In: L. K. Halls, ed.,
Proceedings of the Third National Wild Turkey Sym-
posium 3:6-13. Texas Parks and Wildlife Department,
Austin.
Buttery. R. F., and B. C. Gillam. 1983. Forest ecosys-
tems. Pages 43-71 in R. L. Hoover and D. L. Wills,
eds.. Managing forested lands for wildlife. Colorado
Division of Wildlife, in cooperation with US DA Forest
Service, Rock-v Mountain Region, Denver, Colorado.
459 pp.
Byeks, C. R., R. K. Steinhokst. and P. R. Kjuu.sman
1984. Clarification of a technique for analysis of utili-
zation-a\ailability data. Jouniiil of Wildlife Manage-
ment 48: 1050-1053.
Cochran, W. G. 1963. Sampling techniques. John Wiley
and Sons, Inc., New York. 413 pp.
Jonas. R. 1966. Merriam's Turkeys in southeastern Mon-
tana. Techniciil Bulletin 3. Montana Game and Fish
Depiu^tment, Helena. 36 pp.
LUTZ, R. S., and J. A. Crawford 1989. Habitat use and
selection of home nuiges of Merriam's Turkey in
Oregon. Great Basin Naturdist 49: 252-258.
Mackey, D. L. 1982. EcologyofMerriam's Turkeys in south
central Washington with special reference to habitat
utilization. Unpublished m;ister's thesis, Washington
State University Pullman. 87 pp.
. 1986. Brood habitat of Merriam's Turkeys in south-
central Washington. Northwest Science 60: 108-112.
MOSTELLER, F., and A. Parunak. 1985. Identifying
extreme cells in a sizeable contingency table: probabi-
listic and exploratory approaches. Pages 189-224 in
D. C. Hoaglin, F. Mosteller, and J. W. Tukey, eds..
Exploring data tables, trends, and shapes. John Wiley
and Sons, Inc., New York. 527 pp.
Nenno, E. S., and W M. Healy. 1979. Effects of radio
packages on behavior of wild turkey hens. Journal of
Wildlife Management 43: 460^65.
Neu. C. W., C. R. Byers. and J. M. Peek 1974. A tech-
nique for analysis of utilization-availability diita. Jour-
nal of Wildlife Management 38: .541-545.
Petersen. L. E., and A. H. Richardson 1975. The wild
turkey in the Black Hills. Bulletin No. 6. South Diikota
Game, Fish ;uid Parks, Pierre. 51 pp.
Rose, B. J. 1956. An evaluation of two introductions of
Merriam's Wild Turkey to Montiina. Unpublished
master's thesis, Montana State College, Bozemtui. 37
pp.
Scott, V. E., iuid E. L. Boeker 1975. Ecology of
Merriam's Wild Turkey on the Fort Apache Indian
Reservation. In: L. K. Halls, ed.. Proceedings of the
Third National Wild Turkey Symposium 3:141-158.
Texas Parks and Wildlife Department, Austin.
Shaw, H. G. 1986. Impacts of timber harvest on Merriam's
Turkey populations. Problem analysis report. Arizona
Depiirtment of Game and Fish, Tucson. 44 pp.
Shaw, H. G., and R. H. Smith 1977. Habitat use patterns
of Merriam's Turkey in Arizona. Federal Aid Wildlife
Restoration Project W-78-R. Arizona Department of
Game and Fish, Tucson. 33 pp.
Thomas, J. W. 1979. Wildlife habitats in managed forests:
the Blue Mountains ofOregon and Washington. USDA
Forest Service Handbook 553. U.S. Government Print-
ing Office, Washington, D.C. 512 pp.
Received 24 June 1991
Accepted 15 March 1992
Great Basin Naturalist 52(2), pp. 145-148
POLLINATOR PREFERENCES FOR YELLOW, ORANGE, AND RED FLOWERS OF
MIMULUS VERBENACEUS AND M. CARDINALIS
Robert K. Vickerv, |r.
Abstract. — Red, orange, and yellow niorphs of Mimitltis verhetuwens and M. cardiimlis were field tested for pollinator
preferences. The species are closely similar except that M. vcrhoiaccus flowers ha\e partiallv refle.xed corolla lobes, whereas
M. ccirdinalis flowers ha\e fullv reflexed corolla lobes. On the basis of oxer 6()(X) bumblebee and hummingbird visits, highly
significant (/; < .001) patterns emerged. Yellow, which is the mutant color morph in both species and is determined by a
single p;ur of genes, was strongly preferred bv bumblebees ;uid strongly eskewed by Innnmingbirds in both species. Orange
and, to a lesser extent, red were strongK preferred b\ hummingbirds but eskewed by bumblebees in both sjiecies. Thus,
strong, but partial, reproductive isolation was observed between the yellow mutants and the orange- to red-flowered
populations from which they were derived. Color — yellow versus orange iind red — appeared more iniportant than
shape — piirtiallv reflexed versus fullv reflexed corolla lobes — in determining the preferences of tlu- guild of pollinators in
tliis particular test environment for Mimulus vcrhenaceiis and M. cardinalis.
Kl'ij words: Mimulus, spcciation, flower colors, pollinator preferences, bun^lAcbees, luaninin^hirds.
How mucli of a change in flower color ancl/or
shape is enough to lead to a change in pollinators
and hence to reproductive isolation and poten-
tially to speciation? The flower color and shape
nioq^hs o^ Mimulus verbenaceus Greene and M.
cardinalis Douglas provide an excellent system
for addressing this intriguing question.
Materials
Mimulus verbenaceus and M. cardinalis are
tspicallv bright red flowered and hiuiuningbird
pollinated. However, yellow-flowered morphs
occur in M. verbenaceus, e.g., in a population at
N'assey's Paradise, Grand Canyon, Arizona, and
in M. cardinalis populations, e.g., on Cedros
Island, Baja California, Mexico, and in the
Siskyou Mountains, Oregon. My experimental
hybridizations show that yellow is due to a single
pair of recessive genes that limit the floral
anthocyanins to small dots in the corolla throat.
Intermediate, orange-flowered forms are
known in M. verbenaceus, specificallv the pop-
ulation at Yecora, Sonora, Mexico. And, an
intermediate, orange-flowered form of M. car-
dinalis was obtained bv repeated cycles of selec-
tion. In both cases orange is due to a single pair
of quantitative genes that reduce the amount of
anthocyanin pigments in the corolla lobes.
Thus, parallel series of red, orange, and vellow
color forms are available for both M. ver-
benaceus andM. cardinalis (Table 1).
Mimulus verbenaceus and M. cardinalis are
similar, closely related species in section
Enjthranthe (Grant 1924); however, their flow-
ers differ in shape. Those of M. verbenaceus
have only the upper pair of corolla lobes sharplv
reflexed, giving the flowers a partiallv tubular
aspect. The side pair of lobes and the labellum
curve gently forward forming a bee landing [)lat-
form. Flowers of M. cardinalis have both the
upper and side corolla lobes shaq)K' reflexed,
giving the flowers a fully tubular shape. The
labellum is thrust fonvard and is fokk-tl on itself
forming a ridge instead of a landing platform.
Shapes of the flowers of both species would
seem to invite hummingbirds. Flowers of M.
verbenaceus but not those of M. cardinalis
would appear adaj)ted for bees as well. How-
ever, flowers of all three color moq^hs of both
species showed no reflectance patterns in the
ultraviolet, that is, no putative bee nectar
guides. Thus, flower shapes of A/, verbenaceus
and M. cardinalis are similar in some respects
but differ in others of potential significance to
pollinators.
Department of Biolog\', University of Utah, Salt Lake CAW. Utah 841 IS
145
146
Great Basin Naturalist
[Volume 52
Plan
The effect of flower color and flower shape
on pollinator preferences will be addressed
stepwise. First, pollinator preferences for
color — red, orange, and yellow — will be ascer-
tained for M. verhenaceus plants onl)', holding
flower shape constant. Second, red-, orange-,
and yellow-flowered M. cardinalis plants will be
added to the experiment. Are pollinator prefer-
ences for red, orange, and yellow flowers of M.
cardinalis the same as for those of the M. ver-
henaceus series? Note that the pigments are
identical (Vickery 1978). Or, does the difference
in corolla shape between the two species lead to
a difference in pollinator preferences?
Methods
Seeds for each of the six populations of the
study (Table 1) were collected in the wild or
harvested from transplants brought into the
greenhouse except those of orange M. car-
dinalis, which were obtained by selection. A
large population of red M. cardinalis from
Cedros Island was grown and the most orange-
red flowered plant self-poUinated. Its progeny
included several orange-flowered plants. Prog-
eny of these plants were grown and found to
breed true for orange and were used as the
source of seeds for the orange M. cardinalis
moiph.
Seeds of the sLx populations were sown in
early April 1988 in the University of Utah green-
house, following which seedUngs were trans-
planted into 4" plastic pots and grown to
flowering. Pots were placed in plastic travs to
facilitate bottom-watering, plants being ran-
domly arranged as to flower color.
When plants began flowering, they were
moved outdoors to test pollinator preferences.
Instead of using Red Butte Canyon Natural
Research Area as before (Vickery 1990), with its
relative paucity of pollinators, I scattered the
plants on a lawn adjacent to native gambel oak
clumps at the mouth of Parley s Canyon of the
Wasatch Mountains in an area rich in pollina-
tors. Some 50 to 100 plants of each color morph
of M. verhenaceus made up the artificial popu-
lation of the first part of the experiment. Some
50 to 100 plants of red and of orange M. car-
dinalis plus 20 plants of yellow A/, cardinalis (all
that were available) were added to the M. ver-
Tablf, L Origin of populations studied.
Mimulus verhenaceus Greene
Vasscij's Paradise, Grtuid Caiivon, Arizona, USA, elev.
-650 m
Red moq^h = culture number 14,088
Yellow morph = culture number 14,089
Yrcora, Sonora, Mexico, elev. —1,550 m
Orange = culture number 13,256
Mimulus cardinalis Douglas
Isia Cedros. Baja California, Mexico, elev. -100 m
Red moq^h = culture number 13,106
Yellow morph = culture number 13,2.50
Orange = culture number 13,249
(obtained by selection from the red moiph)
henacens plants for the second part of the exper-
iment.
Pollinator visits to the flowers were observed
and recorded for an average of \Vi hours per
observation period for 15 periods for each of the
two parts of the experiment (Tables 2, 3). Time
of day of the observations was varied to be sure
of noting all the different kinds of \isitors. To
count as a visit, a hummingbird had to thrust its
beak into a flower. A bee had to land on the
flower and crawl into the flower far enough to
brush the stigma and anthers. A fly, butterfly,
etc., had to walk on the reproductive structures.
The numbers of flowers rather than plants of
each color of each species were recorded for
each observation period.
For analvsis of visits, chi-square tests were
Rm for each obseivation period for each part of
the experiment. The null hyj^othesis was that
hummingbirds or bumblebees (very few flies,
butterflies, etc., visited the flowers and were not
listed) would visit the three colors of flowers of
M. verhenaceus in the first part of the experi-
ment and the three colors of M. verhenaceus
and M. cardinalis in the second part of the
experiment in proportion to the mmibers of
those flowers in the experimental population
(Tables 2, 3). If the overall chi-square value for
a period of, for example, bee visits to M. ver-
henaceus or hummingbird visits to M. cardinalis
indicated a significant deviation from expected
values, then the frecjuencv of \isits to each color
was inspected. Those high or low enough that
their term in the chi-square equation was large
enough b\' itself to produce a significant devia-
tion at the 5% level were considered to be
significant (Tables 2, 3).
19921
MiMUIA'S FOLIJNATOH PREFERENCES
14'
TablK 2. Pollinator pifffrencL'S for rt-d. orange, or \x'llow noweis ol Mimulus rcrhciuiccii.s in 19S8.
Numbers of flowers
Bumblebee visits
Hummingbi
ird visits
Month:cla\:time
Red
Orange
Yellow
Red
Orange
Yellow P
Red
Orange
Yellow P
7:26:1630
48
56
70
28i"
523T
1984-
<.001
3
<.100
7:29:0745
56
91
74
30
50
58
<.200
SIT
29
<.010
7:30:0710
46
79
114
24
67
67
<.010
55T
66
70
<.010
8:02:1640
85
77
74
3i
8ST
53
<.001
27
49
27
<.010
8:03:0630
92
101
133
53
99T
81
<.()01
33
79T
36i
<.001
8:03:1.540
120
117
172
3U
74
209T
<.()01
lOOi
24 IT
183
<.001
8:04:0640
S6
73
178
(U
5
52T
<.0()]
S3
145T
170
<.001
8:05:0715
120
UK)
169
33i
71
125
<.()01
9i
77T
28i
<.001
8:05:1645
126
104
174
12i
22i
126T
<.001
36i
149T
92i
<.001
8:05:1830
126
104
174
5i
4i
73T
<.001
75
150T
82i
<.001
8:06:0<S40
126
88
151
74i
159T
291T
<.001
66
lOOT
26i
<.001
8:06:1445
126
9S
150
6i
61
60T
<.001
49
94T
24i
<.001
8:06:1810
130
117
142
50
105
257T
<.001
31
4ST
U
<.001
8:07:1515
130
119
142
Oi
4i
68T
<.001
52
125T
5i
<.001
8:08:0725
118
91
124
12i
32
131 T
<.001
32
67T
5i
<.(X)1
â– "T or J. = significantly high or low; see text.
Table 3. Pollinator preferences for red, orange, or yellow flowers of M. verhenaceus and M. canliiuilis in 1988.
Number of flowers
Month:day:time Red Orange Yellov
Bumblebee visits
Hummingbird visits
Red
Orange
Yellow P
Red
Orange
Yellow P
Mil
mtiltis vc
rhenacens
17i''
29
62T
<.(X)1
23
40T
34
<.001
16i
36
131T
<.001
70
70T
184
<.()01
2i
2U
124T
<.(K)1
171
167T
1.354
<.001
841
129T
190T
<.001
13
10
3
<.001
13i
56
202T
<.(K)1
404
80T
384
<.001
.844-
1()6T
237T
<.(M)1
60
.5()T
04
<.001
mi
97
160
<.0Ol
196
166T
994
<.001
5i
94
120T
<.(K)1
168
147T
163
<.001
54i
66
172T
<.(K)1
115
63T
.564
<.001
3i
4i
160T
<.0()1
44
31 T
24
<.001
27i
37
162T
<.(X)1
71
WT
244
<.()01
39i
36
167T
<.001
54
37
38
<.010
2i
3
50T
<.(K)1
7
2
2
<.300
14i
84,
174T
<.CX)1
21
.3()T
04
<.001
3i
24
128T
<.(M)1
66
72T
264
<.0()1
Mil
<nultis CO
rdinalis
36-L
59
89T
<.001
28
25
14
<.001
2U
37
34T
<.001
61
36
04
<.001
18
8
22T
<.(X)1
137
117T
244
<.001
49
59
26
<.010
4
6
<.1(X)
4S
104
48T
<.(X)1
81
.53
124
<.010
274
62
49T
<.0()1
.34
20
<.020
33
26
34T
<.(X)1
6,3
95T
13
<.001
18
S
10
<.().50
1.30
88
21
<.100
35
40
17
<.010
59
41
04
<.010
28
15
16T
<.0Ol
91
92
14
<.300
39
24
19T
<.(X)I
77
52
34
<.020
20
33
26T
<.(X)1
.55
58
04
<.001
13
4
6T
<.010
11
24
<.010
18i
21
34
<.(X)1
.34
74T
04
<.001
19i
47
12
<.(X)1
204
115T
44
<.001
8:08: 16(M)
8:09:07.50
8:09:1705
8:10:0815
8:10:1640
8:11:0810
8:12:0805
8:12: 17(X)
8:13:08.55
8:1.3:1800
8:14:0815
8:15:0740
8:1.5:17(X)
8:16:08.30
8:17:06.30
8:08: 16(X)
8:09:0750
8:09:1705
8:10:0815
8:10:1640
8:11:0810
8:12:0805
8:12: 17(X)
8:13:08.55
8:1.3: 18(X)
8:14:0815
8:15:0740
8:1.5:1700
8:16:08.30
8:17:06.30
117
92
132
115
73
116
115
73
116
145
90
143
145
90
143
175
83
177
200
111
198
200
HI
198
180
83
175
180
87
175
212
81
165
184
94
183
184
94
183
206
112
1.53
214
S6
177
79
47
61
69
45
32
69
45
32
61
39
23
61
39
23
61
55
12
65
51
18
65
51
18
64
42
14
89
81
14
83
69
15
53
71
15
53
71
15
79
78
21
89
79
18
'T or i = significantly liigh or low; see text.
148
Great Basin Naturalist
[Volume 52
Results
Pollinators showed clear, veiy highly signifi-
cant if) < .001) preference for or avoidance of
yellow flower color, but less clear preferences
for or avoidance of orange or red flower colors.
Bunil)lehees — ^principally Bomlms appositiis and
B. huntii — strongly preferred vellow in both M.
verhenaceus and M. cardinalis. Difference in
shape did not appear to matter. Humming-
birds — principally Selasphonis plati/cenis —
strongly eskewed yellow in both species (Tables
2, 3). Agiiin, difference in shape did not appear
to matter.
Hummingbirds significantly {p < .001) pre-
ferred orange M. verhenaceus flowers and
showed a tendency to prefer orange M. car-
dinalis flowers as well (Tables 2, 3). This prefer-
ence for orange over red flowers should not have
been surprising in view of the fact that orange
and red are equally conspicuous to humming-
birds (Grant and Grant 1968, Raven 1972).
Strong preferences and aversions for yellow
are particularly interesting because yellow is the
mutant color in both species. So, a new yellow
mutant of either species would be preferentially
visited by bumblebees and preferentially
avoided bv hummingbirds, but not in all-or-
none reactions. Apparently then, with the spe-
cies of pollinators tested, we are seeing the
establishment of real, but partial, reproductive
isolation due to the mutation of a single pair of
color senes.
Literature Cited
Grant. A. L. 1924. A monograph of the genus Mimiilus.
Annals of the Missouri Botanical Garden IL 99-.3S9.
Grant, K., and V. Grant. 1968. Hummingbirds and their
flowers. Columbia University Press, New York. 11.5 pp.
Raven, R H. 1972. Why are bird-\'isited flowers predomi-
nantly red? Evolution 26: 674.
ViCKERY, R. K., Jr 1978. Case studies in the evolution of
species complexes in Mimulus. Evolutionary Biology
11:404^506.
. 1990. Pollination experiments in the Mimulus
cardinalis-M. lewisii complex. Great Basin NatunJist
.50: 1.55-159.
Received 21 October 1991
Accepted 1 May 1992
Crrat Basin Xatiinilist 52(2), pp. 149-154
SOIL LOOSENING PROCESSES FOLLOWING THE ABANDONMENT
OF T\VO ARID WESTERN NEVADA TOWNSITES
Paul A. Kiiapp
ABSTRACrr. — Soil compaction was measured at four sites within two abandoned mining camps in the western CIreat Basin
Desert, Ne\ada. Bulk densitv* and macroporositv xiilues were generated from soil samples collected in areas ol different
liuid use intensities in camps that had been abandoned for approximatek- 70 \e;irs. Results show that significant differences
remain in bulk density values bet^\'een abandoned roads and undisturbed areas in both towiis, and that the areas around
foundation peripheries ;u"e still signifkiuitK' more compacted in one towai. There were no significant differences between
liuid use groups as measured bv macroporositv. Estimated soil recoven', based on a linear model using bulk densitv v;iJues,
suggests that appro.ximatek- KX) to 1.30 ve;u-s are necessary for complete loosening to occur for abandoned roads, and that
100 or fewer \ears are necessar\- for complete amelioration of the foundation peripher\- iireas. The wetter towaisite, with
more freeze-thaw davs, finer-grained soils, and greater plant cover, had shorter recoverx estimates. These findings suggest
that die results of human-use impacts in arid areas may still be apparent long ;ilter disturbances cease.
Ki-i/ tcords: soil rccovrn/. soil roinpnrfioii. arid hnuh. Great Basin Desert, g/iosf toiins.
Arid lands are undergoing enxironmental
degradation processes at a rapid rate worldwide
and are being severely disturbed by excessive
soil erosion and salinization (Allen 1988, Goudie
1990). The explosion in human population
levels in the last sexeral decades in arid regions
has been a major cause for land degradation,
especially considering that arid regions are
particularly sensitive to anthropogenic land use
impacts (Goudie 1990). While the greatest
extent of soil degradation has occurred in StJiel-
ian Africa, other arid zones of the world are also
\ulnerable (Goudie 1990).
The arid American West is one such region
where human use impacts liave risen dramati-
cally in the last sexeral decades (Francis and
Ganzel 1984). The increased popularits' of back-
country visits by off-road vehicles, mountain
bikes, backpackers, or horseback riders has had
a considerable impact on the surrounding envi-
ronment, either damaging or altering both the
flora and soils of affected areas (Cole 1983,
1987, 1990, Lathrop 1983, Webb 1983, Prose
andMetzger 1985).
Compaction of desert soils caused bv back-
country activities can decrea.se infiltration rates,
increase nmoff, and impede plant root growth,
which favors further soil degradation processes
(Vollmeretal. 1976, Rowlands and Adams 1980,
Hincklev et al. 1983, Lathrop 1983, Prose et al.
1987, Goudie 1990). While the impacts of back-
countiv activities have been documented over
short time spans (often less than 30 years), little
is known about long-term consequences of
these activities (Knapp 1991). Few studies exist
that document how well a disturbed area recov-
ers following cessation of disturbances, particu-
larly in areas traditionally considered to have
little economic value, such as arid lands.
Recovery processes of compacted soils are
not well understood (Webb et al. 1983, 1986)
and liave been conducted primarily in more
mesic environments (Webb et al. 1983, Knajip
1989). Recovery estimates varv' considerably,
ranging from less than 10 years on Minnesota
forest soils (Thorud and Fris,sell 1976), to 23
years on Idaho forest .soils (Froehlich et al.
1985), to 50 years on forest soils in South Aus-
tralia (Greacen and Sands 1980), and up to 200
years on soils in southwestern Montana (Knap[)
1989).
The few studies that have examined .soil
recoven rates in the arid American West have
been confuuHl to the Mojave Desert (i.e., Webb
and Wilshire 1980, Webb et al. 1983, 1986,
1988, Prose and Metzger 1985). Rates of .soil
recoven from the.se studies of abandoned
mining camps ranged from 80 to 140 years and
Department of GedKrapliy. Uiii\ersit\ of Nevada. Keiio, Nevada S9.557-0()4S.
149
150
Great Basin Naturalist
[Volume 52
-^â– ^^j^!^f:^i'^4^Mtijik
Fig. la. Terrill, ca 1920, looking northwest. Photo by Roly Ham, courtesy Special Collections, University of Nevada,
Reno, Librarv.
â– -"NIC^,
V^
^*^* «dlirHi-
Fig. lb. Terrill, 1990. Photograph I )\ author.
averaged 100 years. Comparable studies have
yet to be conducted in the Great Basin Desert.
Ghost towns abandoned in the earlv twenti-
eth century in the western Great Basin Desert
showcase the long-term effects of soil compac-
tion. Built because of the discovery of valuable
ores such as gold and silver, these towns were
short-lived as the ores became too scarce to
extract profitably (Palier 1970, Carlson 1974,
Shamberger 1974). These towais have been
1992]
Great Basin Soil Kkcon ehy
151
Table 1. Cliinatic ;uicl soils data lor tlie hvo selected Great Basin Desert townisites.
Est.
Est."
p:st.°
annual
mean
mean
Sand, silt
Townsite
Ele\atioii
(m)
precipitation
(mm)
Jan. temp.
{°C)
JiiK temp.
■(°c)
.Soil
type
and cla\
(%) '
Terrill
Wonder
1305
1740
125
2.50
-0.8
-3.9
22.8
20.5
loamy sand
sandy loam
S4/12/4
46/50/4
SoiiR-e of estimate: HouglUon et al. 1975.
ex|X).sed to a xarietv of envdronmental impacts,
including trampling by livestock, humans, and
\ehicles, and lia\e shown a \ariet\' of vegetation
reco\er\' responses (Knapp 1992). The piupose
of this paper is to examine the effects of soil
reco\"er\- in two abandoned mining towns in the
Great Basin De.sert in similar fashion to those
studies conducted in the Mojave Desert.
Study Areas
Two measures of soil compaction, bulk den-
sity' and percentage macroporositv; were gath-
ered from Terrill and Wonder. Terrill (39°05'N,
11S°46'\V) and Wonder (39°35'N, 118°04'W)
were abandoned in ca 1915 and ca 1925, respec-
tivelv (Figs, la, lb). Both sites lie at the base
of north-south trending fault-block mountiiin
ranges in central western Nevada, although
Terrill's elevation ( 1305 m) is substantially lower
than Wonder's (1740 m). Terrill is the drier site,
receiving approximatelv 130 mm of precipita-
tion annuallv with the estimated mean January
and July temperatures being -0.8 G and 22.8 G,
respectively (Houghton et al. 1975; Table 1).
The vegetation in Terrill is a salt desert scrub
habitat txpe (Tueller 1989), and common spe-
cies are the shnibs Sarcobatus bailey i, Atriplex
confertifolia, and Tetradijmio spp.; the grasses
Onjzop.sis Ju/men()i(les and Bromus tectoniin;
and the forb Spliaeralcea ainlji^iia. (iroimd
cover in Terrill is approximatelv 2()9f (Knapp
1992). Wonder receives appro.ximateK' 250 mm
of annual precipitation, has mean January- and
July temperatures of -3.9 G and 20.5 G, respec-
tively (Houghton et al. 1975; Table 1), and sup-
ports a sagebnish/grass habitat t\pe (Tueller
1989) with appro.ximatelv 35% ground cover
(Knapp 1992). Gommon species in Wonder are
the shrub Artemisia tridentata and the grasses
6. tectonim and Sitatiiou hifstrix.
Both towiisites have alluviallv deposited, vol-
canic sandv-loam to loam\' sand soils (Stewart
and Garlson 1978; Table 1). The soils in Terrill
are sandy, mixed, T\pic Galciorthids, while
Wonders soils are fine-loamv; mixed, T\pic C Galci-
orthids (USDA-SGS 1975). Organic matter was
estimated to be less than 1% at both towiisites.
Terrill and Wonder have been subjected to
minimal human -caused impacts since abandon-
ment because of their remote locations. Little
grazing by domestic animals has occurred in
Terrill because of the lack of a nearby water
source. Wonder has experienced greater graz-
ing pressiues by sheep, cattle, and feral horses.
Neither sheep nor cattle have grtized the
Wonder area since 1980 (A. Anderson, District
Range Gonservationist, BLM, personal com-
munication, 1990).
Methods
Soil samples for bulk densitv and macro-
porosit)' measurements were gathered at foiu"
different land use categories at each town. Data
were collected from active roads (to get a theo-
retical upper limit of compaction), abandoned
roads (representing prior high-intensit)' land
use), areas within 5 m of foundation peripheries
(representing prior moderate-intensit\' land
use), and contrf)l plots (an^is of minimal distur-
bance located near [<2 km] the townsite). .All
efforts were made to ensure that the four differ-
ent land use groups within each towaisite were
similar to each other in terms of slope, aspect,
.soil texture, elevation, and parent material so
that accurate coiuparisons could be mack-. Trails
caused bv either (era! liorses or small iiianmials
were avoided.
Soil data from the controls, active roads, and
abandoned roads were gathered using a strati-
fied, imaligned sampHng method. Thirt}' 5-m
line transects were set parallel to both the active
and abandoned roads, and one soil core was
gathered at a random point along each line
transect. Soil cores from control plots were
gathered at a random point on each of fort}- 5-m
line transects. Soil cores were also gathered at a
152
Gre.at Basin Natuh.m.isi
[X'olume 52
T.ABLE
2.
Bulk del
Lsity
and macroporo.sity \
iilues.
and
recovei^' period estim
ate.s
for
abandoned townsites.
Bulk Den.sity"'
(g/cm^)
Macroporosity
(%byvol.)
Recovery period (years)
Site
Bulk densit\' Macroporositv
Terrill
Active road
1.65 ± 0.04''
19.7 ± 2.4''
Abandoned road
1.51* ± 0.05
21.6* ± 2.3
Foundation
peripheries
1.47* ±0.07
21.9* ±2.7
Control plot
1.41*'^ ± 0.03
22.7* ± 2.4
1.59 ± 0.08
Wonder
Active road
17.3 ± 2.0
Abandoned road
1.48* ± 0.07
20..3* ± 1.9
Foundation
peripheries
1.46* ± 0.07
20.8* ± 0.8
Control plot
1.42* ±0.06
21.1* ± 1.1
130
100
100
85
120
100
80
70
â– "Bulk density data with exception of active roads and standard deviation vahies are trom Knapp 1992.
One standard deviation.
= Significantly different {p = .0.5) from active road based on Tukey test.
=; Significantlv different ip = .05) from control plot based on Tiikev test.
= Significantlv different (/) = (15) from fbnndation peripheries based on Tukey test.
random point on each of forty 5-ni line transects
that were set perpendicular to the foundation
periphery sides. The cores were oven-dried
overnight and then weighed for bulk densitv
(Blake 1965). One-fourth of the cores also were
kept intact for macroporosity readings that were
measured under 30 cm of tension using a ten-
sion-table (Orr 1960). Soil te.xture was mea-
sured using the micro-pipette method (Miller
and Miller 1987).
Analysis of variance (ANOVA) was used to
examine whether differences in either bulk den-
sity or macroporosity values existed between
land use categories (abandoned roads, founda-
tion peripheries, and control plots) for each
town (Zar 1984, SAS 1985). Where significant
overall differences existed, Tukey multiple com-
parison tests were used to determine between
which groups these differences occurred (Zar
1984). Soil recovery was considered complete
when no significant differences existed between
disturbed sites and their respective control
plots.
Soil recover)' estimates were based on the
equation (corrected from Webb et al. 1986):
T=[(Ia-Iu)/(Ia-It)]'TA
where It = townsite (either abandoned road or
foundation periphery)
In - undisturbed soils (control plots)
la = active road
Ta = time since abaudomnent of
towTisite
The data collected from active roads were used
only for estimates generated by this equation.
This equation generates rough estimates of soil
recoven.' times. Webb et al. (1986) state that an
exponential deca\' model might gi\e more real-
istic soil reco\eiy estimates, although onlv one
abandonment time per site excludes the use of
the exponential decay model.
Results
Bulk densit\' measurements for the aban-
doned road (1.51 g/cm^) and foundation periph-
eries (1.47 g/cm') were significantl)- greater dian
for the control plot (1.41 g/cm^) in Terrill, but in
Wonder onlv the abiuidoned road (1.48 g/cm^)
had significanth' greater bulk densits' values
than tiie control plot (1.42 g/cm^) CRible 2).
Macroporositv' measurements in botli Terrill and
Wonder were not significantK different between
land use categories.
Estimated recoven' times ranged from 85 to
1 30 years when based on bulk density measure-
ments, and from 70 to 120 years when based on
macroporositv measurements (Table 2). All
measiuvments were greater on abandoned
roads than on foimdation peripheries and were
comparatively longer in Terrill than in Wonder.
While these values are derixed b\ a linear recov-
eiv model, it is most likeh that soil recovery
follows more of a nonhnear path with rapid
reco\eiy early, then recover)' rates slowing.
1992]
Great Basin Soil Hkcon ehv
153
Heinonen (1977) has suggested that the l)ulk
densitv' of soils may decrease to a certain point,
then le\el off without reaching predisturbance
conditions.
Discussion
Soil loosening is dependent upon shrink-
swell, freeze-thaw, and biological acti\it}- pro-
cesses (Larson and Alhnaras 1971, Akrani and
Kemper 1979, Webb 1983, Webb et al. 1986,
Knapp 1989). These processes in turn may be a
function of soil type, climate, and biological
acti\it\. The recover)' times for Terrill and
Wonder show a relationship to all three ot these
processes, with recovery times in Terrill being
longer than those in Wonder.
Soil texture is important because finer-
grained soils are more prone to freeze-thaw and
shrink-swell loosening processes than are
coarser-grained soils (Webb et al. 1986). Fine-
textured soils have more total pore space and
have a higher water-holding capacit\', thereby
pro\'iding the soils of Wonder, that are more
fine-grained than Terrill, more opportunities
for expansion-contraction processes to occur
(Millar et al. 1958). While percentages of clay
mav also be important, particularly if the clay
has a high shrink-swell ratio, total amounts of
clay at the two towns were the same and should
not have a greater effect at one place than at the
other.
Climate plaws an important rc^le in soil loos-
ening processes, particularlv where there is a
high frequency of wetting and drying, freezing
and thawing, or heating and cooling processes.
Three climatic features favor faster soil loosen-
ing processes in Wonder than in Terrill. First,
\V onder is 435 m liigher than Terrill and Wonder
has a shorter freeze-free period by approxi-
mately a month to a month and a half (J. James,
Nevada State Climatologist, personal communi-
cation 1991). Second, Wonder lies at the base of
a bowl-shaped depression and receives maxi-
mum cold-air drainage. Typical diurnal temper-
ature contrasts for Wonder range from 22 to 28
C, with the greatest contrasts occurring in the
summer and the least contrast in the winter
(Houghton et al. 1975, J. James, personal com-
munication, 1991). Terrill, on the other hand,
experiences a 16.5 to 22 C diurnal temperature
range (Houghton et al. 1975, J. James, personal
communication, 1991). These differences in
diurnal temperature range suggest that the
heating-cooling and expansion-contraction pro-
cesses are more pronounced for Wonder Third,
Wonder receiws approximateK twice as much
annual precipitation as Terrill; therefore, the
freezing-thawing and wetting-diving processes
should occur more often in Wonder, facilitating
the soil loosening processes.
Biological acti\it\' through plant root growth
can also ameliorate soil compaction. Cxrasses
and forbs are particularly effective for loosening
of topsoil because they have manv diffuse, shal-
low roots that penetrate the topsoil with subse-
quent minimal increases in soil strength, but
leave behind small channels after the roots die
(Webb et al. 1983). Plants such as shrubs, with
a central taproot, however, cause localized com-
paction around the root, yet have fewer roots
per unit volume and are less effectixe for soil
loosening (Webb et al. 1983). Total plant cover
in Wonder was substantiallv (approximately
20%) greater than in Terrill, especialK* with the
grasses Bromns tectontm and Sitanion Jii/strix,
which both have extensive, shallow root sys-
tems. Therefore, it appears that if soil loosening
can be attributed to biological activity; it would
be more pronounced in Wonder than in Terrill,
although controlled, detailed experiments are
necessai"v for confirmation.
Conclusions
After 75 years of recoveiy, significant dilfer-
ences remain between disturbed and undis-
turbed sites in Terrill as measured by bulk
densitv. Estimates for recoverx based on bulk
densitv' are from 100 to 130 \ears. In Wonder,
after 65 years of recover) , significant differences
remain only between abandoned roads and con-
trol plots. Estimated reco\en- for the aban-
doned road is 100 \ears. These results an^ in
close agreement with similar, previous studies
that examined soil reco\en' times in the Mojave
Desert (e.g., Webb and Wilshire 1980, Wehh et
al. 1986) and suggest that the results of soil
compaction processes that occur in arid en\i-
ronments are long-li\('d. but aiv not irreversible.
Ac K N OW L E D C M E N TS
I wish to thank the Universit)- of Nevada
Graduate School for funding, Louis R. Loftin
for field and laboratorv assistance, and Diana F.
Thran, Can J. Hausladen, and Chris R. Ryan
for comments and suggestions.
154
Great Basin Naturalist
[Volume 52
Literature Cited
Akham, M., and \V. D. Kemper 1979. Infiltration of soils as
affected bv die pressure and water content at the time
of compaction. Soil Science Societ\' of America Journal
43: 1080-10S6.
Allen, E. B. 1988. Introduction. Pages l-Ain E. G. Allen,
ed., The reconstruction of disturbed arid lands.
Westxdew Press, Boulder.
Blake. G. R. 1965. Bulk density. Pages 374-390 m C. A.
Black at al., eds.. Methods of soil analysis: part 1,
physical and mineralogiciil properties. American Soci-
ety' of Agronomy, Madison, Wisconsin.
Carlson. H. S. 1974. Nevada place names. University of
Nevada Press, Reno.
Cole, D. N. 1983. Campsite impact on three western wil-
derness tireas. Environmental Management 7: 27.5-288.
. 1987. Recreational impacts on backcountry
campsites in Grand Canvon National Park, Arizona,
USA. En\ironmental Management 10: 651-659.
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Received 5 Ati^ust 1991
Accepted 16 April 1992
Great Basin NatiinJist 52(2). pp. 155-159
BIOCHEMICAL DIFFERENTIATION IN THE IDAHO GROUND SQUIRREL,
SPERMOPHILUS BRUNNEUS (RODENTIA: SCIURIDAE)
Ayesha E. Gill and Eric Yensen""'
Abstract — SpenmypJdltis hnmncus is restricted to a 90 x 125-km area of west central Idalio, with two distinct (northern
and southern) groups of populations \sithin this limited range. Morphological differences in pelage length and coloration,
e.xtemal and cr;uii;il measurements, imd hacula suggest that these groups are either \er\' distinct subspecies or species. We
used starch-gel electrophoresis to estimate the tmiount of genetic differentiation accompan\ing these morphological
differences bv assaying genetic \ariation at 31 loci in the two geographic groups. Fifteen lcx.i were poKuioiphic (13 in the
northern group, 12 in the southern), and mean heterozNgosits' (H) was high (12.3% northern and lO.S'^ southern). Nei's
genetic distance (0.057) is in the range usualK' associated with subspecific differences. However, Jaccards association
coefficient (0.893) is about the same as that found between se\eral ground squirrel taxa currently recognized as species.
The high levels of heterozygosity suggest that S. hntntwtis is a neoendemic rather than a paleoendemic species.
Ki'ij words: .SjTermophilus bnumeus, Spermophilus, Iddlio <^nniiul s(jiiinvl. <^r(>uiul s<juinTl.s. electrophoresis, taxonoiiiij.
biochemical differentiation.
Spermophilus hnmneus is one of the rarest,
least studied, and most geographically restricted
of the North American ground squirrels. Within
its restricted range of ca 90 X 125 km in west
central Idaho there tire two well-differentiated
subspecies, S. /;. hninneus and S. b. endemicus
(Yensen 1991). Significant differences in pelage
length and color, e.xtemal and cranial measure-
ments, and bacular moq^holog)' suggest that the
two taxa ma)' be close to species-level separation
(Yensen 1991). The northern Spernwphihis
h. hninneus is known from onl\' ca 20 isolated
sites in mountain meadows in Adams and \^alley
counties. These denies consist of <200 individ-
uals and are separated from each other b\- dis-
tances of 1-70 km. In contrast, the southern S.
/;. endemicus is patchily distributed over a con-
tiguous area 70 km long and up to 20 km wide
in the lower-elevation foothills of Gem, Payette,
and Washington counties (Yensen 1991).
Da\is (1939) divided the North American
species of subgenus Spermophilus into "small-
eared" and "large-eared" groups and placed S.
hninneus within the large-eared group. Nadler
et al. ( 1973) found, however, that the karyotyjoes
of S. hninneus and S. townsendii mollis (small-
eared group) differed onlv in the presence or
staining intensity' of minor bands on six chromo-
somes, indicating a close affinity behveen S.
hninneus and the small-eared group. Nadler et
al. (1974) analyzed serum transferrins of S.
hninneus using starch-gel electrophoresis and
concluded that it was biochemicallv "intermedi-
ate" and possiblv ancestral to both the Nearctic
"small-eared" and "big-eared" species groups of
subgenus Spennophilus. Nadler et al. (1982)
extended their analysis to 21 Holarctic species
using 18 loci and concluded that S. hninneus
was a paleoendemic species most closeK' related
to the Eurasian S. dmiricus. Nadler et al. (1984)
revised their phvlogeny to incoq3orate chromo-
somal data and placed the e\'()lutionaril\' con.ser-
vatixe S. hninneus within the S. townsendii group.
The present study was conducted to estimate
the genetic differentiation accompaming the
substantial moiphological differences behveen
the two geographic groups of S. hninneus and
to assess the hypothesis that S. hninneus is a
paleoendemic species with small, rclictual
populations.
Material. s and Methods
Specimens AnaK/.ed
A total of 82 specimens were analyzed from
the following localities: Spermophilus hninneus
^ University of Nevada, Reno, Nevada 89557, Present address: Institute of I leallli Policv Studii
'Museum of Natural History', Albertson College, Caldwell. Idaho 83605.
.\ddress for reprint requests.
versitv of California. San Francisco. California 94143.
155
156
Great Basin Naturalist
[Volume 52
bniniu'iis — Adams Co.: 1 mi NE Bear Guard
Station, 3; Bear Cemetery, 2; Cold Springs Cr.,
1; Little Mud Cr., 5; Mill Cr. 3 mi N Hornet
Guard Station, 2; New Meadows, 12; Price
Vall(n, 2; Lick Cr., 6; Summit Cr., 9. Sper-
mophiliis hninneus endemicxis — Gem Co.:
Sucker Cr. 11 mi N Emmett, 20; 12.6 mi N
Emmett, 1; Payette Co.: Big Willow Cr., 1; Dr)/
Cr. Road, 3; Washington Co.: Lower Mann Cr.,
10; Weiser Cove, 5. These specimens have been
deposited as vouchers in the Albertson College
Museum of Natural History.
Laboratory' Methods
Blood was collected from the suborbital
sinus of living animals (samples sizes were 21 S.
h. hnmneiis, 9 S. b. endemicus). Liver and
kidney tissues were from sacrificed animals ( 10
S. b. bniniietis, 6 S. b. endemicus) or frozen
carcasses collected for other purposes (IS S. b.
hninneus, 31 S. b. endemicus). Carcasses were
stored at -20 C for 1-6 months.
Tissue sample preparation and horizontal
starch-gel electrophoresis follow Selander et al.
( 1971 ) with slight modifications. We used 1 1 .0%
electrostarch for lithium hydro.xide gels and
12.4% for all other gels. Enzyme locus designa-
tions follow standardized Enzyme Commission
(E.G.) nomenclature (Harris and Hopkinson
1976). The enzymes and nonenzymatic proteins
screened in this studv, with tissue and buffer
svstems used, were: alcohol dehydrogenase,
E.G. No. 1.1.1.1 (ADH), liver, tris'-citrate, pH
8.0; glycerol-3-phosphate dehydrogenase, E.G.
No. 1.1.1.8 (GPD), liver, tris-citrate, pH 8.0;
L-iditol dehydrogenase, E.G. No. 1.1.1.14
(IDDH), liver, tris-citrate, pH 8.0; lactate de-
hydrogenase, E.G. No. 1.1.1.27 (LDH), kidney,
tris-citrate, pH 8.0; malate dehydrogenase, E.G.
No. 1.1.1.37 (MDH), liver, tris-citrate, pH 6.3;
isocitrate dehydrogenase, E.G. No. 1.1.1.42
(ICD), kidney, tris-citrate, pH 8.0; superoxide
dismutase, E.G. No. 1.15.1.1 (SOD), kidney,
tris-maleate or tris-citrate, pH 8.0; aspartate
aminotransferase, E.G. No. 2.6.1.1 (AAT), liver,
lithium hvdroxide; hexokinase, E.G. No. 2.7.1.1
(HK), kidney, tris-citrate, pH 8.0; phosphoglu-
comutase, E.G. No. 2.7.5.1 (PGM), kidney, tris-
citrate, pH 8.0; esterase, E.G. No. 3.1.1.1 (ES),
hemolvsate, tris-hvdrochloric acid; peptidase,
E.G. No. 3.4.1 1 orl3.'' (PEP), liver, tris-citrate,
pH 6.3; hemoglobin (HGB), hemolysate, tris-
hydrochloric acid; albumin (ALB), plasma, lith-
ium hydroxide; transferrin (TRF), plasma,
litliium hydroxide; general proteins (GPl and
GP2), hemolysate, tris-hydrochloric acid; and
general proteins (GP3 and GP4), plasma, tris-
hvdrochloric acid. The proteins were numbered
in order of decreasing mobilitv, with the most
anodal labeled 1.
The buffer and stain systems for the proteins
screened in this study were described by Selan-
der et al. (1971), except for stains for'iDDH,
HK, and PEP (Gill et al. 1987). Of the esterases,
only acetylesterases were stained and were
numbered 1 (most anodal) to 5. PEP-G was
detected with L-leucyl-L-alanine. ADH does
not have to be stiiined specifically and is seen on
many dehydrogenase gels. It was read on gels
stained for GPD.
Computational Methods
Gene frequencies, measures of genetic vari-
ation, Nei's (1978) unbiased genetic distance
and unbiased genetic identity, and the average
inbreeding coefficient (Est) were derived from
input on single individual genotypes (elec-
tromoiphs) using the computer program
BIOSYS-1 (Swofford and Selander 1981).
Jaccard's association coefficient, S, - a/(a+u),
where a = the number of matched elec-
tromoq3hs (1:1) and u = the number mis-
matched (1:0 or 0:1) (Sneath and Sokal 1973),
was also calculated for the two groups. Sj
depends only upon the presence ( 1 ) or absence
(0) of alleles, as indicated b\' bands on the starch
gels (electromoq^hs), not on cillehc frequencies
as do measures of genetic distance. Negative
matches were excluded.
Results and Discussion
SpermopJidus b. bninneus was polymorphic
at 13 loci (42%), whereas S. b. endemicus was
polyinoq:)hic at 12 loci (39%). If esterases are
excluded, polvmoq^hism is reduced to 31%,
which is similar to the 29% reported for Mus
rnusculus and Homo sapiens (Lewontin 1974).
Average number of alleles per locus (A) was 1.48
±0.11 (X ± SE) in S. b. bninneus and 1.48 ±
0. 12 in S. b. endemicus. All polymoqjhic loci had
two alleles, except for peptidase and hvo of the
esterases, which had three.
Mean heterozvgositv per individual per
locus in our sample was 12.3 ± 3.7% in S. b.
bninneus and 10.8 ± 3.9% in S. h. endemicus.
These values are much higher than the 2.7%
heteroz)'gosit)' reported b) Nadler et al. (1982)
1992]
Sfehmoi'hiia'sbri'nneus Electropiiokksis
157
for S. h. hniniu'us. The loci coininoii to both
studies, however, were less variable than some
of our 18 additioHcil loci. Even if esterases are
excluded from the anahsis, our measures of
genetic variabilis (S. b. bniiineus, H = 8.2%, A
= 1.35; S. b. endemicus, H = 7.4%, A = 1.38) are
still much higher than theirs. They found H
values of 0.0-10.4% (X = 3.5%) for other species
o{ SpcrniopJiiliis. Cothran et al. (1977) found
high heteroz\gosit\' (9.3%) in the ground squir-
rel subgenus Ictidomys. The average hetero-
z\'gosity for 26 taxa of rodents was 5.4%
(Selander 1975), so Idalio ground scjuirrels ha\e
relativelv high le\els of hetero/Agositv Thus,
the levels of genetic \ariabilit\' are high for a
species postulated to be a paleoendemic
(Nadler et al. 1974, Cothran et al. 1977, Nadler
et al. 1982) with small isolated denies and con-
fined to a small geographic area (Yensen 1991).
Sixteen of 31 protein systems scored for S.
bniiuu'iis were monomoiphic (GPD, LDH-A,
K:D-2, HK-1,2, PGM-1,2, AAT-1,2, iddh,
SOD-B, ADH, ALB, TRF, GP-1,2). Frequen-
cies of alleles in the pol)TOoq3hic systems (the
most common allele <0.99) are shown in Table
1. As in other species (Kojima et al. 1970,
Lewontin 1974), non -glucose-metabolizing
enzymes were more polymorphic than glucose-
metabolizing enzMiies wdth five monomoq)hic
(AAT-1,2, IDDH, SOD-B, and ADH), while
PEP-C, S()D-A and all fixe esterases were poK -
moiphic (Table 1). The two taxa of S. bniniietis
did not differ substantially in glucose- metabo-
lizing enz\ nies, with the majoritv of loci mono-
moiphic, and the sairie allele conuuon in the
poKuioiphic loci.
Nadler et al. ( 1982) found LDH to be mono-
moq:)hic in all 21 North American and Eurasian
Sj)i'nu()j)liilus species examined. Howexer, we
found t\v() indi\iduals of S. b. bntmu'tis that
were homoz)gous for a fast iillele at the LDH-B
locus. Nadler et al. (1982) assayed from LDH in
red blood cells while we used kidney extracts, so
the difference ma\- be between the two tissues.
Both groups of S. bninneus were pol\ nioqihic
for ICD-1 and HK-3, while onl\\S. b. endemicus
was poKmorphic for MDH-1.
Of the enz\nies not invoked in glucose
metabolism, the esterases were the most \ari-
able (Table 1). We also found considerable dif-
ferences between S. b. bninneus and S. b.
endemicus in the other non-glucose-metaboliz-
ing enzymes. Different alleles were conunon for
PEP-C and ES-4 in the two groups of
Tahlk I. Allelic IrcijiR'iicit's oi [Xilvinoipliif
Speniiuphilii.s h ni n iwtis.
Locus"
Allele- "
bninneus
endemicus
Glucose
-METABOLIZINC;
ENZYMES;
LDH-B
a
0.929
L(K)0
b
0.071
0.(X)0
MDH-1
a
0.(X)()
0.018
b
1.000
0.911
c
().(X)0
0,071
ICD-1
a
0.926
0.986
b
0.074
0.014
HK-3
a
0.132
0.097
b
0.868
0.903
NON-GLUCOSE-METABOLIZINC; ENZYMES:
SOD-A
a
0.786
0.957
h -..
0.214
0.043
PEP-C
a
0.365
0.329
b
0.13.5
0..343
c
0.500
0.329
ES-1
a
0.179
0.056 .
b
0.107
0.167 .
c
0.714
0.778
ES-2
a
0.969
l.(X)0
b
0.031
0.000
ES-3
a
0.971
0.944
b
0.000
0.056
c
0.029
0.000
ES-4
a
0.714
0.389
b
0.286
0.611
ES-5
a
0.656
0.944
b
0.344
0.056
Non ENZYMATIC I'KOTEINS
HGB-1
a
0.233
0.667
b
0.767
()..333
HGB-2
a
0.100
0.500
b
0.900
()..500
GP-3
a
0.000
1.000
b
1.000
O.CXK)
GP-4
a
0.962
0.750
b
0.038
0.250
"See text for iicronvm.s of loci.
""Alleles are listed in order of increasing mobilit)'; a is slowest.
S. bninneus. In both cases the differences were
in allelic frequenc)- rather than in the presence
or absence of alleles.
Nonenz\inatic proteins were .scored in both
hemoKsate and plasma, .-\lbumin and transfer-
lin in plasma and hvo general proteins in
hemolysate were monomorphic. We found vari-
ability- at the two hemoglobin loci and at two
general protein loci in plasma (Table 1).
Heterozygosit\ of hemoglobins has been found
in the closely related TowTisends ground s(juir-
rel (S. fownsendii), in which the hvo hemoglo-
bins ha\e identical a-chiiins and differ by only
one amino acid in the sequence of their p-chains
158
Great Basin Naturalist
[Volume 52
(Kleinschniidt et al. 1985). They found no
difference in the oxvgen affinity of the two
lienio2lol)ins.
A general protein in plasma (GP-3) repre-
sented by a band just anodal to albumin distin-
guished the tu'o S. hninneus. A fast allele
apparently has reached fixation in S. h.
hrnnneiis, whereas a slow allele appears fixed in
S. h. cndemiais (Table 1). This is the only locus
that can serxe as a marker gene among the 31
loci scored, although LDH-B and MDH-1 had
alleles that were fixed in one taxon and polymor-
phic in the other. The other presumed loci dif-
fered in allelic frequency only.
Nei's (1978) genetic distance is a measure of
the accumulated number of gene differences
per locus between populations. The genetic dis-
tance of 0.057 found between the two S.
hninneus was within the range associated with
subspecific differentiation (Avise 1974). The
average inbreeding coefficient (Fsr = 0.167)
indicated moderately high genetic differentia-
tion. The two S. hninneus have a genetic iden-
tity of 0.944. By comparison, Cothran et al.
(1977) found genetic identities of 0.808
between S. spilosonia and S. niexicanus, 0.835
between S. spilosonui and S. tridecemlineatus,
and 0.965 between S. tridecemlineatus and S.
mexicanus in the subgenus Ictidonu/s.
To compare our results with other results
from the subgenus Spenriophihis (Nadler et al.
1982), we also calculated Jaccard's association
coefficient. This measure is less sensitive to
sample size and depends on presence or
absence of an allele, rather than on allelic fre-
quencies. Jaccard's coefficient of similarit)'
between the two groups of S. hninneus was
0.893. Judging from Figure 2 in Nadler et al.
(1982:206), the similarity between the two
groups of S. hninneus is about the same as the
similarit)' between S. anmitus and S. heldin^i,
or between some of the putative semispecies in
the S. townsendii complex, the Eurasian S. sus-
licus and S. citeUus, or S. major and S.
enjthro^enijs. SpenmyphUus richanlsoni and S.
elegans are more similar electrophoretically
than the two Idaho ground scjuirrels. However,
direct comparisons are difficult since the simi-
larity coefficients computed by Nadler et al.
(1982) were based on a different, and appar-
ently less variable, set of loci.
The electrophoretic data confirm that the
two Idaho ground s(|uirrels are genetically as
well as moqohologicall\- differentiated taxa. The
evidence does not clearly resolve the question
of whether the two are separated at the subspe-
cies or species level. The presence of one
marker gene and the observed frequency differ-
erices at others could be consistent with either
inteq3retation. The high levels of heterozygos-
ity, however, do not support the paleoendemic
hvpothesis.
Acknowledgments
We thank D. B. Hammond, W. F. Laurance,
and D. A. Stephens for field assistance; P. L.
Packard for specimen shipment; and R. S. Hoff-
man, W. F Laurance, C. F. Nadler, E. A. Rick-
art, P. W. Sherman, O. G. Ward, and two
anonymous reviewers for comments on the
manuscript.
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Systematic Zoologv' 23: 465-48L
CoTHKAN. E. C, E. G. Zimmerman, and C. F. Nadler
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Harris, H., and D. A. Hopkinson. 1976. Handbook of
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Holland Publishing Co., Amsterdam.
Ki.EiN,sc:!iMiDT, T, F. A. BiEBER, C. F. Nadler, R. S.
Hoffman, L. N. Vida. G. R. Honig. and G.
Braunitzeh 1985. The primary structure and func-
tional properties of the hemoglobins of a ground scjuir-
rel (Spcnnophilus townsendii. Rodentia). Biological
Chemistr\-. Hoppe-Sevler 366: 971-978.
KojiMA, K.,J. Gillespie, and Y. N. Tobari 1970. Aprofile
o{ Drosophila species enz\mes assayed by electropho-
resis. I. Number of alleles, heterozygosities, and link-
age di.secjuilibrinm in glucose-metabolising systems
and some other enzymes. BiochemiciJ Genetics 4:
627-6:37.
Lew'ONTIN. R. C. 1974. The genetic basis of evolutionary
change. Columbia Uni\'ersit\ Press, New York, New
York.
Nadler. C. F., L. W. Turner, R. S. Hoffman, and L.
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Spermophilus brunneus Electkopiiorksis
159
Nadlkk, C. F, R. S. Hoffmann. N. N. Vokontson'. J. \V.
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Received 8 JaniKinj 1991
Accepted 18 April 1992
Great Basin Naturalist 52(2), pp. 160-165
NEW GENUS, APLANUSIELLA, AND TWO NEW SPECIES OF -
LEAFHOPPERS FROM SOUTHWESTERN UNITED STATES
(HOMOPTERA: CICADELLIDAE: DELTOCEPHALINAE)
M. W. Nielson' luid B. A. Haws"
Abstract. — A new genns, Aplaimsk'lla (type-species, Aplanusidla utahensis, n. sp.) and bA'o new species, A. ittahemis
and A. calif onriensis , are described antl illustrated. The two species are allopatric and coexist on the same host genus,
(Atriplex) with members of a closely allied leafliopper genus, Aplamis. Notes on distribution of hosts and leaflioppers as
well as leafliopper intergeneric relationships are also given.
Ki'ij words: leaflioppers. new species, new gentis, Cicadcllidae, Aplanusiella, distrihntion. hosts.
In a 1986-89 suney of rangeland leafliop-
pers of Utah (Haws et al. 1989), two populations
were taken from Atriplex spp. and tentatively
identified as members of the genus Aplamis.
One population was later identified as Aplamis
alhidus (Ball) from shadscale, Atriplex con-
ferfifolia (Torn & Frem.) Wats. The other pop-
ulation was collected from four- winged
saltbush, Atriplex canescens (Pursh) Nutt. and
is described herein as a new genus and new
species closely allied to Aplamis. An additional
new species is also described from specimens
collected in California on Atriplex sp. Notes are
given on the ph\1:ogeography of the host genus,
Atriplex, the distribution of the two genera, and
their taxonomic and host relationships.
The general habitus (form and color pattern)
of the component populations are so remark-
ably similar that it is likelv that additional mate-
rial of the new taxa will be found in other
repositories. Only after dissection and examina-
tion of the male genital structures will their tnie
identity be revealed. Moreover, it is probable
that additional new species will come to light
after more thorough collecting is done on
Atriplex spp. in southwestern United States and
northeni Mexico. This assimiption is based on
two additional populations of female specimens
in hand from Nexada and California for which
males are presently unknown and are required
for definitive generic placement. The female
seventh sternal characters appear to place these
populations in the new genus (sensu .stricto).
Populations of these groups are rather rare in
Atriplex host areas of the high- to low-desert
regions of western North America.
Aplanusiella, new genus
T\TE SPECIES. — Aplanusiella utahensis, n. sp.
Small, rather slender species. Related to
Aplamis Oman but smaller and with distinctive
nicile genital characters. General color light
yellow to ivory with numerous, nearly concen-
tric, tiny rufous spots on forewings, spots not
usually forming lines as typically present in
Aplamis, large spots in claviis and in apical
crossveins of costa fonned by aggregation of
smaller spots, pronotum and scutellum some-
times with tiny spots.
Head narrower than pronotum, anterior
margin obtusely angled and rounded to front,
crowii produced mediallv to about one and one-
half times length next to inner margin of eve,
disk somewhat depressed in middle but lacks
transverse depression before apex as in Aplamis;
pronotum and scutellum as in Aplamis; fore-
wings with imier anteapical cell open basally,
appendix well de\ eloped; cKpeus and cKpellus
as in Aplamis.
Male pvgofer with macrosetae in apical half
and with prominent caudoventral spine, some-
times crossing over in caudal view; aedeagus
small, base large in lateral \iew, apical htilf
narrow, tubular, sometimes with smdl angulate
protrusion at base of shaft on dorsal margin,
gonopore subapical on ventral margin; connectiv e
Monte L. Bean Museum. Brigham Young University. Provo. Utah S4602.
Department of Biolog)-, Utah State Universitv-, Logan, Utah 84322.
160
1992]
New Genus, Apianusieija
161
short, Y-shaped, articuhitrd with acdeagus; st\ie
hroad, apophxsis short: plate trianpilate with
row of niacrosetae subinarginalK and row of
microsetae marginally, female seventh sternum
with short projection medially on caudtJ margin.
Two aliopatric species are known that occur
in the southwestern states of Utah and Califor-
nia on desert shrubs of the genus Atriplex.
Aplanusiella can be distinguished from Aplantis
b\ the sniiiller size, by the absence of a preapical
depression on the crown, by the presence of a
prominent caudoventral pvgofer spine, by the
smaller aedeagus that lacks apical processes,
and by the female seventh sternum that has a
more distinctive spatulate process on the
middle of the caudal margin.
Aplanusiella utahensis, n. sp.
Figs, la-ll
Length. — Male 3..5-3.75 mm. female 4.00-
4.20 mm.
General color pale yellow to ivor)' with
numerous, nearly concentric, tiny nifous spots
on forewdngs, large aggregate spots on apex of
clavus and in apical crossveins of costa, some-
times with few similar spots on pronotum and
scutellum. Related to AplamisieUa californien-
sis, n. sp., but with distinctive male genital and
female seventh sternal characters.
Male. — Pygofer in lateriil view with long,
stout caudoventral process that sometimes
crosses its counteipart in caudal view, but usu-
ally closely appressed to caudal margin of pygo-
fer (Fig. lb); plate long, triangulate with
uniserate niacrosetae submarginallv and uniser-
ate microsetae marginallv on outer margin (Fig.
Ic); st)'le in dorsal view long, broad in basal 2/3,
apophysis short, curved and pointed apically
(Fig. Id); connective short, Y-shaped (Fig. le);
aedeagus in lateral view short, ventral margin
abruptly angled near middle, broad basallw
shaft narrow, tubular with basal triangulate pro-
jection on either side of dorsal margin, gono-
pore subapical on ventral margin (Figs. If-lk).
Ft:MALE. — Seventh sternum broadly exca-
vated on caudal margin, with prominent median
spatulate process (Fig. U).
HOLOTYPE (male).— UTAH: Daggett Co.,
Browns park, Pyke plots, roadside, 12.\T.1987,
four-winged saltbush, Atriplex canescens, B. A.
Haws (CAS). Paratvpes, 1 male, Daggett Co.,
Brown's park, 3.5 mi E Jams ranch, 26.\T.19S7,
on four- winged saltbush, Atriplex canescens.
Haws, Nelson (authors collection); 2 males.
1 female, San Juan Co., Div \alley, 8.IX.1987,
four-winged saltbush, Atriplex canescens. B.
Haws, A. Issa (USU); 2 males, 2 females, Uintah
Co., Bonanza, 14.VII. 1975-3.IX. 1976, Afn>/ex
canescens, G. E. Bohart (USU); 1 male. Grand
Co., Jughandle Potash Rd., 19.Vni.l987, four-
winged saltbush, Atriplex canescens, B. A.
Haws, C. R. Nelson (BYU); 1 male. Grand Co.,
Colorado River, Hwy 128, 6 mi NE jet. Uwy 191,
26.V.19H7, Atriplex canescens, B. A. Haws.'C. R.
Nelson (USU).
Remarks, — This species can be distin-
guished from calif orniensis, n. sp., by the longer
caudoventral pygofer process, by the abruptly
angled ventral margin of the aedeagus, b\' the
presence of a small ba.sal triangulate process on
the dorsal margin of the aedeagal shaft, and by
the prominent spatulate process on the middle
of the female seventh sternum.
The species is known from the eastern coim-
ties of Utah bordering Colorado and is likely
present in the western part of that state and in
northern Arizona where the host occurs. Collec-
tion dates suggest that the species has two gen-
erations per \'ear and presuiuabK' cnerwinters as
eggs on its host.
Aplanusiella californiensis, n. sp.
Figs, liii-ls
Length. — Male 3.30-3.50 mm, female
3.60^3.80 nmi.
General color as in A. titahensis, n. sp., and
related to that species but with distinctive male
genital and female seventh sternal characters.
Head similar to utahensis except not as
pointed apicallv
Male.
ately long caudoventral process that usualK
crosses its counteqoart in caudal \iew, not
closely appres.sed to margin of pygofer (P'ig.
Im); plate long, triangulate, with row of mar-
ginal micr().setae and submarginal niacrosetae
(Fig. In); .style in dorsal \iew long, narrow,
apophysis sliort, ol)li(juel\' tnmcate apicalK
(Fig. lo); aedeagus in lateral view short, \entral
margin gradualK' curved, apical third tubular
broad basalK in ventral view, tapered toward
apex, gonopore subapical on ventral margin
(Figs. lj>-lr).
Fe.\L\LE. — Seventh sternum with truncate
caudal margin except for sliort, median process
(Fig. Is).
HOLOTYPE (miile). — CaLIFORNL\: Riverside
Co., Indio, 12.1.1988, Atriplex sp., G. N.
-Pv gofer in lateral view with nioder-
162
Great Basin Naturalist
[Volume 52
Figs, la-ll. Aplfinusk'Ua utahensis, n. sp.: la, head pronotum, and scutellnm, dorsal view; lb, male pygofer, lateral view;
Ic, right plate, ventral view; Id, right style, dorsal view; le, connective, dorsal view; If, aedeagus, dorsal view; Ig, same,
lateral view; Ih, same (enlarged), .showing triangulate process, lateral view; li, same (showing variation), lateral \ie\v; Ik,
same (enlarged), showing apex of aedeagus, ventral view; 11, female seventh sternum, ventral view.
Figs. Im-ls. Aplanmiella calif omiensis, n. sp.: Im, male pygofer, lateral view; In, right plate, ventral view; lo, right
style, dorsal view; Ip, aedeagus, lateral view; Iq, same, ventral view; Ir, same (enlarged), showing apex of aedeagus, ventral
view; Is, female seventh sternum, ventral view.
1992]
New Genus, Aplanusieija
163
Figs. 2a-2f, 2in. Aplanus pauperciilus (Ball): 2a, nude pvgofer, lateral \iew; 21). right plate, ventral view; 2c, aedeagus,
dorsal view; 2d, same, lateral view; 2e, right sUle, dorsal \iew; 2f, connectixe, dorsal \iew; 2m, female seventh sternum'
ventral view.
Figs. 2g-21, 2n. Aplanus albidns (B;ill): 2g, male pvgofer, lateral view; 2h. right plate, ventral view; 2i, aedeagus. dorsal
\iew; 2j, same, lateral view; 2k, right style, dorsal view; 21, connective, dorsal view; 2n, female seventh sternum, ventral view.
164
Great Basin Naturalist
[Volume 52
Oldfield (CAS). Parat)pes, 2 males, 6 females,
same data as holotype (OSU); 5 males, 16
females. Imperial Co., Brawlev, 23.VIII.1983,
Atriplex .sp., J. Williams (OSU, BYU).
Remarks. — This species can be separated
from utahensis by the shorter caudoventral
pvgofer spine, by the smoothly curved ventral
margin of the aedeagus, by the lack of a basal
process on the aedeagal shaft, by the broader
base of the aedeagus in ventral view, and by the
truncate caudal margin and shorter median pro-
cess of the female seventh sternum.
This species is knowTi from southern Califor-
nia on Atriplex (species unknown) at elevations
below sea level. Collection dates suggest that
the species overwinters in the adult stage and
ma\' have as many as three generations per year.
Aplanus Oman
Aplanus Onuui, 1949:138. Tvpe species, Eutctfix
paiipcrctihis Ball.
Only two species are known in the genus,
both assigned by Oman (1949). Crowder (1952)
treated the group with a key to species, rede-
scriptions, and illustrations of the genital char-
acters. The range of^ Aplanus is much broader in
western United States than the presently known
range o{ Aplonusiella.
Characters are given for Aplanus
pauperculns (Figs. 2a-2f, 2m) and Aplanus
aUndus (Ball) (Figs. 2g-2l, 2n) to show generic
relationships between them and species of
AplanusieUa. In Aplanus the pygofer lacks the
caudal spine, and the aedeagus is alxnit twice as
long with distinctive terminal processes. The
female seventh sternum lacks the obvious
median caudal process that is present in
AplanusieUa. Ball (1900) reported that shad-
scale, Atriplex eon feiii folia (Torn & Frem.)
Wats., was the host of Aplanus alhidus. The
specific host of A. pauperculns is yet unknown.
Phytogeographv o{ Atriplex
Four-winged saltbush (Atriplex eanescens) is
endemic to western North America. Its range
extends from southern Canada to northern
Mexico. Shadscale (Atriplex conferiifolia) is also
endemic, but its range is more restrictive within
western United States (Stutz and Sanderson
1979, 1983, Sanderson et al. 1990). Both species
produce hybrids between themselves and other
species of Atriplex. However, autopk)idy is the
most common genetic mechanism in both spe-
cies, which have produced a number of races
throughout their range. These races and other
ecotypes have been identified and mapped by
these workers.
The biogeographical relationships between
Aplanus and AplanusieUa species and their host
species are poorly knouai. Although hosts have
been identified for two leaflioppers (Aplanus
albiclus and AplantisieUa utahensis) of the four
known species, nothing is known about the
others nor has preference, if any, of these leaf-
hoppers for races or ecotypes been studied in
Atriplex. The role of Atriplex in the evolutionary
development and speciation of the group is like-
wise unknown.
Deposition of type specimens
The holotvpe specimens of AplanusieUa
utahensis and AplanusieUa califomiensis are
deposited in the California Academv of Sci-
ences, San Francisco (CAS); parat\pes are in
Oregon State University, Corvallis (OSU), Utah
State University, Logan (USU), and Monte L.
Bean Museum, Brigham Young University,
Provo, Utah (BYU).
Acknowledgments
We thank Paul W. Oman, Oregon State Uni-
versit\', Corvallis, for loan of material of Aplanus
and C. Rilev Nelson, Universitv of Texas, Austin,
for his assistance in collecting material in Utah.
We iilso appreciate helpful comments by H.
Derrick Blocker, Kansas State University, Man-
hattan, and Paul H. Frevtag, University of Ken-
tuck"\', Lexington, who reviewed the paper. This
studv was supported in part by endowanent
funds from the Monte L. Bean Life Science
Museum, Brigham Young University, Provo,
Utah, for which we are grateful.
Literature Cited
Ball. E. D. 1900. Some new Jassidae from the Southwest.
C:anadi;in Entomologist 32: 200-20.5.
(^ROWDKR, II. \V. 1952. A re\ision of some phlepsiuslike
genera of the tribe Delttxephdini (Homoptera,
Cicadellidae) in America north of Mexico. Kansas Uni-
versitv Science Bulletin 35: .309^541.
H.\ws, B.'a., G. E. Boh.'KRT. C. R. Nelson, ;uid D, L.
Nelson 1990. Insects and shnib die-off in western
states: 1986-1989 survey results. Pages 127-151 in
E. D. McArthur. E. M. Romne\. S. D. Smith, and P T.
Tueller, eds., Proceedings — Svmposium on Cheatgrass
Invasion, Shrub Die-off and Other A.spects of Shrub
Biolog)- and Miuiagenient. Las Vegas, Nevada, .5-7
1992] New GE>i{JS, Aflawsieija 165
April iyS9. I'SDA F'ori'st Scnicc. (Jeiieral IVcliiiical Arid land plant tx^sonrces. Pr(K-efdini;s of tlu- Intcrna-
Report INT-276. 351 pp. tional Arid Land Conference on Plant Resonrccs.
Oman, P. W. 1949. The Nearctie leiiflioppers. A generic International Center for Arid and Seini-iirid Land
classification and check-list. Memoirs of the Entonio- Studies, Texas Tech Uni\ersitv. Liihhock. 621 pp.
logical Society of Washington. . 1983. EvoKitionarv studies oi'Atriplcx: chromo-
Sanukkson, S. C'., H. C. Silt/, and E. 1). McArtihh some races of A, confertifolid (Shad.scale), American
1990. Geographical differentiaHon in Atriplcx am- Journal of Botany 70:' 1536-1547.
fertifolia. American Journal of Botany 77: 490-498.
Stutz, H. C, iind S. C. Sanderson. 1979. The role of
pol\ploid\ in theeyolutionofAfn/jfevtYniesren.s-. Pages Received 11 Fehnian/ 1992
615-621 in J. R. Goodin and D. K. Northington, eds., Acccf)te(l 12 May 1992
Creiit Basin Natunilist 52(2), pp. 166-173
SUMMER HABITAT USE BY COLUMBIAN SHARP-TAILED GROUSE
IN WESTERN IDAHO
Victoria Ann Saab and Jeffrey Sliavv Marks"
Abstra(ti" — We shidiecl smnnier habitat use by Columbian Shaip-tailed Grouse {Tytnpuuuchus pJuisiancllus co-
hunhiaims) in western Idaho during 198.3-S5. Vegetative and topographic measurements were recorded at 716 locations
of 15 radio-tagged grouse and at 180 random sites within the major vegetation/cover types in the study area. The mean size
of summer home ranges was 1.87 ± 1.14 km". Of eight cover types identified in the study area, individual grouse used the
big sagebrush {Artcviisia tridentata) cover type more than or in proportion to availability, the low sagebmsh [A. arhusaila)
type in proportion to availability, and avoided the shrubby eriogonum (Eriooonum spp.) tyjie. Characteristics of the big
sagebmsh cover tyj^e that Sharp-tailed Cirouse preferred include moderate vegetative cover, high plant species diversity,
and high stnictural dixersitv. Grouse used areas of dense cover (i.e., mountiiin shrub and riparian cover tyjjes) primarily for
escape cover. Compared with random sites, grouse selected areas with (1) greater horizontal ;uid vertical cover, (2) greater
canopv coverage of forbs tyj^ically decreased by livestock grazing, (3) greater density and canopy coverage of arrowleaf
balsamroot (Balsamorhiza sagittata), and (4) greater canopy coverage of bluebunch wheatgrass (Agroptjron spicatum) in
the big sagebmsh cover type in 1984 ;uid the low sagebrush cover type in 1985. The importance of the native perenniiils
arrowleaf biilsamroot and bluebunch wheatgrass became apparent chiring a drought year when many exotic annuals dried
up and provided no cover. Overall, grou.se selected vegetative communities that were least modified bv lixestock grazing.
Key words: Tympanuchus phasi;uiellus columbianus, C(>luml>itin SJiaiy-taiJcd Gnnise. Idaho, stnnuwr habitat charac-
teristics, nmnaocment .
Coliinibian Shaip-tmled Grouse {Tynipa-
nuchiis phasianellus columbianus) have
declined in both numbers and distribution since
European settlement, currently occupying
<10% of their former range (Miller and Graul
1980). Degradation of native habitat by live-
stock grazmg and agriculture are thought to be
major factors in this decline (Yocom 1952,
Aldrich 1963, Zeigler 1979). Overgrazing
reduced bunchgrasses and perennial forbs that
are important components of nesting and
brood-rearing habitat (Yocom 1952, Jewett et al.
1953, Klott and Lindzey 1990). Conversion of
range to cropland destroyed nesting and brood-
rearing habitat and deciduous shrubs that are
critical for winter food and escape cover (Zeigler
1979, Giesen 1987, Marks and Marks 1988). As
a result, Columbian Sharp-tiiiled Grouse were
designated as a candidate species for listing as
federally threatened/endangered (Federal Reg-
ister 1989).
Quantitative information on home range size
and habitat preferences of (Columbian Shaip-
tailed Grouse throughout their range is lacking.
especially data based on radio-tagged individu-
als during the summer reproductive period (see
Klott and Lindzey 1990). We studied Colum-
bian sharjDtails in areas with eight vegeta-
tion/cover types. The primary objective of our
study was to provide information on summer
habitat preferences by Columbian Sharp-ttiiled
Grouse.
Study Area
The 2000-ha study area is 23 km north of
Weiser in Washington Countv; Idaho. Elevation
ranges from 970 to 1188 m. Annual precipita-
tion averages 39 cm. The springs and summers
of 1983 and 1984 were relatively cool and wet,
whereas those of 1985 were unusually hot and
dry. Sharp-tiiiled Grouse had not been himted
in the study area since 1974.
Vegetation is characteristic of a sliRibsteppe
communitv (Marks and Marks 1987a). The
greatest proportion of the studv area (40%) was
occupied bv the big sagebmsh (Artemesia
trident (it a) cover t\pe; low sagebmsh (A.
arhusndo) and shmbby eriogonum {Eriogonum
Biology Department, Montana .Slate Univt-nsitv. Bo/cmaii, Montana 59717. Present address: USDA Forest Ser^nce, Intermonntain Ke
M)Ttle Street, Boise, Idaho 8.3702.
Di\nsion of Biological Sciences, University of Montana, Missonla. Montana .'59812,
•arcli Station. .316 E,
166
19921
Sharp-tailed Grouse Summer Habitat
16-
sphacrocephaluiii and E. thijiiioidca) Upes
occupied 21 and 20%, respectively. The remain-
ing 19% of the stud\- area was occupied bv' five
other cover tvpes (see below).
The big sagebnish cover t\pe was dominated
bv big sagebrush, with lesser amounts of
bitterbrush {Purshia tridentata) and low sage-
brush. The greatest canopv coxerage of blue-
bunch wheatgrass {Agropyron spicatuin) was
found in this cover type; arrowleaf btilsamroot
(BaJsanwrhiza sogittata)wds the dominant forb.
Bulbous bluegrass was the most common her-
baceous plant in the understor\' of the low sage-
brush co\er t\pe with lesser amoimts of
willoweed {Epilobium paniadatum), blue-
bunch wheatgrass, and Sandberg's bluegrass
{Poa sandhergii). The herbaceous layer of the
shnibbv eiiogonum cover t\pe was relativelv
sparse and dominated bv Sandberg's bluegrass.
The mountain shrub cover type occurred in
dense patches on hillsides; common species
were bittercherr\' {Pnimis emarginatus),
common chokecherr}' [P. virginiana), snow-
brush ceanothus {Ceanothus vehitimis), and
Saskatoon serviceberry {Amelanchier alnifolia) .
The shrub layer of the bitterbrush (Purshia
tridentata) cover type was almost exclusively
bitterbnish, while the herbaceous layer was sim-
ilar to that found in the big sagebrush t)pe.
Riparian vegetation was dominated by Douglas
hawthorn (Crataegus douglasii), with lesser
amounts of wallow (Salix spp.) and Woods rose
(Rosa woodsii). Bulbous bluegrass (Poa
btdhosa), an exotic grass, was widespread
throughout the study area. Plant nomenclature
follows Hitchcock and Cronquist (1976).
Two vegetation t\pes were almost exclu-
sively comprised of nonnative vegetation. A
small portion of the study area contained agri-
culture, composed of dryland wheat and barley,
and monocultures of intermediate wheatgrass
(Agropyron interniedinni) seedings.
The study area was grazed by livestock since
at least 1900. Before about 1940, large bands of
sheep were driven through the area. Since then,
the major land use in the studv area has been
cattle grazing. No livestock grazing occurred
during this study.
Methods
Trapping and Monitoring
Grouse were captured on dancing grounds
using funnel traps, mist nets, and drop nets. Sex
was determined 1)\' examination of crown feath-
ers (Henderson et al. 1967) and age by exami-
nation of outer primaries (Ammann 1944).
ThirtA-eight grouse (28 males and 10 females)
of 46 captured were fitted with solar-powered
radio transmitters attached to Herculite pon-
chos (Marks and Marks 1987b). Radios weighed
between 13.5 and 14.5 g. Fifteen (13 males and
2 females) grouse provided data for home range
and microhabitat analyses. The other 23 grou.se
with radios were relocated for two months or
less as a result of mortality (Marks and Marks
1987b) or dispersal from the stud\' area. Data
from these birds were used in the microhabitat
analyses but not in the calculation of home
range size. Sample sizes were not large enough
to compare habitat use or home range size
between male and female grouse.
Radio-tagged grouse were monitored from
May to September 1983—85. Each time a grouse
was located, it was flushed (hereafter these loca-
tions are called flush sites). Flush sites sened as
focal points for habitat sampling and for calcu-
lation of home ranges. Grouse were located
throughout the day and locations were stratified
into four time intervals: sunrise to 0800, 0801 to
1100, 1101 to 1700, and 1701 to sunset. On
average, each radio-tagged bird was flushed
four days a week, once in each of the four time
intervals.
Habitat Sampling
The stud\' area boundaiv was determined by
grouse movements during 1983. C^cn-er tvpes
were digitized and areas calculated for each t) pe
using GEOSCAN (Software Designs 1984), a
geographic information program. Flush sites
were plotted and home range sizes (Mohr 1947)
were calculated using the compute^- program
TELDAY (Lonner and Burklialter 1986). '
Use vs. availabilih' of cover t\pes (i.e.,
macrohabitat) was assessed by (1) using the
proportion of cover t\pes within each bird's
home range, and (2) using the proportion of
cover tyj:)es within a 1.2-km radius of the danc-
ing ground at which each bird was captured.
The 1.2-km radius around each of three dancing
grounds (upper, middle, and lower) encom-
passed 90% of all grouse locations. Flush sites
within 50 m of a dancing ground during the
spring and autumn display periods were omitted
from macrohabitat analyses.
We measured \egetation at each flush site
(i.e., microhabitat) to estimate plant .species
168
Great Basin Naturalist
[Volume 52
composition, frequency, percent canopy cover-
age, and bare ground using a 20 X 50-cm frame
(Daubenniire 1959). Five frames were read at
each flush site: one at the approximate center
and one in each of the four compass directions
at randomly chosen distances of 2, 4, 6, or 8 m
from the center location. Vertical structure of
the vegetation was evaluated by a coxer board
that was a 16.5 x 49.5-cm rectangle. The cover
board was placed at the center of the flush site
and read twice from 5 m away in each of the four
compass directions while the observer was
prone and standing, respectively. A total reading
of 150 squares was possible from each compass
direction. In total, five canopy coverage and four
cover board measures were taken at each site.
Other variables recorded at flush sites included
(1) cover type, (2) distance to water, (3) percent-
age of slope, (4) distance to nearest riparian or
mountain shrub cover type, and (5) cover type
where flushed grouse landed (landing site).
We recorded vegetative and topographic
measurements at randomly located sites to
assess microhabitat avmlability in the cover
t\pes used most bv grouse. Habitat characteris-
tics were sampled with similar methods as
described at flush sites. A total of 180 random
sites were sampled during the study, 30 each
month during Mav through Julv in 1984 and
1985. The number of random sites located in
each cover type was based on the percentage of
area occupied by that cover type in the study
area. Canopy coverage and cover board read-
ings were recorded at the origin and at points
every 10 paces along a straight line until 20
readings were completed. Slope and distance to
the nearest mountain shiub or riparian cover
type were recorded onlv at the first, tenth, and
twentieth frames of each random site.
Data AnaK'sis
Data were anab'zed with the Statistical Anal-
ysis System (SAS Institute, Inc. 1982). Use-
availabilit)' analyses of cover types were
conducted with chi-s(juare goodness of fit tests
(Neu et al. 1974) and Bonferroni z-tests (B\ers
et al. 1984). Data were analyzed separately for
each year and pooled when differences were not
significant. For analyses of canopy coverage,
each plant species was placed into one of 10
categories: ( 1) big sagebrush, (2) low sagebrush,
(3) bitterbrush, (4) other shrubs, (5) arrowleaf
balsamroot, (6) other composites, (7) non-
composite forbs, (8) bluebunch wheatgrass, (9)
bulbous bluegrass, and (10) other grasses. Non-
parametric statistics (Mann-Whitney U- and
Kruskal-Wallis tests) were used to anaK'ze
canopy coverage and vertical stmcture because
these data were not nonnally distributed (Con-
over 1980). Vegetative measurements at flush
sites from May through July were combined by
cover tvjje and month for comparisons with data
collected at random sites for the same period.
All multiple comparisons were computed with
Tukey tests (Zar 1974). The Shannon-Wiener
index was used to calculate plant species diver-
sity (Hill 1973). Proportions entered into the
diversit)' formula were derived from the total
number of plant species occurrences within the
frames used to estimate canop\' coverage. The
significance level for all tests was P < .05, and all
tests of means were two-tiiiled. Means are fol-
lowed by ± one standard deviation.
Results
Home Ranges and Macrohabitat Selection
The mean size of summer home ranges was
1 .87 ± 1 . 14 km- (N = 15, range = 36-68 locations
per grouse). Based on habitats within home
ranges, three trends emerged from the use-
availabilit)' analysis of coxer t\pes: (1) grouse
used the big sagebrush cover t}pe more than or
in proportion to availabilitv; (2) the low sage-
brush cover t\pe was used in proportion to
a\ailabilif\', and (3) the shrubby eriogonum and
intermediate wheatgrass coxer txpes xvere
avoided (Table 1). These trends xvere similar
xvhether use-aviulabilit\' xvas assessed xxithin
estimated home ranges or xxithin a fixed radius
aroimd the upper and lower dancing grounds
(Table 1). In addition, a single grouse from the
middle dancing ground used the big sagebiTish
coxer t\pe more than that expected bx' chance
xxithin its home range and the fixed radius.
Grouse were seldom found in the denser cover
txpes, i.e., riparian and mountain shrub habitats.
Hoxx'exer, thex used these coxer txpes as escape
coxer in 77% of the cases xxhere the landing site
of a flushed radioed bird xx'as obsened (N =
338).
Microhabitat Selection
Mean distance to xvater did not differ signif-
icantly bet\x'een flush (.V = 297.6 ± 183.3 m) and
random (.v = 295.9 ± 211.7 m) sites (F < .40),
and no evidence xvas found that Shaip-tiiiled
Grouse sought free xvater. The range of slopes
1992]
Sharp-tailed Grousk Summer Habitat
169
Table 1. Sunnner hahitat usf-a\ailal)ilih analysis showing tli(- iniinl)er oi raclio-taggml (Columbian Sliarp-tailed Crouse
using the major cover types more than ( + ), less than ( — ), or in projxjrtion to (NS)'' that expected by chance ', 1983-85.
Cover types
Home range
NS
1.2-km fixed radius
NS
Upper dancing ground
Big sagebnish
Low sagebrush
Shnibbv eriogonuni
Mountain shrub
Number of grouse
Lower dancing ground
Big sagebrush
Low sagebrush
Intermediate wheatgrass
Number ot grouse
Total number ot grouse
2
3
1
4
5
1
4
N = 5
7
2
3
6
2
'
N = 9
N = 14
5
5
5
1
4
8
I
1
8
6
3
â– 'Not sigiiitkant,
'T < .05.
used by grouse was 0-47%. Grouse used three
classes^ of slope intervals (0-9%, 10-29%,
>30%) in proportion to their availabilit\; with
>95% of the use occurring on slopes <30%
(Marks and Marks 1987a).
Grouse did not show a strong preference for
sites that were close to nioinitiiin shmb or ripar-
ian \egetation except in 1985, the drought year.
The mean distance to mountain shrub and ripar-
ian habitats measured at flush sites {x = 151.5 ±
156.5 m) was farther than that measured at
random sites (.v = 120 ± 99.7 m) in 1983 and
1984 (Mann-Whitney L'-test P < .04) but signif-
icantly closer (flush sites, x - 84.4 ± 90.9 m) in
1985 (F< .0001).
Vertical cover measured at random sites dif-
fered significantly among cover types (Kruskal-
Wallis P < .001). Mean cover board readings
indicated that the bitterbrush cover tyjie pro-
vided the greatest cover; big sagebrush, inter-
mediate wheatgrass, and low sagebrush tyjoes
])ro\ided intermediate cover; and eriogonmn
sites had verv little cover (Fig. 1). A drought
during 1985 resulted in significantly less vertical
cover in 1985 than in 1984 (Mann-Whitney li-
test P < .01 ). However, the rank order of cover
availabilitv was the same among all cover t\pes
except intermediate wheatgrass, which de-
creased substantiiillv in 1985.
Eight\'-three percent of the flush sites for
which microhabitat measurements were taken
occurred in big and low sagebrush cover t\pes.
\'egetative data on microsite use vs. availability-
were evaluated onlv for big and low sagebrush
cover types because sample sizes were too small
for the other types.
Vertical cover measured at flush sites dif-
fered among years within big and low sagebrush
cover types (Kmskal-Wallis P < .05). As noted
at random sites, there was significantly less
cover in 1985 than in 1984. A comparison of
grouse flush sites with random sites revealed
that grouse selected denser cover than that mea-
sured at random sites (Fig. 1).
The cover types used most by grouse, big and
low sagebnish, had a higher diversitv of shrub,
forb, and grass species than the otlier cover
tvpes (Fig. 2). The big sagebrush cover type had
the highest diversit)' of shnibs and grasses, and
the low sagebrush cover tvpe had the highest
diversity of forbs. Overall, the big sagebnish
cover tyjDe had the highest stnictural heteroge-
neity (measured as the coefficient of variation of
canopv coverage and cover board readings).
During 198.3-85, canopv coverage oi shnibs
at grouse flush sites averaged about 9% in both
big and low sagebnish cover types. Forb cover-
age averaged about 30%, and grasses ranged
from 28% to 32% canopv coverage in low sage-
brush and big sagebrush cover tvpes, respec-
tively. Overall, canopy coverage at flush sites
was significantlv greater than at random sites
due largelv to greater total forb coverage at flush
sites (Table 2). (^onverselv, percentage of bare
ground was less at flush sites than random sites
in all cases (Table 2). Sites chosen by grouse in
1984 and 1985 had significantlv- higher
arrowleaf balsamroot cover than did random
sites. There was significantly higher canopy
170
Great Basin Naturalist
[Volume 52
O RANDOM
D FLUSH
t
6
^
ARAR ERIO
COVER TYPES
Fig. L Mean (± SD) cover board readings at random
sites and Sharp-tailed Grouse flush sites in the major cover
types (big sagebnish [ARTR], low sagebnish [ARAR],
shrubby eriogonum [ERIO], intermediate wheatgrass
[AGIN], bitterbrush [PUTR], 1984-85 (° = F < .001).
Vertical tixis represents the number of boxes visible on the
cover board (see Methods).
26-
24-
22
X
3. 20
>-
(/) '
EC ,
liJ
I 1^
LLI
CL
cn 6
(66)
(67)
(45)^5
.{24)
COVER TYPES
Fig. 2. Plant species diversit) (e" ) at random sites for
shrubs, forbs, and grasses in the major cover tyjies (big
sagebnish [ARTR], low sagebnish [ARAR], shrubby
eriogonum [ERIO], intermediate wheatgrass [AGIN]),
1984-85. The total number of plant species sampled in each
cover type is in parentheses.
coverage of bhiebunch wheatgrass at grouse
flush sites than at random sites in the big sage-
brush cover ty|3e in 1984 and in the low sage-
brush cover ty|)e in 1985.
Canopy coverage at grouse flush sites in the
big sagebrush type differed among years in five
of six vegetative categories (Fig. 3). Bare ground
increased while bulbous bluegrass, other forbs,
and other composites decreased during the
drought of 1985 as compared to 1983 and 1984.
However, bluebunch wheatgrass increased in
1985, while the cover of arrowleaf balsamroot
was not significantly different among years.
Bluebunch wheatgrass and arrowleaf
balsamroot are native perennials that are con-
sidered decreaser species (Bhiisdell and
Pechanec 1949, Evans and Tisdale 1972); i.e.,
they tvpicallv decrease or are eliminated under
heaxy livestock grazing (Dyksterhuis 1949).
Canopy coverage of decreaser forbs was signif-
icantly greater at flush sites than at random sites
in the big and low sagebrush cover types ( IVIarks
and Marks 1987a).
Discussion
Summer home ranges for this subspecies in
Colorado (Giesen 1987) and for other subspe-
cies (Artman 1970, Christenson 1970, Ramhar-
ter 1976, Gratson 1983) were sniiiller than we
observed in this study. Differences in home
range size were probably a reflection of habitat
condition; larger home ranges were obsened in
western Idaho, where decreaser forbs were lim-
ited and historic livestock grazing apparently
had a greater influence on the vegetation.
From spring to fall, >90% of all grouse loca-
tions were within 1.2 km of a dancing ground.
Similarly, locations of Sharp-tailed Grouse in
other studies were within 1.0 and 2.5 km of
dancing grounds (Pepper 1972, Oedekoven
1985, Giesen 1987, Nielsen and Yde 1982).
These results suggest that maintiiining habitats
within 2.5 km of dancing grounds will provide
summer habitat recjuirements for Shaip-tiiiled
Grouse.
Compared with other cover tyjoes, big sage-
brush sites had a high diversit)' of shrubs, forbs,
and grasses; the highest structural diversity; and
the greatest canopx' coverage of perennial bimch-
grasses. The sharptails" overall preference for
the big sagebrush cover type indicated that they
likely selected for habitat diversitv relative to
surrounding areas.
1992]
Sh.\hp-tailed Grouse Summer Habitat
171
T.^BLK 2. Mean can<)p\- coverage (%) of\'cgetativecateg()rie.siiil)igsagebnish (ARTR) and low sage! )ni.sli (ARAR) cover
tvpes at Columbian Sliarp-tailed Grouse flush sites vs. random sites.
Year
1984
1985
ARTR
ARAR
ARTR
ARAR
Vegetative
Flush
Random
Flush
Random
Flush
Random
Flush
Random
category'
(io7r
(42)
(21)
(24)
(107)
(42)
(21)
(24)
Big sagebrush
3.43
4.03''
0.02
0.07
4.97
6.. -32
0.22
0.33''
Low sagebnish
0.21
0.49''
5.45
7.. 84
0.55
0.79''
7.03
7.88
Bitterlmish
1.52
1.02
0.86
0.17
2.76
l.w''
1.15
0.88
Otlier shmbs
1.73
0.89
0.14
0.59''
2.21
2.69''
1.36
0.40
Total shrubs
6.89
6.43
6.47
8.67
10.49
11.84
9.76
9.49
Arrowleaf balsamroot
13.60
6.55''
12.21
3.91''
13.06
7.40''
11.91
5.28
Other composites
7.05
3.78'-
5.14
2.95''
2.90
3.33
3.02
3.19
Otlier forbs
12.76
15.3l''
12.83
14.24
9.70
7.87
14.97
7.22
Total forbs
33.40
25.64''
30.18
21.10''
25.66
I8.6O''
29.90
15.69
Bluebnnch wheatgrass
2.93
2.56''
1.02
0.85
5.18
2.91
4.72
0.46''
Bulbous bluegrass
35.87
24.59''
36.83
23.09
15.97
16.52
13.20
22. a3''
Other grasses
3.76
4.32
2.52
3.32
3.01
2.02
3.33
3.29
Tcjtal grasses
42.56
31.47
40.37
27.26
24.16
21.45
21.2,5
26.08''
Bare ground
23.93
35.93''
28.05
42. 30''
40.23
48.62''
39.31
48.94''
â– 'Sample .size (number ot trausecLs conducted in each tspe).
'Indicates significant difference (P < .05) in mean canopy coverage behveen flush and random sites withi
ver tjpes.
Slinibb\' eriogonum sites, which were
strongK' axoided by grouse, contained a low
di\ersit\ of forbs, and even in the absence of
grazing proNided Rttle cover. Exchiding dancing
grounds, Shaip-tailed Grouse studied else-
where have exliibited similar selection against
areas of sparse cover (Pepper 1972, Ziegler
1979, Klott and Lindzey 1990). The intermedi-
ate wheatgrass cover type also was avoided by
grouse. Grouse were particularly absent from
intermediate wheatgrass during years with rela-
ti\elv low numbers of grasshoppers.
Mountain shrub, riparian, and bitterbnish
habitats were used primariK as escape cover
during spring and summer. Beginning in late
siunmer, moimtaiu shrub and riparian plant spe-
cies produced fniits that became an important
part of the grouse diet (Marks and Marks
19S7a). Proximity to this shrubby vegetation
ma\- not have been critical during earK" to mid-
summer when the cover types preferred by
grouse were providing adequate food and cover.
Grouse were found closer to mountain shrub
and riparian habitat than expected l)y chance
only in the drought year (1985), when xertical
cover decreased significantK' in all cover t\pes
that were measured.
Shaq^tails apparentK' selected areas least
modified by lixestock grazing. Grouse locations
were characterized by greater herbaceous cover
and less bare ground than random sites. Studies
of plant communities with and without gnizing
indicate that areas with relati\eK- little bare
ground are least modified b\- li\estock (.see
40 -
a
e
/
S.BAGR
/
30 ^
y
^^
y?\
\
20 -
\
\
>POBU
10 -
'^""'g'"--.,^
.
~-^OTFO
-AGSP
~"OTCO
^'
_^^Z-=-'
"^
-
k
k
1
1
1
1984
YEARS
Fig. 3. Comparison of canopy coverage at Sharp-tailed
(Jrouse flush sites in tlie big sagel)nish co\er t%pe in western
IiliJio. 198.3-85. On each line different letters indicate that
corresponding means are significantK' different at P = .05.
(BAGR = bare ground. POBU = bulbous bluegrass, BASA
= arrowleaf b;i]samroot, OTFO = other forbs, AGSP =
bluebiuich wheatgrass, OTCO = otlier composite forbs.)
172
Great Basin Naturalist
[Volume 52
Holechek et al. 1989). When eoinpared with
random sites, grouse locations had significantly
higher proportions of forb species that decrease
from overgrazing (Dyksterhuis 1949). In partic-
ular, grouse preferred microhabitats with
greater abundances of arrowleaf btilsamroot
and bluebunch wheatgrass, two plant species
that ty}3icallv decline with overuse by livestock
grazing (Blaisdell and Pechanec 1949, Evans
andTisdale 1972, Muegglerand Stewart 1980).
These native perennials are major components
of later serai stages (Hironaka et al. 1983).
The presence of arrowleaf balsamroot and
bluebunch wheatgrass as cover plants during a
drought year is especially noteworthy. These
plants are particularlv drought resistant (Tisdale
and Hironaka 1981, \Vasser 1982). Bulbous blue-
grass, the most abundant and widespread grass
in the study area, is an introduced perennial
with root systems that die each year; it is virtu-
ally nonexistent during years of low moisture
(Monsen and Stevens, in preparation). Indeed
bulbous bluegrass contributed lower cover
values in 1985 than in 1983 and 1984 (years with
average moistiu-e) (Table 2). In contrast, cover
of bluebunch wheatgrass was similar among
those years. In the absence of nati\e perennials,
grouse would not have had as much cover dining
drought years. The loss of these important cover
plants may have contributed to the disappear-
ance of Columbian Shaq:)-tailed Grouse from
large portions of their historic range.
CONCLUSlOxNS AND MANAGEMENT
Implications
Ciiven the widespread decline of the Colum-
bian Sharp-tailed Grouse and the fragmented
nature of extant populations, consenation of all
potential sources of genetic variation should be
a critical concern to managers. Maintenance of
shniljsteppe coiumunities in advanced serai
stages is especially important for con.servation of
summer habitat in the Intermountain region.
Habitat features that characterize occupied
habitats in western Idaho are flat to rolling
rangeland in relatively good condition with a
diversity of native shmbs, forbs, and grasses.
Native perenniiils arrowleaf balsamroot and
bluebunch wheatgrass are critical for cover
during a drought year. Also important is riparian
vegetation and numerous patches of mountiun
shrubs for escape cover and late summer food.
These habitat characteristics suggest that
Columbian Sharp-tailed Grouse are an indica-
tor of good range condition in the mesic
shnibsteppe of the Intermountain region.
Federal land management agencies are
directed to conserve candidate species and their
habitats and to avoid actions that mav cause the
species to become listed as federally threat-
ened/endangered. Conservation efforts for
Columbian Shaqo-tailed Grouse, a candidate
species, must include protection and enhance-
ment of habitats that are occupied by the sub-
species throughout their range, especially
disjunct populations in jeopardy of extirpation.
The success of attempts to improve their cur-
rent status is dependent on reducing distur-
bances that may damage the natural diversity of
shrubsteppe habitat (e.g., overgrazing by live-
stock and agricultural development).
Protecting habitats within 2.5 km of dancing
grounds is critical for mmntainence of summer
habitat. Suitable habitats for reestablishment
within their historic range need to be identified.
However, reestabHshment efforts for this native
species should not take precedence over pres-
ervation and restoration of habitats that cur-
rently support sharptails (cf. Griffith et al. 1989).
Acknowledgments
We thank A. Sands, L. Nelson, S. Mattise, R.
Eng, T Lonner, R. Autenrieth, R. Nelson, and
J. Connelly for their contributions to the study,
and the G. Tarter and T Nelson families for
granting access to their lands. We are also grate-
ful for the field assistance provided by B. Czech,
J. Berr)'hill, S. Lisle, and R. Morales. J. Craw-
ford, C. Groves, K. Giesen, and T. Martin pro-
vided useful suggestions for manuscript
improvement. The research was funded bv the
U.S. Bureau of Land Management; additional
support was provided bv the Idalio Department
of Fish and Game and Montiuia State Universit)'.
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HoLECHEK, J. L., R. D. Pieper, andC. H. Herbel 1989.
Range niiinagement: principles and practices. Prentice
Hall, Englewood (.'liffs. New Jerse\-, .501 pp,
Jewett, S. G., W R Taylor. W. T' Shaw, and J. W.
Aldrk.tl 19.53. Birds of Washington state. Universit)'
of Wiishington Press, Seattle. 767 pp.
Klott J. H., and F. G. Lindzey 1990. Brood habitats of
sympatric Sage Grouse and (Columbian Shaq)-tailed
Grouse in Wyoming. Journal of Wildlife Miuiagement
.54: 84-88.
LoNNKH T. N., and D. E. Burkhalter 1986. Users
manual for the computer program TELDAY. Montana
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Marks J. S., and V. S. Marks 1987a. Habitat selection by
Columbian Shaip-tailed Grouse in west-central Idaho.
Bureau of Laiul Management Re^xjrt, Boise, Idaho.
115pp._
. 1987b. Influence of radio collars on snr\i\al of
Sharp-tailed Grouse, journal of Wildlife M;uiageinent
51: 468^71.
. 1988. Winter habitat use In (Columbian Shaip-
tailed Carouse in western Idaho |ournal of Wildlife
Management 52: 74.3-746.
Miller. G. C, and W. D. Graul 1980. Status of Shaq>
tailed Grouse in North America. Pages 18-28 in P. A.
Vohs and F. L. Knopf, eds.. Proceedings of the Prairie
Grou.se S\'mposinin, Okhilioma State University Still-
water
Moll R C. (). 1947. Table of e(]ui\ alent populations of North
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USDA Forest Senice General Technical Report INT-
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Nielsen, L. S., and C. A. Yde 1982. The effects of rest-
rotation griizing on the distribution of Shaq)-tailed
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eds., Proceedings of the VV'ildlife-Livestock Relation-
ships Symposium, Universitv of Idaho, Moscow.
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population distribution and habitat use in south central
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Pepper, G. W. 1972. The ecolog\ of Sharp-tailed Grouse
during spring and summer in the aspen parkkmds of
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DepiU^tment of Natural Resources, Regina.
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versity of Idaho, Forestn, Wildlife, and Riuige
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cies useful in re\egetating disturbed lands in the West.
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oecetes pliasianvUus coliiinhianus) in the state of Wash-
ington. American Midland Natiualist 48: 18.5-192.
Z\R. J. H. 1974. Biostatistical ;xnalvsis. Prentice-HiJl, Inc..
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Received 25 September 1991
Accepted 16 March 1992
Great Basin NatiinJist 52(2), pp. 174-178
CHARACTERISTICS OF SITES OCCUPIED BY SUBSPECIES OF ARTEMISIA
TRIDENTATA IN THE PICEANCE BASIN, COLORADO
Thomas R. Cottrell and Chiules D. Bonham"
Kit/ words: Artemisia tridentata. Colorado, sagehnisli, clirointifoij^raplu/, factor analysis, sod.
Artemisia tridentata, big sagebiiish, is the
dominant plant species in the Piceance Basin of
western Colorado and displays great morpho-
logical variabilitv between sites. The existence
of at least three subspecies is widely accepted
(McArthur et al. 1981, 1988). These are A.
tridentata spp. tridentata Beetle, A. tridentata
spp. ivi/oming^ensi.s Beetle and Young, and A.
tridentata spp. vaseyana Beetle.
Despite extensive research in the Piceance
Basin (Redente and Cook 1986), we have found
only one study referring to intraspecific taxa of
sagebrush (Ward et al. 1985). This work
referred to subspecies tridentata but did not
indicate where this taxon was found. Because
the taxa are known to respond differentiiilly to
soil and climate factors (Hironoka 1978, Sturges
1978) their existence in the basin should be
recognized. The present study was designed to
identify the subspecies o{ Artetnisia tridentata
present in the Piceance Basin and to describe
soil characteristics of sites occupied by sub-
species.
Study Site
The Piceance Basin comprises about 3()()()
km" in Garfield and Rio Blanco counties of
northwest Colorado (Fig. 1). The cUmate of the
Piceance Basin is semiarid and shows extreme
variability in monthlv precipitation (Wymore
1974). Consecutive months often receive little
precipitation. The mean annual precipitation
for eight weather stations in the region for the
period 1951-70 was 35.3 cm, with a 95% confi-
dence intei-val of ±18.7 cm. About one-half of
the total precipitation falls as snow. The average
annual temperature ranges from 7 C at 1800 m
to - 1 C at 2700 m.
The strong influence of topography on tem-
peratiu'e and precipitation results in a complex
of habitats in the basin (Tiedeman and Terwilli-
ger 1978). Generally, soil development is corre-
lated to elevation. At higher elevations, except
ridge tops, soils are dark brown, shallow
mollisols. At mid-elevations, aridisols are
common on deep loess. The lowest elevations
are characterized by entisols developed on
heavy clays and deep, sandy alluvial soils.
Methods
Six sites dominated by sagebrush were
selected for this study (Table 1). These sites
spanned the environmental extremes of sage-
brush habitat in the Piceance Basin. Two sites
were selected from each of three broad topo-
graphic regions. High mounttiin sites were
about 2000 ni; upland terraces and valley
bottom sites were below 2000 m.
Sagebrush subspecies were identified bv the
combined information of three techniques and
verified by A. H. Winward, regional ecologist
for Range and Watershed Management, USFS
Intermountain Region, Ogden, Utah. The first
technique involved field identification using
moqihological characteristics based on keys by
A. H. Winward and Tisdale (1977). Leaf sam-
ples were taken for the other two procedin-es.
Two-dimensional chromatography, as described
by Hanks et al. (1973), was done on persistent
overwintering leaves from three plants at each
site except site 5, where the moqihologiciil vari-
abilit)' of the sagebnish plants was greater than
at the other sites. At this site five plants were
^Department of Biology, Colorado State University, Fort C^ollins. Colorado 80523.
Range Science Department, (Colorado State University, Fort Collins, Colorado 80.52,3.
174
1992]
Notes
175
m.
''830
WYOMING
PICEANCE
BASIN
C OLORADO
NEW MEXICO
2130-
24 30-
Meeker
P'il
e^^
40°N
/
PICEANCE BASIN
COLORADO
SCALE IN MILES
SCALE IN KILOMETERS
CONTOUR INTERVAL 300m
Fig. 1. The shulv iirea of the distribuHcMi ,A Anemisia trklcutata subspecies i„ northuest Colorado.
tested by chromatography. Results were com- matelv IS other plants in the stucK- sites Leaves
pared with representative chromatograms. The were crushed In hand and placed in glass con-
hird procedure was a leaf extract in uater. This tainers for four hours. These were viewed under
Uitter method was performed on all plants tested long-wave ultraviolet light and compared to
by chromatography and on a total of approxi- descriptions by Stevens and McArthur (1974)
176
Great Basin Naturalist
[Volume 52
Tabi.K 1. Location, elevation, ;uid sagebnisli subspecies of stutly sites. VAS — ssp. id.sci/ana: TRI — ssp. tridoitata: WTO
= ssp. wuoimngensis. Selected soil characteristics are listed for ()-L5 cm and 16^30 cm soil samples for each site.
L()cation
Site
Elev.
ssp.
Depth
in cm
% sand
% silt
pH
C:aCO^
est.
llji^h mountain
I
2.365
VAS
0-15
54
26
6.9
low
1.5-30
52
26
6.8
low
2
2585
VAS
0-15
40
33
6.8
low
1.5-.30
36
33
6.5
low
Viilley bottom
3
1987
TRI
0-15
74
13
8.2
med
1.5-.30
67
20
8.1
med
4
2057
TRI
0-15
56
30
8.1
med
1.5-.30
52
32
8.2
med
Upland terrace
5
1920
WYO
0-15
46
27
8.2
med
15-30
51
27
8.3
med
6
2070
WTO
0-15
32
45
7.7
med
1.^.30
28
44
8.2
med
In each site, soil samples were collected at
two random locations from two depths, 0-15 cm
and 16-30 cm. These were analyzed for pH,
organic matter, electrical conductivity, esti-
mated CaCOa, sand, silt, clay, K, Mn, Zn, Cu, P,
and Fe. These data were used in a factor analysis
as described by Affifi and Clark (1984). the
factor scores for each site and depth were then
graphed. This graph was used to inteqDret the
axes that usually represent some environmental
characteristic associated with plant species.
Results
Sites 1 and 2 were high mountain sites. Sage-
brush plants averaged less than 50 cm in height.
(Common associated plants were Liipimis .sp.,
ChnjsotJiamnus viscidiflonis, Erio<s<^iuiin iimlwl-
latum, Stipa letfentuinii, and Si/nifjlioricaiyo.s
oreophilus. Soils were all deeper than 40 cm and
dark in color. Near these sites Populus
tremuloides stands were c-onunon in favorable
microenvironments.
Sites 3 and 4 were in a valley bottom in the
Yellow Creek area. The sagebrush in these sites
commonly reached heights greater than 2 m.
Associated vegetation included the moss Tor-
tula niralis, CJwnopodium prdtcricohh and
Lepidhun laiifolhnn. Soils were light in color,
and depths greater than 40 cm were common.
Sites 5 and 6 were at similar elevations to 3
and 4, but away from streams. Sagebrush plants
averaged 40 cm in height. Site 5 soils were
approximately 10 cm in depth. Bromiis fec-
tonim, Gutierrezia sarodirac, Ah/ssuni ahfssoidcs.
and Ort/zopsis Ju/moioides were the common
plant species. Site 6 soils averaged 20 cm deep.
Common understoiy species were Koeleria
crisfafa, A^ropijwn smitliii, and PJilox hoodii.
Site 6 was surrounded by forests ofPintis ednlis
and Jiinipenis osteospemw.
Locations of A. fridentata subspecies in the
study area relate generalK' to elexation. The
lowest elevations supported both ssp.
wijominfj^ensis and ssp. fridentata. Factor anal-
ysis results indicate that soil texture and chem-
istiy differences existed between the sites (Fig.
2). Subspecies tiidentata was found in sandier
soils and wijomin^ensis in siltier soils. The tex-
ture differences were generally related to topo-
graphic position. Subspecies tiidentata was
most common in xallev bottoms, and ssp.
wyoniin^ensis was tvpicallv dominant awaN'
from streams, at the lowest elexations to approx-
imately 2100 m. Sites above 2100 m supported
ssp. vasei/ana. Soil te.xtures in vaseijana site 1
were similar to those in tridentata sites, while
vaseijana site 2 textures more closely resembled
those of the wt/oinin^iensis sites. Soil pH was
lower in the vasetjana sites than sites with the
other siibspecies.
Moiphologiciil identification of ssp. vasei/ana
and tridentata generalK- agreed with the results
from two-dimensional chromatograph\ and the
leaf extracts. Subspecies wyomingensis chro-
matograms were not consistently separable
from those of subspecies tridentata. None of
the wi/oniin(^en.si.s chromatograms closely
matched published chromatograms. Leaf ex-
tracts from ssp. ivt/innin^ien.si.s showed almost no
1992]
Notes
177
LJ
<
Ld
u
O
<
Q
<
1.5
0.5 -
- -0.5
ct:
O
I—
u
<
-1 -
-1.5
-1.5
3s
tridentata
3d
4d
^=d
5s
vaseyana
6s
2d
wyomlngensis
2s
6d
1 1
1 1 1
-1 -0.5 0.5 1 1.5
FACTOR 2: ELEV. INCREASES, pH DECREASES
Fig. 2. Results of factor analysis on all soil data. Y axis corresponds to increasing sand:silt ratio; X axis corresponds to
decreasing pH. Stand numbers are as in Table 1; s indicates sample from 0-15 cm; d indicates sample from 15-30 cm.
Names show approximate region of ordination occupied bv each subspecies.
fluorescence and were not separable from ssp.
tridentata. Moqohologicallv, howev'er, this sub-
species was separable from vasci/ana and
tridentata bv the ke\s of Winward and Tisdale
(1977).
Discussion
Three subspecies of A. tridentata were iden-
tified in the Piceance Basin b\ reference to
moiphologv; chromatography, and leaf extracts.
The subspecies identified were wyomingensis,
tridentata, and vasei/ana. Two-dimensional
cliromatograpliN and leaf extracts \ielded pre-
Hininary evidence to suggest that ssp.
wyominiiensis in the Piceance Basin is chemi-
cally different from those previously identified.
The distributions of A. tridentata subspecies
are generaliv related to soil moisture, tempera-
ture, depth, and parent material (Hironaka
1978). The overall tendency seems to be for ssp.
tridentata to occupy deep, somewhat sandv
soils. Although subspecies wijcnningensis occurs
in an overlapping zone \\A{\\ tridentata, it is more
common in shallow, silt\' soils where moisture
stress is greater. Subspecies lasei/ana occurs in
cool, moist sites, usutilly above 2100 m, but
lower elevations have been documented (Good-
rich et al. 1985).
Each subspecies was found at eJexations and
in soil textures similar to those reportcnl in the
literature. Soil texture, expressed as a ratio of
sand to silt, explains the first factor in the factor
analvsis and distinguishes i)etween sites of ssp.
tridentata and wyomingensis (Fig. 2). That is,
the vertical axis in Figure 2 corresponds to this
ratio. It appears that the relative proportion of
sand and silt determines whether ssji. tridentata
or ssp. wt/oniingensis will be dominant. Barker
and McKell ( 1983) reported similar results and
suggest that the characteristics of soils associ-
ated with these subspecies are different. Fine-
textured soils ha\e been implicated in increased
water stress in ssp. wyoniingensis sites (Shumar
and Anderson 1986). This might indicate a
differential adaptation to water stress and.
178
Great Basin Naturalist
[Volume 52
consequently, different life history' strategies in
the subspecies (Bonham et al. 1991).
Soils at sites with ssp. vaseyana are distin-
guished from the other sites by factor 2 of the
factor analysis. This axis represents both an ele-
vational and soil pH gradient (Table 1, Fig. 2).
Sites with ssp. vaseyana were at a higher eleva-
tion, and soils were lower in pH and CaCO^
values. The textures at these sites did not differ
substantiiillv from the other sites.
No previous study in the area has identified
these taxa or characterized their habitats. The
great differences in habitat preference among
these subspecies suggest this is a major over-
sight.
Acknowledgments
The research was supported jointly by the
U.S. Department of Energy, Contract No. DE-
AS02-76EV04018 to Colorado State Universit)-
and the Agricultural E.xjoeriment Station, Colo-
rado State University, Project 660(4242).
Literature Cited
Affifi. a. a., and Clark. V. 1984. Computer-aided multi-
variate analy.sis. Lifetime Learning Publications, Bel-
mont, California. 458 pp.
Bakker, J. R., and C, M. McKell 1983. Habitat differ-
ences between basin and Wyoming big sagebrusb in
contiguous populations. Journal of Range Manage-
ment 36: 450-454.
BoNiiAM. C. D., T. R. CoTTHELL. and J. E. Mitchell.
1991. Inferences for life history strategies of Artemisia
tridentata subspecies. Journal of Vegetation Science 2:
339-;344.
GooDRiCM, S., E. D. McArthur, and A. H, Winward
1985. A new combination and a new variety oi Artemi-
sia tridentata. Great Basin Naturalist 45: 99-104.
Hanks, D. L., E. D. McArthuk. R. Steven.s, and A. R
PiAJMMER. 1973. Chromatographic characteri.stics and
phylogeiietic relationships of Artemisia section
Tridentatae. USDA Forest Service Research Paper
INT-141.24pp.
HiRONAKA. M. 1978. Basic svnecological relationships of
the Columbia River sagebrush type. Pages 27-.32 in
The sagebnish ecosystem: a symposium. Utah State
University, LogiUi.
McARTHUR, E. D., C. L. Pope, and D. C. Freeman 1981.
Chromosomal studies of subgenus Tridentatae ofArte-
misia: evidence for autopoKploidv. American Journal
of Botany 68: 589-605.
McArthur, E. D., B. L. Welch, and S. C. Sanderson.
1988. Naturd and artificial hybridization between big
sagebnish {Artemisia tridentata) subspecies. Journal of
Heredit)' 79: 268-276.
Redente, E. F., andC. W. Cook, eds 1986. Structural and
functional changes in early successional stages of a
semiarid ecosystem. Progress Report to U.S. Depju't-
ment of Energy. Depiutment of Range Science, Colo-
rado State University, Fort Collins. 67 pp.
Shumar, M. L., and J. E. Anderson 1986. Gradient anal-
ysis of vegetation dominated by two subspecies of big
sagebnish. Journal of Range Miuiagement 39: 156-
159.
Stevens, R., iuid E. D. McArthur 1974. A simple field
technique for identification of some sagebnish taxa.
Journal of Range Management 27: .325-.326.
Sturc:es. D. L. 1978. Hydrologic relations of sagebrush
lands. Pages 86-100 in The sagebnish ecosystem: a
symposium. Utah State Universitv', Logan. 251 pp.
TiEDEMAN. J. A., and C. Terwilliger. Jr 1978. A phy-
toedaphic classification of the Piceance Basin. Colo-
rado State University, Department of Range Science,
Science Series 31. 265 pp.
Ward. R. T, W L. Slauson, and C. W. Weldon 1985.
Response of shnib ecotvpes to mining waste material
in soil profiles and competitive interactions of woodv'
species under experimental and naturiJ conditions.
Pages 87-94 in E. ¥. Redente, C. W. Cook, J. M. Stark,
and C. L. Simmons, eds., Semiarid ecosvstem de\eIop-
ment as a function of resource processing and alloca-
tion. Progress Report to U.S. Department of Energ\-.
Colorado State University, Department of Range Sci-
ence, Fort Collins.
Winward, A. H., and E. W. Tisdale 1977. Titxonomv of
the Artemisia tridentata complex in Idiilio. Uni\ersit\'
of Idiilio, Forest, Wildlife and Range Experiment Sta-
tion Bulletin No. 19. 15 pp.
WvMORE, I. F. 1974. Estimated average annual water bal-
ance for Piceance and Yellow Creek watersheds. En\a-
ronmentiJ Resources Center, Colorado State
University, Fort Collins. Technical Report Series No.
2. 60 pp. â–
Received 20 November 1991
Accepted 4 May 1992
Great Basin Natunilist 52(2). pp. 179-184
USE OF LAKES AND RESERVOIRS BY MIGRATING SHOREBIRDS IN IDAHO
)aiiiil \l.Ta\I()r and ( iharlfs il.lVo.st
Kit/ uonh: shorrhird.s. habitat ii\c. iiiiiilfldts. uaU'r (Irandoini. irri<^atiiiii rcsi'troirs. ini^ratin^ liinl.s
ShorebircLs migrating long cli.stances are \iil-
lu^able because their wetland .stopover sites are
limited in number and susceptible to distur-
bance or destniction b\' humans (Senner and
Howe 1984, Myers et al. 1987). It is therefore
critical to know which wetland areas migrating
shorebirds use, and the factors making the.se
sites attracti\e to shorebirds.
\\ e conducted shorebird censuses at numer-
ous wetland sites in Idaho with these objectives:
(1) to identify t>pes of lakes and reservoirs that
are important for migrating shorebirds, (2) to
identih' habitat characteristics at these wetlands
used b\ shorebirds, (3) to determine the inilu-
ence of mudflat exposure and water le\el
cliang(^s on shorc^bird use.
Study Ahk.\s and Methods
A total of 19 lakes and resenoirs were cen-
sused at least once in 1989 (Table 1). Nine
high-ele\'ation lakes were visited in the Saw-
tooth Wilderness in earlv September 1976, and
three high -elevation lakes in the Seafoam area
of the Frank Church River of No Return Wil-
derness in earlv .August 1990. Additional obser-
vations from Lake Lowell were made in 1986,
1987, and 1990. All shorebirds were censused
within 100 m of the shoreline in and out of the
w ater at all sites; thus, evei-v 500 m of transect
censused was equal to 0.1 km". We estimated
birds per 500 m of shoreline for our densitv
estimates. The Springfield area of American
Falls Reservoir had over 15 km of mudflat
exposed by drawdown during the study period
and also included numerous seep areas awav
from the main shoreline; because of this, it was
not possible to make density- estimates from this
site. Four of the lakes and reservoirs visited in
1989 had mudflat areas that were censused at
least six times at roughly weekly inteivals from
mid-Julv to earlv September, the time of peak
shorebird abundance in Idaho (Tavlor et al.
1992). We used ANOVA and Newman-Keuls
tests (Zar 1974) to compare differences in
shorebird numbers at these four sites. Birds
were censused bv walking from 10 to 1 00 m back
from the shoreline and using binoculars and a
25X spotting scope. Care was taken not to dis-
turb birds. If birds moved, their numbers were
kept track of, or the entire coimt was restartc^d
to avoid counting birds more than once.
Results
The natural lakes at high elevations we cen-
sused in 1989 (Table 2) had onfy 0-2 Spotted
Sandpipers (see Table 3 for cill scientific names).
Only a single Spotted Sandpiper was found at
nine high-elevation lakes visited in the Sawtooth
Wilderness in September 1976. No shorebirds
were found at three high-elevation lakes in the
Seafoam area in early August 1990.
At the Lowell, Walcott, American Falls, and
Carey areas we found significant differences in
the densities of total shorebirds (ANO\'A, F2(3)
26 = 88.76, P < .001). Lake Lowell had signifi-
cantly the most shorebirds, American Falls had
significantlv more than Carey Lake, but Carey
Lake's higher mean was not significantlv more
than Lake Walcott s (Newnian-Keuls, q = 29.89
to 7.47, for significant differences P < .05 or
greater; (j = 2.04, P = :2 for Carey Lake-Lake
Walcott). These differences in shorebird num-
bers reflect the amount of mudflat available at
the different sites; the larger the mudflats, the
greater the number of shorebirds.
The pattern of more shorebirds being
attracted to larger mudflats is further supported
In shorebird numbers at different Lowell sites
Department of Biolopcal Sciences. Idaho State University, Pocatello. Idalio 83209
179
180
Great Basin Naturalist
[Volume 52
TaBI.f: 1. Characteristics of hkilio lakes ;uicl reservoirs sur\'eyecl for shorehirds in 19S9.
Transect
Elevation
length
Name
County
(m)
(m)
Habitat
Reservoirs and lakes with mudflats
American Falls
Power
1321
900
500 m mudflat
Lowell
C;inyon
757
4600
1200 m mudflat
Walcott
Minidoka
1279
1500
20 m mudflat
Carey
Blaine
1453
2200
200 m mudflat
Little Camas
Elmore
1502
800
120 m mudflat
Dry
Ciuiyon
818
15(X)
50 m mudflat/700 m grass
Mackay
Custer
1849
1400
200 m mudflat
Palisades
Bonneville
1708
1600
1000 m mudflat
Reservoirs and lakes
without mudflats
Cascade
ViJley
1472
2600
1-2 m sandy or muddy shore
Wilson
Jerome
1224
1800
dirt or grass shore
Boulder
Valley
2127
900
2 m mud or rocky shore
Bnineaii
Owyhee
763
23(:K)
1 m mud or sandy shore
High-elevation lakes
Alice
Blaine
2622
1000
herb or rocky shore
Toxaway
Custer
2539
9(K)
herb or rockv shore
Edith
Custer
2611
6(X)
herb or rocky shore
East
Valley
2373
1100
herb or rocky shore
West
Valley
2361
900
herb or rocky shore
North
Valley
2367
7(X)
herb or rocky shore
Payette
Valley
1522
700
herb or rocky shore
responding to changes in mudflat conditions in
1989 (Fig. 1). In July Public Access No. 1 had
verv' few shorebirds, and nearly all of its
mudflats were submerged by water (Fig. lb).
The New York Canal site was submerged at this
time and had no birds (Fig. la). When the large
mudflats of the New York Canal site became
exposed in August, thousands of shorebirds
appeared there (Fig. la). Numbers of shore-
birds at some of the other sites declined (Fig.
lb), which may have been due in part to birds
shifting to the New York Canal site. The reflood-
ing of Lowell in late September 1989 com-
pletely eliminated shorebirds from census areas
by 27 September (Fig. 1), although American
Falls Reservoir had over 500 shorebirds at this
time. On 27 September 1990, uith Lake Lowell
very low due to dam reconstniction, there were
extensive mudflats at the New York Canal site,
and 926 individuals of 10 species of shorebirds
were present. In earlv Julv 1986 there were
hundreds of shorebirds on the exposed mudflats
at Public Access No. 1, but in early July 1987,
with high water flooding into riparian vegetation
at this site, there were no shorebirds.
The reservoirs we counted once or a few
times in 1989 usually supported the pattern of
total shorebird numbers declining with decreas-
ing mudflat size, but there were some excep-
tions (Table 2). Wilson, Boulder, and Cascade
reservoirs all had zero or onh' a few meters of
exposed shoreline, and thev had only 1 or 2
shorebirds. Mackay Reservoir had onlv 2 shore-
birds on 3 July when no mudflats were exposed,
but 351 two weeks later when there was 200 m
of mudflat. The Drv^ and Little Camas reservoirs
supported hundreds of shorebirds (Table 2),
and these sites had mudflats of 50-120 m. How-
ever, Bruneau had onlv 1-2 m of mud or sandy
beach, and it had 79 individual shorebirds. An
even stronger anomalv was Palisades, a reservoir
which had exposed mudflats of about 1000 m
and water drawdown continually exposing new
areas, but practicallv no birds (Table 2).
Black-bellied Plovers, Lesser Golden-Plo-
vers, Sanderlings, Pectoral Sandpipers, and Stilt
Sandpipers were found only on mudflats with
>500 m of exposed mud (Table 3). Ten other
shorebirds species were most abundant at sites
with >500 m of exposed mudflat. Eight shore-
bird species had similar-sized peaks at sites with
>500 m or between 20 and 200 m of ex'posed
mudflat. The onlv species with a maximmn peak
on mudflats between 20 and 200 m was the
uncommon Long-billed Curlew. No individual
shorebird species had maximum numbers at
1992]
Notes
181
Table 2. Total numher and, in parentheses, densit\ per 0.5 km of transect ol sliorehirds counted at lakes and reservoirs
in Idaho in 1989.
Count area
Mean
SD
Range
Springfield
American Falls
9
9
Lowell
8
Wiilcott
9
Carey
6
Little Camas
4
Da'
4
Macka\-
2
Palisades
4
Cascade
Boulder
2
1
Wilson
Bnmeau
Alice
Payette
Edith
Toxawav
West
East
North
2296
578.1
1698-3252
209
87.2
92-337
(105)
(43.6)
(46-168.5)
3061
1839.6
752^5739
(323)
(230.6)
(7^717)
54
40.6
17-153
(18)
(13.4)
(6-.50)
254
111.9
80-393
(58)
(25.4)
(1^89)
294
161.5
117^46
(184)
(101.0)
(7;3-279)
132
28
93-158
(44)
(9.3)
(31-53)
177
2^51
(62)
(1-125)
18
23.6
0-70
(6)
(8.3)
(0-18)
1
(0.6)
79
(17)
1
(1)
1
(0.7)
sites with <5 m of mudflats or rocWlierh shore-
lines.
Discussion
Tlie \irtiial absence of shorebirds from the
19 hi<rh-ele\ation lakes we \isited in 1976, 1989,
and 1990 is similar to the findings of the only
previous study of a high-elevation lake in Idaho.
Visits annually to Fish Lake, Idaho Co., from
1923 to 1929 found only a few Solitary Sandpip-
ers and Spotted Sandpipers, and one or two
indi\iduals of four other species (Hand 1932).
Burleigh (1972) reported no large numbers of
shorebirds at any high-elevation lakes in Idaho.
Further investigation may reveal some high-ele-
vation lakes to be important for migrating shore-
birds, but the lack of mudflats at most of these
lakes probabK' limits their use by most shorebird
species.
The concentration of most shorebirds at
large mudflats is consistent with our previous
findings at American Falls Resenoir, where we
foimd verv few shorebirds on sand\', clay, or
boulder beaches or bedrock (Taylor et al.,
unpublished data). Shorebirds also concen-
trated on mudflats at inland studies done in
Nevada (Hainline 1974), Missouri (Rundle and
Fredrickson 1981), and Saskatchewan (Colwell
and Oring 1988), although the latter study also
had some shorebird species associated with dif-
ferent habitats. Our stud) also shows that small
and moderate-sized mudflats of both natunil
lakes and reservoirs mav attract some shore-
birds, especially those that often feed in water.
Shorebird species that primarily or com-
pletel) feed b)- probing in or gleaning off land
surfaces or very shallow water almost always had
higher peaks on the larger mudflats, or were
foiuid there exclusi\ely. An exception was
Baird's Sandpiper, which had a similar peak
between large and moderate mudflats. Five of
the shorebird species with equal-sized peaks on
large and moderate mudflats, the Black-necked
182 Great Basin Naturalist [Volume 52
Table 3. Slioiehird sjx-cies (ouiid at 19 tfsenoiis and hikes in Idalio in 1989.
Species Abundance'' and habitat use '
Black-bellied Plover Uncommon on large mudflats.
Pluiialis scjiiatawla
Lesser Colden-Plover Rare on large mudflats.
Fluvialis dominica
Semipalmated Plover Uncommon on large mudflats; rare on moderate nmdflats luid muddy shores.
Cluirad lilts scmipaliiuitiis
Killdeer Common on large and moderate mudflats; uncommon on muddv shores; occasional on
Charadriiis vocifcnis rocky/lierb shoreline.
Black-necked Stilt Uncommon on kri'ge and moderate nmdflats; rare on muddv shores.
Himantopus mexkamis
American Avocet Abundant on Luge mudflats; uncommon on moderate mudflats and muddv shores.
Real m irostra anie rica 1 1 a
Greater Yellowlegs Uncommon on laige and moderate mudflats; occasion^ on muddy shores.
Tringa melanoleuca
Lesser Yellowlegs Conuiion on large nmdflats; uncommon on moderate mudflats; occasional on muddy
Tringa flavipes shorelines.
Solitary Siuidpiper Occasional to rare on all shore t\pes.
Tringa solitaria
Willet Unconunon on Uirge mudflats; occasional on moderate nmdflats; rare on muddv shorelines.
Catoptwphonis scmipahnatus
Spotted Sandpiper Uncommon on large and moderate mudflats, muddy shorelines; occasional on rock)'/herb
Actitis maadaria shorelines.
Long-billed Curlew Occasional on moderate mudflats; rare on large mudflats.
Numeii iu.s aincricanus
Marbled Godwit Common on large mudflats; occasional on moderate mudflats.
Lhnosa fedoa
Sanderling Uncommon on huge mudflats.
Calidris alba
Semipalmated Sandpiper Uncommon on large mudflats; occasional on moderate nuidflats.
Calidris pusilla
Western Sandpiper Abundant on huge mudflats; common on moderate mudflats; uncommon on mudd\' shores.
Calidris iiuiuri
LeiLst Sandpiper Unconunon on hu^ge muiiflats; occasional on moderate mudflats.
Calidris minittilla
Baird's Sandpiper Conunon on large and moderate mudflats; occasional on nnidd\' shores.
Calidris hairdii
Pectoral Sandpiper Uncommon on large mudflats.
Calidris mclaiiotus
Stilt Sandpiper Riu-e on large mudflats.
Calidris hiiiuiiitopus
Short-billed Dowitcher Occasional on large and moderate mudflats.
Limnodromus grisciis
Long-billed Dowitcher Conunon on large uuuinats; uncommon on moderate nuidflats and nuidd\ shores.
Limnodromus scolopaccns
Common Snipe Uncoiumon on huge nuidflats; occasional on moderate nmdflats and uurIcIv shores.
Gallinago gallinago
Wilson's Phiilarope Conunon on large and moderate mudflats; uncommon on uuuldv shores.
Phalaropiis tricolor
Red-necked Pluilarope Conunon on large and moderate nuidflats; occasional on imidd) shores.
Phakiropxis lobatiis
â– 'A species was considered abundant if it had a single peak count over 10(K) at a siiecific site, cuinnion u-itli a peak o\er 100, iiiiconinion wUh a peak <)\er 10, ocxiisional
with a peak under 10, and rare if only one or two individuals were found
Uirge mudflats include American Falls, Springfield, Palisades, and Lowell, and all had water drawdnwii exposing mndllats of distances >.5tK) in. Moderate mudflats
include Carey, Little Camas, Di\' (in part), Mackay, and V\'alcott. and had water drawdouii exposing 20 2IKI ni ol mudflat. Muddy shores included Dry (in part),
Bnmeau. Oiscade, Boulder, and Payette (in part), and tliese included small muddv shorelines or iiiudflais of .5 m widtli or less and also sandy or dirt shorelines.
Rocky/herb shorelines included Alice. Dry(in])art), Kitst. F.ditli. North. Pavette l in parti, Toxawav, and Wil.son.
19921
Notes
183
shorebirds
1500 -
Public #1
Public #2 /\
1000 -
Public #3 / \
B
Q / 1
500 -
\
j\l \
•
^
:^^^*=*-fcJl
Q' ' T '^ T
■-r ■1 ■1 ' 1 ■1 ' T '^P •
isades Reservoir in tliis stiicK, indicates there are
additional factors inflnencing shorebird use.
This could include food abundance (Harrison
1982, Myers et al. 1987), which is important at
American Falls Resenoir (Mihuc 1991), tradi-
tional use (Myers et al. 1987), and in the case of
Palisades Reservoir possible difficulty of shore-
birds locating it because it is enclosed by high
mountains in all directions (personal observa-
tion). Steep-sided resenoirs, such as C. J.
Strike, Hells Canyon (personal observation),
and Lower Granite Creek (Monda and Reichel
1989) on the Snake River, and stretches of the
Columbia River subject to water level fluctua-
tions (Books 1985), supported few shorebirds
even with water drawdown in summer and tail.
The absence of shorebirds at Lake Lowell
and Mackay Reservoir from sites when high
water covered mudflats shows the importance
of water drawdown exposing these areas during
migration. At American Falls Resenoir we have
previously found shorebird numbers to be cor-
related with rate of drawdown (Tavlor et al.,
unpublished data). Water levels at reservoirs in
this region are usually determined by irrigation,
power generation, recreational activities such as
boating, or waterfowl management. It is impor-
tant that controllers of water levels at reservoirs
and lakes (1) become aware of the potential or
real use of shorebirds in their area and (2)
manage water levels for shorebirds whenever
feasible.
Fig. 1. W'eeklv counts of the total number of shorebirds
at four sites at Lake Lowell, Canvon Co., Idaho, in 1989. (A)
New York Canal Mouth site, with both total number of
shorebirds and the amount of mudflat exposed. (B) Open
circle is Public Access No. 1 site; open triangle is Public
Access No. 2 site; vertical line is Public Access No. 3.
Stilt, Greater Yellowlegs, Short-billed Dow-
itcher, Wilsons Phalarope, and Red-necked
Phalarope, along with the Long-billed Curlew,
all often feed in water. The two remaining spe-
cies with similar-sized peaks between large and
moderate mudflats, the Killdeer and Spotted
Sandpiper, were the most widespread.
This study indicates that most reservoirs and
lakes in Idaho and the Intermountain West can
provide habitat for shorebirds in fall migration
if they have moderate to large mudflats that can
be exposed by water drawdown during summer
and fall. The absence of shorebirds at some
reservoirs with large mudflats, in particular Pal-
ACKNOWLEDCMENTS
We would like to thank E. Stone, S. Bailey,
S. Hart, and two anonvmous reviewers for their
comments on earlier drafts of this paper This
studv was funded in part by the Department of
Biological Sciences, Idalio State Universit}'.
Literature Cited
BooK.s, G. G. 1985. Avian interactions with mid-Columbia
River level fluctuations. Northwest Science 59: 304-
312.
Bl KLF.ICH, T. D. 1972. Birdsof Idaho. The Caxton Printers,
Ltd., CiJdwell, Idaho. 467 pp.
CoLWELL, M. A., and L. W. Ohinc; 1988. Habitat use by
breeding and migrating shorebirds in soudicentral Sas-
katchewan. Wilson Bulletin 100: .554-566.
H.MNLINE. J. L. 1974. The distribution, migration, and
breeding of shorebirds in western Nevada. Unpub-
lished master's thesis, Universit)' of Nevada, Reno. 84 pp.
lt\ND, R. L. 19.32. Notes on the occurrence of water and
shorebirds in the Lochsa region of Idaho. Condor 34:
23-25.
184
Great Basin Naturalist
[Volume 52
H\RRisoN, B. A. 1982. Untviiig the enigma of" the Red
Knot. Living Bird Quarterly 1: 4-7.
MlHUC, J. 1991. An experimental study of tlie impact ot
shorebird predators on benthic invertebrates in Amer-
ican FiJls Reservoir, Idaho. Unpublished master's
thesis, Idaho State University, Pocatello, 61 pp.
MONDA, M. J., iuid J. D. Reichel 1989. Aviiui connnunitv
changes following Lower Granite Dam construction on
the Snake River, Washington. Northwest Science 63:
13-18.
Myers. J. R, R. I. G. Morrison. R Z. Antus. B. A. Har-
rington. T. E. LovEjOY, M. Sallaberry, S. E.
Senner. and A. Tarak. 1987. Conservation strategy
for migratorv shorebird species. American Scientist 75:
18-26^
RUNDLK VV. D., and L. H. Fredrickscjn 1981. Miuiaging
seasonally flooded impoundments for migrant rails and
shorebirds. Wildlife Societv Bulletin 9: 80-87,
Senner, S. E., imd M. A. Howe 1984. Con.servation of
Nearctic shorebirds. Behavior of Marine Animals 5:
379-421.
Taylor, D. M., C. H. Trost. and B. Ja.mison 1992. Abun-
dance iind phenology of migrating shorebirds in Idaho.
Western Birds 23:49-78.
Zar J. H. 1974. Biostatistical ;malvsis. Prentice-Hall, New
Jersey. 620 pp.
Received 15 September 1991
Accepted 1 May 1992
Great Basin Natuidist 52(2), pp. 185-188
DISPERSAL OF SQUARROSE KNAPWEED
{CENTAUREA VIRGATA SSP SQUARROSA)
CAPITULA BY SHEEP ON RANGELAND IN JUAB COUNT\', UTAH
Cind\ Talbott Roc-he ', Ben F. Roche, Jr. , and G. Allen Rasniu.s.sen
Key words: Centaurea \irgata ssp. s(|uarrosa, sciiiarrosc knapweed, weed dispersal. ranf^eUnid weeds, wool, sheep.
Among Centaurea species naturalized in
western North America, squarrose knapweed
(Centaurea virgata Lam. ssp. sqiiarrosa Gugl.)
has a unique dispersal mechanism. The seeds
(achenes) of other CentourtY/ species (C. diffusa
Lam., C. maculosa Lam., C. solstitialis L., C.
jacea L. x C nigra L.) disperse either as indi-
viduals with crop seed, vehicles, and gravel, or
as branches or entire plants moved by wind or
vehicles, or in hay. Squarrose knapweed involu-
cral bracts recurve or spread outward with a
short tenninal spine about 1-3 mm long. The
entire head (capitulum) is deciduous via an
abscisson laver at the base of the capitulum.
Thus, the capitula of squarrose knapweed func-
tion like burs clinging to passing animals as
l)urdock {Arctium minus (Hill) Bemh.), cockle-
bur {Xantliium strumarium L.), or buffalobur
{Solanum rostratnm Dunal). Soon after the dis-
covery of squarrose knapweed in California
(1950) and in Utah (1954), its occurrence was
linked to the practice of trailing rangeland sheep
from one area to another (Bellue 1954, Tingey
1960). On Utah rangeland squarrose knapweed
is more abundant along sheep trails and on
bedgrounds than in other areas (H. Gates and
T Roberts, personal communication). Wool is
idealK suited to catching and holding the
burlike capitula, but squarrose knapweed along
trails and in sheep bedgrounds may have been
carried by vehicles or other means and estab-
lished in soil disturbed b\' sheep. The objective
of this study was to determine if the distribution
of squarrose knapweed in Utah is due to seed
carried in the wool of rangeland sheep.
Methods and Materials
In mid-April 1990, sheep examined in this
study were trailed from winter range west of
Tintic Junction, Juab Comity, Utali, and sheared
before being mo\'ed to spring range. We
received permission from the owoiers to collect
wool samples during shearing of a band that had
wintered on rangeland known to have squarrose
knapweed. We had predicted that sheep would
pick up the "burs'" by lying on or brushing
against knapweed plants growing on their
bedgrounds. However, we saw no obvious knap-
weed capitula in bellv wool or on the sides of
sheep being sheared. One shearer pointed out
several ewes with a profusion of kiiapweed
capitula around their faces and on top of their
heads (Fig. 1). We then collected samples of
topknot wool (that shorn from the top of the
head) from 458 randomly selected white ewes
from a band of approximately 2500 ewes at the
Jericho shearing station in Juab Count); Utah.
Black ewes were not sampled. Samples from
individual ewes, averaging 10 g, were kept sep-
arate in small plastic bags. Squarrose knapweed
capitula were sorted bv hand from each sample,
and the number of achenes per capitulum was
recorded. Filled achenes (hard, plump, dark tan
or browni achenes) and light aclienes (.softer,
flatter, pale tan or whitish achenes) were
recorded separately. Presence or absence of
insect o^AhiUropJiora ajfinis Frauenfeld and U.
(juadrifasciata [Meigen]) in the knapweed
capitula was noted.
Achene viabilit)- was determined with germi-
nation trials nm for 10 da\s at 20 C, 12 hours
^Department of Natural Resource Sciences. Washington State University. Pullman, Washington 99164-6410.
"Present address: Department of Plant. Soil, and Entomological Sciences, University of Idaho, Moscow. Idaho 8.3843.
' Department of Range Science. Utah State University, Logan. Utah 84.322-5230.
185
186
Great Basin Naturalist
[Volume 52
Fig. 1. Numerous squarrose knapweed capitula were caught as burs in the topknot wool of sheep that Iiad wintered
where squarrose k-napweed occurred on rangekind in Juab Count\', Utah.
T.^BLE 1. Proportion ol capituki containing 0-6 aclienes
per capituhnn, comparing all capitula from iui intact plant
with sheep-gathered capitula removed from topknot wool,
in Juab Count)', Utah.
Achenes/capituluni Intact pkuit
'7c
Extracted from wool
14
75
1
12
IS
2
19
6
3
35
1
4
17
trace
5
3
6
trace
light alternating with 12 hours dark. Seeds were
placed in germination bo.xes on wetted blotter
paper. Filled and light achenes were tested sep-
arately. We germinat(Hl 30 filled achenes in four
replications in each of two trials. Two trials of
light achenes were run with 20 seeds in each of
two replications.
In August 19<S9, a scjuarrose knapw eed plant
with all ol its capitula was collected in a bag. We
dissected the capitula and recorded the number
of achenes per capitulum. These \alues were
compared to capitula and achenes found on
sheep.
Results
We determined that sheep on rangeland
infested with squarrose knapweed picked up
and carried its burhke capitula. Squarrose knap-
weed capitula were present in topknot wool
samples from 73% of the ewes. A total of 2469
knapweed capitula were reco\ered from the 458
ewes, an average of 5.5 capitula per 10 g wool.
Most capitula were on the wool surface,
although a few were embedded deeply and
appeared to have been there longer as the\" were
satmated with lanolin and spines had worn off
the in\()lucral bracts.
Sevent)'-five percent of the sheep-gathered
capitula were barren, compared with 14% of the
capitula produced on a whole plant (Table 1).
()iil\- 49% of the wool samples that contained
capitula had one or more achenes. Barren capit-
ula in this study were not the result of biocontrol
insects because we foinid no insect galls.
The nimiber of knapweed capitula on sheep
1992]
Notes
18'
< V'
•/>"
Fig. 2. Squarrose knapweed phuits along the sheep trails
west ot the Jericho she;iring station were grazed in mid-April
1990. A few capitula remain on the npper right side of the
plant.
heads would lead a casual obsener to couclude
that the sheep carty more achenes than we
found by dissecting the capitula. Among all ewes
sampled, only 36% carried achenes in the sam-
pled topknot wool. These seed-carriers aver-
aged 4.5 filled achenes per 10 g wool. Those
filled achenes averaged 69% germination. In
addition to the filled achenes, 5% of the light
achenes germinated. Light achenes composed
only 23% of the total numbc^r of achenes.
Discussion
Sheep carried squarrose knapweed capitula
but not as many achenes as the ninnber of
capitula woidd indicate if the proportion were
the same as that estimated in August. This find-
ing could indicate one of two conditions: ( 1 ) the
capitula were picked up in late winter or early
spring, when only the lighter capitula remained
on the plants, or (2) some achenes were lost
from capitula lodged in the wool during late
summer or fall. In late summer heavier capitula
are more easil\- dislodged from plants than are
the lighter capitula. Capitula do not open wideK'
at maturity-; instead, achenes sift out throush a
small opening created as the dried flowers fall
from the capitulum. The proportion of empt)'
capitula increases with time following maturity
as plants are shaken b\- wind, animals, or \ehicles.
Sheep acquired knapweed capitula in a
manner different from what we had predicted.
Although some capitula clung to sheep brushing
against plants or King upon them, the numerous
knapweed capitula in the wool aroiuid their
faces suggest that ewes searched out squarrose
knapweed as a food source. We observed that
scjuarrose knapweed plants along the sheep
trails had been grazed (Fig. 2). This relationship
was nuitually beneficial for knapweed and
sheep, providing propagule dispersal for the
knapweed and nourishment for the sheep.
Previousl) reported to be poor forage
(Tingev 1960), squarrose knapweed rosette
leaves may be an excellent source of protein in
late winter and early spring. Nutrient content of
spotted knapweed rosette leaves is comparable
to native forage plants with 9-18% crude pro-
tein (Kelsey and Mihalovich 1987). Similar
values have been obtained for diffuse knapweed
and yellow starthistle rosette leaves (Roche,
unpublished data). In the stud\' area, Septem-
ber 1989 through Mav 1990 was unusualK' dry
(Utah State University' Tintic research site
weather station, unpublished data), and the
normal growth of cheatgrass {Bromus tectorum
L.) was not present on the winter range.
Squarrose knapweed, a deep-rooted perennial
forb, was one of the few plants exhibiting new
growth at the time sheep would normalK forage
on cheatgrass.
Although we found that sheep carr>-
squarrc:>se knapweed seeds as they move across
rangeland, they are by no means the only dis-
persal mechanism for squarrose knapweed.
Other animals, both domestic and wild, may
carry knapweed seeds. In addition, these
rangelands are hea\il\- used b\- off-road \ehicle
recreationists. Mining traffic, railroad acti\it\'.
and militar\' maneuxers are important in certain
areas. Hunters, rockhounds, and other
recreationists also frequent the area.
Shearing limits the dispersion of scjuarrose
knapweed b\- sheep. It is unlikeK that knapweed
achenes remained on sheep after shearing.
These ewes had not yet lambed, and so all sheep
in this band left the knapweed-infested winter
range shorn of seeds. Seeds in the wool are
remox ed at the woolen mill, which has been one
of the fates of squarrose knapweed seed for
188
Great Basin Naturalist
[Volume 52
centuries, as evidenced by squarrose knapweed
found at Juvenal Gate, a woolen mill in France
where imported wool was washed for 200 years,
beginning in 1686 (TheUung 1912).
Acknowledgments
This study was made possible by the cooper-
ation of H. Gates and T. Roberts (Bureau of
Land Management), S. Dewey (Utah State Uni-
versity), and the ranchers who permitted us to
sample wool during their shearing operation. J.
Miller, Universit)' of Idaho, was consulted con-
cerning vegetable matter in wool. E. Evans and
D. Scamecchia reviewed the manuscript and
provided valuable suggestions.
The project was fimded in part by the
Renewable Resources Extension Act through
Washington State University Cooperative
Extension.
Literature Cited
Bkllue, M. K. 1952. Virgate stiir thistle, Ccntaurca virgata
vai". squarrosa (Willd.) Boiss. in California. California
Dep;xrtnient of Agriculture BuOetin 41: 61-63.
Kelsey, R. C, and R. D. Mhialovich. 1987. Nutrient
composition of spotted knapweed {Centatirea
maculosa). Journal of Range Management 40: 277-
281.
Roche, C. T, and B. F. Roche, Jr 1989. Introductory-
notes on squarrose knapweed (Centatirea virgata Lam.
ssp. squarrosa Gugl.). Northwest Science 63: 246-2.52.
Thellung, a. 1912. La flore adventice de MontpelLier.
Memoires de la Societe Naturelles et Mathematiques
de Cherbourg 38: 57-728.
TiNGEY, D. C. 1960. Control of squarrose knapweed. Utixli
State University Experiment Station Bulletin No. 432.
11 pp.
Received 11 March 1991
Accepted 31 March 1992
Great Basin Natunilist 52(2). 1992. pp. 189-193
VEGETATION ASSOCIATED WITH TWO ALIEN PLANT SPECIES IN A FESCUE
GRASSLAND IN GLACIER NATIONAL PARK, MONTANA
R()l)in W. Tyser
Ki'i/ icord.s: alini flora. CJacicr F<nk. FesiwcA grasslands.
The presence of alien flora in natunil area
grasslands of the Great Basin and surrounding
areas is of considerable concern, given the wide-
spread success of alien flora and associated
decline of nati\e species in this region (Young et
al. 1972, Mack 1986, 1989). Suiveys of indige-
nous bunchgrass communities in northern
Roclcs Mountain national parks have detected
the occurrence of several alien plant species
(Koterba and Ilabeck 1971, Stringer 1973,
Weaver and Woods 1985, 1986, Tvser and
Worley 1992). In addition, alien species have
commonK' been used to revegetate human-dis-
turbed sites such as roadsides and housing areas
in national parks. Livestock-related introduc-
tion of Eurasian pasture grasses by private out-
titters is also known to have occurred (Glacier
National Park Records 1939). Ho\ve\er, in spite
of these observations and practices, the effects
of alien vegetation in natural area grasslands of
this region remain poorly studied.
This study compares the indigenous \ascular
flora and crvptoganiic ground cover associated
with two cdien species, Centoiirea ituicidosa
Lam. (spotted knapweed) and Phleiim pratense
L. (common timothv), that ha\'e in\'aded a
fescue grassland in Glacier National Park, Mon-
tana. C. nmcitlosa, now a noxious rangeland
imader throughout the Pacific Northwest
(Watson and Renney 1974, Lacey 1989), was
first detected in the park in the mid- 1960s (R.
Wassem, personal communication). Earlier
obserxations have shown that this species is
expanding into grasslands adjacent to roadsides
in the park and reducing species richness (Tyser
and Key 1988). The impact of C. nuiculoso on
the cr\ptogamic ground crust — of potential
importance in soil stabilization, moisture reten-
tion, and nitrogen fixation (Rvchert and Skujins
1974, Anderson et al. 1982, Brotherson and
Rushforth 1983) — has not yet been considered,
nor has the impact of C. maculosa been com-
pared to that of other alien species. P. pratense
is widelv distributed throughut the park s grass-
lands. Unlike C. maculosa, this species appears
to have been intentionally seeded in grasslands
by outfitters before the 1940s and along road-
sides by park personnel before the 198()s (D.
Lange and J. Potter, personal comnumication).
Study Site and Methods
The ca 10-ha stud\- area, located adjacent to
Going-to-the-Sun Highway in the St. Mary
drainage of Glacier National Park, Montana, is
fairly topographically homogeneous, i.e., clearK'
defined drainage channels are absent, and
slope, exposure, and substrate texture are rela-
tively uniform. The study area includes a large
(ca 5 ha), irregularly shaped stand dominated bv
PJileum pratense and a small (ca 0.1 ha) stand
adjacent to the roadside ditch dominated b\
Centaurea maculosa. The Centaurea stand
extends >20 m away from the ditch and is not
part of the road-cut corridor. The remaining
portion of the studv site is composed of a stand
of natixe Festuca grasses and associated \egeta-
tion in which inxasion by alien species has been
minimal. Though no homesteading is known to
ha\ e occurred in the .studv area before establish-
ment of the park in 1910, this area was likel\
used as summer pasture for concession trail
horses from approximately 1915 to 1940 (B.
Fladmark, personal communication). The study
area has not been used for stock grazing since
that time. Othenvise, there is no indication that
any of the areas sampled in the three stands have
been subjected to anthropogenic disturbance
Department of Biolog\ and MicTol)iolog\, I'niversitv of Wisconsin-La Oosse, La Crosse. Wisconsin 546()L
189
190
Great Basin Naturalist
[Volume 52
since the park was established. In addition, no
fires have been recorded in or near the study
area since 1910. A cnptogani ground layer com-
posed of small lichens and mosses covering
undisturbed soil surfaces is commonly present.
Qualitative observation suggests that mosses are
the dominant element in this layer. Mean annual
precipitation in the study area is ca 65 cm
(Finklin 1986).
In each stand, vegetation was sampled using
20 X 50-cm quadrat frames along two transects
placed in representative areas. Within each
quadrat, presence of all vascular species was
determined, and the canopy cover of each vas-
cular species and the surface cover of the cry|3-
togamic ground crust were estimated to the
nearest percentage. A stratified random sam-
pling procedure was used in which quadrats
were randomly placed along transect segments
of fixed length. For the Centaurea stand, tran-
sects were 20 m long, and one quadrat was
randomly placed within each 2-m segment (N =
20 quadrats). For the Plilcmn and Festuca
stands, transects were 100 m long, and one
quadrat was randomly placed within each 5-m
segment (N = 40 quadrats per stand).
Five vegetation measures were calculated
for each individual quadrat: (1) vascular species
cover diversity using the Shannon -Wiener index
(H' = -S Pi log p,), (2) vascular species richness,
(3) cumulative canopy cover of native forb spe-
cies, (4) cumulative canopy cover of native grass
species, and (5) surface cryptogam cover. One-
way ANOVAs were used to assess among-stand
differences for each of these quadrat measures.
With the exception of the diversity measures,
data did not meet parametric assumptions
(normal distributions, homogeneous variances)
and could not be transformed using standard
logarithmic, arcsine, or square root transforma-
tions. Therefore, data were analyzed by the
Kruskal-Wallis nonparametric one-way
ANOVA procedure as described by Conover
and Iman (1983). The Tukey multiple compari-
son procedure, applicable to both parametric
and nonparametric ANOVAs (Conover and
Iman 1981), was used to make pair-wise among-
stand comparisons. Species nomenclature fol-
lows that of Hitchcock and C^rontjuist (1973).
Results and Discussion
Prominent graminoid and forb species in the
Festuca stand included Achillea millefolium.
Carex spp., Festuca idahoensis, F. scabrella,
Gaillardia aristata, and Lupinus sericeus (Table
1). Species composition of this stand was similar
to prairie communities described elsewhere in
the Pacific Northwest, e.g., the Festuca
scabrella/F. idahoensis association of western
Montana (Mueggler and Stewart 1980), the
Festuca/Danthonia prairie of Alberta (Stringer
1973), and the Washington Palouse prairie
(Daubenmire 1970). Estimated surface cover of
the cryjDtogam layer in this stand was relatively
high, characteristic of western bunchgrass prai-
ries (Daubenmire 1970, Mack and Thompson
1982). Three alien species were sampled within
the Festuca stand, though total cover of these
species was <1.0%.
Significant among-stand variation occurred
for all community measures (Table 2). Each of
these measures was lowest in the Centaurea
stand. Canopy cover of native forbs and crypto-
gam ground cover in this stand were particularly
low, only ca 18% and 4%, respectively, of the
corresponding Festuca stand measures. Thus,
effects of the Centaurea macidosa invasion on
the native flora in the study site appear to be
relatively severe. The impact of this species is
even more impressive considering its relatively
recent entry into the park.
All but one of the Phleum stand measures
were significantly lower than those of the
Festuca stand (Table 2). Canopy cover by native
graminoids in the Phleum stand was reduced to
about 50% of its level in the Festuca stand.
However, forb cover differences between these
two stands were not statisticiilly significant
(Table 2). Three species {Achillea millefolium,
Agoseris glauca, and Lupinus sericeus) were
among the four forb species with highest canopy
cover in each stand, suggesting that the forb
components of these two stands were relatively
similar. These observations suggest that Phleum
invasion has affected natixe graminoids more
than native forbs. It should also be noted that
while mean quadrat richness was lower in the
Phleum stand (Table 2), eight more species were
recorded in this stand than in the Festuca stand
(N = 59 vs. N = 51; see Table 1). Thus, different
Phleum vs. Festuca richness patterns may occur
if comparisons are derived from sampling units
larger than the 0.1-m~ quadrats used in this
study.
Cryptogam cover in the Phleum stand was
approximately 50% lower than in the Festuca
stand (Table 2). Mack and Thompson (1982)
1992]
Notes
191
Table 1. Canopv cover (%) estimates of six^cies within the Fcstuca, Phlcitin, and Ccnidurcd stands. ° = iJien sjieeies.
Species
Festuca
Phleuni
Centaurea
GlUMINOIDS
Agropijron caninum
0.4
0.6
Agroptjwn spicatum
0.3
0.3
Carex spp.
12.3
5.6
9.3
Danthonia intcmivdia
4.2
0.9
Fcstuca iclaliocnsi.s
9.2
4.3
0.2
Fcstuca saibrclla
7.1
4.1
2.1
Hclictotrichon hookcri
0.9
<0.1
Kitclcria crlstata
1.4
0.4
<0.1
Flileum pratense'
0.2
38.4
0.7
Fodjuitcifolui
<0.1
Poa pratcnsis'
<0.1
0.9
1.0
Stipa occidentalis
3.7
2.1
Stipa rirhnrrlxonii
0.1
0.8
FOKBS
Achillea millefoliuiu
11.7
8.6
0.8
Agoscri.'i glaiica
4.0
4.3
Allium cenmiim
0.1
<0.1
Atnehmchicr alnifolid
0.3
0.9
0.5
And rosace septenthoiidli.s
1.0
0.3
Anemone midtifida
1.4
1.0
<0.1
Antennaha inicroplujlld
O.S
0.3
1.7
Arahis <^lahra
<0.1
Arahis nuttallii
0.2
0.1
<0.1
Arctostaplujlos u vd-tt rsi
0.4
0.2
Aster laevis
1.8
0.9
Berheris repens
0.1
0.6
0.3
Campanula rotundifolid
0.5
1.0
<0.1
Castilleja aisickii
0.3
<().!
Centaurea nmculosa'
62.0
Cerastiu m arven.se
4.0
3.1
0.7
Colloniia linearis
<0. 1
Comandra nmtjeltata
0.5
0.3
Species
Festuca
Phleum
Centaurea
Epilohium angustifoliu m
0.5
F,rigeron suhtrinervi.s
1.5
Erysimum inconspicuum
0.3
Fru'^a ri a v i r^i niana
<0.1
0.7
Gaillardia aristata
1.9
0.6
<0.1
Galium horeale
0.6
1.8
0.2
Gentiana amarella
1.3
0.7
Geran iu m viscosissimum
<0.1
1.2
Hedijsanun horeale
0.5
Heuehera cijlindhca
0.1
0.2
0.2
Hieracium umhellatum
0.2
Jiinctts haltieus
1.0
Latlujriis oehroleucus
0.2
Lithospermu m niderale
1.9
3.9
0.7
Lonmtium tritematum
1.0
2.4
0.3
Ltipinus serieeus
5.6
6.0
<0.1
Monarda fistulosa
0.6
Orthocarpus tenuifolius
1.2
<().!
Oxijt n )p is cam pest ris
2.8
0.9
Oxtjtropi.s splendens
0.3
Penstenum confertus
0.8
1.9
0.7
Potentilla arguta
<0.1
1.1
Potentilla is^racilis
<0.1
0.4
0.3
Potentilla hippiana
0.5
Pninus vir<i^iniana
0.1
Fdnnanthus cri.Hta-<icdli
0.9
0.4
Rosa woodsii
1.3
2.3
0.2
Silene pamji
0.4
0.1
<0.1
Sisijrinchium an^iiistifolium 0.2
0.4
Soliddfio missouriensis
1.6
1.8
0.2
Taraxacum officinale °
0.2
1.4
0.3
Traoopogon duhius °
<0.1
0.4
Vicia americana
1.6
1.0
Zioadcnus venenosus
<0.1
Tablk 2. Among-stand compari.sons of quadrat means for five vegetation measures. N = 40, 40, antl 20 cjnadrats,
resjDectively, for the Festuca, Phleum, and Centaurea stands.
H'
Richness
Native
graminoids
Native
forbs
Cryptogam
crust
Festuca "
Phleum
Centaurea
F2.97
P
0.966"
0.872''
0.385''
90.084
<.001
14.8-'
12.9''
7.2'
41.1.50
<.001
.39.. 5^'
19.2*'
11.6'
.53.807
<.001
48.4"
55. r'
8.8''
40.896
<.001
28.^
15.1''
1.3''
31.835
<.001
°\\ ithin each vegetation me;Lsurt-. means with clitferf nt letters differ significantl\' Fn
otlier (P < ,05, Tiikey multiple conipi
suggest that the extensive rhizome-tiller mats of
Eurasian grasses limit cryptogam colonization
sites, which may account for the reduced cryp-
togam cover observed in the Phleum stand. A
large elk herd overwinters in the St. Mar\' \alley
grasslands in which the study area was located
(Martinka 1983). Thus, it is possible that elk
trampling/grazing may reduce cryjotogam cover
and facilitate Phleum invasion.
The role plaved bv pre- 1 940 horse grtizing in
the occurrence of Phleum in the study site
cannot be assessed. However, the prominence
of this species some 50 years after the cessation
of horse grazing ck)es indicate that ongoing live-
stock griizing is not necessarv for its persistence.
The more recent Centaurea maadosa invasion
in the study site and in other fescue grasslands
in the park (Tyser and Key 1988, Tyser and
192
Great Basin Naturalist
[Volume 52
Worley 1992) suggests that livestock grazing is
not a prerequisite for successful invasion of nat-
ural areas by this species. The success of both P.
pratense and C. maculosa in the study site sug-
gests that mechanisms proposed for the success
of alien flora in agro-systems, e.g., rapid coloni-
zation of disturbed sites, structural and physio-
logical adaptations to grazing and trampling,
and low piilataliilit}' (Mack and Thompson 1982
and references therein, Lacey et al. 1986,
Locken and Kelsey 1987, Kelsey and Bedunah
1989), may also operate in natural area systems.
In addition to overwintering elk, diggings of
other native herbivores, especially ground
squirrels (Sperniopliilus Columbia mis), were
common throughout the study area. At any rate,
though the status and impacts of C. maculosa
and P. pratense require additional research, this
study shows that the potential effects of these
species — particularlv that of C. maculosa — in
natural area bunchgrass prairies need to be seri-
ously contemplated.
Reduction of Plileum pratense is not a real-
istic option in Glacier National Park or other
natural areas in which this species is now widely
established. Perhaps the most reasonable rec-
ommendation for this species and other Eura-
sian grasses is simply that resource managers not
use these species for revegetation (see also
Wilson 1989). Centaurea maculosa, though
potentially more ecologically disruptive than P.
pratense, is at an earlier stage of invasion in the
park and probabK' in other natural areas in this
region as well. Thus, the fate of this species may
yet be influenced by appropriate resource man-
agement actions.
Acknowledgments
I thank Tom Jacobsen and Andy Tyser for
their assistance with fieldwork, Andy Matchett
for statistical advice, and Glaciers research and
resource management staff for support and
assistance with this stud\'. The study was funded
by a grant from the Universit\' of Wyomincj-
National Park Service Research Center.
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Received 26 April 1991
Accepted 16 April 1992
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GREAT BASIN NATURALIST voi 52 no 2 June 1992
CONTENTS
Articles
Red Butte Canyon Research Natural Area: history, flora, geology, climate, and
ecology James R. Ehleringer, Lois A. Arnow, Ted Arnow,
Irving R. McNulty, and Norman C. Negus 95
Influences of sex and weather on migration of mule deer in California
Thomas E. Kucera 1 22
Diatom flora of Beaver Dam Creek, Washington County, Utah, USA
Kurtis H. Yearsley, Samuel R. Rushforth, and Jeffrey R. Johansen 131
Stratification of habitats for identifying habitat selection by Merriam's Turkeys
Mark A. Rumble and Stanley H. Anderson 139
Pollinator preferences for yellow, orange, and red flowers of Mimulus
verbenaceus and M. cardinalis Paul K. Vickery, Jr. 145
Soil loosening processes following the abandonment of two arid western Nev-
ada townsites Paul A. Knapp 149
Biochemical differentiation in the Idaho ground squirrel, Spennophilus brun-
neus (Rodentia: Scuridae) Ayesha E. Gill and Eric Yensen 155
New genus, Aplanusiella, and two new species of leafhoppers from south-
western United States (Homoptera; Cicadellidae: Deltocephalinae)
M. W. Nielson and B. A. Haws 160
Summer habitat use by Columbian Sharp-tailed Grouse in western Idaho . . .
Victoria Ann Saab and Jeffrey Shaw Marks 166
Notes
Characteristics of sites occupied by subspecies of Artemisia tridentata in the
Piceance Basin, Colorado . . Thomas R. Cottrell and Charles D. Bonham 1 74
Use of lakes and reservoirs by migrating shorebirds in Idaho
Daniel M. Taylor and Charles H. Trost 1 79
Dispersal of squarrose knapweed {Centaurea virgata ssp. squarrosa) capitula by
sheep on rangeland in Juab County, Utah Cindy Talbott Roche,
Ben E Roche, Jr., and G. Allen Rasmussen 185
Vegetation associated with two alien plant species in a fescue grassland in Gla-
cier National Park, Montana Robin W. Tyser 189
H E
MCZ
LiijRARY
V mo
HARVARD
UNIVhHSlTY
GREAT BASIN
MURAUST
VOLUME 52 NO 3 - SEPTEMBER 1992
BRIGHAM YOUNG UNIVERSITY
GREAT BASIN NATURALIST
Editor
James R. Barnes
290 MLBM
Brigham Young UniversiU'
Provo, Utah 84602
Associate Editors
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Museum of Southwestern Biolog)', Universit)' of
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Mailing address: Box 3140, Hemet, California
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Depiirtment of Biology, John Carroll University,
Universit)' Heights, Ohio 44118
Paul C. Marsh
Center for En\ironmental Studies, Arizona State
University; Tempe, Arizona 85287
Brian A. Maurer
Department of Zoolog); Brigham Young Uni\ersity,
Provo, Utah 84602
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Copyright © 1992 b\ Brigluuii Young University
Official publication date: 18 December 1992
ISSN 0017-3614
12-92 750 2473
The Great Basin Naturalist
Plblishkd atPhono, Utah, by
Bricham Young Um\ersi'it
ISSN 0017-3614
Volume 52 September 1992 No. 3
Great Basin Naturalist 52(3), pp. 195-215
PLANT ADAPTATION IN THE GREAT BASIN AND COLORADO PLATEAU
Jonathan P. Comstock antl James R. Elileringer
Al5STlU(X — Adapti\e features of plants of tlie Great Basin are reviewed. The combination of cold winters and an arid
to semiarid precipitation regime results in the distinguishing features of the vegetation in the Great B;isin and Golorado
Plateau. The priniaiy effects of these climatic features luise from how the\ structure the hvdrologic regime. Water is die
most limiting factor to plant growth, and water is most reliabK axailahle in the earl\- spring after winter recharge of soil
moisture. This factor determines main characteristics of root moipholog\, growth phenolog\- of roots and slKX)ts, and
photos\ndietic physiolog): Since winters are hpicallv cold enough to suppress growth, and drought limits growth during
the summer, the cool temperatures characteristic of the peak growing .season are the second most importiuit climatic factor
influencing plant habit luid perform;uice. The combination of several distinct stress periods, including low-temperature
stress in winter and spring and high-temperature stress combined with drought in summer, appears to have limited plant
habit to a greater degree thiui found in the warm de.serts to the south. Nonetheless, cool growing conditions and a more
reliable spring growing season result in higher water-u.se eiiiciencv and productiv ih" in the vegetation of the cold de.sert
than in warm deserts with equiv;ilent total rainfall amounts. Edapliic factors are also importimt in structuring communities
in these regions, and halophvtic connnunities dominate main landscapes. These haloph\-tic communities of the cold desert
share more sj^ecies in common with warm deserts than do the nonsdine communities. The Golorado Plateau differs from
the Great Basin in having greater amounts of smniner rainfall, in some regions less predictable riiinfall, sandier soils, and
streams which drain into river .systems rather than closed basins and salt plavas. One result ofthe.se climatic and edapliic
differences is a more important summer grov\ing seasf)n on the (Colorado j'lati'au and a sonu-wliat <ii"eater di\c'rsilication
of plant habit, phenolog), and physiolog)'.
Key icords: cold desert, plant adaptation, water stress, phenalo'^ij. salinitt/. Great Basin. Colorado Plateau.
Several features arising from climate and Nevada and increase both to the north and ea,st,
geolog)' impose severe limitations on plant and to the southeast moving into the Colorado
gro\\i:h and activit)^- in the Great Basin and Col- Plateau (Fig. 1, Table 1). The fraction of annual
orado Plateau. The climate is distinctlv conti- precipitation during the hot sununer months
nental with cold winters and warm, often dn (|une-Se[)tember) varies considerabh; from
sununers. Annual precipitation levels are low in l()-2()9f in northern Nevada to 30-40% along
the basins, ranging from 100 to 300 mm (4-12 the boundaiAof theCold and .\Iojave deserts in
inches), and t)piciilly increasing with elevation southwestern Nevada and southern Utah, and
to 500 mm (20 inches) or more in the montane 35-50% throughout much of the Colorado Pla-
zones. Precipitation levels are lowest along tlu^ t(^au. Winter jirecipitation falls primariK' as
southwestern boundan' of the Great Basin in snow in the Great Basin and liii£her elevations
Department of Biologv-. Universit)- of t'tali. Salt Lake Cit^â– . Utiih 84 1 12.
195
196
Great Basin Naturalist
[\blume 52
TaHI.K 1. Srlec'tec! climatic data tor l(m-elc'\ation sites in different regions of the Great B;Lsin, Moja\e Desert, and
Colorado Platean. Viilues are based on a\erages for the U.S. Weather Bureau stations indicated. The tliree dixisions of the
year presented here reflect ecologically relevant units, but are unequal in length. The fixe months of October-Februan
represent a period of temperature-imposed plant dorm;uicy and winter i-echarge of soil moisture. The spring mondis of
March-Mav represent the potential growing period at cool temperatures immediately follcnving winter recharge. The
summer and ei\r\\ fall from Jiuie through September represent a potential warm growing season in areas with sufficient
summer rain or access to other moisture sources.
Map #
Weather E
levation
Total precipitation
Mean
temperature
Region
Annua
1 Oct-Feb
Mar-Ma\ J
ini-Sep
Annual Oct-Fe
'b Mar-Ma\
Jiui-Sep
(Fig. 1)
station
(m)
(mm)
(%)
(%)
(%)
(°C)
(°C)
(°C) '
(°C)
Northern
I
Fort Bidwell
1370
402
63
24
13
9.0
3.0
8.0
17.3
Great Basin
r>
l^eno
1340
1S2
61
24
15
9.5
3.3
8.4
18.0
3
Elko
1547
230
52
29
19
7.6
0.1
7.1
17.5
4
Snowxille
1390
300
43
33
24
7.4
0.7
6.2
18.4
Southern
5
Sarcobatus
1225
85
45
22
33
13.5
6.4
12.5
23.1
Great Basin
rs
C;iliente
1342
226
47
24
29
11.7
4.1
11.2
21.5
'
Fillmore
1573
369
44
34
•1-1
11.0
3.0
10.0
21.7
Moja\e Desfc
'rt S
Trona
517
102
70
19
11
19.0
11.3
18.4
29.0
9
Bea\erdam
570
169
50
23
28
18.3
11.0
16.9
28.6
Colorado
10
Hanksxille
1313
132
36
19
45
11.4
2.1
11.5
22.8
Plateau
11
Clrantl Junction
147S
211
39
25
36
11.3
2.4
10.9
22.9
12
Blanding
1841
336
48
19
.33
9.7
2.1
8.7
19.9
13
TnbaCit\'
1504
157
38
21
41
12.6
4.8
12.0
22.8
14
(]haco Canvon
1S67
220
35
20
45
10.3
2.6
9.4
20.6
of the Colorado Plateau, which is thought to be
a critical feature ensiuins soil moisture recharge
and a reliable spring growing season (West
1983, CakKvell 1985, Dobrowolski et al. 1990).
During the winter period, precipitation is gen-
eralK' in excess of potential exaporation, but low
temperatures do not permit gro\\1:h or photo-
sxnthesis, and exposed plants may experience
shoot desiccation due to dry winds and frozen
soils (Nelson and Tienian 1983). Strong winds
can also cau.se major redistributions of the snow-
pack, sometimes rexersing the expected
increase in [)recipitation with ele\ation and
having important consecjuences to plant distri-
butions (Branson et al. 1976, Sturges 1977, W'e.st
and Caldwell 1983). The important growing
season is the cool spring when the soil profile is
recharged from winter precipitation; growth is
usualK' curtailed b\- dning soils coupled with
high temperatures in earl\- or mid-smnmer. A
clear pictiu-e o( this climatic regime is essential
to an\- chscussiou of plant adaiitations in the
region.
A second major feature affecting plant per-
formance is the prevalence of saline soils caused
l)\' the C()ml)ination of low precipitation and the
internal drainage txpical of the Great Basin. In
this paper we address the salient morphological,
physiological, and phenological specializations
of nati\ e plant species as the\' relate to siua i\al
and growth tmder the constraints of these
potentialK stressful limitations. We emphasize
(1) edaphic factors, particularK soil salinit\ and
texture, and (2) the climatic regime ensuring a
fairlv dependable, deep spring recharge of soil
moisture despite the overall ariditv; as factors
molding plant adaptations and producing the
uni(jue aspects of the regional plants and vege-
tation. The majoritv of the Great Basin lies at
moderatelv high elevations (4000 ft and aboxe),
and it is occupied bv cold desert plant comnm-
nities. Reference to "the Great Basin" and its
environment in this paper will refer to this high-
{^]e\ation region as distinct from that corner of
the Mojave Desert that occupies the southwest-
em corner of the Cireat l^asin geographic unit
(Fig. 1 ). Our emphasis will be placed on these
cold desert shnib communities in both the
Great Basin and the Colorado Plateau ranging
from the topographic low points of the saline
plavas or cauNon bottoms up to the pinvon-juni-
per woodland. The lower-elevation, warmer.
19921
Plant Adaptation
197
Great Basin
Mojave
Colorado Plateau
Fig. 1. Distribution of the major desert vegetation zones
ill tlie Great Bitsin and Colorado Plateau. Numbers indicate
l(K'atioiis of climate stations for which data are presented in
Table 1. Most of the Mojave Desert indicated is geologically
part of the Great Basin, but, due to its lower elevation and
warmer temperatures, it is climaticallv distinct from the rest
of the region.
antl drier Mojax'e De.sert portion of the Cjreat
Basin will he considered primariK as a point of
e()inj)arison, and for more tlioronii;h coxerage of
that region we recommend the reviews h\
Ehleringer (1985), MacMahon (1988), and
Smith and Nowak (1990). For the higher mon-
tane and alpine zones of the desert mountain
ranges, the reader is referred to rexiews l)\
\'asek and Thome (1977) and Smith and Knapp
( 1990). We are indebted in onr own c()\erag(^ of
the cold desert to other rec-ent rexic^ws. includ-
ing Caldwell (1974, 19S5). West (19SS).
Dobrowolski et al. (1990), and Smith and
Nowak (1990).
The Great Basin and the Colorado Plateau
share important climatic features such as overall
ariditv; frequent summer droughts, and conti-
nental winters; yet the\^ differ in other ecjualK
important features. Temperatures on the Colo-
rado Plateau are similar to ecjuixalent elexatioiis
in the southern (Ircat liasin. hut suiiiiiici' pre-
cipitation is suhstantially greater on the Colo-
rado I'lateau (Tahk" \). Soils and drainage
patterns also differ in crucial wa\s. The high-
lands of the Colorado Plateau generally drain
into the Colorado Hixer sv'stem. In manv areas
extensive exposure of marine shales from the
Chinl(\ \hmcos, and Morrison Brnshv-Basin
formations wc^ithcr into soils that restrict plant
diversitv and total cover due to high concentra-
tions of NaSOa, and the formation of clavs that
do not allow water infiltration (Potter et al.
1985). In other areas massive sandstone out-
crops often dominate the landscape. Shrubs and
trees mav root through ven shallow rock"v soils
into natural joints and cracks in tlie sub.stratum.
Deeper soils are generallv aeolian deposits
forniinti sands or sandv loams. In contrast, high
elevations of the Cireat Basin drain into closed
\alleys and evaporative sinks. This results in
greater average salinitA' in the lowland soils of
the Great Basin, with NaCd being the most
common salt (Flovxers 1934 ). and a more exten-
sive development of haU)ph\ tic or salt-tolerant
vegetation. Soils tend to be deep, especialK at
lower elevations, and van' in texture from clav
to sandv loams. Summer-active species with
different photosvnithetic pathwavs, such as C4
grasses and CAM succulents, are poorlv repre-
sented in nuich of the Crreat Basin, but the
combination of increased summer rain, sandier
soils, and milder winters at the lower elevations
of the Colorado Plateau has resulted in a greater
expression of phenological diversit\.
The interactions of edaphic factors and cli-
mate are complex and often subtle in their
effects on plant performance. Furthermore,
[)Iant distributions are rarelv determined bv a
single factor throughout th(ir geographic range.
e\en though a single factor mav appear to con-
trol distribution in the context of a local ecosvs-
tcMii. Species-spcxific characteristics generally
do not inqxirt a narrow re(|uirement for a spe-
cific environment, but rather a unique set of
"ranges of tolerance" to a large arrav of enxiron-
mental j)arameters. In different enviromncntal
contexts, different tolerances mav be more lim-
iting, both abiotic and biotic interactions may be
altered, and the same set of species may .sort out
in different spacial ])attenis. A further conse-
(juence of this is that a local combination of
species, whicli we might refer to as a Great
Basin plant communitv. represents a region of
oxerlap in the geograpln'calK more extensive
198
Great Basin Naturalist
[N'oluiiie 52
and treiieralK miicjue tlistrihutioiis ot each coni-
ponenf species. In fact, the distributions of spe-
cies commonly associated in the same Great
Basin connnunitv' may be strongly contrasting
outside the Great Basin. This is an essential
point in attempting to discuss plant adaptations
with the implication oi cause and effect,
because few species discussed will have a strict
and exclusixe relationship with the environment
of interest. Unless we can show local, ecot\pic
differentiation in the traits discussed, we need
to take a broad view of the relationship between
environment and plant characters. In a few
instances, a small number of edaphic factors and
plant characters, such as tolerance of veiy high
salinity in soil wdth shallow groundwater, seem
to be of overriding importance. In most cases we
need to ask, what are the common elements of
climate over the diverse ranges of all these spe-
cies? One such common element, which will be
emphasized throughout this re\iew, is the
importance of reliable winter recharge of soil
moisture in an arid to semiarid climate. B\- iden-
tifying such common elements and focusing on
them, we do not fully describe the autecologv of
an\' species, but we attempt a cogent treatment
of plant adaptations to the Great Basin environ-
ment, and an explanation of the unicjue features
of its plant connnunities.
Climate, Edaphic Factors, and Plant
Distribution Patterns
Typical zonation patterns observed in spe-
cies distributions around playas (the saline flats
at the bottom of closed-drainage basins) are
quite dramatic, refl(^cting an o\ erriding effect of
salinit)' on plant distribution in the Cireat Basin.
Moving out from the basin center is a gradient
of decreasing soil salinity often correlated with
progressively coarser-textured soils. Along this
gradient there are conspicuous species replace-
ments, often resulting in concentric zones of
contrasting vegetation (Flowers 1934, Vest
1962). In the lowest topographic zone, saline
groundwater may be very neav the surface. Soils
are ven' saline, fine textured, and subject to
occasional flooding and anoxic conditions, in
this enxiromnent the combination of available
moisture with other poteutiallv stressful soil
characteristics seems to be more important than
climatic factors of temperatiu'e or seasonal rain-
fall patterns. Speci(>s found here, such as desert
saltgrass {Distichlis spic<il(i), pickleweeds
(Allciirolfia occich'ittdlis and Salicontia spp.),
and greasewood (Sarcobatiis vcniiicitkitiis),
may themselves show zonation due to degrees
of tolerance. They tend to occur in close prox-
imitv, however, on the edges of salt plavas, saline
flats with shallow water tables, and near saline
seeps over a wide range of elevations, tempera-
tures, and seasonal rainfall patterns in both the
Great Basin and southern warm deserts
(MacMahon 198S). This relative independence
of distribution from prevailing climate is a spe-
cial characteristic of strongly halophytic plant
communities throughout the world and reflects
the overriding importance of such extreme
edaphic conditions. Species found on better-
drained, moderately saline soils, where groimd-
water is not found within the rooting zone,
include winterfat {Ccratoidcs laiuita) and
shadscale {Atiiplcx confeififolia). These species
are, in turn, replaced at higher elevations and on
moister, nonsaline soils bv big sagebnish iAiic-
inisia tridcntatd), rabbitbrush [Chnjsoiluntinus
sp.), bitterbnish {Piirsliia sp.), and spinv hop-
sage {GiYHfia spinosa). Shadscale is often fcnmd
in areas where soils are notably saline in the
lower half of the rooting zone, but not in the
upper soil lavers. Such a tolerance of mt)der-
ately saline soils seems to control its distribution
around playas, especially in the wetter, eastern
portion of the Great Basin (western Utah) and
lower elevations in the warm Mojave Desert. In
the more arid regions of western and central
Nevada, however, the shadscale connnunitv
occurs widely on nonsaline slopes lower in ele-
vation, warmer, and drier than those dominated
by big sagebrush. These higher bands of
shadscale have been variously inteipreted in
terms of ariditv tolerance and climate (Billings
1949) or an association with limestone-derived
calcareous soils (Beatlev 1975). The latter
author points out that even on nonsaline soils
percent cover in the shadscale connnunitv is
lower than expected for the level of precipita-
tion, and argut^s that this indicates stress from
ecUiphic factors. Thus, shadscale distribution is
most correlated with salinitv tolerance in some
regions and other eckiphic and climatic tolcMan-
ces in other regions.
Where the higher elevations of thc> Cyreat
Basin conu^ in contact with the lower-elevation,
generallv drier, and warmer Mojave Desert
region, comminn'ties ck)minated by creosote
(Larrca tridfufafa) replace sagebrush commu-
nities on nonsaline slopes and bajadas.
19921
Plant Adaitxtion
199
Shadscak' can ht' toiiiul liotli on saline soils at
\en low t'k'\ ations in tlu^ Mojaw and as a tran-
sitional band at liigli ekn ations l)et\\een creo-
sote and sagebmsh. Elements of the cold desert
shnib conimnnities, adapted to continental win-
ter's and a cool s[)ring growing season, can be
tonnd throughout the winter-rain-doniinated
\h)ja\"e Desert region as a high-elexation band
on arid mountain ranges. They also extend to the
southeast at high ele\ations into the strongK-
bimodal precipitation regime of the Colorado
i'latean, and northward at low elexations into
Idaho. Washington, andexen British (-oluinbia.
Nhning up from bajadas of the southern warm
deserts, there appears to be no suitable combi-
nation of temperature and precipitation at an\'
elevation to support floristic elements of the
cold desert. As precipitation increases with alti-
tude, zones with equivalent total precipitation
do not \et ha\e cold winters and are occupied
In warm desert shnib connnunities grading into
chaparral composed of unrelated ta.xa. Higher
ele\ ations with cold winters have sufficient pre-
cipitation to support forests. Other elements
coimnon in shadscale and mixed-shrub connnu-
nities of the Great Basin, such as winterfat and
budsage (Ai-tcmisia spiiiosa), are often found
outside the Great Basin in cold-winter but
largel\- summer-rainfall grasslands.
f^rom these patterns of communitv- distribu-
tion within the Great Basin and Colorado Pla-
teau, and also from a consideration of the more
extensive ranges and affinities of the component
species, we can isolate a few ke\- features of the
environment that are largely responsible for the
unique plant features seen in the Great Basin.
The most obvious features are related to the
patterns of soil salinitv and texture generated bv
the (Aerall ariditv combined with either internal
drainasie basins or tlie in situ weathering of
specific rock tvpes. The most important climatic
variables are slightlv more subtle. There is
cknulv not a requirement for exclusivelv winter
rainfall, but there is a re(|uirement for at least a
substantial portion ol the annual rainfall to come
dniing a continental winter This permits v\inter
(iccitninlatioit of precipitation iod 'greater depth
in the soil profile than w ill occur in response to
less predictable sunnner replenishment of
dning soil moisture reserves. Unck'r an overall
arid climate, winter n^charge maintains a pre-
ilictablv favorable and ck)minant spring growing
season even in manv areas of strongly bimodal
rainfiill. Most of the phvsiological. moqihologi-
cal. and plieiiological traits lonnd in llie (k)mi-
nant shrubs rell(^ct the [)riman importance of
such a cool spring growing .season.
PlI()T(lSY\'THKSIS
Piiotosyxtiiktk; I'ATIIWAVS. — The pro-
cess of photosvnthesis in plants relies on the
acquisition of CO2 from the atmo.sphere, which,
when coupled with solar energ\', is transformed
into organic molecules to make sugars and pro-
vide for plant growth. In moist climates plant
communities often achieve clo.sed canopies and
1(){)% cover of the ground surface. Under these
conditions competition for light may be among
the most important plant-plant interactions. In
the deserts total plant cover is much less than
100%, and in the Great Basin closer to 259f.
Photosviithesisisnotgreatlvlimitcxlbv available
light, but rather bv water, mineral nutrients
needed to .synthesize enzAines and maintain
metabolism, and maximum canopv leaf-area
development.
Three biochemical pathwavs of photosvii-
thesis have been described in plants that differ
in the first chemical reactions associated with
the capture of CO2 from the atmosphere. The
most common and most fundamental of these
pathways is referred to as the C3 pathway
because the first product of photosynthesis is a
3-carbou molecule. The other two pathways, C4
and CAM, are basically modifications of the
primaiy C3 pathway (Osmond et al. 1982). The
C4 pathwav (first product is a 4-carbon mole-
cule) is a morphological and biochemical
arrangement that overcomes photorespiration,
a process that results in reduced photosviithetic
rates in C3 plants. The C.i pathway can confer a
much higher temperature optimum for photo-
.synthesis and a greater water-use efficiency. As
water-use efficiencv is the ratio of photosvn-
thetic carbon gain to transpirational water loss,
C4 plants have a metabolic advantage in hot, dn^
environments w4ien soil moisture is available. In
grasslands C4 grasses become dominant at low
elevations and low latitudes where animal tem-
]x^ratur(\s are warmest. In interpreting })lant
distribution in deserts, the .seasonal pattern of
rainfall usuallv restricts the periods of plant
growth, and the temperature during the rainy
season is thus more important than m(\ui annual
temperature. The C4 pathwav is ofti'u associated
with smnmei-active species and also with plants
of saline soils. While C3 grasses pre(k)minate in
200
Great Basin Naturalist
[\'olunie 52
most of the Cireat Basin, C4 grasses beeonie
iiicreasinglv important as summer rain increases
to the south, and especiaHv on the Colorado
Plateau. Halophvtic plants are often C4, such as
saltbush iAfrij)Icx spp.) and saltgrass (Disticlilis
spp.), and tliis mav gixe the plants a competitixe
advantage from increased water-use efficienc\-
on saline soils.
The third photo.sMithetic pathway is CAM
photosMithesis (Crassulacean Acid Metabolism).
CAM plants open their stomata at night, capture
COo and store it as malate in the cell \acuole,
and keep theii stomata closed dining the dav
(Osmond et al. 1982). The CO2 is then released
from the vacuole and used for photos)aithesis
the folloxxing da^'. Because the stomata are open
onl\ at night when it is c()t)l, water loss associ-
ated with CAM photosNuthesis is greatlv
reduced. This pathwa\' is often found in succu-
lents such as cacti and agaxe, and, although
C^AM plants are present in the Great Basin, they
are a i-elati\eK- minor component of tlie vegetation.
Photosxnthetic rates of plants, like most met-
abolic processes, sho\\' a strong temperatm-e
dependence. UsualK, photosvnthetic rates are
reduced at low temperatures because of the
temperature dependence of enz^'uie-catah'zed
reaction rates, increase with temperature mitil
some maximum \alue (which defines the "tem-
perature optimum"), and then decrease again at
higher temperatures. The temperature optima
and niiuimum photosxnthetic rates of plants
show considerable variation, and the\' generalK
reflect the growing conditions of their natural
environments.
PHOTOSYNTHETIC adaptation. — In the
spring the photosynthetic temperature optima
of the dominant shrub species are tvpicalK' as
low as 15 C (40 F) (Caldwell 1985), correspcMid-
ing to the prevailing en\ironmental tempera-
tures (mi(kla\- ma.xima generally less than 20 C).
As ambient temperatures rise 10-15 C in the
summer, there is an upward shift of only ,5-10 C
in the photos\iithetic temperature optima of
most shrubs, coupled with a slower decline of
photosynthesis at high temperatures. The max-
imimi ph()t()s\nithetic rates measunxl in most
Great Basin shrubs under either natural or irri-
gated conditions range from 14 to 19 jjluioI ClO^
m- s' (DePuit and Caldwell 1975, Caldwell et
al. 1977, Evans 1990). These rates are (|uite
mode.st compared to t\ie high maxima of 25 to
45 jjLmol CO2 m " s ' ob.sened in man\- warm-
dc^sett species adapted to rapid growth at higher
temperatures (Ehleringer and Bjorkman 1978,
Mooney et al. 1978, Comstock and Ehleringer
1984, 1988, Ehleringer 1985). This presumably
reflects the specialization of these Great Basin
shiiibs towards utilization of the cool spring
growing season. Positive photosynthetic rates
are maintained even when leal temperatures
are near freezing, which permits photosvnthetic
activity to begin in the very early spring (DePuit
and Caldwell 1973, Caldwell 1985).
Unusuallv high maximmn photosvnthetic
rates of 46 ixmol CO2 m ~ s ' have been reported
for one irrigated Great Basin shnib, rabbitbrush
{Chnjsothamnus nauseosus) (Da\is et al. 1985).
This species is also unusual in having a deep tap
root that often taps groundwater, unusuallv high
levels of summer leaf retention (Branson et al.
1976), and continued photo.sxnthetic activitx*
throughout the summer drought ( Donoxan and
Ehleringer 1991). All of these characters sug-
gest greater photosvnthetic activity during the
warm summer months than is found in most
Great Basin shrub species.
Shoot ACTTIMTY' and phenology. — Gener-
ally speaking, there is a distinct drought in early
summer (June-|ulv) in the Great Basin Cold
Desert, the Mojave Desert, and the Sonoran
Desert. All of these deserts ha\e a substantial
winter precipitation season, but they differ in
the importance of the summer and early fall
rainy seas(jn (|ul\-October) in supporting a dis-
tinctive period of plant growth and acti\itv
(MacMahon 1988). The relationship between
climate and plant growing season is complex and
includes total rainfall, seasonal distribution of
rainfall, and predictabilitv of rainfall in different
seasons as important \ariables. Fmthermore, in
\en arid areas the seasonalih' of temperatures
may be as important as that of rainfall. In the
Great Basin, cold winters allow the moisture
from low lexels of precipitation to accumulate
in the soil for spring use, while hot summer
temperatiu'es cause rapid evaporation from
plants and soil. High temperatures can prevent
wetting of the soil profile bevond a few centime-
t(Ms depth in response to sununer rain, even
when sununer rain accounts for a large fraction
of the animal total (Caldwell etal. 1977). As total
annual rainfall decreases, the relative impor-
tance of the cool spring growing season
i I icreases as the oiiK potential growing period in
which available soil moisture approaches the
evaporative demand (Thornthwaite 1948, Com-
stock and Ehleiintier 1992). Finally, reliabilih
19921
Plant Adaitation
201
of nioisturc is important to [XTcnnials, and as
axerage total precipitation decreases, the \ari-
ance bet\veen \ears increases (Ehleringer
1985); \ariabilit\' of annuiil precipitation is dis-
cussed in more detail later in the section on
annuals and life-histor\' dixersitv. Summer rain
is more likel\- to be concentrated in a few high-
intensit\ storms that max not happen e\eiA' \ear
at a gi\en site and ma\' cause more nmoit when
the\ do occur. The abilits' of moisture from
w inter rain to accumidate o\er several months
greatly enhances its reliabilits' as a moisture
resource. Thus, most plants in the Great Basin
have their priniar\- growing season in the spring
and earl\- summer. Some species achie\e an
e\ergreen canop\' b\' rooting deepK; but few
species occur that specialize on growth in the
hot summer season (Branson et al. 1976, Cald-
well et al. 1977, Everett et al. 1980). Ehleringer
et al. (1991) measured the abilitv of common
perennial species in the Colorado Plateau to use
moisture from summer convection storms.
The\- obserxed that less than half of the water
uptake b\- wood\' perennial species was from
suriace soil laxers saturated b\' summer rains.
Prexalence of summer-active species increases
along the border betxveen higher basins and the
southeast Mojaxe Desert xvhere summer rain is
more extensixe, and especialK' on the Colorado
Plateau xx'here summer rain is greatest. Summer
temperatures are also lower on the Colorado
Plateau than in the eastern Mojaxe (Table 1),
alloxxing more effectixe use of the moisture.
Most phenolog)- studies, especiallx' from the
more northern areas, emphasize the directional,
sequential nature of the phenological phases
(Branson et al. 1976, Saner and Uresk 1976,
Cambell and Harris 1977, West and Gastro
1978, Pitt and W'ikeem 1990). A single period of
spring vegetative groxvth is usually folloxved by
a single period of floxxering and reproductix'e
groxx'th. Manx- species produce a distinct cohort
of ephemeral spring leaves and a later cohort of
exergreen leaxes (Daubenmire 1975, Miller and
Schultz 1987). For most species, multiple
groxxth episod(\s associated xxith intermittent
spring and summer rainfall exents do not occur.
In xears xxith unusually heavy August and Sep-
tember rains, a distinct second period of xegeta-
tixe growth may occur in some species (West
and Gastro 1978). Climatic xariations from xear
to xear do not change the primaty importance
of spring gro\xi:h or the order of phenological
exents. In particular \ears, thex' ma\- cause
expansion or contraction of xc^gt^tatixc pluuses
and exen the omission of reproductix-e pha.ses.
Most species initiate grox\th in earlx' spring
(March) xvhen maximum da\time temperatures
are 5-15 C and xx'hile nighttime temperatures
are still freezing. Delaxed initiation of spring
groxxth is generally associated xxith greater
summer actixit\- and max- be related to an exer-
green habit, a phreatophxtic habit, or occupa-
tion of habitats xxith greater sununer moisture
axailabilitx from regional rainfall patterns,
nmoff, or tirovmdxx'ater. Higher-than-ax-erase
xxinter and spring precipitation tends to prolong
vegetatixe growth and delax- reproductive
groxx'th till later in the sununer ( Saner and U re.sk
1976, Cambell and Harris 1977). Strong xegeta-
tive dormancy ma\' be displayed in mid-summer
(Everett et al'. 1980, Evans 1990), although root
groxx'th (Hodgkinson et al. 1978) and increased
reproduction (W'est and Gastro 1978, Exans,
Black, and Link 1991) max' still occur in
response to rain at that time. In xears with
beloxx'-axerage spring and svunmer precipita-
tion, leaf senescence is accelerated and floxx'er-
ing may not occur in man\- species.
The time taken to complete the full annual
groxxth cxcle including both xegetatixe and
reproductixe stages is stronglx related to rooting
depth in most of these conmumities, xxith deep-
rooted species prolonging actixit\' further into
the summer drought (Pitt and Wikeem 1990).
The exact timing of floxx'ering and fniit set shoxvs
considerable xariation among different shrub
species. Some, especiallx those that are
drought-deciduous, lloxxer in late sprin>j; and
earlx summer just prior to or concurrent xxith
the onset of summer drought. Manx- exergreen
shRib species begin floxxering in midsummer
(Artonisia) or in the fall {Gutierrczia and
Chn/sothainntts). These late-flowering species
generallxdo not aj)pear to utilize" stored reserx'es
for floxx'ering. but relx on current photo.sxnthe-
sis during this least fax-orabk" period. In the case
(){ Aticmisia fridoitafa. it has been shoxxn that
earlx )lix-drates used to fill fruits arc dcrixcd
exclnsixi'lx from the inflorescences theniselxes,
xxhile photosxnthate from xegetatixe l)ranches
presumablx continues to support root groxx'th.
Summer rain during this time period does not
promote xegetatixe shoot groxxth but does
increase xvater use by and the ultimate size of
inflorescences (Exans 1990). Exans, Black, and
Link (1991) haxe argued that this pattern of
floxx'ering, ba.sed on residual deep soil moisture
202
Great Basin Naturalist
[Volume 52
and the unreliable summer rains, ma)' contrib-
ute to competitixe dominance within these
comnumities. The more favorable and much
more reliable spring growing season is used for
\egetative growth and coiupetitive exploitation
of the soil \olume, while reproductive gro\\i:h is
delayed until the less favorable season, and may
be successful only in years with adequate
su mmer precipitation .
Most grasses in the northern part of the
Great Basin utilize the G,5 pathway and begin
growth very early in the spring. These species
complete fruit maturation by early or mid-
sunnner, often becoming at least partially dor-
mant thereafter. On the Colorado Plateau, with
much greater amounts of summer precipitation,
there is a large increase in species number and
cover by C4 grasses such as bluestem
(Andropogon) and grama {Bouteloua), espe-
cial K at warmer, lower elevations and on deep
sandy soils. Many of these species occur in
mixed stands with the C3 species but delay ini-
tiation of growth until May or Jime; they can be
considered suiumer-active rather than spring-
actix'e. In contrast, some C4 grasses such as sand
dropseed {Sporoholii.s cri/ptcmdnis), galleta
grass (Hilaria jainesiii), and three-awn {Arisfkla
purpurea) are widespread in the Great Basin
where sunuuer rain is only moderate in long-
term averages and veiy inconsistent year to year.
Spring growth of these widespread species
tends to be slighth' or moderately delayed com-
paied to co-occurring C5 grasses, but they are
still able to complete all phenological stages
based on the spring moisture recliarge. The\'
show a greater abilit)' than the G,; species to
respond to late spring and simuiier rain witli
renewed growth (Everett et al. 1980), however,
which compensates in some years for their later
developuKMit. Other C4 grasses in the Great
Basin may be associated with seeps,
streamsides, or salt-marshes, and show a
summer activity' pattern. G4 shrubs such as salt-
bush (Atriplex) show similar, spring-actixe
growth patterns to the (v; shrubs, but may show
slightly greater tolerance of sunuuer drouglit
(Caldwell et al. 1977).
Phenolog)' in the Mojave Desert shows both
similarities and strong contrasts to the Great
Basin. Although rainfall is largeK biiuodal in the
eastern Mojave, absolute amoimts are vvw low.
The sunuuer is so hot that little growth normally
occurs at that time unless more than 25 nun (1
inch) comes in a single storm (Ackerman et al.
1980). Fall and winter precipitation is the mo.st
important in promoting good spring growth of
perennials (Beatley 1974). Comstock et al.
(1988), looking at one years growth in 19
Mojave species, described an important cohort
of twigs initiated during the winter period that
accounted for most vegetative growth during
the following spring. Although new leaves were
produced in response to summer rain, summer
growth in many of the species was largeK' a
further ramification of spring-initiated floral
branches. In most species summer growth made
little contribution to perennial stems. Despite
the preferred winter-spring orientation of many
shmbs, winter recharge is much less effective
and reliable in the Mojave Desert than in the
Great Basin. Due to warmer temperatures,
winter dormancy may be less complete, but
vigorous growth rarely occurs until tempera-
tures rise further in the early spring. Rapid
growth luay be triggered by rising spring tem-
peratures or may be delayed until major spring
raius provide sufficient moisture (Beatley 1974,
Ackenuan et al. 1980). Furthermore, a shal-
lower soil moisture recharge often results in
fluctuating plant water status and multiple
episodes of growth and flowering during the
spring and early fall. Some Great Basin species
that also occur in the Mojave, such as winterfat
and shadscale, commonly show multiple growth
and reproductive episodes per year under that
climate (Ackennan et al. 1980) but not in the
Great Basin (West and Gastro 1978). The
degree to which this difference is due entirely
to environmental differences as opposed to eco-
t\pic differentiation does not appear to have
been studied.
Water Relations
Ai:)APTATION TO LIMITED W.ATER. — Stoma-
tal pores provide the pathvx'av by which atmo-
spheric COo diffuses into the leaf replacing the
CO2 incorporated into sugar molecules during
photosynthesis. Because water vapor is present
at \eiy high concentrations inside the leaf,
opening stomata to capture COo inevitably
results in trauspi rational water loss from the
plant; thus, leaf water content is decreased,
resulting in a decrease in plant water potential
(^). Thus, plant water status, transpiration, and
ac(juisiti()n of water from the soils have a tre-
mendous impact on photosynthetic rates and
overall plant grovxth.
1992]
Plant AnAPTYnox
203
Main soils in the (Ticat Basin arc liiu^ t(^\-
tured, which has botli atKantagcs and disadxan-
tages for plant growth. Infiltration of water is
slower in fine-textured soils, increasing the like-
lihood of runoff and reduced spring recharge,
especialK' on steeper slopes. They are also more
prone to water-logging and anoxic c-onditions.
The deep root systems of Cireat Basin sluMihs are
ver\' sensitive to anoxia, and this can be a strong
determining factor in species distributions
(Limt et al. 1973, CiroeneN'eld and Crowley
1 9S8). Unnsualh' wet \ears ma\' e\en cause root
dieback, especially at depth. Once water enters
the soil profile, the extremely high surface areas
of fine-textured soils with high clav and silt
content gi\e them a much higher water-holding
capacit\' than that foimd in sandy, coarse-tex-
tured soils. Much of this water is tighth' bound
to the enormous surface area of the small
particles, howe\er, and is released onl\ at \en'
negatixe matric potentials. Plants nuist be able
to tolerate low tissue water potentials to make
use of it.
As soil water is depleted during sunuuer,
plants reduce water consumption b\ closing sto-
mata (DePuit and Caldwell 1975, CambeJl and
Harris 1977, Caldwell 1985, Miller 1988) and
reducing total canop\' leaf area to a minimum
(Bran.son et al. 1976). Evergreen species shed
only a portion of the total canop\, however,
maintaining the youngest leaf cohorts through-
out the drought (Miller and Schulz 1987).
Although plnsiological actixit)' is greatK'
reduced b\' water stress, exergreens such as
sagebnish can still have positive photosviithetic
rates at leaf water potentials as low as —50 bars
(Exans 1990) and may surxive even greater
](nels of stress. In contrast, crop plants can
rareK" sunixe prolonged M^ of less than - 15 bars.
Remaining functional at loxx' xx'ater potentials
requires the concentration of solutes in the cell
sap, and this appears to be accomplished b\
several mechanisms. In manx mesic species,
increases in organic solutes may account for
most of the change in osmotic potential. In other
species, and especialK' tho.se that experience
xeiy loxv leaf xvater potentials, a large fraction of
the solutes is acquired by the uptake of inor-
ganic ions such as K+ (Wvii fones and (^orhani
1986). High concentrations of inorganic ions
may l)e toxic to enzx'me metabolism, hoxxexer.
and they are xxidely thought to be se(juestered
largely in the central vacuole, xvhich accoimts
for 90% of the total cell xolume. exen thoush
much of the exick^ice for this is (|uite indirect.
Nonetheless, the osmotic potential of the cxto-
plasm irnist also be balanced or suffer dehxdra-
tion. The cytoplasmic .solutes must haxe the
special propeitx of lowering the osmotic poten-
tial of the cell sap xxathout dismpting enz\nne
function or cellular metabolism, and are hence
termed "compatible" solutes (W'xii Jones and
Gorham 1986). The use of specific molecules
shows considerable^ xariation across species
apparentlx' due to both species-specific xaria-
tions in cell metabolism and taxonomic relation-
ships. Some frecjuentlx encountered molecules
thought to function in this manner include
amino acids such as proline and also special
sugar-alcohols. Soiue plants appear to generate
low osmotic potentials bx' actixeK" manufactur-
ing larger quantities of dissolxed organic mole-
cules per cell in response to water st^^ss. a
process referred to as "osmotic adjustment."'
This process ma\' be costh; hoxx'exer, recjuiring
the inxestment of large amovmts of materials
(nexv solutes) at a time xx'hen xx'ater stress is
largely inhibiting photosvnthetic activitv'. An
alternatixe method seems to inxolve dramatic
changes in cell xx'ater xolume. Sexeral desert
species haxe been obserx'ed to reduce cell xx'ater
xolume bx' as much as 80% xx'ithoutxxiltingxx'hen
subjected to extremelx' loxx' soil xxater potentials
(Moore et al. 1972, Meinzer et al. 1988, Evans
et al. 1991). This alloxx'ed the leaf cells to have
sufficiently loxv osmotic potentials for xx'ater
uptake exen though solute content })er cell xx'as
actually reduced. Although total solutes per leaf
(and presumablx per cell) decreased, the rela-
tix'e concentrations of specific solutes changed
(Evans et al. 1991) such that "compatible"
solutes made larger contributions to the osmotic
potential untk'r stress. Thus, tolerance of loxv
leaf xxater potentials was achieved bv a combi-
nation of anatonncal and phxsiological special-
izations. The anatomical mechanisms inxolxed
in this magnitude of xolume reduction and the
im]ilied cell elasticitx' haxe not been studied, but
tlie process has been shown to be fnllx rexcrsible.
Soil texture^ is also an important factor in
determining the abilitx' of plant connnunities in
a coId-x\int('r climate to respond to summer
rain. In areas xxith moderate lexels of precipita-
tion, sandx' soils, because of their loxx- xxater-
holding capacitx. nsuallx' hold a sparser, more
drought-adapted x (^getation than finer-textured
ones. In xvarm, arid areas, however, what has
been called the "rexerse texture" effect results
204
Great Basin Naturalist
[\'oliime 52
ill the more liisli xegetation oceiirrintj; in tlie
coarse-textured soils. This occurs because the
high water-holding capacit)' of fine-textured
soils allows them to hold all the moisture
deri\'ed from a single rainfall event in the upper-
most layers of the soil profile, where it is liigliK
subject to direct e\aporation from the soil. The
same amount of rainfall entering a sandv soil,
precisely because of its low \\'ater-h()lding
capacity', will penetrate to a much greater depth.
It is also less likeK' to return to the dning surface
b\' capillaiv action. Less of the rain will exapo-
rate from the soil surface, and a greater fraction
of it will await extraction and use by plants. This
inverse-textiu-e effect further restricts the effec-
tiveness of summer rains in the fine soils of the
Great Basin. The effect is less operative in
respect to winter precipitation in the Great
Basin, however, because of the gradual recharge
of the soil profile to considerable depth under
conditions where surface e\aporation is mini-
mized by cold temperatures. The combination
of much sandier soils and greater amounts of
summer rainfall in the region of the Colorado
Plateau is largely responsible for the major flo-
ristic and ecological differences bet\\'een the
two regions. At lower elevations on the south-
east edge of the plateau, shiid^-dominated
desert scnib mav be replaced by grassland dom-
inated by a mix of spring-active C5 and summer-
active C4 species.
ROOTINC; DEPTH, MORPHOLOGY, AND PHE-
NOLOGY. — One of the unique and ecologicallv
most important features of the Great Basin
shmb communities is not apparent to abo\e-
ground obseners. This is the investment of the
vast inajorit\- of plant resources in the growth,
maintenance, and tunioxer of an enormous root
system. The dominant slinibs of the Great Basin
usually root to the full depth of the winter-spring
soil moisture recharge. Depending on soil tex-
ture, slope aspect, and elevation, this is gener-
ally between 1.0 and 3.0 m (Dobrowolski et al.
1990). Although this represents a wide range of
absolute ck^pths, nianv of the ([ualitatixe pat-
terns of rooting behaxior are similar on most of
these sites. Ratios of rootishoot standing bio-
mass iang(^ from 4 to 7, and estimates of
root:shoot annual carbon inxe.stment are as high
as 3.5. Most of the shrubs ha\e a flexible, gen-
eralized root system with dexelopment of both
deep taproots and laterals near the surface.
Moreover, it is the categon of fine roots < 3.0
mm in diameter that constitutes 50-95% (Cald-
well et al. 1977, Sturges 1977) of the total root
biomass. The veiy large annual root inxest-
ments, therefore, are not primariK- related to
large storage compartments, but to the tunioxer
of fine roots and root respiration necessan- for
the acquisition of water and mineral nutrients.
The fine root network thoroughK' permeates
the soil x'olume. Root densities are grenerallv
quite high throughout the upper 0.5 meters of
the profile, but depth of maximum root devel-
opment \aries with depth of spring soil-mois-
ture recharge, species, and lateral distance from
the trunk or crowai. A particularly high densit)'
in the first 0.1 m ma\' often occur, especially
immediateh under the shmb canopx (Branson
1976, Caldwell et al. 1977, Dobrowolski et al.
1990). AlternatixeK', maximal densit) mav occur
at depths between 0.2 m and 0.5 m (Sturges
1980), and sometimes a second zone of high root
densit}' is reported at depths of 0.8 m
(Daubenmire 1975) to 1.5 ni (Reynolds and
Fralev 1989). Regardless of the precise depth of
maximum rooting, sliRib root densit\' is usualK'
high throughout the upper 0.5 m and then
tapers off, but max continue at reduced densi-
ties to much greater depth. The radius of lateral
spread is usuallx' much greater for roots ( 1-2 m)
than for canopies (0.1-0.5 m). Percent plant
coxer abox'eground is often in the neighborhood
of 25% xxdth 75% bare ground, but beloxvground
the interspaces are filled xvith roots throughout
the profile, and root sxstems of adjacent plants
xxdll overlap. The perennial grasses that are
potentiallv co-dominant xxith shnibs in manx of
these communities, such as xxheatgrass
{A^ropi/roii sp.), xx'iklne (Eh/nui.s sp.),
squirreltail {Sitaiiioii liisti-ix). Indian ricegrass
(On/zopsis lu/i)icii()i(h:s). and galleta grass
{Hilaiia iainesii), generallx haxe somexxhat shal-
loxxer root .sxstems than the shrubs (Branson et
al. 1976, Rexiiolds and Fralex- 1989, Dobro-
xvolski et al. 1990). Root densities of grasses are
often as high as or higher than those of shrubs
in the upper 0.5 m but taper off more rapidlx
such that shnibs usuallx haxe greater densitx at
depth and greater maximum penetratit)n.
The moisture resource supporting the great-
est amount of plant groxx'th is usuallx- the xx'ater
ston^l in the soil profile during the xxinter. This
j)r()(ile usuallx has a positixe balance, xxith more
XX ater being added bx precipitation than is xxith-
draxxn bx' exapotranspiration bet\xeen October
and March. As temperatures xx-arm in March,
exergreen .species nia\' begin draxxing on this
19921
Plant Ai:)\rT\TK)\
205
iiioistiiiT resent", ami most species l)eii;iii aetixc
growth ill March or ApriL Due to both plant
water use and soil-surface exaporation, soil
moisture is depleted first in the shallow soil
hncM's. As the upper layers dr>', plants withdraw
moisture from successively deeper soil hners, a
proc(^ss that continues in e\ergreen species
throughout the summer until soil moisture is
depleted throughout the profile. This progres-
sion of seasonal water use beginning in superfi-
cial la\'ers and proceeding to deeper soil layers
is facilitated In the pattern of fine root growtli,
w liicli shows a similar spatial and temporal pat-
tern (Fenuindez and (Caldwell 1975, C'aldwell
1976). Root growth generalK precedes shoot
growth in the earl\- spring and continues
throughout the summer in e\ergreen species,
which mav appear quiescent abo\egroiind. In
annual budgets of undisturbed communities,
.soil moisture withdrawal almost alwaxs
approaches measured precipitation o\ er a wide
range of annual fluctuations in total precipita-
tion, and yew little moisture is lost to runoff or
deep drainage beneath the rooting zone
(Daubenmire 1975, Caldwell et al. 1977,
('ainbell and Harris 1977, Sturges 1977). Calcu-
lati( )ns of the portion of exapotranspiration actu-
alK' used b\" plants in transpiration are quite high
for a desert enxironment with low percent
co\er; they range from 50 to 75% (Caldwell et
al. 1977, Cambell and Harris 1977, Sturges 1977).
Competition for soil moisture is not neces-
saril\- limited to any single season. Plant water
stress is highest in late sunuuer, and siir\i\al is
most likeK to be influenced at this time, espe-
cialK if one plant can deplete residual soil mois-
ture below the tolerance range of another. This
has been sliown in sexeral cases with regard to
seedling establishment (Harris 1977, DeLucia
and Schlesinger 1990, Reichenberger and Pvke
1990). Growth and productivits" are more likel\-
to be affected in the spring and earl\ summer
growing season. This is because most of the
years water resource is alread\- stored in the soil
in earK spring, and all plants are drawing on a
dwindling resene which ultimateK determines
growing season length. Competitixe abilits' is
often found to be associated with an abilit\ to
begin using the limiting water resource earlier
in the spring (Eissenstat and Caldwell 19<SS,
Miller 1988), but spatial partitioning is also
important. Competition ma\ be most intense in
shallower soil la\ers because grasses and
drought-deciduous shrubs, wiiicli are actixe
oiiK ill the spring, are shallower rooted, and
lateral root spread of tlu^ e\ergreeii species is
greatest in the shallower soil la\ers. The more
dominant perennials usualK use more water
o\er the whole season 1)\- tapping deeper soil
la\ers ((>line et al. 1977, Sturges 1980) and are
characterized b\ higher water-use efficiencies
(DeLucia and Sclilesinger 1990, Smedlev et al.
1991).
Shnibs appear to be better than grasses at
withdrawing water from deep soil laxers for
several reasons. In shallow soils underlain by
fractured or porous bedrock, the flexible, mul-
tiple taproot structure of a shrub root sxstem
ma\" be better suited than the more diffuse,
fibrous root system of grasses for following
cliinks and clea\age planes to indeterminate
depths. This should allow shnibs to better cap-
italize on deep, localized pockets of moisture.
Unfortunatelv such d\iiamics are rareK studied
quantitatixeK because of the difficult\" of mea-
suring soil moisture budgets or root distribu-
tions under such conditions, but the\' are
implicated b\' plant distribution patterns in
man\ areas. Indeed, despite the \isiial austeiit\'
of such habitats, rooting into major joints and
cracks in bedrock outcrops can create sucli a
fa\"orable microsite b\' concentration of ninoff
in localized areas that ele\ational limits of tree
and shrub distributions may be substantiallv
lowered as would be expected along riparian
corridors or other unusnalK' niesic liabitats
(Loope 1977). Even in deep soils, shrubs tend
to ha\e deeper root svstems than grasses, but, in
addition to this, shiTibs may be more efficient
than grasses at extracting deep water. Shiiibs are
sometimes able to draw on deep soil moisture
to a greater extent than would be predicted from
root biomass distribution alone (Sturges 1980),
and this is due in part to an intriguing phenom-
enon reported b\- Richards and Caldwell ( 1 987),
and named b\- them "Indraulic lift." During the
Iat(^ spring and earK summer most ol the
remaining soil moisture is present in ckn-per soil
layers wheic rooting (lensit\ ma\ be relati\eK'
low. l^ue to low (k'usities, deep roots alone ma\'
be unable to absorb water as (juickl\- as it is lost
l)\ the tiaiisi)i ling shoot. During the night, water
is actnalK ic^distributed within the soil, flowing
from deep soil lavers through the roots and back
out into shallower soil laxcrs. This is the phe-
nomenon named â– indraulic lift." and the
adxantage to the plant is thought to be a reduc-
tion in the rootiii'i densitN necessar\' to fully
206
Great Basin Naturalist
[N'olume 52
exploit tlie resources of the deepest soil lavers.
During the dav, rates of water uioxeiuent from
the soil into the roots can be limiting to shoot
activit)'. Moistening the upper soil lavers bv noc-
turnal h\draulic lift increases the root surface
area for al^soiption during the periods of high
transpiration. Davtinie water use can he sup-
ported by the entire root system and not just b\'
the low-densitv deep roots (Caldwell and Ricli-
ards f989).
The root s\'stems of Great Basin shrubs and
Mojave Desert shrubs differ strongly in several
ways. (1) Mojave Desert shiiilis often have max-
imal rooting densities at soil depths of 0.1-0.3
m, and maximmn rf)ot penetration of 0.4-0.5 m
(Wallace et al. 1980). These shallower roots are
due to lower rainfall and warmer winter temper-
atures resulting in shallower wetting fronts in
the soil, and the de\ elopment of shallow caliche
layers. (2) Great Basin species have more roots
in the shallowest 0.1 m soil laver (Wallace et al.
1980, Dobrowolski et al. 1990). Differences in
soil temperatures ha\"e been used to explain
these patterns, but the link betvveen cause and
effect is less ob\ious in this case, and conjec-
tures should be \iewed cautiouslv. Much hotter
soil temperatures in tlie Moja\e may be detri-
mental to roots in the uppermost few centime-
ters during summer, and the rapidly di"ving soil
surface may be too ephemeral a moisture
resoiu'ce to favor large investments in roots. In
contrast, snowmelt and cooler spring tempera-
tures may result in less rapid evaporation from
the soil surface in the Great Basin and thus fax or
more rooting l^v perennials in that zone. (3)
Because of the greater soil volume exploited, as
well as the high root densitv of Great Basin
species, their ratios of rootishoot biomass are
al)Out twice that of Moja\e Desert species
( Bamberg etal. f 980, Dobrowolski et al. 1990).
Adaptation to salinity. — When annual
precipitation levels are much lower than poten-
tial evaporation, salts are not leached to an\
great depth, and they can accumulate within the
root zone. This is especialK important in associ-
ation with particular bedrock outcnps, such as
the Nhuicos and Ghinle shales, which release
high concentrations of salts during weathering
(Potter et al. 1985). Precipitation increases with
elexation, and. lollowing major storms or spring
snowmelt, there may be surface runoff and
recharge of groundwater sy.stems that trans[)ort
water from high elexations into closed basins.
Streams in the Great Basin usualK terminate in
evaporati\e pans where salinities reach extreme
le\els and salts precipitate forming a surface
crust. The water table near these evaporative
pans is often at or ven near the sin-face through-
out the \'ear, l)ut, if there is no groundwater flow
out of the basin, it too is often quite saline
(Dobrowolski et al. 1990). Salinities are lowest
on slopes and at higher elevations where precip-
itation exceeds evaporative loss, and they
increase on more level terrain and in lower-ele-
\ation basins where exaporation exceeds pre-
cipitation. Sahnities may also be higher in areas
where wind-borne materials are transported
from saline playas to surrounding slopes (Young
and E\ans f 986). These patterns of soil salinitx'
are important in determining plant distribu-
tions, with more specialized salt-tolerant spe-
cies (halophvtes) replacing less-tolerant species
repeatedh along gradients of increasing salinit)'.
In general, species diversity is low on saline
soils. The vast majorit)' of tolerant shrub species
in our deserts, and all the shrubs specifically
mentioned in this section, lielong to a single
plant family, the Chenopodiaceae (goosefoot
famiK). Most other important taxa in the saline
connmmities are grasses.
In the most extreme case of h\persaline salt
flats and pans there may be standing water in
the wet season with saturating salt concentra-
tions. Under such conditions, only microflora
consisting of a few species of photosMithetic
flagellates, cyanobacteria, and halobacteria are
commonly found. The halobacteria appear to be
unique in having adapted in an obligate manner
to the high salinities of these environments.
Thev not only tolerate, but require, high
cvtoplasmic salinities for membrane stability
and proper enzymatic function (Brown 1982).
In strong contrast to this, algae and all higher
plants growing in hvper-saline environments
show severe inhibition of enzvnne fvmction at
high salinity, and thev must compartmentalize
salt-sensitive metabolic processes in celhdar
regions of low ionic strength ( Muuus et al. 1982).
The best definition of a liahphvte is simply
a [)lant tolerant of soil salinities that would
reduce the gi'owth of unspecialized species. This
is .somewhat circular, and that reflects our lim-
ited understanding of how halophv tes do what
thev do. Halophv tes are more likely to use Na+
in their tissues for osmotic adjustment, while
glvcophvtes are more likely to have high K+
( Ilellebust 1976, Flowers et al. 1977), but there
are munerous exceptions. Other differences
1992]
Plant Ai:)aptatk)n
207
max he nunc (juaiititatixc than (|iialitati\ c \ ar-
ious aspects of mineral nutrition in halophx tes
are less sensitixe to high soil salinities, hut
unique mechanisms to achiexe this tolerance
ha\e rareK' heen identified. It is wideK held that
the ahilitv to compartmentalize salts and restrict
high Na+ concentrations to the \acuole is of
crucial importance (Cakh\'ell 1974, Flowers et
al. 1977, linens and I.arhtM" 1982). This conclu-
sion is hased primariK- on indirect e\idence of
low enz\nne tolerance of salinitv; howexer,
rather than direct measurements of actual salt
compartmentalization (Munns et al. 19S2,
Jefferies and Rudmik I9S4).
Haloplntes differ in which ions reach high
tissue concentrations when all plants are grown
on the same medium (Caldw^ell 1974). Some
will concentrate C1-, for instance, while others
concentrate S04~'. These differences do not
necessarih' determine plant distrilnitions, such
as occurrence in soils dominated h)' NaCl \'ersus
NaSOa, but rather seem to reflect different reg-
ulatoiA' specializations in plant metabolism
(Moore et al. 1972). A strong requirement for a
uni([ue composition of soil salts is the exception
rather than the mle, and the most important
effect of soil salinitv' seems to be a disniption of
plant water relations from low soil osmotic
potentials rather than toxic effects of specific
ions. Halophvtes tolerate these conditions bx'
ha\ing better regulatoiA' control o\er ion mo\e-
ment within the plant, ion compartmentaliza-
tion at both tissue and subcellular lexels, and
better homeostasis of other a.spects of mineral
nutrition in the presence of ver\' high Na-K.
Salinit\ poses three major problems for
higher plants. First, salts in the .soil solution
contribute an osmotic potential depressing the
soil water potential, and this ma\' be aggra\"ated
as salts become concentrated with soil drving.
E\en when sul)stantial moisture is present,
[)lant tissues must endure \ t-n low water poten-
tials to take it up, and this recjuires a specialized
metabolism. Second, an\' salts entering the plant
with the transpiration stream will be left behind
in the leaf intercellular fluids as water ('\a])()-
rates from the leaf. This can result in salt
buildup in the intercellular solution causing
water moxement out of the cells and leading to
cellular dehxdration. Third, salts entering the
cxtoplasm in high concentration will disrupt
enz)ine function. Haloph\1:es are able to deal
with all of these factors over a wide range of soil
1. . .
salinities. Haloph\tes show a greater capacit\'
for osmotic adjustment, and positixe phot()s\n-
thetic rates can be measured in the leaxes of
man\ haloplntes at leaf water potentials as low
as -90 to - 120 bars (Caldwell 1974), well below
the range that would result in death of e\en
desert-adaj)ted gl\coph\tes. Tliis is accom-
plisluHl in part 1)\' transforming the available
salts in the enxironment into a resource and
using them for osmotica in j)lant ti.ssues (Moore
et ak 1972, Bemiert and Schmidt 1984). Many
haloplntes actualK show stimulation of growth
rates at moderate^ en\ ironmental salt levels.
Halophvtes too must deal with the problem
of salt buildup in the leaves, and the\' do so by a
wide \ariet\' of processes. There is a great deal
of interspecific \ ariation in which method! s ) are
used. All the methods appear to incur substan-
tial energetic costs associated with maintaining
high ion concentration gradients across key
membranes (Kramer 1983). Exclusion of salts at
the root is possible; this is the method most
employed by winterfat (Ceratoides Janata). Salt-
bush (Atriplex spp.) has specialized hair-blad-
ders on the leaf surface into which e.xcess salts
are actively pumped. The hairs e\'entualK' nip-
ture, excreting the salts to the outside. Other
plants may transport salts back to the root \ia
the phloem. Man\- plants exhil)it increased leaf
succulence when growii under high salinit\; and
this increase in cell xolume can create a sink for
salts within the leaf without raising salt concen-
trations or furtlier lowering leaf osmotic potential.
hi strong contrast to the exident importance
of temperature and rainfall pattern in favoring
C:5 versus C4 grasses, Ci shnibs tend to belong
to edaphic comnumities as.sociated with saline
soils. The same species ma\' occur in both warm
and cold deserts, and in areas with both winter
and summer rainfall patterns. This is an intri-
guing difference, but the phwsiological basis
linking C, shrubs with high salinitv' is less well
understood than the tradeoffs associated with
temperature and controlling C5 and C^ grass
distributions. Sjx'cies number and percent
cover b\ shrubs sucli as saltbush {Ahiplcx spp.)
and inkA\'eed (Siicda spp. ), wliich possess the C4
pathwav, usualK inc-rc^ise drainaticallv with
increasing salinitv on w(41-drained soils. In
marshx' habitats or soils with a shallow, saline
water table, howex er. haloplntic (>-, species such
as pickleweeds {Allen rolfia spp. and Saliconiia
spp.) and greasewood [Sarcohatus ver-
micnloicles) regain dominance. It has been sug-
gested that hitrher water-use efficiencv bv C4
208
Great Basin Naturalist
[\ nluiiie 52
species niav be acKantageous on saline soils to
help avoid salt bnildnp in the leaf tissues. How-
ever, it has not been showii that transpiration
rate is an important factor controlling salt
buildup in the leaves of halopln tes when com-
pared wath other regulaton' mechanisms
(Osmond et al. 1982), nor does this Inpothesis
explain the dominance of C3 species in wet
saline soils. In the greasewood and pickleweed
commimities, soil salinities are extreme, but
soils remain wet throughout the growing season,
or else groundwater is available within the root-
ing zone (Detling 1969, Hesla 1984). As a con-
sequence, plant water potentials do not reach
the extreme low values of the saltbush commu-
nities. Over a wide range of soil salinities, plants
such as greasewood appear to draw on readily
available deep soil moisture, and high leaf con-
ductances are maintained throughout the
summer (Hesla 1984, Romo and Hafercamp
1989). The highest whole-plant water-use rates
may occur in the presence of high soil salinitv" in
mid-summer (Hesla 1984). The communities in
which C4 shRil:)s are most prevalent have
summer stress from both high soil salinitv and
mid-sunnner soil water depletion combined.
These species reach much lower plant water
potentials during summer than either nonsaline
communities or wet-saline ccnnmunities. The
role of C4 pliotosviithesis in tolerating these
conditions remains to be determined, but it
could he related to avoiding excessively low leaf
water potentials either liy (1) retarding soil
moisture depletion, which both lowers the soil
matrix potential and concentrates soil salts, or
(2) avoiding exacerbation of low soil water
potentials due to high flux rates and large water
potential gradients between the leaf and root.
Water mo\ement in the x-xlem occurs under
tension, and anatomical features that avoid cav-
itation in the xylem at extreme]\ low water
potentials usually reduce the hydraulic conduc-
tivity of the x"\'lem per unit cross-sectional area
(Davis et al. 1990, Speny and Tyree 1990). Low
specific c()nducti\it\' of the xTlem will, in turn,
predispose the plant system to large water
potential gradients between roots and shoots,
causing an even greater depression of leaf water
potential. This problem could be ameliorated
either by increased cross-sectional area of the
xylem by increased allocation to wood growth,
or by features such as C.| photos\Tithesis that
reduce the flux rate of water associated with
photosN nthetic acti\it> under warm conditions.
Nutrient Relations
Acquisition of mineral nutrients. —
Apart from the veiy high elevation montane
zones, water appears to be the most limiting
resource in the Great Basin and Colorado Pla-
teau communities. Productixit) is usualK well
correlated with yearlv fluctuations in precipita-
tion and spring moisture recharge over a wide
range of \alues (Daubenmire 1975, Kindschy
1982), and competitive success has more often
been associated with soil water use patterns
than nutrient budgets. Nonetheless, addition of
mineral fertilizer sometimes does result in
modest increases in producti\it\', and studies
ha\e shown strong effects of neighboring plants
on nutrient uptake rates (Caldwell et al. 1987).
These dynamics have been less studied than
have plant water budgets, and broad ecological
relationships are just now being worked out in
detail. Nutrient acquisition has been showni to
be a major factor determining communits' com-
position only in veiy special habitats such as
large sand dunes (Bowers 1982) or unusual bed-
rock (DeLucia and Schlesinger 1990).
MiCROPHYTIC CRUSTS. — Throughout the
Great Basin and Colorado Plateau, it is common
for the exposed soil surface to l)e covered by a
complex connniuiit\' of nonvascular plants
including dozens of species of algae, lichens,
and mosses (West 199()). These organisms often
form a biotic ciTist in the upper few centimeters
of the soil and, when undisturbed, may result in
a vei"y conx'oluted microtopograplu' of the sur-
face. While a detailed discussion of the
microplutic crusts is bcNond the scope of this
review, it is important to realize that percent
cover by such crusts often exceeds that of the
vascular plants, and their contribution to total
ecosvstem prochicti\itA' is consitlerable. Perhaps
most important to co-occurring \ascular plants
are the nutrient inputs to the soil b\' nitrogen-
fixing cnist organisms (c\anobacteria and
lichens). These inputs will be particularlv
important in the cold deseit where fewxascular
plants form sMiibiotic relationships with nitro-
gen-fixing bacteria.
Nurse plants and fertile islands. — In
man\ des(Mt areas, including both the Mojave
and the Great Basin, establishment of newindi-
\iduals may occur preferentialK' under the exist-
ing canopies of alreadv established indi\iduals.
Tliese pre\iousl\' established indixidnals mav
tlieu be referred to as nurse plants. Preferential
1992]
Pl.WT Ai:)MT\TION
209
estahlisliiiK'nt inulcr iiiirsc plants nia\ ocfiir in
spite of the fact that 759ic or more of tiie gromid
area nia\' he liare interspaces b(^t\\'een plant
canopies. The phenomenon can he important in
both steadx-state commnnitA cl\ namics and also
snccessional patterns following distnrbance
(Wallace and Ronme\- 1980, Exerett and Ward
1984). Two .somewhat distinct factors contiibntc^
to the nnrse-plant phenomenon. The first has to
do the with beneficial effects of partial shading
and rednced wind nnder existing canopies
resulting in cooler temperatnres and possibK'
moister soil conditions in the snrface huers.
These interactions depend directk- on the pres-
ence of the nnrse plant in creating a fa\orable
microsite, and ha\e been studied with particular
reference to pin\on and juniper establishment
in the Great Ba.sin. A second factor inxoKes the
creation of fertile islands bv the prolonged occu-
pation of the same microsite b\' man\' genera-
tions of plants; this can make the fertile island a
preferred site even if the previous occupant is
alreacK deceased. This microsite impro\'ement
occurs due to preferential litter accumulation
and more e.xtensixe root growth directK under
a plant canopw and deposition of aeolian mate-
rials under reduced wind speeds in plant cano-
pies. In time, soils nnder existing plants mav
come to be slightK' raised above the interspace
level, have a lighter, loaniier texture, higher
organic matter content and better soil structure,
less surface compaction, better aeration and
more rapid water infiltration, and/or higher
l('\els of available mineral nutrients than
immediatelv adjacent interspace soil (A'est 1962,
Wood et ai 1978, Homnev et al. 1980, Hesla
1984, West 1989, Dobrowolski et al. 1990).
Direct effects of nurse plants and indirect
effects of fertile islands should complement and
reinforce each other in maintaining selective
spacial patterns of seedling establishment. Sur-
face soil nnder haloplntes mav also show-
increased salinitv (Richard and (]line 1965) due
to excretion ol excess salts bv the canopv or
translocation and re-excretion Ironi the roots.
DiXER.SITY OP^ Ghowtii Foinis
One of the striking features of the cold desert
vegetation is the uniformlv low stature of the
vegetation. This is undoubtedlv due to several
factors, and few studies have specificallv
addressed the role of plant stature in these com-
munities. Since low temperatures mav limit
photosx nllicsis in tlic cool spiiirj;. and earlier
growth on limited soil moisture resen(\s mav be
c-()mp(titi\c'l\ advantageous, occupving warm
microsites mav be favored. Substantial increases
in air temperature and reductions in wind speed
will exist in the lowest meter next to the ground,
and especiallv in the lowest decimeter. Low
cushion plants oi- low. dense shrub canopies
should have vvarmei" spring leaf temperatures by
virtue of being short and bv virtue of leafing out
first in a dense clump of old dead leaves and
twigs ( Smith et al. 1983, Wilson et al. 1987). This
advantage mav be partiallv offset by overlv high
temperatures in summer for species remaining
active all sununer. Stature is also likelv to affect
aeolian deposit of materials under the shrub
canopies (W^ood et al. 1978, Young and Evans
1986), snow accumulation (Branson et al. 1981,
West and Caldwell 1983), and the likelihood of
winter desiccation under cold, windv conditions
(Nelson and Tienian 1983). All of these could
be important factors, but few detailed studies
have been done.
Having considered tlie relationships of the
dominant plant habits and phenologies to cli-
mate, it is perhaps instructive to consider whv
.some of the other famous desert life forms are
so poorlv represented in this region. Three life
forms vvliich are prominent features of the warm
desert but inconspicuous elements of the cold
desert are (1 ) large CAM succulents (e.g., cacti
and agave), (2) opportunistic drought-decidu-
ous shnibs specialized for rapid knif-flnshing,
and (3) animals. Definitive work explaining the
structural nnilormitv of the vegetation is not
available, but the environment is well enough
understood to identifv at least some of tlu^ likelv
causes.
CAMSrcci'I.KXTS — Most of the large C.\.\I
succulents are not tolciant ol freezing temper-
atures, and most extant species would be
excluded from the (jrcat Basin bv this factor
alone. Thei'e ai"e, however, a sulfitienl mimbcr
of species which have adapted to tolerate cold
temperatures that we are justified in asking whv
thev have not undergone more adaptive radia-
tion, or claimed a more prominent role in these
communities. The most important factor limit-
ing this life form is probably the importance of
the cool spring growing season. CAM succu-
lents generallv ( 1 ) allocate ven little biomass to
root (root/shoot ca. 0.1), (2) are shallow rooted,
(3) store moderate-sized (compared to soil
v\ater-liolding capacitv ) water resenes inside
210
Great Basin Naturalist
[Volume 52
their tissues wheu water is available in the sur-
face soil layers, and (4) use their stored water in
photosynthesis with unparalleled water-use effi-
ciency by opening their stomata only at night
when temperatures are cool (Nobel 1988). They
are fa\ored bv (1) very warm days (30-40 C),
which allow them to have higher photosyiithetic
rates and cause competing species to ha\-e very
low water-use efficiencies; (2) large diunial tem-
perature fluctuations allowing for cool nights
(10-20 C) which allow them to have high rates
of CO2 uptake with high water-use efficiency;
and (3) intermittent rainfall \\'hich onl\' wets the
upper soil layers so that the limitations of their
shallow roots and water-hoarding strategy are
compensated foib\ the ephemeral natiu-e of the
soil water resoiu'ce. These conditions are some-
what poorK" met in the cold desert. The impor-
tant water resource is one of deep soil recharge
that favors deep-rooted species and confers
much less advantage on internal water hoarding.
Freezing tolerance in CAM succulents appears
to be associated with low tissue water contents,
and this mav inhibit uptake of water when it is
plentiful in the siu-face layers in the thermalK'
x'acillating eark' spring (Littlejohn and Williams
1983). Furthermore, water-use efficiencies of
C3 and C4 species are quite high in the cool
spring.
Nonetheless, even moderate amounts of
summer rain in the southern and eastern por-
tions of the Great Basin result in numerous
species of cacti. Due to the open nature of the
understoiy, many of these species ha\e a large
elevational range, and they are often more
common in the pinvon-juniper or even the mon-
tane zone than on the desert piedmont slopes.
Almost all of these cacti are small, usually 5-20
cm liidi, with a small, globose (e.g., Pediocactiis
siinpsonU), prostrate (e.g., Opiintia pohj-
cantha), or low, caespitose habit (e.g.,
Echinoccreus tn<4ochidi(itus). This allows them
to take acKantage of the warmer da\time tem-
peratures near the ground in the sj^ring and
facilitates an insulating snowcover during the
coldest winter periods. The number of and total
cover by cacti increase considerabK with
increased summer rainfall on the Colorack) Pla-
teau, but oulv in the eastern Mojave with both
summer rain and warm spring tempcratun^s do
we find the larger barrel-cactus (e.g., Fcrocddiis
acanfhoidcs) and tall, shnibb)^ chollas (e.g.,
Opinilid (ic(i)ithiH'arpa).
Opportunistic drought-deciduous /
MULTIPLE LEAF-FLUSHING SPECIES. — This
habit, like that of the succulents, is favored by
( 1 ) intermittent rainfall wetting only shallower
soil layers, and (2) warm temperatures allowing
for rapid leaf expansion in response to renew/ed
soil moisture. Again, these requirements are not
well met in the Great Basin. The priman' mois-
ture resource is a single, deep recharge in the
winter. Most shiaib species are deep rooted, and
rather than experiencing \acillating water avail-
abiHtv', they have actixe root grow1:h shifting to
deeper and deeper soil kners during the season,
thus producing a gradual and continuous
change in plant water status. This allows manv
spring-active shrubs to remain partially ever-
green throughout the summer, and, in regions
where it occurs, the\' are able to make rapid use
of anv moisture availalole from simimer precip-
itation without the need for renewed leaf pro-
duction. The only shrub reported to ha\'e
)iiultiple leaf flushes in response to late spring
or summer rain in the Great Basin is the dimin-
utive and shallow^- rooted Artemisia spinescens
(Everett et ak, 1980). Some species found in the
Great Basin are reported to have multiple
growth c\'cles/year where they occur in the
Mojave (Ackerman et ak 1980).
ANNUALS AND LIFE-HISTORY DIVERSITY. —
The spectacular wildflower show^s displayed in
favorable years in the Mojave Desert do not
occm- in the cold desert of the Great Basin
(Ludwig et ak 1988). Annu;il species are few in
nimiber, and, except in earK" succession after
fire in woodlands or on \en disturbed sites, the)'
rarely constitute a major fraction of total com-
munit>' biomass. This is undoubtedly related to
sexeral complex factors, but various aspects of
precipitation patterns are likeK' to be among the
most important. To begin with, the paucits' of
summer rain in some parts of the Great Basin
ma\ largeh' eliminate an entire class of C4
siuumer annuals important in the floras of other
regions including the Cok)rado Plateau. Other
aspects than seasonalih' are also cnicial, how-
e\er. Ver\ low means oi annual precipitation are
conunonh' associated with large annual floras,
but correlated with low mean precipitation is
high \ear-to-\ ear \ariation in precipitation
which some authors have argued is equally
important. The coefficient of xariation (CV) in
precij)itation shows a r(4ationship to mean pre-
cipitation in the (wvat Basin and Colorado Pla-
teau (Fig. 2) veiv similar to that found in warm
19921
Plant Ad AinviioN
211
0.7
o
0.6
t â–
m
';z
O.S
ro
>
B
0.4
, â–
r
CD
0.3
o
â– ,
CI)
0.2
o
O
0.1
, 1 - T 1
3
r2 = .57
-
O
-
\oO cj^
s °
o
^
o o°o ^ — ~--
o
o
-
"
0.0
100 200 300 400 500
Mean annual precipitation, mm
Fig. 2. The relatioii.ship hctween mean precipitation and
the \ariabilit\' of rainfall between years as nie;isured h\- the
coefficient oi \ariation in annual precipitation. The data
inchide points scattered throughout the Great Basin in Utali
and Xe\ada and the Colorado Plateau in Utah and Arizona.
The line shown is the least squares best fit for the data: C\'
= 1.27 - 0.403 ° log(niean annual precipitation, mm) {ii =
69 sites./) < .001).
deserts (Ehleringer 1985). Although mean pre-
cipitation has tlie greatest single effect, there
are. aclclitionalK; important geographic influ-
ences on the CV of precipitation which are
independent of mean precipitation. A multiple
regression of the CV of precipitation on
logdnean annual precipitation), latitude, and
elexation in the Great Basin has an r of .81 and
indicates that each \arial)le in tlie model is
highl\- significant (j) < .001 or better). For a
given mean precipitation, the C\^ increases with
decreasing altitude in the Great Basin, but an
ind(^p(Mident effect of elevation was not signifi-
cant in the Colorado Plateau. The CV also
increases from north to south in the Great Basin
and increases from south to north in the CJolo-
rado Plateau, which results in a latitudinal band
of greatest annual \arial)ility nuniing through
southern Nevada and Utah. This l)and is related
to two major as]:)ects of regional climate. Mo\ing
southward in the Great Basin, temperatures
gradually increase, favoring moister air masses
and more intense storms, but sites are morc^
remoxed from the most common winter storm
tracks, and the number of rainv daws per Near
decreases (Houghton 1969). Moving northward
from Arizona and New Mexico, the southern
Nevada and Utah band of highest precipitation
variabilit\- also corresponds to the northenunost
extent of summer storms associated with tlie
2 3 4
CV for mean annual
precipitation
F"ig. 3. The relationship between relial)ilit\- of annual
precipitation and life-histon' strategy of herbaceous plants.
The site with greatest representation of annuals is Deadi
\'allev in the Moja\e De.sert, the second highest is C^auNon-
lands in the Colorado Plateau of southeastern Utah, and the
other three sites aie Great Basin Cold Desert or shrub-
steppe (data were collected b\' Kim Haiper and pnniously
published in Schaffer and Gadgil 197.5).
Arizona monsoon, and the region where the
fraction of summer rain increases substantially
moving southward. This zone also has some of
the most arid sites of the entire region located
along the transition to the Mojave Desert in
southern Nevada and the canyon countiy of
southeastern Utah, and these sites can be
expected to have the highest variabilit\ due to
both low mean rainfall and geographic position
correlated with rc^gional weather patterns.
Becau.se the (ireat Basin and Colorado Plateau
are only .semiarid, the CV ol annual precipita-
tion is not usually as high as in manv of the more
arid warm deserts (Beatley 1975, Ehleringer
1985), but particular sites may be both arid and
highlv unpn^dictable.
Ilaiper (cited in Schaffer and Ciadgil 1975)
found that the prevalence of annuals was posi-
ti\eK associated with the C\' in annual precipi-
tation for five sites located in the Great Basin,
Colorado Plateau, and Moja\e Desert (Fig. 3).
The largest annual populations occurred in
Death Valley (Mojave), followed by Canyon-
lands (Colorado Plateau in southeastern Utah).
One inteipretation of this relationship is that
high \ariabilit\- in total precipitation between
vears mav be associated v\ ith high rates of mor-
tality and therefore favor earlv reproduction and
an annual habit (Schaffer and Gadgil 1975).
Manv desert annuals are facultativ elv perennial
in better-than-averaee vears, and some have
212
Great Basin Naturalist
[Volume 52
perennial races or sister species (Ehleringer
1985). The dynamics and distributions of these
closely related annual and perennial taxa should
receive further study in regard to their expected
life span, reproductive output, and relationships
to climatic predictability'. Another perspecti\'e is
to ask how competition between very distinct
shnib and annual species is affected by precip-
itation variability. While in many respects com-
plementaiy with the optimal life histoiy
arguments, this approach emphasizes how large
differences in habit affect resource capture and
competition rather than focusing on subtler dif-
ferences in mortalit)' and reproductive sched-
ules. The lower variability of precipitation in
much of the Great Basin compared to the
Mojave and Sonoran deserts, as well as the more
reliable accumulation of moisture during the
winter-recharge season, may favor both stable
demographic patterns and growth of perennials.
Annuals tend to be shallow rooted (most roots
in upper 0.1 m depth), and they are poorly
equipped to compete with shrubs for deep soil
moisture. If shrub density is high, and years of
unusually high mortality' are rare, then shiaibs
may largely preempt the critical water and min-
eral resources and suppress growth of annuals.
The dominant shrubs of the warm deserts do not
have high root densities in the upper 10 cm of
the soil profile (Wallace et al. 1980), have lower
total root densities, and have lower total cover
when compared with Great Basin perennials.
Annuals are therefore likely to experience more
intense competition from shnibs in the Great
Basin. This conjecture is finther supported by
considering that perennials in the Great Basin
generally transpire 50% or more of the ammal
moisture input over a wide range of yearly vari-
ations. In the Mojave this fraction may average
only 27% and vary between years from 15 to
50% at the same site (Lane et al. 1984), or even
be as low as 7% (Sanimis and Gay 1979). The
reduced overlap in rooting profiles and the
greater availability of unused moisture
resources may have favored the development of
annual floras in the Mojave Desert more than in
the Great Basin. With severe distin-bance from
grazing and other anthropogenic activities,
exotic annual species have invaded many Great
Basin communities. Once established following
distiubance, these annuals are not always easily
displaced by short-tenu shrub succession. While
this discussion has been presented in the con-
text of annuals versus perennials, tradeoffs
lietween short- and long-lived perennials may
be influenced by very similar climatic parameters,
sometimes operating over different time scales.
Other factors that may be important in the
ecolog)' of Great Basin annuals include the
effects of the very well developed ciyptogam
soil cRists or vesicular horizons on seed preda-
tion (abilit)' of seeds to find safe sites), seed
germination, and seedling establishment. The
restriction of winter growth by cold tempera-
tures could also be of crucial importance, inhib-
iting the prolonged establishment period
enjoyed by winter annuals in warm deserts. Fall
germination followed by low levels of photos)ii-
thesis throughout the mild winter is essential for
\igorous spring growth of winter annuals in the
Mojave, and, while heavy spring rains may cause
germination, such late cohorts rarely reach
maturity (Beatley 1974). Annuals are common
in transition zone sites of the ecotone between
Mojave Desert and Great Basin plant commu-
nities in southern Nevada, but associated with
changes in perennial species composition along
decreasing mean temperatiu'e gradients in that
region are decreases in annual abundance
(Beatley 1975).
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Spkkkv. J. S., and M. T. Tm?f.f, 1990. W'ater-stress-induced
embolism in three species of conifers. Plant ('ell and
Environment 13: 427-436.
Sturges. D. L. 1977. Soil water wididrawal and root char-
acteristics of big sagebrush. American Midland Natu-
ralist 98: 257-274.
. 1980. Soil water withdrawal and root distribution
under gnibbed, spraved, and undisturbed big sage-
brush vegetation. Great Basin Naturahst 40: 157-164.
TiiORNTiiWAlTF.. C. \V. 1948. An approach to a rational
cliissification of climate. Geographicbil Re\iew38: 5.5-94.
Vasek, F. C. iuid R. F. TllORNE. 1977. Transmontane conif-
erous vegetation. Pages 797-832 in M. G. Barbour and
J. Major, eds.. Terrestrial vegetation of Califf)rnia. John
W'ilev ;ind Sons, Inc., New York.
\'e.st, E. D. 1962. Biotic communities in the Cireat Salt Lake
Desert. Institute of Emironniental Biological
Research. Ecolog\ and Epizoolog\- Series No. 73. Uni-
versit)' of Utah, Salt Lake Cit\.
Wallace. A., and E. M. Ro.mney 1980. The role of pioneer
species in revegetation of disturbed areas. Great Basin
Naturalist Memoirs 4: 29-31.
Wallace A., E. M. Romney. and J. \\'. Cha 1980. Depth
distribution of roots of some perennial plants in the
Nevada Test Site area of the northem Mojave Desert.
Great Basin Naturalist Memoirs 4: 199-205.
West, N, E. 1983. Overview of North American temperate
deserts ;uid semi-deserts. Pages 321-330 in N. E.
West, ed., Temperate deserts and .semi-deserts. Elsev-
ier, Amsterdam.
. 1988. Intermountain deserts, shrub steppes, and
woodlands. Pages 209-230 //( M. C;. Biubour and
1). W. I^illings, eels.. North American terrestrial vege-
tation. CJanibridgc Uni\er.sitv Press, New York.
. 1989. Spatial pattern-function;il interactions in
shrub-dominated plant connnunitics. Pages 28.3-306
in G. M. McKell ed.. The biologv and utilization of
shnibs. Academic Press, New York.
U»0. Shnichirc ;uid f miction of microphvtic soil cnists in
wildliuid ectjsv'Stems of arid to semi-arid regions. Pages
179-22;3 in .Advances in ecological research. \'ol 20.
Academic Press, New York.
West, N. E., and M. M. Galdw ell 1983. Snow as a factor
in Siilt desert shnib vegetation patterns in Curlew
Valley, Utali. Americtui Midkuid Naturalist 109: 376-.378.
West, N. E., and J. Gastro 1978. Phenologv- of aerial
portion of shadscale and winterfat in Curlew X'allev,
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Wilson, C., J. Grace. S. Allen, and F. Sl.\ck. 1987.
Temperature and stature: a study of temperahires in
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Wood, M. K., E. H. Blackburn, R. E. Eckert, Jr , and
F. F. Peterson 1978. Interrelations of the phvsical
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Wyn Jones. R. G., and J. C^jrham. 1986. Osmoregulation.
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York.'
YouNc;, J. A., and R. A. Evans 1986. Erosion and deposi-
tion of fine sediments from plavas. |ournal of .Arid
Enviromaents 10: 10.'3-115.
Received 17 August 1992
Accepted 25 October 1992
Creat Basin Naturdist 52(3), pp. 216-225
LIFE HISTORY, ABUNDANCE, AND DISTRIBUTION
OF MOAPA DACE (MOAPA CORIACEA)
G. Gaiv Scoppettone , Howard L. Biirt^e ", and Peter L. Tuttle ' '
Abstract — Moapa dace {Moapa roriaccii) is a teder;iliv listed endangered fish endeniie to the spring-fed iieadwaters
of die Muddv River, Clark Connty, Nevada. Speeies life history; abundance, and distribution were studied from March 1984
to JanuiUT 1989. Reproduction, which was obsei"ved yetu-round, peaked in spring and was lowest in fall. It occurred in
headwater tributaries of the Muddy Ri\er, within 150 ni of warm water spring discharge in water temperatures ranging
from 30 to 32 C. Feni;iles matured between 41 and 45 mm in fork length (FL). Egg abundance increased with female size
(r" = .93); counts ranged from 60 for a 45-mm-FL female to 772 for one 90-mm FL. The oldest of eight fish, aged by the
opercle method, was a 90-mm-FL, 4+-year-old female. Adults are omnivorous but tended toward caniivory'; 75% of matter
by N'olume consumed was invertebrates and 25% pkints and detritus. Fish size was generally commensurate with flow, the
largest fish occurring in the greatest flow. Adults were near bottom, in focal velocities ranging from to 55 cm/s. Jn\'eniles
occupied a narrower range of depths and velocities thim adults, and lai^vae occupied slack water. From December 1984 to
September 1987, the total adult population ranged from 2600 to 2800. Although these numbers are higher than prexiouslv
believed for Moapa dace, they are still sufficiently low to warrant its end;uigered status. The dependency of Moapa dace s
different life histoiy stages to \arious areas and habitat t\pes of the Warm Springs area suggests that all remaining habitat
is necessary for their sumval.
Ki'i/ icord.s: Moapa coriacea, Moapa dace, life liislonj. rcpnulniiioti l)iolo^y.jccmi(littj. agc-i^n>ictli,Jo(Hl habits, habitat
use, bodij size, Mitdch/ Riiei; Nevada.
Tlie Moapa dace [Moapa coriacea) i.s a tlier-
mophilic niiniiow endeniie to the Mndd\' Ri\ er
system, Clark Counts, Nexada. First collected
in 1938, it has lustorically been relegated to the
headwater area where the Miiddv River origi-
nates from a series of warm springs (Hubbs and
Miller 1948). La Rivers (1962) cafled the Moapa
dace and its coinhabitant, Moapa White Ri\er
springfish [Crenichthijs baileiji nioapac), ther-
mal endemics becanse of their apparent affinit\
for warm water. Rarely exceeding 12 cm in iork
length (FL), Moapa dace ha\e moiphological
similarities to ronndtail chnb (Gila roJ)iista) and
speckled dace (Rliinichtlujs osctiln.s), wliich also
inhabit the Muddy River (Hubbs and Miller
1948). They are more similar, however, to the
genus Agasir/, which occurs in other lower Col-
orado River drainages; the two genera are spec-
ulated to have a conunon ancestor (Hubbs and
Miller 1948). Moapa dace are distinguished In
small embedded scales and a bright black spot
at the base of the caudal fin.
Little was known of Moapa dace life histor\
prior to this studv La Rixers (1962) identified
them as methodical schoolers; a curson' gut
examination bv him indicated that they foraged
primariK' on arthropods and some vegetative
matter. In a systematic sampling effort, Deacon
and Bradlev (1972) collected Moapa dace in
28-30 C water; one specimen was collected in
19.5 C water. Within the confines of its limited
distribution, Moapa dace ha\e been captured in
a variety of habitats, including spring pools and
slow- to fast-mo\ing water, and in association
with \arious substrates and submergent \egeta-
tion (Hubbs and Miller 1948).
l^ast ichtlnofaimal siuvevs suggested a
declining Moapa dace population (Deacon and
Bradle\' 1972, Cross 1976). These suivevs were
([ualitatixe and produced neither an estimate of
the number of dace remaining nor the relati\e
population decrease between suneNS. Ono et al.
(1984) tliought that ouK sexeral lumdred
M()a[)a dact^ persist(xl and that their distribution
had been hirtlier restricted within the alread\
liiiiited historic habitat, conlininsj; them to the
nj.S. Fish and Wildlife Senitc, Nation.i
^Present address: U.S. Tisli and W iMIil,-
â– ^Present addirss; U.S. Kisli and W ildlil,
(<-s.'aivhC:rii
.(.rsli.ik FisKr
â– at H.isin ( ni
. Siilistatliai, H.-iio. Nrvada. L'S.X Sm02.
.tancvOrinr. Misalika. Idalui. L'S,\ S:«2().
•nn, NiA.id.i, i:s\ S9.5()2,
216
19921
MOAPA Dace
217
main stem of the upper Muddy River and a
semi-isolated headwater spring system about
130 m long. The puq)ose of this study is to
expand information on Moapa dace life histon\
abundance, and distribution. Life histoiy infor-
mation includes reproductiye biologv', habitat
use, food habits, and age and growth.
Study Area
The Mudd\' River is at the northern edge of
the Mohave Desert, where average annual pre-
cipitation is 15 cm usualK^ in the form of rain.
Caipenter (1915) described historic terrestrial
vegetation which included greasewood
{Sarcohatus vennicidatiis), shadscale (Atriplex
confei'fifolia), creosote bush {Larrea triclen-
tata), and mescjuite (Prosopis .sp.). Stream banks
were lined with willows {Salix sp.), screw-bean
(Prosopispubescens), cottonwood (Populus sp.),
and mesquite (Carpenter 1915, Harrington
1930). Prior to the completion of Hoover Dam
(aka Boulder Dam) in 1935, the Muddy (aka
Moapa) River was about 48 km long and dis-
charged into the Virgin River, which joined the
Colorado Rixer (Hubbs and Miller 1948).
Today, it is about 40 km long and discharges into
the Overton arm of Lake Mead (Fig. 1). Source
springs of the Mudd\ River probably originate
from Paleozoic carbonate rocks (Garside and
Schilling 1979) and occur within a 2-km radius.
As is t}pical of warm springs, the water is rela-
ti\ely rich in minerals. Garside and Schilling
(1979) list sodium and calcium as predominant
cations, and carbonate and sulfate as predomi-
nant anions; total dissolved solids were 854 ppni
and pH was 7.7. Water emerges at 32 C and
cools and increases in turbidit)' downstream
(Cross 1976). Although spring discharge is rela-
tively constant at about 1.1 mVs, the Mudd\
Rixer flow fluctuates because of rain, agricul-
tural diversions, e\aporation, and transpiration
(Eakin 1964). The headwater region, the his-
toric range of the Moapa dace, is known as the
Warm Springs area (Fig. 1). During our stud\'
the area was used primariK' for agriculture, and
up to 0.25 m Vs of river discharge was being
diverted to irrigate alfalfa, barley, and pasture.
Spring outflows had been channelized, and se\-
eral were converted into irrigation ditches,
some lined with concrete. Earthen tributan-
channels had scant to thick riparian corridors of
fan palm {Washingtonia filifera), tamarisk
(Tamarisk sp.), ash trees {Frazinus sp.), and
arrow weed (Pluchea sericea). Two nonnative
fishes successfulK established in the Warm
Springs area: mosquitofish {Ganihiisia affinis),
present when Moapa dace were discovered in
1938 (Hubbs and Miller 1948), and shortfin
moUy (Poecilia mexicana), introduced in the
earlv 1960s (Hubbs and Deacon 1964). Besides
Moapa dace and springfish, roundtail chub and
speckled dace are the only native fishes occur-
ring within the Warm Springs area, but they are
rare and in greater abundance downstream
(Cross 1976, Deacon and Bradley 1972).
In 1979 the Moapa National \Vildlife Refuge
(NW^R) was established in historic habitat at the
southern edge of the Warm Springs area for the
preservation and peipetuation of the Moapa
dace (Fig. 1 ). The refuge stream originates from
five small springs occurring in a radius of 70 m
and having a cumulative discharge of abut 0.09
mVs (Fig. 2). Fan palms are the predominant
riparian vegetation. In 1984 Moapa dace larx'ae
and adults were reintroduced into the upper
Refuge Stream, and by Januaiy 1986 there was
a stable reproductive population of 120 adults
(authors, unpublished data). Thev' were isolated
by a 75-cm-high waterfall. Springfish were the
only other fish present, and they were abundant.
Materials and Methods
RepR0DUCTI\'E BIOLOCY. — Among our
objectives was to quantify' duration of the repro-
ductive period and the season of peak laivae
recruitment. To this end, a segment of the upper
Refuge Steam system was snorkeled at 30- to
90-da\ intenals from Febnian" 1986 to |amiar\'
1989 and laivae were enumerated (Fig. 2). This
is the area in which virtually all reproduction on
the Moapa NWR occurred. Dace 7-15 mm TL
were considered larvae. This range approximates
the proto- to metalanae stages of the similar-
sized speckled dace (Snyder 1981). Snorkeling
enabled us to locate reproduction sites in the
headwater Muddv River .system and to deter-
mine the abundance and distribution of adult
Moapa dace as well as to (juantifv hal^itat u.se for
all life stages. Areas with lanae close to swim-up
size (about 7 mm TL) were considered repro-
duction sites. Fish used for food habit analysis
and aging were also used to detemiine fecundit)^
H\BITAT use. — We defined habitat use in
terms of stream depth and velocitv' at foraging
sites and at suspected spawniing areas. Depth
measurements included focal and total, while
218
Great Basin Naturalist
[\blume 52
Colorado
River
115
Fig. 1. Map showing relatioiisliip of the Miiddv to the Xiigin River and Lake Mead, Ne\'ada, ;uid relationship of the
Warm Springs area to tlie Mnd(l\- liixer (helowV \\'ann Springs area or headwaters of the Muddv River showing tribntaiy
streams to tlie upper Mnildv Ri\er and relationship ot the- Moapa National \\ildlile Rehige (above).
1992]
MoAPA Dace
219
Upper
Refuge
Stream
Fig. 2. Map of Moapa Nationd VMldlife Rcriigc: shaded site indicates the reaeli of the upper Refuge Streaiu where
liUAae snorkel counts were made from Februan 19S6 to January 1989.
\ elocih' nieasurenient.s included focal and mean
water column, as prescribed b)' 13()\ee (1986).
DissoKed oxxgen and temperature were also
measured. Fish were located using mask and
snorkel. A Marsh and McBiniex model 20 ID
digital flow meter mounted on a calibrated rod
was used to measure depth and velocitN-, and a
Yellow Springs Instrument model 57 dissolved
oxvgen meter for temperature and dissolved
owgen. Sampling occurred from 1984 to 1986.
Adult habitat was also defined b)' contrasting
bod\- size with (juantitv of stream flow; it was our
subjective evaluation tliat larger fish were
inhabiting lariier water \ olumes. We tested this
220
Great Basin Naturalist
[Volume 52
200
150 -
_
1
c
1
c
1
1
1
c
1 1
1
1
1
c
1,
1.
1
il
i
1
1
1
L
c
-L
1
1
c
1
1
1
c c c
1 1 1 1
I
c
1
T5
CO
^^
0)
E
C
LU
(D
CO
_l
o
FMAMJ JASONDJ FMAMJ JASONDJ FMAMJ JASONDJ
to i~- 'o 9J
00 CO CO CO
o> en CT) O)
Month / Year
Fig. 3. Abundance of Moapa dace laivae from Februaiy 1986 to janiiaiy 1989 in tlie Muddy River system on the Moapa
National Wildlife Refuge, Nevada. Bars represent a single dav's count for the month. NS indicates not sampled.
h)^othesis in the summer of 1986 when samples
of adults \\'ere minnow-trapped from the
Muddy River, Muddy Spring Stream, Refuge
Stream, and Apcar Stream and their length fre-
quencies compared. Discharge for each stream
was measured usino; standard U.S. Geological
Survey methods (Rantz et al. 1982) near each
fish sample. A one-way factorial ANOVA was
used to test whether there was a significant
difference between length frequency among
fishes and different water volumes.
Ace and GROVNTH. — The opercle bone was
used for estimating age as described by Cassel-
man (1974). Eight specimens, collected in
summer 1985 and 1986, were aged. Flesh was
scraped with a scalpel and the bone allowed to
dry'. Glycerin was used to highlight the more
transparent region of the bone, which was
assumed to have the greatest calcium concen-
tration and to have been formed in the winter
when food is scarce. The more opaque region
signifies greater concentration of protein asso-
ciated with growth (Casselman 1974).
Food habit— Food habit anaKses were
made from 10 Moapa dace taken 9-1 1 Novem-
ber 1984 from each of three uppc-r Muddv Riv(M-
tributaries (Apcar, South Fork, and Muddv
Spring). They were captured by seining and
with unbailed minnow traps fished no longer
than 10 minutes. Ranging from 42 to 71 mm FL,
they were preserved in 10% formalin solution.
Contents in the anterior third ol the gut were
examined using a dissecting microscope and
quantified by frequency of occiuTence (Windell
1971) and by percent composition (H)iies 1950).
Abundance and distribution. — The
abundance and distribution of adult Moapa
dace (>4() mm FL) were determined by snor-
keling the upper Muddy River svstem begin-
ning from 200 m downstream of Warm Springs
Road bridge (Fig. 1). Except for 1984, the sur-
veys included 5.3 km of the upper Muddy River
and 7.5 km of its spring- fed tributaries (Refuge
Stream svstem, Apcar Stream, Muddv Spring,
South Fork, and North Fork). In 1984 the
survev area was the same except that only the
upper 130 m of the Apcar Stream was snorkeled
rather than its entire stream length. Snorkeling
was conducted over periods of foiu" to six days
when turbidit)' was low (between 1.4 and 5.0
NTU) because no agricultural return flows were
entering the stream. Coimts were made 6-10
December 1984, 6-10 June 1986, and 16-22
September 1987. Each observer enumerated
Moapa dace twice at three areas of relatively
high concentrations (30-60 fish), and the range
of results was then calculated. These sites were
chosen because the greatest variation among
obsei-vers was expected among them. For the
three sites, variation was less than 15% in counts
1992]
MoAPA Dacb:
221
1,000
800 -
c/5
LU
CD
n
E
13
600
400 -
200 -
30
-
r ^= .93
n =23
D
D^ /^
â–
/-"m
y^
-^D
D
-
f^
D
D
-
â–¡ 02^
1 1
1
1
1 1
40 50 60 70 80
Fork Length (mm)
90
100
Fig. 4. Moapa dace fecundit)- iis a tuiictioii of fork length.
between indmduals; thus, we consenathelv
estimated a 15% \ariation in o\\\ population
counts.
Results and discussion
Reproducti\e Biolog)'
Moapa dace lanae were found vear-round,
iudicating\ear- round reproduction. On the Moapa
NW'R peak lanal reciiiitment was in spring, tlie
low in autumn (Fig. 3). Fish at other reproduc-
tive sites in the Warm Springs area exliibited this
same general trend. Seasonal fluctuation in lanal
recruitment was probabh" linked to a\ ailabilitv'
of food. In the upper Mudd\ Ri\ er system the
abundance of benthic and drifting invertebrates
is much lower in winter than in spring (Scop-
{)ettone, un]:)ubli,shed data). Naiman (1976)
documented substantial seasonal fluctuation in
primar)' producti\it\' in anothei" southwestern
warm springs where production is lowest in
winter; presumably most invertebrate popula-
tion fluctuates with priman' production.
Recenth' emerged lanae were found within
150 m of spring discharge over sandy silt bot-
toms in temperatures of 30-32 C and dissoKed
ox)gen of 3.8-7.3 mg/L. Whether spawning
occurs only at these head\\'ater sites or is suc-
cessful onl\- at these sites is unknown. Visual
cues such as sexual dichromatism, pronounced
male spawiiing tubercles, or o\ertly gra\id
females were not readily apparent, and spawiiing
was not observed during our stud\". Howexer, we
indirectly identified and quantified spawning
habitat. The presence of hundreds of proto-
lanae in a concrete irrigation channel
immediately downstream of the Baldwin
springhead (Fig. 1) indicated that reproduction
had taken place. Progenitors apparentlv came
from the South Fork, entering Baldwin Spring
outflow through a diversion channel (Fig. 1).
The concrete irrigation channel had homoge-
neous water depth and \elocit\', and substrate
was sandv silt. Se\eral depressions in the sand
were similar to "redds" described for longfin
dace (Agcw/V/ clm/so'^aster; Minckle\' and Wil-
lard 1971). Depth and \ elocity at the suspected
redds were representative of the outflow chan-
nel and similar to other suspected spawning
areas in the Warm Springs area. Depth ranged
from 15.0 to 19.0 cm, near-lx^d \elocities from
3.7 to 7.6 cm/sec, and mean water colunm veloc-
it\- from 15.2 to 18.3 cm/sec.
Similar to the longfin dace, which repro-
duces during much of the year (Kepner 1982),
eggs in the skein of Moapa dace were in differ-
ent stages of development. All visible eggs were
counted, but because they are intermittently
deposited and develop throughout a gi\ en year,
our counts do not represent absolute annual
fecundity. How^ever, egg production increased
with fish size (r = .93, n = 25; Fig. 4). Counts
999
Great Basin Naturalist
[\^olnme 52
80
60
40
20
gso
>»60
O
O 40
3
Li.
80
60
40
20
80
60
40
20
^80
O 60
c
40
S"20
III
80
60
40
20
Dace
Adults
; n = 564
H,..
Dace
Juvenile
n = 148
I
Dace
Larvae
n = 201
.â– , ,,
80 -
60-
40
20
10 20 30 40 50 60 70 80 90 100110
Total Depth (cm)
Dace
Juvenile
n = 148
' ' * I L.
Dace
Larvae
n = 201
-1 1 * *
Dace
Juvenile
n = 147
Dace
Larvae
n = 199
10 20 30 40 50 60 70 80 90 100110
Focal Depth (cm)
10 20 30 40 50 60 70
Mean Water Column Velocity (cm/sec)
80
60
40
20
80
60
40
20
80
60
40
20
Dace
Juvenile
n = 147
i
Dace
Larvae
n = 201
10 20 30 40 50 60 70
Focal Velocity (cm/sec)
Fig. 5. Mean water coliiinn and loeal point xelocitics, total depth, and local point depth nsed l)\ Moapa daei' adn
juveniles, and larvae in the upper Mudd\ Rixcr system (Warm Springs area), Ne\ada. UiS-l through liJSB.
1992]
MoAPA Dace
223
Tabi.K 1. Fork IcMigth, sex, ;iiul cstiinatecl age of" eis^ht TaBI.K 2. Fckk\ items ingested In 2] Moapa dace b\'
Moapa dace collected from the upper Miiddv Hi\ cr s\ stem. percent conn^xjsition ( H\iies 1950) and percent frequence
Ne\ada. in 1985 and 19S6. Age was (Ictcnniind 1)\ the ol occnrrence (W'indell 1971). Nine odier guts examined
opercle method. w tre empty.
FL
(nnn)
Sex
Collection
date
Food items
Age
45
55
61
67
69
SO
90
Unknown
4/86
Unknown
7/86
Unknown
7/86
Female
4/86
Female
04/22/86
Unknown
10/09/85
Unknown
10/11/85
Female
10/08/85
0+
1+
1+
2+
2+
3+
3+
4+
ranged from 60 in a 45-nini-FL indiviclnal to 772
ill a 9()-nini-FL dace. Eggs were just developing
in a 41-nnn-FL female and were matnre in a
45-mm-FL fish, suggesting that females mature
at lengths in this range.
Habitat U.se
Again, Moapa dace larvae were found exclu-
sixely in the upper reaches of spring-fed tribu-
taries, while juveniles occurred primarilv in
tributaries but were more far-ranging. Adults
were present in tributaries and in the main ri\er,
with larger fish generalK' found in the larger
water volumes. There were significant differ-
ences in length frequencies among adults from
different water \olumes {p < .006). In the
MudcK Rixer, in a flow of about 0.50 m Vs, mean
FI. was 73 mm (/] = 78, SD = 16 mm); Muddy
Spring had a flow of 0.20 m Vs, and the mean FL
was 64 mm (n = 72, SD = 14 mm); the Refuge
Stream flowed at 0.17 m'/s, and mean FL was
56 mm (/; = 64, SD - 8 mm); the Apcar Stream
llowed at 0.06 mVs, and mean FL was 51 mm (n
= 89, SD = 5mm).
Lar\ae occurredand fed in tlu^ mid- to uppc^"
region of the column. They were found most
frequentk' in zero water velocit\" ( Fig. 5). As size
increased, individuals tended to occupy faster
water and occur lower in the water column,
juvenile Moapa dace occupied focal and mean
water column velocities ranging from to 46
cm/s. Adults were found in a wide range of water
depths and velocities, but they tended to orient
at the bottom in low to moderate current. Water
column depth ranged from 15 to 113 cm and
focal point depth from 9 to 107 cm. Mean water
column \elocit)- ranged from 2 to 77 cm/s and
focal point velocit)' from to 55 cm/s. Water
temperatures within adult habitats ranged from
% composition % of occurrence
Gasthopoda
Tt/ronui clathnita
1.1
Olk;()(:iiaf.tk
27.0
AMI'IIII'ODA
Hi/dlli'la aztcra
1.7
IlKMIITKHA
Pclocoiis shoshoiw
4.5
HOMOITKIU
Apiiiidac
9.0
Tkiciioitf.ha
Dolophilodcs
5.1
Necfop.si/clie
4.5
LEPlDOn'F.HA
Para'^i/ractis
4.5
COLEOPTKKA
Steucluiis ralicla
1.1
Dijtiscidae (lan'ae)
9.0
DiPTElU
Chlronomidae
4.5
Unidentified insect parts
3.3
Filamentous algae
18.5
N'ascniar plants
3.4
Detritus
2.8
4.8
23.8
9.5
4.8
4.8
9.5
9.5
9.5
4.8
4.8
4.8
9.5
42.3
9.5
14.3
27 to 32 C and dissoKed o.wgen from 3.5 to 8.4
mg/L.
Age Crowth
Annulus formation is t\picalh' associated
with an annual period of slower growtli cau.sed
bv seasonal changes in environmental condi-
tions such as temperature or food resources
(Tesch 1971). Although seasonal water temper-
atures do not change sul)stantiall\ in the Warm
Springs area, there is an apparent reduction of
potential food during the winter (Scoppettone,
unpublished data). We were unsuccessful in
aging Moapa dace bv the scale method because
scales were small, embedded, and extremely
difficult to remove from live specimens. Also,
einiroiinuMita! conditions in waters of the Warm
Springs area were sufficientlv constant that
aiinuli were not readilv apparent. Assmiied
annuli on opercular bones were presumed to be
associated with slow(^r growth dining the winter.
Ages of the eight fish examined ranged from O-l-
for a 43-mm-FL individual to 4+ for a 90-mm-
FL female (Table 1).
Food Habit
Nine of 30 guts examined were emptv' and
the remainder generalK contained few items,
224
Great Basin Naturalist
[Volume 52
Table 3. Estimated number of Moapa dace adults in six tributan- streams in the Warm Springs area, Muddv Ri\er
system, Nevada, 6-14 December 1984, 1.3-18 June 1986, iuid 16-22 September 1987.
Stream
December
X'ariation
June
\'ariation
September
Variation
name
1984
in count
1986
in count
1987
in coimt
Muddy River
475
±71
1230
±185
1165
±175
Refuge System
370
±56
406
±61
806
±121
Apcar
200
±30
565
±85
475
±72
South Fork
300
±45
185
±28
100
±15
Nortli Fork
15
±2
30
±5
60
±9
Muddv Spring
1450
±218
160
±24
200
±30
Total
2810
±422
2581
±387
2806
±421
"Onh till- iippi-r loO in of stiva
but what had been consumed indicated Moapa
dace to be omnivorous tending toward caniiv-
ory; 75% by composition was invertebrates
while 25% was plant material and detritus
(Table 2). Among 21 dace guts, oligochaetes
represented the largest \'ohune (27.0%) of food-
stuffs consumed, followed by filamentous algae
(18.5%). In terms of frequency of occurrence
filamentous algae occurred in 42.3% of the guts
while oligochaetes were in 23.8%. The stmcture
of the pharyngeal teeth also suggests an omniv-
orous diet; they are strongly hooked but ha\e a
well-developed grinding surface (La Rivers
1962). The presence of detritus and gastropods
indicates at least some foramne; from the ben-
thos, and we obsened fish in the field occasion-
ally pecking at substrate. However, the greatest
time in foraging is expended on drift feeding
(authors, unpublished data), although our data
set does not strongh' support this obsenation.
Abundance and 13istribution
Moapa dace were more widespread and
numerous than had been previously rej)orted
(Ono et al. 1984); they were in five headwater
tributaries and the upper Muddy River to about
100 m downstream from the Warm Springs
Road bridge (Fig. 2). Numbers ranged from
about 2600 in 1986 to 2800 in 1984 and 1987.
The numerical distribution for the three years
suggests movement by the adult population
(Ttible 3). In f984 the Muddy Spring stream
supported about 50% of the population (1450
adults), with only ]6%> (450 adults) foimd in the
river In June 1986 we could account for only 7%
of the population in the Muddy Spring stream,
while almost 50% of the total was in the river. In
1987 the mainstream river again supported
most adult Moapa dace (1200). The distribution
of adult Moapa dace was patchy and clumped.
For example, during the snorkel suive)' in
summer 1986, 79% of the observed dace in the
main stem Muddy River were in groups of 10 or
more, and 37% were in groups of 30 or more. In
tributaries, groups were generally smaller, with
52% of the adults in groups of 10 or more and
only 13% in groups of 30 or more.
Conclusion
Moapa dace are dependent upon the link
between the upper ri\'er and its tributaries. The
main stem river typically harbors the largest,
and presumably the longest-lived, and most
fecund fish; yet tributaries are important for
reproduction and as lanae and juvenile nurser)^
habitat. Age and growth information suggests
that three years is the mean age of fish in the
river and that adults in smaller tributaries are
one to two vears old.
Although the Moapa dace population is
more widespread and abundant than previously
beliexed, its existence remains in jeopardy.
Widespread movement and obligator' spawTi-
ing near warm water spring discharge suggest
that species survival depends on access to the
entire headwater Mudd\' River svstem (Warm
Springs area), river and tributaries alike. Everv
effort should be made to presene all of its
remaining habitat.
Ac: K N OW L E D C; M E N TS
William Burger and Dana Winkleman
assisted in snorkel surveys, and Michael Parker
and Nadine Kanim assisted in estimating fish
populations. Peter Rissler helped to determine
habitat use. Michael Parker conducted gut iuialv-
sis. Glen Clemmer, Randy McNatt, and Tom
Strekal reviewed the manuscript. Linda Hallock
1992]
MOAPA Dace
225
ln'Iped with editing and Steplianic Byers with
graphics.
Literature cited
R()\i:i:. K. D. iy<S6. Dexelopmcnt andexaluatioii oi iiabitat
siiitahilih' criteria for use in the instreani flow iiicre-
iiieiital mcthodologv-. Instreani F'low information
Paper 21. U.S. Fish and W'ildHfe Service Biological
Reixjrt 86(7). 235 pp.
C.\RPENTKR. E. 1915. Ground water in .southern Nevada.
U.S. Geological SunevWater-Supply Paper 365: 1-86.
CasSELMAN, J. M. 1974. Analysis of hard tissue of pike Esox
lucins L. with special reference to age and growth.
Pages l.'^27 in T. B. Bagenal ed.. Proceedings of tui
international .symposium on the ageing of fLsh.
European Inland Fisheries Commission of FAO, The
Fisheries Societ\ of the British Isles and The Fish
Biological Association. Unwin Brothers.
Choss. J. N. 1976. Status of the native fauna of the Moapa
River (Clark Count}-, Nevada). Transactions of the
American Fisheries Society 105: 503-508.
Deacon. J. E., andW. G. Bradley. 1972. Ecological di.s-
tribution of the fishes of the Moapa (Muddv) Ri\er in
Clark County, Nevada. Transactions of the .American
Fi.sheries Societ>- 101: 408-419.
Eakin T. E. 1964. Ground-water appraisal of Coyote
Springs iuid Kiuie Spring \'allevs and Mnckh' River
Springs Area, Lincoln and Clark counties, Nevada.
Nevada Department of Conseivation and Natural
Resources, Ground-Water Resources — Reconnais-
sance Series. Report 25.
Garside. L.J. .and J. H. Schillinc: 1979. Thermal waters
of Nevada. Nevada Bureau of Mines and Geolog\-,
Bulletin 91. Mackav School of Mines, Universitv of
Nevada, Reno. 163 pp.
Harrington. M. R. 19.30. Archaeological e.xploration in
southern Nevada. Southwest Museum Papers No. 4.
Reprinted in 1970. 126 pp.
IIlbbs, C, iind J. E. Deacon 1964. Additional introduc-
tions of tropical fishes into southern Nevada. South-
western Naturalist 9: 249-251.
HuBli.s, C;. L.. and R. \\. Miller. 1948. Two new relict
genera o( c\prinid iislies from Nevada. University o(
Michigan Musemn of Zoology Occasional Papers 507:
1-30. '
Hynes. H. B. N. 1950. The food of freshwater sticklebacks
{Gastewsteus aculeatus and Pijgpsteiis puii<iitiii.s) with
a review of metliods used in studies of the food of
fishes. Journal of/Vniniiil Ecologv- 19: 3.5-58.
Kepner, \V. G. 1982. Reproductive biolog) of longfin dace
{A<^osi(i chnjso'^dstcr) in a Sonoran Desert stream,
Arizona. Unpublished masters thesis, Arizona State
University, Tucson.
La Ri\ ers, I. 1962. Fishes and fisheries of Nevada. Nevada
Fish and Game Commission, Reno. 782 pp.
MiNCKLEY, W. L., ;uid W. E. WiLLARD. 1971. Some aspects
of biologv' of the longfin dace, a cyprinid fish character-
istic of streams in the Sonoran Desert. Southwestern
Naturalist 15: 459-464.
Naiman. R. J. 1976. Primarv' production, standing stock,
and export of organic matter in a Mohave Desert
tlieniial stream. Linniologv and Oceanographv 21: 60-
7.3.
Ono. R. D., J. D. WiLLLWis, and A. Wagner. 1984. \'an-
ishing fishes of North America. Stone Wall Press, Inc.
R\NT/. S. E. and Others. 1982. Measurement and com-
putation of streamflow: volume 1. Measurement of
stage and discharge. CJeologiciil Suivev Watcr-SuppK
Paper 2175.
Snyder. D. E. 1981. Contributions to a guide to the cvprini-
form fish Itirvae of the Upper Colorado River Svstem
in Colorado. U.S. Bureau of Liuid Management.
Denver. Colorado Contract \'.'\-5612-CTS-129. 81 pp.
Tesch. F. W. 1971. Age and growtli. In: W E. Ricker. ed..
Methods for assessment of production in fresh waters.
IBP Handbook No. 3. Black-v\ell Scientific Publication.
Oxford and Edinburgh.
Windell. J. T 1971. Food analvsis and rate of digestion.
In: W. E. Ricker, ed., Methods for assessment of pro-
duction in fresh waters. IBP Handbook No. 3. I^lack-
well Scientific Publication, Oxford and Edinburgh.
Received 1 August 1991
Accepted 15 September 1992
Creat Basin Natur;ilist 52(3), pp. 226-231
CONDITION MODELS FOR WINTERING NORTHERN PINTAILS
IN THE SOUTHERN HIGH PLAINS
Lorcn M. Smith , Douglas G. Sheelev", and Da\i(l B. Wester
AbstiucT. — Three condition models ior wintering Northern Pintails (Anas acuta) were tested for their abiiit) to predict
fat mass, logarithm of fat mass, or a condition index (CI) incoiporating fat mass. Equations generated to predict fat mass
and the logarithm of fat mass accounted for more than 69% of the variation in these dependent variables. Log transforma-
tions of body mass, wing length, and total lengdi explained at least 60% of the variation in CI. All models performed better
on an independent data set. Mean prediction error was minimal (<8% of measured variables) and negative for all models.
Regression models apply to live and dead pintails and thus represent tools that have utilit)' in a wide variet)- of studies on
pint;ul condition.
Ki-if words: Niiillirni Pintails. Anas acuta, Ixxli/ n>ii(iitii>)i. predictive models. Texas, ivateifoiel.
Biologists have used variotis indices for
assessing waterfowl nutritional status. Initially,
only body mass was used (Hanson 1962, Folk et
al. 1966, Street 1975, Flickinger and Bolen
1979), but later stmctural variables were incor-
porated to adjust for individual size differences
(Oven and Cook 1977, Bailey 1979, Wishart
1979). Ringelnian and Szvmczak (1985) and
Johnson et al. (1985) re\"iewed a\ian condition
indices and noted the value of an accurate index
of lipids in migratoiy^ bird management. These
studies noted that scaling moiphological \ari-
ables with body mass provided tiseful indices to
avian body condition.
Northern Pintails {Anas aciifa)M-e one of the
most widespread waterfowl species in North
America (Bellrose 1980), but recently their pop-
ulations have declined, making them a species
of special concern (Smith et al. 1991). Our
objectives were to pro\ide an ecjuation to pre-
dict total carcass fat (b()d>- condition) of North-
ern Pintails and to test that index on an
independent data set. The auatonn'cal \ariables
tested are suitable for field studies.
Study Area
The stud\ was conducted in the Southern
High Plains (SUP) of Texas, an 82,88()-km- area
that is one of the most intensixel)- cultivated
regions in the Western Hemisphere (Bolen et
al. 1989). Twents' thousand pla\as are present in
the SHP providing winter habitat for waterfowl
(Haukos and Smith 1992). At least one-third
(>300,000) of the Northern Pintails wintering in
the Central FK'w^ay wdnter on the SHP (Bellrose
1980).
Methods
Northern Pintails were collected using
deco\'s and b)' jtmip-shooting on plavas and
associated tailwater pits in the SHP from Octo-
ber through March of 1984-85 and 1985-86.
Tarsal length (measured from the junction of
the tibiotarsus and tarsometatarsus to the point
of articulation bet^veen the tarsometatarsus and
middle toe, 0.01 mm), flattened wnng chord
(measured from the insertion of the ahila to the
tip of the tenth priman', 0.1 cm), and total body
length (measured from the tip of the bill to the
end of the p\'gost\le, thus avoiding complica-
tions due to tail feather growth, 0.1 cm) were
recorded for each bird. During 1985-86 an
additional wing measurement was recorded
from the insertion of the alula to the tip of the
ninth priman' because the ninth primary maybe
slightK- longer than the tenth. Birds were
])lucked and frozen.
Ingesta and intestinal contents were
remoxed in the laboratoiy. Birds then were
^ Department of Range and Wildlife Management, Texas Tecli Ui\iversi(\ . I .iilihoek. Texas T9K)9.
-Box 464, Eldora, Iowa .50627.
226
1992]
PlNTAII,C:()\'niTK)\ MODKLS
00'
TaHI.K 1. X'arialile.s u.st'd in prcdictixc models ol IxxK condition lor Xortlicrn TiiitaiLs [Anas aciitd} on tlie Soiitlicrn IIi<j;li
Plain.s, Texas.
Adult
Adult
Ju\en
ile
Jmenile
miJes (n
=
140^
females in
= 69 !
males
in
= 58)
lemales
(11 = 49)
Variable
X
SE
X
SE
.V
SE
X
SE
Ma.ss (g)
963.93
10.94
8.35.07
12.60
911.97
16.41
786.68
14.90
Tarsal length
(mm)
41.15
0.17
.38.68
0.23
41.13
0.25
38.90
0.30
Wing len
igtli (
cm)
26.5S
O.Ofi
24.69
0.08
25.92
0.10
24.23
0.09
Total len
gth (
em)
49.72
0.12
43.37
0.14
49.53
0.22
4.3.14
0.19
1 ,ipid mass (g
:)
171.57
6.27
173.20
8.33
147.93
11.07
148.21
9.56
reweighed (neare.st 0.01 g) to determine a net
carcass mass and refrozen (Table 1 ). Frozen
hi i-ds were sectioned with a meat saw and passed
twice through a meat grinder. The homogenate
was dried to a constant mass in either a forced-
air o\en (60 C) or freeze dner. Dried pintails
were regronnd to insure a uniform mixture.
Lipid was extracted from 10-15 g samples using
petrolevmi ether soKent in a Soxhlet apparatus
(36-48 hrs). Fat-free diy mass (FFDM) was
calculated by subtracting water and lipid from
total carcass mass (body mass minus feathers
and ingesta). Total carcass mass minus water
mass \ielded diy mass (DM).
Three models were e\aluated to predict (1)
fat mass, (2) a condition index (CI) incorporat-
ing fat mass, and (3) the logarithm of fat mass of
wintering Northern Pintails. First, pintails were
sorted b\' sex (age was not significant; multiple
regression, P > .05). A predictive model for fat
was generated ff)r each sex using total bodv
length (TOTAL), wing length (WING), tarsal
length (TARSAL), and bocK mass (MASS) as
cxplanaton- \ariables.
In model 1 , regression coefhcients of cxpian-
atoiy variables between sexes w(m-(> iu)t different
(P > .05). A predictixe e(juati()n ap[)licable to
I)oth .sexes was therefore constnicted which
included a dunun\- xariable for sex (DSFX) as
well as stnictuial \ariables.
Th(^ second model was constnictcnl follow-
ing John.son et al. (1985); a Lipid Index was
dehncd:
Lipid Index = Fat / FFDM.
Fat-free dn' mass is included to correct for size
tlifferences between indi\iduals. Lipid Index
was transformed to:
CI = log (Lipid Index + 1)
because the structural measurements are allo-
nietric and because logarithms can be used to
linearize ratios (Johnson et al. 1985). The con-
stant 1 was added to smooth tlie function. CI can
be simplified to:
CI = log(DM/FFDM)
because
DM = Fat + FFDM.
Log FFDM was modeled as a function of the
logarithms of structural variables (LTOTAL,
LWING, and LTARSAL) and log DM as a hmc-
tion of these plus the logarithm of bodv mass
(LMASS) (Johnson etal. 1985). Unlike Mallards
{Anasplatyrhynchos; Ringelman and Szxniczak
1985) and Canada Geese {Brantn canadensis;
Ra\'eling 1979), water content of wintering
Northern Pintails fluctuated widel\- (Smith and
Sheeley 1993). Therefore, we did not test fat-
free mass as an index to structural size (Ringel-
man and Sz)'mczak 1985).
Johnson et al. (1985) used k)garithms of
structural \ariables to model logarithms of car-
cass fat mass (log fat). A separate equation was
estimated for each age/sex group (model 3)
using dummv \ariables for age (DACE) and sex
(DSEX) because regression coefficients for
explanaton \ariables differed (P < .05) among
these four groups.
Predictixe equations were \alidated on a
data set of 40 randomlv selected pintails not
inchuknl in the generation of models. Percent-
ages of each age/sex class of pintails in the inde-
pendent sample were consistent with their
occurrence in the sample collection.
Pn^liction (MTor (PF) was calculated as an
additional test of model jx'riormance. PE is
defined as:
PE = .Measured \' - Predicted Y,
where Y is the dependent \ariable. Mean PE is
an axerage \alue for all members of the \alida-
tiou data set. Finallv, predicted fat, CI, and log
fat wen^ correlated with Lipid Index in the
\ alidation data.
228
Great Basin Naturalist
[Volume 52
T.\BLE 2. Regression equations and associated statistics tor predicting carcass fat (model 1) content (g) in Northern
Pintiiils (Anas acuta) collected on the Soudiem High Plains of'Texas, October-Mtuch 198-1— S6.
r'
Exjilanatory variables
Equation
Intercept
MASS
WING
TOTAL
DSEX
1.1
.779
P;xrameter estimate 191.854
0.560
-13.386
-4.136
_
(Male; n = 198)
SE —
0.022
3.894
1.901
—
\'ariance inflation factor —
1.181
1.231
1.221
—
Piuti;il R- —
0.741
0.013
0.005
—
1.2
.711
Parameter estimate 145.570
0.570
-9.516
-4.953
—
(Female; »i = 118)
SE —
0.035
5.561
2.994
—
Variance inflation factor —
1.125
1.212
1.174
—
PartiJ R- —
0.691
0.007
o.oor
—
1.3
.757
Parameter estimate 190.494
0.563
-12.068
-4.409
-22.513
(ComJDined; tt = 316)
SE —
0.018
3.178
1.600
10.536
Variance inflation factor —
1.492
3.164
6.842
5.987
Partial R' —
0.726
0.011
0.006
0.004
•'Not .significant (P > .0.5).
T.\BLE 3. Regression equations and associated statistics for predicting Condition Index (model 2) in Nortliern Pintails
(Anas acuta) collected on the Southern High Plains of Te.xas, October-March 1984—86.
r2
Expl
anatory' v
txriables
Equation
Intercept
LMASS
LWING
LTOTAL
DSEX
2.1
.673
Parameter estimate —0.816
1.371
-1.025
-0.909
_
(Male;;) = 198)
SE —
0.069
0..343
0.312
—
Vari;uice inflation factor —
1.190
1.233
1.229
—
Partid R- —
0.656
0.015
0.014
—
2.2
.599
Parameter estimate —0.725
1.316
-1.179
-0.710
—
(Female;/) = 118)
SE —
0.101
0.512
0.486
—
Variance inflation factor —
1.123
1.206
1.176
—
Partiiil R~ —
0.595
0.019
0.008
—
2.3
.657
P;u"ameter estimate —0.761
1.350
-1.080
-0.834
-0.041
(Combined; n = 316)
SE —
0.057
0.286
0.264
0.016
Varimice inflation factor —
1.496
3.207
7.035
6.141
P;utiiil R- —
0.610
0.016
0.011
0.007
Stepwise multiple regression (iiiiLximum R'
improvement technique) was used to generate
and test all models (SAS Institute, Inc. 1985).
Variables were eliminated that did not contrib-
ute significantly (F < .05) to a model. Partial R-
values were calculated for each variable in a
model. A sum of scjuares (Ty|:)e II) for each
model variable was divided by the total sum of
squares in the model. A partial R- value for a
given variable represents the uni(|ue contribu-
tion of that variable when all other \ariables are
already present in the model. Partial H" values
are not additive, and, therefore, their sum will
not equal the total model K~. Differences in
variation accounted for by ninth \ersus tenth
primar\- length were evaluated using the K" pro-
cedure (SAS Institute, Inc. 1985).
Results
In model 1 (Table 2) bodv mass ex|:)lained a
major portion of \ariation in carcass fat content
in males (equation 1.1) and females (equation
1.2). Total length did not account for a signifi-
cant (P > .05) portion of variation in fat content
for females as it did males. Based on low vari-
ance inflation factors (\TF), regression coeffi-
cient estimates for each sex were stable. When
sexes were combined through use of a dummy
xariable (equation 1.3), the \TF for TOTAL and
DSEX were relatixeK' high; this is largelv attrib-
utable to the hitrh correlation between length
and sex of bird (point biserial correlation coeffi-
cient ecjual to 0.91).
LTOTAL, LWING, and LTARSAL ex-
plained variation in log FFDM. For modeling.
1992]
Pintail Condition Models
229
Table 4. Regression equations and associated statistics for predicting log carcass fat (model 3) in Nortlieni Pintails (Anas
acuta) collected on the Southern High Plains of Texas, October-March 19S4-86.
Explanator)' variables
Equation
Intercept
LMASS
LWING
3.1 .727
(Adult male; /) = 140)
3.2 .693
(Adult female; it = 69)
3.3 .722
( |u\eiiile male; ii = .58)
3.4 .745
(Ju\enile female; n = 49)
Parameter estimate —3.410
SE —
N'iiriance inflation factor —
Partial R- —
Parameter estimate — 1 .61 1
SE —
\'iU"iance inflation factor —
Partial R- —
Parameter estimate —11.066
SE —
Vtiriance inflation factor —
Partial R- —
Parameter estimate —.5.444
SE —
\'ariance inflation factor —
Partial fi" —
3.412
-3.209
0.182
0.993
1.156
1.156
0.697
0.021
3.687
-4.998
0.303
1.472
1.034
1.034
0.687
0.054
5.028
-1.223
0.422
2.009
1.015
1.015
0.719
().()()2
3.968
-2.834
0.348
1.844
1.109
1.109
0.720
0.013
Table 5. Coefflcients of determination (R'} and predic-
tive error estimates from the xiilidadon (n = 40) of predic-
ti\e equations to measured \ariables and Lipid Index for
wintering Northern Pintails (Anas acuta) on the Southtni
Iliiih Plains of Texas, October-March 1984-86.
Mean prediction''
Lipid Index
Equation
R-
error (± SE)
R-
1.1 Mid 1.2
.785
-9.921 ± 5.850''
.662
(fat)
6.16%
1.3
.765
-9.043 ± 5.853
.659
(fat)
6.24%
2.1 and 2.2
.697
-0.0192 ± 0.0091
,671
(Condition 1
Index'
I
7.87%
2.3
.7(X)
-0.019 ± 0.0()92
.675
(Condition ]
[ndexl
1
7.79%
3.1-3.4
.7.33
-0.050 ± 0.()()()9
.634
(log fat)
2.41%
â– "Prediction error expressed ;is a percentage of the mean in the validation data
set.
Negative prediction error iiuhcates o\erestimation of the true \alue.
log DM, LMASS, LWING, and DSEX were
significant (F < .05). TlnLs, CI was modeled with
LTOTAL, LWING, LTARSAL, and LMASS for
sexes separately and combined (Table 3). As in
model 1, regression coefficient estimates were
stable in equations 2. 1 and 2.2; when se.xes were
combined, mnlticollinearitv betsveen TOTAL
and DSEX resulted in relati\cly high MFs for
these variables.
Age and sex effects were significant when log
fat was regressed on the same e.xplanaton' \ari-
ables used in model 2. Furthermore, the struc-
tural \ariables LMASS and LWING were the
onlv variables that contributed siguilicantK
(P < .05), but they were not homogeneous
(F < .05) between age/sex groups. Therefore,
four equations were estimated (Table 4). DAGE
explained variation in log fat but not CI.
Given other model variables, bodv mass
(MASS and LMASS) consistentlv accounted for
the largest portion of variation in carcass fat
(Table 2), CI (Table 3), and log fat (Tiible 4) of
wintering Northern Pintails. Wing length
(WING and LWING) explained 1-5% of the
variation in carcass fat, log fat, and (>I when
other variables were itlready in the models.
TARSAL did not contribute to any model. Vari-
ation accounted for b\' ninth and tenth pri man-
lengths always differed by less than 1%. Conse-
quently, ninth priman' length was not tested in
any model.
In the \alidation data set all models
accounted for 69% or more of \ariation in car-
ca.ss fat mass, (]I, and log fat (Table 5). All
models explained less than 70% of the xariation
in Lipid Index for \ alidation data-set birds. Bias
in all models was relatively low and negative.
Predictive e(|uations overestimated fat mass,
CI, and log fat of \ alidation data-set pintails.
DISCUSSION
A useful condition index will sa\e fimds b\
eliminating the need for expensive laboraton
analyses and will lessen the need to sacrifice
birds for direct nutrient anaKses. The problems
230
Great Basin Naturalist
[Volume 52
associated with using body mass alone as an
index to condition of migratory birds have been
noted (Bailev 1979, Wishart 1979, Iverson and
Volis 1982. Johnson et al. 1985). Because indi-
viduals \aiy in stnictural size, bod\' mass will
reflect that \ariabilitA' in muscle and bone, in
addition to variation in lipids.
Models have been dexeloped that predict fat
content in waterfowl, but these require sacritice
and dissection of the bird (Woodall 1978, Chap-
pell and Titman 1983, Thomas et al. 1983,
Whvte and Bolen 1984). These equations may
incorporate skin (subcutaneous), abdominal
(omental), and/or intestinal (visceral) fat mass,
and often account for most of the variation in
total body-fat content. Our study was designed
to develop models using explanatoiy variables
that could be applied to live as well as dead
pintails.
Miller (1989) developed regression models
to predict carcass fat on live pintails from Sac-
ramento Vallev, California, but cautioned
against their use outside that region. Our regres-
sion models for carcass fat provided better esti-
mates of fat (K" > .71) for live pintails than those
developed for California birds IR' < .66). How-
ever, similar to Millers (1989) studv, bodv mass
alone accounted for most of the variation (R' >
.69) in pintail carcass fat.
The possibility of a condition bias among
water-fowl captm-ed in baited traps versus the
general population has been addressed
(Weatherhead and Ankney 1984, 1985,
Buniham and Nichols 1985). Hypothetically,
birds in poor condition may be hungrier, less
wary, and more likely to enter a trap contiiining
food. Condition models could be used to test for
evidence of a body-condition bias, given that
samples of pintails captured both in baited traps
and bv presumably less-biased methods (e.g.,
net gun) are available.
Models could be u.sed to test for annual
variation in bodv condition and for chans;es in
condition across the winter. Ringelman and
Szymczak (1985) demonstrated the potential of
condition indices in determining spatial differ-
ences in body condition and the preferabilitv of
condition indices to use of body mass alone.
Heppetal. (1986) also used condition indic-esto
docmnent a po.sitive relationship betAxcen con-
dition and sun ival in mallards.
The.se pintail condition models should be
useful to waterfowl biologists. However, models
should be verified when used outsick^ the eeo-
graphical range in which they were developed.
For comparisons between age and sex classes
we encourage use of model 3. Research also may
refjuire knowledge of absolute fat content.
Importance of accuracy and precision will affect
model selection. Care should be exercised to
restrict model use to winter when changes in
bod)' mass primarily reflect fluctuations in fat,
not fat-free diy mass (i.e., protein and mineral
fractions).
Acknowledgments
We offer our thanks to A. R Leif, CD.
Olavv/sky, D. G. Cook, R J. Grissom, and R N.
Gray for field assistance. E. G. Bolen, L. D.
Vangilder, D. H. fohnson, and C. B. Ramsey
provided comments on the manuscript. The
project was supported by the Caesar Kleberg
Foundation for WildHfe Consen'ation and the
Texas State Line Item for Noxious Brush and
Weed Control. This is manu,scriptT-9-488 of the
College of Agricultin-al Sciences, Texas Tech
Universitv'.
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Hanson 11. C. 1962. The dviiamics of condition factors in
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ilMKds D. A., ;uid L. M. Snutii 1992. Ecolog\- of plava
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PiN'IAII, CoxniTIOX MODKLS
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\iilii('ral)ilit\. |()iinial ol \\ ildlifc Maiia<4ciiH'iit 30: 177-
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assiunption of band-reco\ en models mav often be
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Received 1 fuiic /.9.9/
AreepU-d 22'}\nie IW2
Great Basil) Natunilist 52(3). pp. 232-236
EVALUATION OF ROAD TRACK SURVEYS
FOR COUGARS (FELIS CONCOLOR)
\\ alter D. \ an Sickle aiul Frederick Ct. Lindzev
.'Kli.sTlucr — Road track sui"\'e\s were a poor index of eoutjar ileiisitA in .sontlieni LUiili. The weak rekitionship we iound
betsveen track-finding frequency and coug;u" den.sit}' imdouhtedK' resulted in piut from the fact that aviiilable roads do not
sample properK' from the nonuniformlv distributed cougar population. Howe\er, the significantly positi\e relationship ir"
= .73) we found between track-finding fre(jueuc\ and number of cougar home langes crossing the sur\('\ load suggested
the technique may be of use in monitoring cougar populations where road abundance and location allow tlie population to
bv sampled properK. The amount of variance in track-finding frequency unexplained 1)\ number of iiome r;uiges o\erlapping
suiACN roads indicates the index ma\ be useful in demonstrating onl\- relati\'el\' large changes in cougar population size.
Kci/ uiirds: cDiti^cir. Felis coneolor. truck .s»rrr//. l^tali.
Sign left by animals ha.s been connnonly
u.sed b\- wildlife managers to make inferences
about population characteristics (Neff 1968,
Lintlzevet al. 1977, Novak 1977). This approach
is appealing becan.se it .seldom recjuires special-
ized equipment and is usually nnich less costlv
than other, more intensive techni(jues. The
approach requires, however, that the relation-
ship between sign and the population character-
istic of interest (e.g., size, composition) be
understood.
Track counts have been used to indicate
cougar (Fclis co)}color) abundance or change in
abundance, but population estimates were
seldom available to evaluate the validitv of these
indices (Koford 1978, Shaw 1979, Fitzhugh and
Smallwood 1988). \'an Dyke et al. (1986)', how-
e\er, conducted road track sunews in an area of
known cougar densit\ and found a weak rela-
tionship (/â– " = .18) between track-finding fre-
(jueucv and densitv. Because of the potential
value ol this techni(}ue to agencies charged with
management of cougars, our objectixe was to
test again the relationship between track-find-
ing f re(|U(Micy and cougar densit\ follow iug [)ro-
cedures of Van l>ke et al. (1986). AdditionalK.
we examined the influence cougar distribution
patterns, as measured b\ cougar home ranges,
had on track-finding lre<|U(mc\.
Study Aiuv\
The Bonlder-Escalante stud\' area comprises
4500 knr of Garfield and Kane counties in south
central Utah. Boulder, Escalante, and Canaan
moimtains dominate the area topographicallw
and elevation ranges from 1350 m to 3355 m.
Hot, dry' weather is characteristic of )nne and
July, with rains beginning in August and contin-
uing through September. Annual precipitatit)n
ranges from 18 cm at low elexations to 60 cm at
high elevations; axerage temperatures for
Escalante in januan and juK' are -2.8 C and
24.5 C, respectixcK" (U.S. Department of Com-
merce 1979).
Desert grass and shrub communities domi-
nate the \egetation with a sparse o\"erstoi^' of
piu\()n pine {Finns cdiilis) and juniper
ijunipcrus (isti'Dspcniui) between 1350 m and
1800 m. Dense pinxon-juniper stands with a
sagebmsh {Aiicinisid tridciitatti) underston'
dominate the xegetation between 1800 m and
2400 m. Ponderosa pine {Finns poiidcro.sa) and
oakbrush {Qucrciis <j^(ii)ihclii) are pn)miuent
abo\ (' 2400 m w here rock"\, \ertical-w ailed can-
Nons with large areas of bare sandstone charac-
terize the topographv Subalpiue meadows with
suuill stands of Eugelmann spruce (Picea
cn<i('hn(nni), ([uaking aspen (Popnhis li\-ninl(>i(lcs).
WyoiningCoi.iu-ratiw Fisli unci Wildlife- Hi-sr.ucli I'liil. I5i)\ .-^Kifi, Uiiivcrsitv St.itiiiii. 1„
. \\voiiinii;.S207r
O-T,--)
19921
CoucarThack Slknkv
233
a\id w liitc lir (Al)i('s concolor^ occur ahoxc 2700
111. Hi\ er camons transxcrse the area with asso-
ciated \e<z;etatioii consistinti; primariK' of Fre-
inout Cottonwood {Fopulus jrcniojitii) and
willow (.SV///.V sppJ (Ackcrnian 19S2. Heinker
19S2).
The human population of about 800 is con-
centrated in the towns ot Escalante and Boul-
der. I.i\estock grazing, timber harvesting, and
energx" exploration are tlie priman^ land uses in
the area. Road densit\' is about 25 km oi road
per 100 km- (\an Dyke et al. 1986). Hunting of
cougars is prohibited on the stud\' area.
Methods
Capture and \h)nitoring Procedures
Cougars were tracked on horseback, treed
with the aid of trained hounds, and immobilized
with an intramuscular injection of ketamine
Indrochloride and wlazine h\ch"ochloride
(Hemkeretal. 1984). Each immobilized cougar
was fitted with a collar containing a motion-sen-
sitixe radio transmitter (Telonics, Inc., Mesa,
.\rizona). Radio-collared cougars were moni-
tored with portable radio telemetrv equipment
on the groimd and from the air. All radioloca-
tions were assigned UTM coordinates and
recorded to the nearest 100 m. An attempt was
mad(^ to locate all radio-collared cougars a min-
iiiiuin of once each week.
Tlie Bould(M--Escalante stucK area, including
areas occupied b\ collared cougars, was
searched periodicalK' for sign of new cougars
(e.g., tracks, scats, scratches). When detectetl,
uncollared cougars taking up residence and
tnuisi(^nts were captured and radio-collared
Hoad IVack Surxcws
C-ougar densit\ was measured as both the
number of known cougars per km" in the siua ex
area and the number of home ranges ol inde-
pendent cougars oxcrlapping the suiacx road.
We conducted both s\stematic (Fitzliugh and
Smallwood 1988) and random-svstematic (\'an
Dyke et al. 1986) road tiack sune\s. Onl\' dii1
roads were sunexed.
For the sxstematic sunev the study area was
dixided into three sune\' areas spatialK' and
behaxiorally (home range boundaries) isolated
from the others. One 11.3-kni secticMi of road
xx'as chosen in each area; roads xx'ere similai" in
elexation change, habitat t\pe. and condition
(substrate, surface condition). Suncx aieas dil-
lered in cU^isitx (indejH'Uck'ut adult cougars per
km") and the number ol home ranges that inter-
sected road sc^'tions: 2-3 in the first, 4—5 in the
second, and 6-7 in the third.
Roads xx'ere sunexcnl from a pickup truck at
8-12 kph. Each road including both shoulders
x\ as dragged xvitli a conifer tree pulled from the
rear of the tnick. The folloxving dax" both sides
of the road xvere searched for cougar track sets
bx drixing on one side and returning on the
other. A track set xxas defined as a continuous
set of tracks created bx one cougar on a single
occasion. Three to 10 days later each road xxas
again sin"xexed and dragged. We felt that after 3
daxs the effect of dragging xxould be minimal,
antl moxements of cougars in the area (Heniker
et al. 1984) suggested this intenal xxould be
sufficient to proxide independent sampling
periods. Dust ratings, determined from imprint
characteristics of the obseiver's shoe (\'an D\ke
et al. 1986), xxere conducted ex^ery km before
and after dragging to (juantif\" road surface con-
dition. At each stop the obsener took 10 steps,
5 on each shoulder; then each impressic^i xxas
given a point xalue from 1 to 4. Simple regres-
sion anaK'ses xvere used to examine the relation-
ship betxx'een track sets per km surveyed and
both measures of densitx. Track .sets per km
surxexed xx'ere considered the independcMit
xarial)le because onlx'these (kita xxould be ax ail-
able to the manager.
The random-systematic road track suiaox-
inxolxed dixiding the studx aica into four sun ex'
areas. Again, the four ai"(^as were spatially and
behaxiorallx isolated from eacli other. Txx'o
sune}" areas had 2 — f cougar liome ranges oxer-
lapping roads and t\x'o had 5-7. Each area had a
different dcnsitx' of cougars (0.0 1 7, 0.032, 0.042,
0.057 cougars/km"). A 16-km stretch ot roadxx-as
landonilx selected in each area, and the first
ai"ea to be suiACxcd xxas randomix chosen. Sur-
x"exs xx'ere run as described lor sxstematic sur-
xexs except that an all-terrain xeliicle xxas used
and onlx' on(> shoulder ol the road xxas dragged
Once ;ill tour Avvds had be(^n surxexed, xx'e
returned to the first aica, randomly selected
different 16-km suncx routes for each area and
l)egan the se(juence again. Sunex ed roads xxere
not eligible for resampling until all dirt roads
xxithin an area had bcx'u sampk'd once. For
analxses. each 16-km section ot road xxas
di\ ided into segments xai"xing in length from 1
to 10 km depending on the numlx^r of home
234
Cheat Basin Natlhalist
[\i)l
r2=0.73
df=3
P = 0.066
0.005 007 0075 0.02
TRACK SETS PER KILOMETER SURVEYED
Fig. 1. Relationship behveen cougiu' track sets per kilo-
meter and cougars with home rmiges overlapping tlie snr\i\
road on the Boulder-Escakuite stuch' area, lltah, 19SS.
ranges o\erlapping the segment. Each segment
tlien had a home range oveHap \'ahie (2-7) and
was assigned one of the ionr densitv xahies.
We examined the relationship between traek
sets found per km sunexed and the t\vo niea-
sin"es of densit\"\\dth simple regression anaKsis.
Road segments with the same home range o\ er-
lap values were eombined to obtain km sur-
veyed, as were road segments representing the
same densities. Data points entered into the
regression etjuations were the siun of traeks
found in eaeh of the six home range overlap or
four densitv categories divided bv the sum of km
surveyed in the respective categories.
We evaluated whether dragginti would
improve suivev roads with a simple regression
of pre-drag dust ratings against post-drag rat-
ings. Data from both road track sune\\s were
combined to increase sample size, and regres-
sion slopes were tested against 1. The number
of track sets found on dragged and undragged
roads was also compared b\' dividing the total
luuiiber of track sets in each by the total km
searched in each.
Multiple regressiou analysis was used to
examine the effect of rainfall and traffic on
one-day, post-drag dust ratings. I're-drag dust
ratings, rainfall, and traffic were the in(k^pen-
dent variables considered. We used two indica-
tor variables to code the three levels of rainfall
and two to code the three levels of traffic. The
three road surface categories related to increas-
ing rainfall intensit)' were: unchanged, dimpled
(individual raindrop impressions distinct), and
deformed. Traffic categories were: no traffic,
traffic on one-h;i]f the length of the road, and
traffic on more than one-half the length.
Results
The systematic njad track siuAevs were con-
(hicted Mav-june 1988. During this period 407
km of road v\'as surveyed and two track sets were
found. One-hundred thiitv-five km (12 surveys)
of road was sun'ev'ed in an area where 2-3
ranges overlapped the suney road, 146 km (13
sui"vev's) where 4-5 ranges overlapped, and 126
km (11 sunevs) where 6-7 ranges overlapped
the survey road. Unequal survev numbers
resulted from weather or ecjuipment problems
precluding surveys being run. Each road (11.3
km) was sruveyed in three hoiu's, v\ith tv\'o areas
being surveyed the first day and the third the
next da\\ The two track sets were found on a
road overlapped bv 4-5 cougar home ranges.
Because of the small number of track sets found,
these results were not regressed against either
measure of densits'.
Random-sv'stematic road track siu-veys were
nni in Inly and August 1988. During this period
684 km v\as siu'veved and seven cougar track
sets v\'ere found. Three hundred fiftv km (37
road segments) was located in an area of lovv-
home-range/road overlap and 334 km (42 road
segments) in high. The number of km searched
per day was 16.
We identified no relationship between den-
sitv, as measured in cougars per km", and track
finding frequency (r = .00, P - .886, n = 4).
However, the relationship (Y = 2.23 + 197X, r
= .73, P = .066, ROOT MSE = 1, /i = 5) betvyeen
number of cougars knov\ni to have home ranges
overlapping die road and track-finding frequency
was positive (Fig. 1 ). Tlu^ data point associated
v\ ith the home range ov erlap value of 7 was drop-
ped because <20 km of road v\'as suneved.
Results from both one-dav periods and three or
more days were combined for these analvses.
Because of the small number of track sets
lound, we did not statisticalK evaluate the rela-
tionship beh\'een track-tinding frequency and
dust rating categories or dragged and
undragged roads. We found a positive relation-
ship between post-drag dust ratings (Y) and
pre-drag ratings after one (AT) and three or
more (X2) days (r = .54, Y = 6.05 + 0.875X1. P
< .001, ROOT MSE = 10.4, n = 43) (r = .34, Y
= 3.14 + 0.707X2, P < .01, ROOT MSE = 4.6,
n = 20). However, we ftiiled to reject the null
1992]
CorcAH Track SrH\i:Y
235
li\ potlicsis islopi' = 1 Hii both cases, iiidicatiiisj;
that our iiictluKl ol road drasfs^iiiu; did little to
iiiiproM' ttaekiiiij; inediuiii or that dust iatiu'j;s
were uot sensitixe enough to detect changes in
the tracking medium. Data associated with
heaxA rainfall \\ t're omitted Irom these anaKses.
Multiple regression anaKsis (onc^ da\ ) relating
[)()st-drag dust ratings to pre-drag dust ratings,
lain tall, and traffic Nielded a three-variable
model that contained onl\- pre-drag dust ratings
(A'l 1 and rainfall (X2, X3) as the independent
\ ariables (r = .67, Y = 7.65 + 0.838X1 + 0.76X2
- 5.65X3, P < .000[X1], P < .583[X2], P <
.001 [X3]. ROOT MSE = 9, /i = 43). Moderate
rainfall had little effect on post-drag dust rat-
ings. Howexer, heaxA' niintall resulting in road
surlace deformit\" had a deleterious effect on
post-drag dust ratings. The effect of traffic on
post-drag dust ratings was not signiHcant(F> .05).
location in determiiu'ni^ umuberof tracks found,
use of index \alues to compare cougar density
betx\eeu areas in tenuous. The probabilitx of
existing road net\\'orks in t\\T) area.s sampling
similarh' from tiu^ tA\'o po])ulations seems small.
U.se of track suiacns to document cougar pres-
ence is feasible, but again, the approach ulti-
mateK relies on loads intersecting a cougar
home range.
IdealK; roads with suitable trackin*! surface
o
should be abundant, as in paits of the Northwest
where logging is connnon, and located .so that
the home range of each cougar would be inter-
cepted. Even in an ideal situation, howe\(M\ the
index maxpnne sensitixe onlv to relativeK' large
clianges in cougai" [lopulation size. Twentx-
sexen percent of the xariauce in number of
tracks found xx-as unexplained bx' number of
cougar hoiiu^ ranges ()xerlap[)ing sunex* roads.
Discussion
The ntilitx of road track sunex's for monitor-
ing cougar abundance is limited bx' the generallx'
])()()r relationship betxveen cougar density and
track-finding frequencx'. Both our results {>" -
.00). although based on a small sample, and
tho.se of\'an Dxke et al. (1986) {r = .18) inilicate
a weak relationship bet\xeen cougar densit) and
track-finding frequencx'. The strongest signifi-
cant relationship found bx \'an Dyke et al. {r -
.61 ) resulted from a nuiltiple regression model
with track-finding frecjuencx' the dependent
\ ariable and female densitx; good tracking con-
ditions, aud proxiuiitx of cougars to sunex road
the iud(q)endent \ariables. As the authors
noted, hoxxexer. a biologist xvould sekUjin haxe
kuoxxledge of cougar distribution in regard to
sunex' roads.
The poor relationship documented betxx'een
track-finding frequencx and cougar densitx
appears tlie n^snlt of sampling problems, largelx
bexond the coutiol of the biologist. (Cougars an^
rarelx uuiloruiK distribut(nl (Hemkc^r et al.
I9S4!. and axailable roads, the sampling sti'ata.
are sekkim abundant enough or optimalK
located to sample from a nonnniforui distribu-
tion. .\xailable roads, for example, could fail to
intercept anx' cougar home ranges or could be
found ()ul\ in the areas occupied bx cougars, in
both scenarios, the index (tracks found) could
easilx proxe to be a poor measure of change in
cougar numbers o\-er time in an area. Likexxise,
because of the potential importance of road
AcKNow i.i:i:)(;.\iENTS
This research xvas fimded bv the Utali Dixi-
sion of Wildlife Resources and administered bx'
the \\\"oming and Utah Cooperatixe Fishen
and Wildlife Research Units. We thank W. j.
Bates for coordinating oiu" project thnigh the
UDWR. H. J. Harioxx; R. A." Poxvell, L. L.
McDonald, aiul D. G. Bonett rexiexved initial
drafts of the manuscript. \\^e offer special thanks
to C. S. .Mecham and M. (]. Mecham for field
assistance and fimctioniug as houndsmen.
LlTl'lHATlM^I-: ClTK.n
.AcKKHMAN, H. H. 19S2. (^ouiiar pivdation and ecological
energetics in sontliem Utiili. Unpublished masters
tliesis, Utah State University, Logiui. 95 pp.
FiTZHUcar E. L., and K. S. Smai.i.uood 198S. Teclnii(jue.s
for monitoring monntmn lion population le\eLs. Pages
69-71 /â– ;/ H II Smith, ed.. Proceedings of the Third
Moimtaiii I, ion Workshop, .\rizona Came and Fish
Department.
Ill Mkl-K T. v. 19S2. Pi )[)nlat ion characteristics and mo\e-
ment patterns (jf cougars in southern Utah. Unpub-
lished masters thesis. Utah State Unixersitv Logan. 66
pp.
ill Aikii; T. P.. 1' (;. I,lMr/.l•,^ and B. B. Ac kkhnian 19S4.
Population characteristics and moxement patterns of
cougars in .southern Utdi. journal of Wildlife Manage-
ment 48: 1275-12S4.
KoiOlU). (J. B. 1978. The welfiueol the puma in (!aliloniia.
Camixore I: 92-96.
I.ixnzKY. F. C. S. K. Thompson and J. 1. nou(a:s 1977.
Scent station index of black bear abundmice. Journal of
Wildlife .Management 41: 151-1.53.
.\kfk D. ). 1968. The pellet-group count technique for big
game trend, census, and distribution: a re\iew. Joumal
of Wildlife Manauement .32: .597-614.
236 C;reat Basin Naturalist [\oIiime52
NOXAK.M. 1977. Determining the a\erage size and coinpo- presence. |ounKiI (.rWildlifr .Management 50: 102-
.sition of beaver familie.s. |()urnal of Wildlife Manage- 109.
ment41: 751-754. U.S. Dhpahtmiat OF Conlmkiu:!'. 1979. CJimatokweal
Sll.uv. II. C. 19,9 A monntam lion Held gnid<>. .Arizona data annual snmnian. Climatologieal Data Utdi
Game and Fi.sli Department Special Report No. 9. 27 Sl( 1.3)
pp.
\\\ DvKK. F. G., R. II. Bkockk and II. G. Shaw 19S6.
U.se ol road track connts a.s indices of monntain lion Received U) \oiemh â– â– IMl
Accepted 16 April 1992
Great 13asiii .\atiir;Ji.st 52(3), pp. 237-244
LEAF AREA RATIOS FOR SELECTED RANGELAND PLANT SPECIES
Mark A. Welt/,', Wilhcrt H. Blackhuni". and J. Hosier Sin laiitoii'
AHSTKACr — Leaf area estimates are re([iiiretl In Indrolojiie, erosion, and 'j;ro\\ tli A ii'kl siniukition models and are
important to the nnderstanding ol trtuispiration, interception, COo fixation, and tlie energ\ balance for native pkmt
connnnnities. Leaf l)iomas.s (g) to leaf area (nim") linear regression relationships were e\alnated for 15 perennial grasses,
12 shruhs, .md 1 tree. The slope coefficient ((So) of the linear regression eqnation is a ratio of leaf area to leaf hiomass and
is definetl as the leaf area ratio [LAR = one-sided leaf area (nim~)/()\en-dr\- leafweight (g)]. LAR represents (3(1 in each
regression eqnation, where Y = P{|(X). Linear regression relationships lor leaf area were compnted (r~ = .84-.9S) for all
28 natixe nuige species after fnll leaf extension. Within-pkint estimates of leaf lU'ea for niesquitc iProsojns ^Idiidulosa Torn
\Ar.<^hni(liiIosa [Torr.] Cockll.) or liinepricklviish (Zanthoxt/hnn fag^ara [L.\ Sarg.) were not significantK' different (P< .05).
LARs for three of the shnibs and the tree were established at fonr different phenological stages. There were no significant
differences {P < .05) in LARs for lime prickh- ash, niesqnite, and Texas persimmon {Diospijras texana Scheele) after fnll
leaf extension dnring the growing season. The LAR relationship forTe.xas persimmon changed significantly after fnll leaf
extension. LAR relationships for Texas colnbrina (Cohtbrina texemis [T & G.] Gray) changed in response to water stress.
Kct/ tcanls: h'tij diva index, drought response. Icafhioiiiaw
Eighh" percent of the world's rangeland is
classified as arid or seniiarid (Branson et al.
UJSl I. i.e.. precipitation is less than e\"apotrans-
piratioii. Under these conditions water axail-
al)ilit\' is tile most important en\ironniental
factor controlling plant production and snni\ al
t Brown 1977). E\apotranspiration (ET) is the
major component of the water balance and is
estimated to accomit for 96% of annnal precip-
itation for rangeland ecos\stems (Branson et al.
1981, C^arlson et al. 1990), with surface rinioff
accounting for most of the remaining 4%
(Gifford 1975, Lauenroth and Sims 1976.' Carl-
son et al. 1990).
Ex'apotranspiration has Ixn^n irieasiired for
selected rangeland plant coimnunities with
Ksimcters and tlu^ Bow en ratio method (\\'ight
1971, Hanson 1976, C;av and Frit.schen 1979.
Carlson etal. 1990). Estimates of ET for mnnea-
sured rangeland plant connnmuties are usualK'
simulated from hydrologic models (Lane et al.
1984, \\'ight 1986). For luclrologic simulation
models to be biologicalK' meaningful, inipnned
metliods of sinnilating exapotranspiration from
rangeland plant connnnnities are needed. Two
different approaclies are currently being used.
One approach is to use a crop coefficic^nt (Kc)
(W'ight 1986). Kc is defined as the ratio of actual
exapotranspiration to e\apotranspiration when
water is nonlimiting. This empirical method is
extremeh' difficult to parameterize for range-
lands because water is often limiting and esti-
mates of transpiration are confounded h\ soil
water exaporation (Wight and Hansen 1990).
Thus, \Vight and Hansen (1990) reporied that
Kc \alues were not transferable across range
sites. The second method is based on leaf area
inde.x (LAI) (Ritchie 1972). LAI is defined as the
foliage area per unit land area (Watson 1947).
The LAI method is uiore process-ba.sed than the
Kc approach and has Ikhmi siiccesshdK used in
se\eral rangeland Indrologic, erosion, and
growtli/\ield sinnilation models (Wight and
Skiles 1987, Lane and Nearing 1989, Arnold et
al. 1990).
A limitation in using natural Resource
models, like the \\'ater Erosion Prediction Proj-
ect (WTPP) (Lane and Nearing 1989), is in
dexeloping L.\I c-oefficients for rangeland
[)lants. LAI is difficult to measiu-e because of the
drought-deciduous nature of certain shrubs, in
wliicli sexcral c\cles of leal initiation and defo-
liation occur within a single growing season
(C;anskoi)p and Miller 1986) and seasonal
.,USD,\. .Xgriciiltural Rp.search Senice. Southwest V\atersliecl Researcli Center. 2()()() F,;Lst Allen Road. Tucson. .Arizona 8.57194.596.
"Northern" Plains Area Adniinistratne OITice. 2625 Redwing Road. Suite ^50. Fort Collins, Colorado 80.526.
231
23(S (;hkat Basin Naturalist [Volume 52
TaBI.K 1. DfSfriptioii of studv sites, raii^e sites, and soil series oC species exaliiated (or leaf area to leaf hioiiuiss
relationsliips.
Frost-
.Mean
i>p'r
In 'c
[)eriod
Location
Range site
(mm)
(days)
Soil series
Soil famiK
T<)iiihstoii(\ AZ
I,iine\ upland
.â– 35(i
239
Stronghold
('oarse-loaiiiN, mixed
thermic, Ustollic Calciorthid
Meeker. CO
(;Ia\('\ slopes
200
ISO
Degater
Clav, montmorillonitic,
mesic, Tvpic Caniborthid
Sidnev. MT
Siltv
.300
130
\ida
Fine-loamv, rui.xed, T\pic
Argboroll
Chickaslia. OK
Loani\ praiiie
927
200
(;rant
Fine-silty. nii.xed, Udic
.\rgiustoll
Cliiekaslia. OK
iM'oded prairie
927
200
Eroded
C;rant
Fine-siltv, niLxed, Udic
Argiiistoll
Ft. SiippK. OK
Dnne
.597
200
Pratt
Sandy, mixed, thermic,
Psammentic Haplustalf
Wooilward. OK
Shallow praii'ie
5S4
200
Oiiinlan
Loam\-, mixed, thermic,
shallow T\pic Ustochrept
Alice. TX
Fine sand\ loam
710
2S()
Miguel
Fine, niLxed, h\perthermic,
Udic Palenstalf
Soiiora, TX
Shallow
009
240
Punes
Fine-loam\-, mixed, thermic.
T\pic Calciustoll
cliaiiti;c.s ill leaf .size, shape, antl/or tliickues.s i.s with the leaf area ratio (LAR) method (Rad-
re.siilt IVoni water, nutrient, and chemical ford 1967). LAR is defined as the ratio of leaf
.stresses ((>utler et al. 1977, (>urtis and Luchli areaper unit weight ofplant material. The slope
19S7). Foliar surface area of irregular-shaped coefficient On) of the linear regression e(juatit)n
tree leaxes has l)een estimated b\- coating the is a ratio of leaf area to leaf biomass and is
Iea\es witli a monolayer of glass heads and mea- defined as the leaf area ratio [LAR = one-sided
suring displacement (Thompson and Le\ton leaf area (nnn-)/oven-diy leaf weight (g)]. LAR
1971) and 1)\ estimating from photographs represents Po in eacli regres.sion equation,
(Miller and Scliultz 1987). Miller et al. 0987) where Y = P(,(X). LAI can be calculated as the
estimated total surface area of juniper foliage product of LAR and live biomass per unit area,
from projected leafar(\i determined from a leaf Tlie objectixe of this study was to determine
area meter. Miller et al. suggested this method LARs for selected rangeland species,
underestimated leaf area by 10% diic to leaf
owrlaj). Cregg (1992) reported that knif area MATERIALS AND METHODS
could be satisfactoriK' (\stimate(l from leaf
weight or xolume ior Juiiipcnts vir^iiiiaiia and The study area included nine range sites in
J. .sco})til<>niiit. llowexcr. leafar(>a r(>lationships fUe states and was part of the USDA Water
differed In crown position and seed source. Erosion Prediction Project (WEPP) (Table 1).
Sapwood area, stem diameter, trec^ height. The dominant plants on each range site were
canopy area, and canopy \()lume ha\e been exaluated. LARs for 15 grasses, 12^ shrubs, and
correlated to total .shrub biomass and leaf bio- 1 tree were deyeloped (Table 2). Selected
mass (Ludwiget al. 1975, Brown 1976. Ritten- rangeland .species were sampled once during
house and Snexa 1977. Whi.senant and Burzlaff the sununcM- of 1987 near Tombstone, Arizona;
1978. Cianskopp and Miller 1986, Hughes etal. and in 1987 near Meeker, Cok)rado; Sidney,
1987). In contrast, onl\ a few studies ha\('esti- Montana; Chickaslia, Ft. Supply and Wood-
mated leaf area and LAI for rangeland plant ward, Oklahoma; and Sonora, Texas, sites. Sea-
communities ((;olf 1985, (;au.skopp and Miller sonal fluctuations in LAR for du'ee shrubs and
1986, and Ansley et al. 1992). oiu^ tree were exaluated near Alice, Texas, in
An eftecti\(> method is needed to iinpro\e 1 985 and 1986.
LAI estimates lor natural resource models. One Vov k^af area (k'termination grass leaf bioma.ss
potential a[)pr()acli lor impnning LAI (>stimates from 10 raii(k)mK located ().25-nr (jnadrats was
19921
Ranc;ela\u Leaf Area K.vnu.s
239
TMii.!-; 2. Ixjcation of stucK' sites, sample dates, Iieitjlit class, iiniiiher of samples, and species exaliiated for leaf area to
at liiomass relatiousliips.
Height class (iii)
Species
Location Sample 0-11-2 2-3 3—4 >4 (ionimon name Scientific name
date
ihston
Meeker, CO
AZ Aucr. 1983
Au>i. 1983
An<i. 1983
Aug. 1983
Ano;. 1983
Aug. 1983
fmie 1987
June 1987
6 6
7 8
Si(lnc\. M r |uK 1987
|iil\ 1987
Chiekaslia. OK |une 1987
I line 1987
"liiiie 1987
Chifkaslia. OK "|mie 1987
June 1987
I line 1987
10
15
15
10
10
10
10
10
10
10
10
10
10
Ft. SuppK.OK |nne 1987
10
|une 1987
10
|une 1987
10
Woodward. OK |inie 19S7
10
|une 1987
10
Mice. TX .\Ia\ 1985
4
4
4
4
Aug. 1985
2
2
2
No\-. 1985
2
-)
2
â– ->
Jan. 1986
NA'
Apr. 1986
2
2
2
2
Ma\- 1985
5
5
5
5
Aug. 1985
3
3
3
3
Nov. 1985
3
3
3
3
Jan. 1986
3
3
3
3
Apr. 1986
\la\- 1985
3
5
3
5
3
3
Aug. 1985
5
5
Nov. 1985
5
5
Jan. 1986
5
5
Apr. 1986
.\Ia\- 1985
5
5
5
5
.\ug. 1985
5
5
Nov-. 1985
5
5
Jan. 1986
NA
A]K-. 1986
5
5
Soiiora, TX |niie 1987
10
June 1987
10
June 1987
10
Little l("af sumac
Tarbusli
Hrooiii snakeweed
Creosotel)usli
Desert /.iimia
Mariola
Shatiscale saltl)usli
\\'\()ming big sagehnisli
Needle-and-tl u'ead
Western wheatgrass
Indiangrass
Big hluestem
Little bluesteni
Buffalograss
Seribners dicliai 1 1 1 lel i 1 1 m
Sand paspalnm
Sand sagebrusli
Tall dropseed
Sand lo\egriiss
Haii^v grama
Sideoats grama
I lonex mesiiuite
Hliiis inicr()j)liijll(i Kngelm.
Floiireiisia ccmiia DC.
(Uiticrrczia sarotlirae (Pursh)
Hritt. 6c Rusb\.
Ijirrcd tridoitata (DC.) Coxille
'Aiimhi puiuila Cra\
hnihciiiiDu iiicanitm H.B.K.
.\lri])lcx cotifeiiifolki (Torr. & Frem. ) Wats.
Aiicinisia trklenlala sulwp.
ut/(>inin<iensis Beetle & Young
Stipa coDKita Trin. &c Hupr.
.\<^r()j)i/r()ii fiinithii R\db.
S(>r<i^lui.slniin iiiitmis (L.) Nash
.\iiclr<>i)i><^()n gcrardii Vitnuui
Scliizaclii/hiiin scoparium (Mich.x.) Nasli
Biichlof (Idcti/loklcs (Nutt.) Fngelm.
niclunilhcliitm olif^osaiUlies (Scluilt.)
(aiikl \ar. scrihiicrianuin (Niish) Could
I'dspaliim sctdcenm Miclrc. \ar.
strdiniiu'iiiii (Nash) D. Banks
.A livmisid jihfolia Torr.
Sporoholus aspcr ( Michx. ) Kunth
Erogro.s/Z.s- tiicliodcs (Nutt.) Wood
Boiitcloiid liirsuta Lag.
Boittchnui aiiiipc'iulula (Michx.) Torr.
Prosopis gidiululosd Torr. \iu".
"laiuhilosd (Torr.) C-tx-kll.
5 Lime priekK ash Zdiillioxi/liiin faodra (L.) Sarg.
Texas colubrina Colnbrind tcxciisi.s (T. 6c C.) Gray
Texas persimmoTi Diospyms texaiui Scheele
White tridens Tridtiis dlhe.sccns (N'asev) \Vo<it. & Standi.
(>"urK mescjuite llilarid l)clan<ieria (Steud.) Nash
Texas wintergrass Slijxi Iciicotiiclui Trin. & Riipr.
'No. sample
cted (or dtxidiions shrubs and tp
240
Great Basin Naturalist
[\ oluine 52
Tablf, .3. Mean and standard error oflcat hioniass and leaf ;irea. and linear regression'' model slope eoettieients (LAR '
relating leafiu-ea to leafbionuLss for selected rangeland grasses and shnibs sampled after f'nil le;if' extension.
Species
Grass Ks
Needle-and-tliread
Western wiu-atgrass
Indiangrass
Little l)ln(^stem
Big hlnesteni
Buff;ilo grass
Scrihners dicliantheliiuu
Sand paspdnm
Tiill dropseed
S;uid lovegrass
Hain grama
Sideoats grama
White tridens
Texas wintergrass
(JurK' mesqnite
SllKlBS
Desert zinnia
Mariola
Broom snakeweed
Little leaf snmac
Tarhnsh
Oeosott-husli
Sand sagebrnsh
Shadscale saltbnsli
Wyoming big sagebnish
Leaf biomass
3.6
2.0
S..5
2 7
1.3
1.5
1.3
1.5
0.9
0..S
0. r
o.r-)
0.7
1.2
o.s
1.6
3.5
•3.7
3.9
3.7
.3.0
3.2
3.9
5.3
SE
O.SO
0.33
1.56
0.3S
0.45
0.22
0.21
0.23
0.15
0.12
0.13
0.22
0.16
0.24
0.15
0.10
0.40
0.51
0.71
1 .00
0.19
0.58
O.Sl
0.S.3
Leaf area
(nmi")
SE
3,580
5,760
82,670
28,030
11.290
6,820
15,300
7,580
8,500
8,650
4,360
5,240
.3,980
8,.32()
5,270
9,440
19.410
11.160
22,0.50
2.3,.360
16,790
5,9.50
10,5,30
18,220
900
902
1,3.50
4,710
2,213
1,091
2,601
1,1.36
1,3.34
1,3.S3
769
2,8.36
1,007
1,.361
925
580
1,2.S(I
920
331
20.3
910
1 .257
2,047
2,715
LAR
{nim"g )
1.040
.98
2,910
.98
9,440
.96
10,780
.98
12,970
.86
5,680
.97
16,110
.96
6,890
.95
9,.390
.99
11.380
.98
5,890
.99
10,210
.98
5,8.30
.98
6,720
.95
6,620
.99
5,700
.89
5,690
.84
2,700
.96
4.700
.91
6,100
.97
3,660
.86
2,010
.98
2.640
.98
3,340
.97
''.All areuiweiglit regressions were sigiiilkaiil at /' «
'Ix'af area ratio ( 1.AR) represents ^n in eaih ici^re
■V = p„iXi,
used. Cirass hioinass in each (juadrat was clipped
to a 2()-mni stubble height and separated by
species into lix'e or dead leaves. Li\'e lea\"es were
placed in plastic bags on ice for later determina-
tion ot leal area. The lea\es were flattened and
[)laced between clcnir plastic sheets and then
processed tlnough a leaf area meter. Leaf area
was determined with a Li-Cor .3()()()' leaf area
meter to the nearest 1 mm"^. The samples were
then oven-dried at fSO C for threc^ da\s and dn
mass determined.
To ensure that .samples ol' shrubs and trees
represent(Hl the full range of size of plants pres-
ent, a .stratific^d random sampling procedure w as
used. Height classes of 1 m were adiitrariK
chosen, and plants were selected randoniK from
each class. As a result, total number of plants
.sampled \ aried among .speci(>s depending upon
the range of plant heights (Table 2).
An open-ended cul)e (250 mm on a side) was
used to sample shrub and tree leaf biomass. The
The ii.se ot a trade or linn name in this papi-r is tor reader inlornialion .ind
does not in)pl\ endorsement In the U.S. Department of .-Vgrienhnre ol .in\
prixlnet or service.
sample cube was placed in an area considered
representatixe of the entire canop\', and the
lea\es within the area were remoxed In* hand.
LARs were determined in the same manner as
for grasses.
Within-plant \ariabilit\' of LARs was e\alu-
ated for four mes({iiite trees and foiu" lime
prickh' ash shnibs in Mav 1985 near Alice,
Texas. Fifteen sample cubes were randomlv
located and sampkxl from each of the four raes-
(jiiite trees. For the lime prickK ash shrubs 12
sample cubes were hanested from each of the
four shrubs. LAR was determined in the same
inanntM- as pre\ionsl\ described. A one-wax'
anal\ sis of \ariance was used to test for differ-
ences (F < .05) among the slopes of the regres-
sion e(juations within plant canop\' b\ species
(Ste(>l and Torrie 1980). Within-plant LARs
were not significanth' different for lime prickly
ash and mescjuite hi May 1985. Based on these
relationships, one sample per plant was utilized
during the reinaind(M- of th(> stiuK.
Three shrubs, lime prickK ash, Texas per-
simmon, and Texas colubrina, and one tree.
19921
Raxcklam) Li-:af Area Kviios
241
Table 4. Mean and standard error of" leaf biomass and leaf area, and linear regression'' model slope coefficients (LAU
lating leaf iUX'a to leaf bionuiss for selected rangeland shrubs and trei' on a line sandv loam range site near /Vlice. Texius.
Species
Date
Leaf liiomass
SE
Leaf area
(mm")
SE
LAR
(nnn"g' )
r'
Unie priekK ash
Max- 19S5
4.7
0.73
45,180
1,450
8,760 a''
.99
Ang. 19S5
4.2
0.63
40,330
1,530
8,730 a
.98
N(n-. 1985
5.6
0.89
43,360
1,460
8,670 a
.98
Jan. 19S5
4.9
0.76
44,310
1,450
8,870 a
.98
Apr. 19S6
5.3
0.65
52,730
1,580
8,690 a
.98
\h'S<jnite
Mav 1985
6.5
0.87
57,830
1,610
8,990 a
.98
Aw^. 1985
5.7
0.64
56,040
1,470
8,780 a
.98
Nov. 1985
5.5
0.70
48,460
1,410
8.630 a
.98
Jan. 1985
\A''
Apr. 1986
6.4
0.81
59.100
1,470
9,290 a
.98
Texas persimmon
Max I9S5
4.6
0.64
49.960
1,940
10,590 b
.96
An<j;. 1985
4.1
0.65
41.670
1,780
10,.360b
.98
No\-. 1985
4.8
0.59
51.060
1.790
10,1.30 b
.98
Jan. 1986
4.6
0.68
44.720
1,900
10,020 b
.98
Apr. 1986
4.7
0.69
64, 150
2,070
12,660 a
.97
Texas colnbrina
Max- 1985
4.9
0.78
55,070
2,020
10,310 b
.98
.-\ng. 1985
5.2
0.89
57,010
1,720
10.110 b
.98
Nox-. 1985
3.8
0.65
55.380
2.090
13.360 a
.98
Jan. 1986
NA
Apr. 1986
4.1
0.71
41,760
1,880
10.230 1)
.98
''.\11 area: ueiglit regres.sions were signilitaiit at P < .05.
Leaf area ratio (LAR) represents |3ii in each regression, w'liere V = 3ii(X).
'Parameters in the columns by species sliaringa common letter are not signilicaiitK different if < .0.51 lia.sctl <
' No sample was collected for deciduous slirnbs.
;)f slope test.
honex" nie.s({uite, were .selected for e\ iiluation ot
.seasonal fluctuation in LAR. Hone\ mesquite,
Texas j3ersininion, and Te.xas colnbrina are
drought-deciduous while lime prickK ash is an
exergreen. Sauij:)le dates were selected to cor-
resjiond to the jihenological stages of ( 1 ) niaxi-
niuni leaf area, (2) peak drought defoliation, (3)
autunui, just prior to winter leaf fall and dor-
mancy, and (4) after winter leaf fall for the
ck'ciduous shnili.
The Statistical Analxsis Sxsteni (SAS 1982)
was utilized to tnaluatc linear regression rela-
tionships, Y = Pi, + Pi(,X), between leaf biomass
and leaf area. Where Y is estimated leaf area
(nmr), p,, is the intercept, pi is the slojoe (LAR
coefficient as defined bv Radford 1967 in mnr
g ), and X is leaf biomass (g). The intercept was
tested to determine if it w^as significantK differ-
ent (P < .05) from zero. The intercej^t was not
significantK ditfert^nt from zero for all sjoecies.
Therefore, tht^ data were reanal\y,ed and pre-
.sented using a linear regression model, Y =
P(i(X). similar to that reported by Coombs et al.
(1987) and Ansley et al. (1992) for estimating
LAR. All statistical tests were judged significant
at P < .05 unless otherwise stated. A homogene-
ih of sloj^e test was used to test for differences
among the slojx^s of the regression equations
(LAR) between sample periods within spcH'it^s
(Steel and Torrie 1980).
Results a.\d Discussion
Leaf area of graminoids was highl\- corre-
lated with leaf biomass for all species within
samj3le dates (Table 3). The LAR for perennial
grass leaf area ranged from 2910 to 16,110 mnr
g~'. The LAR for shnibs and trees ranged from
2010 to 13,360 nun- g '. Goff (1985) also
reported significant lin(^ar regression relation-
sliips (r = .83-. 97) for LAR for 1 1 natixe grass
species in southern .\rizona. Golf rejiorted that
the lintnir regression coefficients for stem area
to stem biomass (SAR) ranged from 32 to 739f
of the LAR and the mean SAR was 44% of the
mean L,'\R.
There was no significant seasonal \ariation
in L\R for lime prickK- ash and mesquite (Table 4 ).
Although there was no significant sea.sona] dif-
ference between mescjuite L.\R relationships, a
gradual decrease in the LAR from May through
Noxember xx'as apparent in 1985. Furthermore,
the LAR xxas larger in April 1986, though it xxas
not significantK- different from 1985 sampling
dates. Moonex- et al. ( 1977) found that the sj)ecific
242
Giiivvi" Basin Naturalist
[\ olume 52
leaf densitv' (nig innT") of niesquite leuM's
increased over the growing season. The densit}'
ranged from 0.{)()()4 nig nnn " in the spring to
0.01 7 nig mm' in die fall. This corresponds with
a leaf area change of 5880 to 25,000 nnii' g '.
Ansle\- et al. ( 1992), working in north central
Texas, reported that LAR of niescjnite ranged
from 9916 to 5944 ninr g'. Mesquite LAR
declintnl from Ma\ throngli Angust 1987, hnt
stabilized from Angnst through September fol-
lowing substantial precipitation. In 1988 precip-
itation was substantialK' less than in 1987, and
the mean I.AR was significantly lower than in
1987. LAR followed the same pattern in 1988,
declining from a high of 6877 in the spring to a
low of 4996 mm" g' in October. Anslev et al.
(1992) speculated that the decline in LAR was
caused b\- cell-wall thickening in response to
dning conditions, based on the work of Kramer
and Ko/.k)wski (1979).
The siinilarit\' in LAR across sampling dates
f ron 1 th is stud\' may be partially explained in th at
sam[)ling was not initiated until all leaves were
lulK expanded (or approximatelv lour weeks. In
addition, Ajiril, Ma\; June, and September pre-
cipitation was significantlv above the k)ng-terni
average ])recipitation and no noticeable water
stress was apparent in the trees sampled. Nilsen
et al. (1986) indicated that relati\e leaf area of
phreatoplntic mesfjuite {P. olanchilosa \ar. tor-
rcijana) in tlu^ Sonoran desert of southern (iai-
ifoniia remained nearK constant from Ma\
through \o\ember. Maximum leaf area was
maintaiiK^d throughout the liottest and driest
months ol the wav \ia access of deep stored soil
water by taproots. When water availabilit}' to the
normally phreatophytic mesquite was reduced,
total leaf area was reduced (Nilsen, Virginia, and
Jarrell 1986). We hvpothesi/xMl that nies(juite
lea\-es reach a stable weight at niaturit\ and the
lack of water stress during the growing season
prevents the changes in leaf weight (o leaf area
reported by Ansley et al. (1992). Changes in leaf
weight as a result of translocation ol' sugars,
.starches, other compounds, and insect damage
could not be detected or .separated from cell-
wall thickening from water stress witliin the
precision of sampling in om- stud\.
Texas persimmon LAR in April 1986 was
significantl)- greater than for sampling dates in
1985. Meyer (1974) reported that Texas persim-
mon produces two tvpes of leaxcs: a large leaf in
the center of the canopy and a smaller leaf
around the i^erimeter of the plant. The leaxes
arc* initialK light green in color and become
glabrous after elongation ceases. As the leaf
matures, the x)'leni and bundle fibers become
increasingly lignified and the leaf tunis dark
green, with the underside becoming densely
covered with trichomes. Leaf modification is
complete by early July. The lower LAR of Texas
persimmon leaxes in 1986 was attributed to the
leaxes not being fullv elongated, with
incomplete development of trichomes and lig-
nification.
LAR relationships forTexascolubrinaxaried
seasonally. LAR was similar during the early
growang seasons in May 1985 and April 1986,
and in August 1985. In November the LAR was
33% greater than during other sample dates
(Table 4). Rasal leaves of Texas colubrina are
approxiniateK 10 times larger than the outer
canop\ leaves. In response to an extended diy
period in fuK and August, Texas colubrina
dropped 95% of its leaves. The onl\- leaves
I'etained during this diy period were the large
basal leaves in the center of the shnib. The
significant difference in LAR between the
sample dates was attributed to the different
proportion of leaf tspes and not the change in
specific weight of the leaves.
(Tunskopp and Miller (1986) reported sim-
ilar significant seasonal changes in LAR for
Wxoming big sagebrush. Tlie\' speculated that
the greatest proportion of seasonal \ ariation was
due not to the development or alterations in
starch and sugar accuniulations but rather to
changes in the proportion of larger persistent
leaves to smaller ephemeral leaves.
Shiiib leaf biomass to leaf ai'ea was liighlv
correlatetl for the nine other shrubs sampled
(Table 3). The LAR for slinib leaf area ranged
from 2010 to 6100 nuir g"'. Other researchers
have also reported satisfacton results in relating
l(\il biomass to leaf area (Schilesinger and
(^habot 1977, Kaufmann et al. 1982, Ganskopp
and Miller 1986) within sample date. Based on
the seasonal xaiiabilitvin LAR for Texas persim-
mon and Texas colubrina in this stiuK and the
findings ol (Tunskopp and Miller (1986) in ea,stern
Oregon tor Wvoming big .sagebni.sh, we c;ui state
that s(\i.s()nal \ariabilit\ in tlie.se and other
(h'ought-deciduous shmbs is an important source
of xaiiation tliat needs to be accounted for when
simulatinti LAI owv the entire* tirowiii<i sea.son.
19921
Raxcelam) Lkaf Ahka Ratios
243
Conclusion
For tlic spt'cics saiiipli'd. leal hioiiiass is a
reliable^ estimator of leaf area, llowexer, for
some slinil) species, seasonal differences in
cle\('loi)m(Mit and shedding of different t\pes of
l(^a\es and leal nioiphological de\elopnient c-an
prodnce significant temporal flnctnations in
LAR. Caldwell et al. (198f) reported that for
semiarid hnnciigrasses, leaf blades of regrowing
tillers had grc\it(^r photos\nthetic capacit\' than
blades on nnclipped plants. This resulted in
greater carbon gain for clipped plants and an
increased photosMithesis/transpiration ratio.
Nowak and Caldwell (1984) reported that the
photosvnthetic rate for both clipped and nn-
clipped plants decreased with age of the lea\es.
Cnrrent rangeland Indrologic simnlation
models do not account for changes in LAR or
exapotranspiration rates as a function of age of
the leaf. [)r()poition of leaf t\pe, or compensa-
ton photosNiithesis rate increases following
defoliation due to grazing. Models currently
utilize a fixed coefficient for calculating LAI. If
significant adxances in modeling e\'apotranspi-
ration on langelands are to b(^ made,
improxements in the relationships used to sim-
ulate exapotranspiration that incoiporate these
processes will he needed. The LAR method of
calculating LAI exaluated in this studx" proxides
a fast, reliable method of estimating LAI neces-
san to [)arameterize these hydrologic simula-
tion modc^ls. To account for the seasonal
diffen^nces in L.\R for Texas persimmon and
Texas colubrina, a xx'eighted average based on
season of xear is recommended for parameter-
izing tlu^WT.PP model. For plants like m(^s(|uite
and lime piicklx ash, one LAR xalue can be u.sed
in non-drought vears. For xears xxith significant
dn periods, a decrease in LAR of 10^0% max
need to be accounted for xxith non-phreato-
piixtic nies(|nite. as indicated bx this xxork and
thatof Ansleyetal. U992j.
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Great Basin Naturalist
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Received r, June 19^1
Accepted U) Septeniher 1992
Cwrat Basin NatiinJist 52(3), pp. 245-252
ECOLOGY AND MANACiEMENT OE \IEi:)US AHEAD
(TAENIATHERUM CAPUT-MEDUSAE SSP. ASPERUM [SIMK.J MELDEKIS)
|ames A. Youiuj;
AhsTKaci'. — Mediisahcad is aiiotlicr in the cxlcnsixc list ol annual herbaceons spciirs to invade the temperate desert
raniielands ol tlie Great Basin. Medusaliead is not preferred 1)\ lar^e herl)i\i)i(s and apparentK' is not preferred In'
'j;rcnn\ores. I lerl)age of tills annual grass enlianees ignition and spread ofwildl'ii'es. Mcdus.ilie.id is liigliK eonipetitixc witli
the seedlings of native species and is prohabK' tlie greatest threat to thi' l)iodi\crsit\ of ihc natnral \fgetation that has \et
been aeeidentalK introduced into the Great Basin. Hespite the ob\ions biologii-al disni|)tions that are associated with
niedusahead imasion. the species offers awealtli ol oppoit unities for stndents to exaniiiie the nieehanisni b\- which this
species is so successful. Stutlents of cNoIntion. |ilant pli\ sioli>g\. and ecologx max find this species to be an excellent model
lor colonization.
Kci/ uonl.s: uwdusdhfdd. Tat'niatherum ca]")ut-mednsa(^ aiiiiiKil tr^rass. coloiiiziit^siwrii's. uihlfircs. grr/z///g.
Ill the nianagemeiit of natural resources
tlieii' are certain problems that 1)\ their persis-
tence, inagnitnde of ecological disniption. and
eeononiic impact refuse to dissipate as a result
ol being ignored and neglected. UnFortunateK
tor range management, niedusahead
iTaoiidtlicrunt c(ipnt-})icdus(ic [L.] Nevski) is
that t\pe ot problem. During the 1950s
niedusahead was considered among the most
pressing problems on the rangelands of C^alifor-
nia, Idaho, and Oregon. A great deal of research
effort was dexoted to solving the niedusahead
problem, \aliiable information was learned
about the ecoplnsiologs and s\niecolog\" of
iiiechisahead. (Control methods were dexeloped
using herbicides. The fatal link in integrated
])rograms for the suppression of niedusahead
populations pro\ed to be artificial rexegetation
technologies after niechisahead was controlled.
The nature of tlu^ sites infested had more to do
with this lailiire tliaii the weed itself, especialK
in the bitermountain area. The recent discox'en
ol niedusahead in northern Utah has renewcxl
interest in suppressing tilis rangeland weed.
M\ purpose in this review is to relresh our
c()llecti\-e memories about medusaliead ecologx
and management.
Ta.xoxomv
As is olten the case with an introduced s])e-
cies, there has bec^i coulusion about the c'orrect
scientific taxon lor medusaliead. Tlie first
description ol niedusahead in a Nortli American
flora used the tiixon Eh/iims caput-nwdiisac L.
(Howell 1903). There is apparent agreement
that niedusahead is a member of the trilx^
Triticeae of the grass tamil\-. There is also appar-
ent agreement among moiphologists and c\ to-
geneticists that niedusahead does not fit in the
genus Elipmis. N'arious autliors haxc placed
niedusahead in Hordciiin or Hordcli/iitits.
Newski (1934) proposed tliat medusaliead was
tniK" a different genus and published the name
Tdcnidfhcrum. jack Major ol the I iii\ cisilx ol
(California suggestcHl in 19f-)()tliat material intro-
duced to the United States was Taeiiiathcnini
(ispcriiiii (Major et al. 1960). Based on the
European and Hussian literature. Major
reported tliat I'dcuidlUcnitn contained three
geograpliic and moiphologicalK tlistinct ta\a, T.
cdjntt-incdnsdc. T. dsfxTiiiii. and T. crinituiii.
Tlie.se three sjx'cies are loiiiid in the Mediterra-
nean region and extend eastward into central
.\sia. Alter examiiiiiiij; the European material,
growing in place. .Xhijor decided the I iiited
States introduction was T. d.spcruin.
The Danish scientist Signe Frederikseii
rexi.sed the genus in 19Sfl He kept the same
three taxa, but reduceil them to subspecies of
Tdciiiddici'iini cdpul-nicdusdc. Positixe identifi-
.\griciiltmal Hesearcli Senice. U.S. Dep.irttm-iit c>IAi;ntiiltun-. 920 \all<-\ Hoad. Hi-iio. \\-\a(la Sy.5I2.
245
246
c;riv\t Basix Naturalist
[Volume 52
cation to the lowest le\el possible is ahsolntely
essential for am proposcxl biological control
program for medusaheacl. According to
Frederiksen's revision, subspecies crinituiii has
a \'er\' strict spike. Subspecies captif-nicdiisae
lias a large open spike with straight awais. The
spike of subspecies a.spcniDi is intermediate
with angled awns. Subspecies (ispcnim is die
only one of the three witli pronounced barbs
coated with silica on the awns. Apparently, the
correct taxon for the medusahead of western
North America is Tacniatlicnuu capiit-nicdiisac
ssp. aspeniin (Simk.) Melderis (Frederiksen
1986).
Taeniathenun caput-niedusae ssp. capiit-
nicdiisae is mostK restricted to Portugal, Spain,
southern France, Morocco, and Algeria. It has
been collected outside this area in Europe and
Asia, but Frederiksen considers it adxentitions
in the.se areas. Subspecies chnitnt)i is found
from (ireece and Yugosla\ia eastward into Asia.
Subspecies aspcniiii completely overlaps the
distribution of the other two subspecies. All
three subspecies integrate with each other.
ApparentK' onlv the one subspecies occurs in
Nortli America. Does this indicate one or vev\
limited introductions?
.Mechi.saliead is predominanth' self-polli-
nated. Genetically the genus appears to stand
alone in genomic relations within the Triticeae
(Schooler 1966, Sakamoto 1973). ApparentK
Tacniafhcrinii has a genome that is distinct, but
faintK' related to those of Fsadii/rostachi/s,
Dasijpi/nitii, Erciiiopiptim, or Hordcmii
(Frederiksen and Hot hue 'r 1989).
IIlSTOm- IN NOHTII Amkhica
.Medusahead was first collected in the
United States near Roseburg, Oregon, on 24
June 1 887 by Thomas Jefferson Howell ( 1903).
It was next collected ncnu- Steptoe Butte in east-
ern Washington in 1901 b\- George Xixsex (Piper
and Beattie 1914), followed by a collection n(>ar
Los Gatos, California, in 1 908 In Charles I litch-
cock (Jepson 1923). Medusahead certaiuK
attracted the noted agrologi.st. McKell. Hobin-
.son, and Major (1962) commented on diis
.strange initial distribution reaching 390 miles
north and 450 miles south from the point of
initial collection. EaH\ lied)arium .specimens
show a rapid spread to the .south into California.
J. F. PechantH- made die first collection in
Idaho in 1944 near Payette or about ISO miles
.south ol Steptoe Butte (Sharp and Tisdale
1952). Fred Rennertold jack Major he had seen
medusahead near Mountain Home, Idaho, as
early as 1930, and Lee Sliaq) had reports from
ranchers that the species occurred in Idaho as
early as 1942. The medusahead infestation in
Idaho increased to 30,000 acres b\' 1952. Min
Hironaka estimated that 150,000 acres were
infested by 1955, and the Bureau of Land Man-
agement estimated 700,000 acres were infested
by 1959. At that rate of spread it appeared that
all of Idaho would be infested by the end of the
next decade. The spread of medusahead slowed
and nearly continuous infestations remained
confined to Gem, Payette, and Washington
counties in southwestern Idaho. There were
several spot infestations in surrounding counties
(Hironaka and Tisdale 1958).
Medusahead spread soutli in California to
Santa Barbara on the southern coast and Fresno
Count\' in the interior vallexs. The rapid spread
from southwestern Oregon through northern
and central California occiuTed in annual-dom-
inated grassland, oak {Qtierciis) woodland, and
chaparral commimities. These areas lia\e a
Mediterranean t\pe climate with hot, di")' sum-
mers and cool, moist falls, winters, and springs.
Germination occurs in the fall and flowering
and seed set in the spring.
In northea.steni California, east of the Sierra
Ne\ ada-Cascade rim, medusahead inxasion
occiuTed at a much slower rate. In the Pitt Ri\'er
drainage, vegetation is an intergrade of Oregon
white oak (Qucrciis ^(irnjaiui) woodlands,
cismontane California species, western juniper
ifiiiupcnis occidental is), ponderosa pine {Pi)uis
pondcrosa) woodlands, and sagebrush {Aifeini-
.s7V/)/buncli grass communities more tspical of
the Intennountain area.
Medusahead was discoxt'red in the Great
Basin at \erdi, Nevada, in the earK 1960s. Iso-
lated inf(\stations were subsequentK found
along the eastern front of the Sierra Ne\ada in
ar(>as wliere range sheep bauds used to concen-
trate^ wliile waiting for mountain summer pas-
tures to be Iree of snow.
In northeastern C'alifoinia in the CTreat Basin
duiing (h(" earl\' 1960s, tluM'e were two small
inlestations in citv lots in Snsanxille and a small
infestation at the old slu^ep-shearing site of
\iew land along the niilroad above Wendel, Cal-
ilornia. .Another isolated infestation occurred at
die mouth of Fandango Pass in Suiprise Valley.
B\ the earK 1970s, medusahead was uearK"
19921
ECOlXKiY AND MANA(;KMENT()F MKDI SAIlKAl)
247
continuous ox'er al)out 60. ()()() at'ics of tlic
Willow (]reek-Tal)l('Ian(ls northeast ol Susan-
\illc. ('uncntK. alter lour \ears ol extreme
(lrouu;iit. uiedusahead s[)()t iutestatious occur
o\tM- [)erliap.s an additional uiillion acres on the
westcM'u maitjiu ol the (weat Basin.
HlOl.OCV OF MEI^USAIIIvM;
Medusaliead. in some wavs, is a rerun of
clieatgrass {Bronms tectoniin) imasion.
(dieatgrass dominates secondan' succession in
a majorit)' of sagehnisli/bunchgrass communi-
ties in the Great Basin and proxides a significant
portion of the forage base for lixestock grazing.
Howe\er, there are hiiihK' si(j[nificant differ-
ences in the ecolog\- of the t^vo grass species
(Harris and Wilson 1970, Al-Dakheel 1986).
Germination. — The canopsis of medusa-
head is less than a millimeter wide with a \en
shaip callus and an elongated, non-geniculated
awii. The medusaliead caiyopsis is covered with
small barbs of silica. \^cious is the best descrip-
tion for this grass canopsis. Bo\e\" et al. (1961)
determined that medusaliead had a much
higher ash content (o\er 10%) than other grass
species and the ash was about 7o7c silica. Hea\A'
deposition of .silica occurs on the barbs of awns
and the epidermis of leaxes.
For the \ast majorit\ of collections of
cheatgrass from the Intermountain area, seeds
are ready to germinate when tlun are mature.
No pregermination treatments are necessar\
(Young and Exans 1982). For collections from
the Great Plains and perhaps the Columbia
Basin, seeds may have a brief afterripening dor-
mancy. In contrast, seeds of medusaliead have a
temperature-related afterripening, and germi-
nation will not occur except at cold incubation
temperatures for about 90-120 daws after matn-
ritx (Young et al. 1968). Nelson and Wilson
( 1969) found this (loi-manc\ was eontiolled In
niat(M-ials located in the awn.
The high silica content on the herbage of
medusahead makes the litter xen slow to
decompose. Harris (1965) described Hie chok-
ing accumulations of medusahead litter that
built up for sexeral \ears. We exalnated the
germination of seeds of \arious annual grass
species in medusaliead litter (Young et al. 1971a).
Allelopathy was not suspected, but rather the
ph\ sical holding of seeds out of contact with the
surface of the seedbed. Medusahead seeds ger-
minate \-er\- well without the callus end of the
seeds touching a moisture-supplving substrate,
bi this situation, germination of medusahead
seeds is controlled In' the relatixe humiditx
within the litter and tlie incubation tempera-
ture, which of course influences the relatixe
humidity. The needlelik(\ xitreous carxopses of
medusahead appear hxdrophobic rather than
hygroscopic. Not ouK' can medusahead seeds
germinate under diese conditions, but thex can
be dried until the priman- root is dead; then,
lolloxxing remoistening. a nex\- adxcntitious root
xvill dexelop.
Raxuiond Exans and I demonstrated x\ hat a
great modifxing influence litter coxer can be to
the surface of seedbeds on temperate desert
rano;elands in terms of n^dncing extremes in
temperature and consening moistm-e (Exans
and Young 1970, 1972). (^anopses of
s(juirreltail {Eh/nuis In/strix) are xcn- similar in
moqihological appearance to those of
medusahead. As I xxill discuss later, s(juirreltail
seedlings are one of the fexx- natixc species that
can become established in undisturbed
medusahead stands. Both Tacniaflicniin and
Ely nuts are members of the tribe Triticeae, but
thex" do not share the same genome.
Medusahead populations easiK- exceed 1000
plants per square foot, and thex- are phenotxpi-
callx' plastic enough that a population of 1 plant
per square foot can exceed the seed production
of 1000 plants per square foot (unpublished
research, ARS, Reno, Nexada). Huge seed
banks dexelop in medusahead conunuuities in
the litter and .soil. Medusahead seetl accjuires a
dormancx in the field similar to that of
cheatgrass (see Young et al. 1969). The.se dor-
mant seeds respond to eiuichment of the seed-
bed xxith nitrate and gibberellin (Exans and
^bnug 1975).
Life cycle. — Medusahead seeds can ger-
minate in the fall, xxinter, or spring; and seed-
lings liom all seasons can j^roduce fioxx'ers and
seeds earix in the sunnner. The striking thing
about the medusahead life cxcle is that it
matures from 2 to 4 xveeks later than other
annual grasses. All those famous botanists and
range scientists xx'ho xxere out on the range di.s-
coxering nexx- infestations of medusahead xx'ere
led to the populations In the bright green color
xx'lien all other aimuals in either cisniontane
Galifoniia or the Great Basin xvere broxxn.
R. L. Piemeisel recognized the dominance
of alien plant species in the secondan succes-
sion of disturbed satiebrush communities in the
248
(;rk,\t Basin Naturalist
[\ oluiiie 52
InterniounUiin area (PicMiieisel 1951). Wbrkiiiii;
on the Snake Rixer plains of Idaho (hirin<j; the
1930s. Piemeisel enumerated (k)niinance honi
Russian thistle (Salsola austral is) to tumble
mustard {Sisipnhriuni altissiinuin) to eheat-
grass. Continued disturhanee tended to per-
petuate cheatgrass donn'nance. According to
Piemeisel, the animal species that germinates
first, reaches nuL\imum growth and maturit\
first, lias the capacit\' to withstand crowding,
and has high seed production is the one that will
occup\' and persist in serai sagehmsh plant com-
munities. Piemeisel always noted that no one
species had a clear tlominance on all these char-
acteristics, hut on balance cheatgrass was the
clear winiKM'.
Medusahead contradicts sexeral of
Piemeisels criteria. Medusahead seeds are ini-
tiall\- ck)rmant with temperature-related
afterripening requirements, while cheatgrass
seeds ha\e no such restraints. This works only
for initial establishment because once seed
banks are established with seeds with ac(|uired
dormancy our research indicates that
cheatgrass and medusahead seeds ha\e e(jual
chances of germination with the initial moisture
exent in the tall. Medusahead does take iruich
longer to mature than cheatgrass and perhaps
tumble mustard. Min Hironakaand his students
hax'c conducted a series of excellent experi-
ments comparing the cumulatiye growth cunes
for roots and aerial structures of medusahead
and otlier grasses (Hi ronaka 1961. Hironakaand
Sindelar 1973. 1975). Dr. Ilironaka concluded
from these studies that the comparati\e growth
phenokjgx restricts medusahead to areas with
suiplus .soil moisture alter cheatgrass normally
matures.
Soils
Ha\ni()nd Eyans noted in the 195()s when
medusahead first inxaded Glenn and Colusa
counties in the northern Sacramento \alle\- of
C^alifoniia that medusalunid appeared to be
restricted to clay-textured soils (personal com-
munication). Malloiy (1960) reported on this
relationship at the 1960 meeting of the California
.section of the Societx' for Range Manag(Mn(Mit.
Burgess Kay made the cliilling obsenation that
after a cotiple of decades this relationship disap-
peared and medusahead occupied many sites
with coarser-textured soils (personal communi-
cations).
In the Intermomitain area. Ma\narcl
Fosbergof the Unix ersit\'of Idaho reported that
the medusahead infestations along the Colum-
bia l^ixer in Washington, Idaho, and Oregon
were restricted to clay-textured soils (Fosberg
1965). He suggested that the greater soil mois-
ture-holding capacity of these soils allowed
medusahead to complete its life c\cle.
Building on the work of Fosberg and
Ilironaka, I sampled the plant communities in
the medusahead in\asion area along the western
edge of the Great Basin (Young and E\ans
1970). Medusahead was foimd on the margins
of man\' degraded meatlows where moisture
relationships probabK fa\ored it oyer
cheatgrass. A much larger area of infestation
was sagebnish/grass communities. The sage-
brush communities consi.sted of mounttiin big
sagebrush (Aiiciuisia tridcntata ssp. vaset/ana)
on .soils with sand\ loam to loam-textured sur-
face horizons and often well-dexeloped argillic
horizons. A second series of sagebrush commu-
nities consisted of low sagebiTish (A. arbuscida)
growing on soils wdth clay-textured surface hori-
zons. Harn" Simimerfield (retired soil scienti.st,
Soil Consenation Senice and Forest Senice,
USDA) suggests the low sagebrush soils share
the same development as the big sagebrush
soils, but the surface horizons have been
removed by erosion (personal commimication).
On the Modoc Plateau of northeastern Califor-
nia these two series of plant conununities divide
the landscape about ecjuallv (Young et al. 1977).
In the northern Crreat Basin low sagebnish con-
stitutes onK about lO^f of the total sagebrush
vegetation.
On the western edge of the Great Basin,
medusahead. in nonmeadow situations, is
largely restricted to low sagebrush potential
plant coimnunities. Would this restriction to
cla\ soils change over time as appears to have
happened in cismontane California? Remem-
ber the studies of Raymond Evans tliat showed
competition in the cismontane portion of the
Califoiuia annual grasslands is initiallv for light,
while in chcMtgrass communities of the Inter-
mountain area, competition is oyenvhelmingly
foi-soil moisture ( Pa aiis ct al. 1970. 1975).
WiLDFlHKS
Accumulations of litter, on areas where
medusahead is t\stablished, will bum. McKell,
Wilson, and Kav (1962) had initial results tliat
19921
Ecoi.ocv A\i) M \\ \(:i:\ii:ntof Mkdusaiikad
249
seeiiu^d to iiulicatc that hiirniiiii; \\;is tlic answer
to the control of nu'diisalicad. Ilic idea was to
hui'ii stands wliilo coinpctinij; annnal (2;rassrs
were tulK mature and niedusiiliead seeds were
still in the inflorescences. This stucK' showed
hm^ned seeds would not (germinate. Ilowexer,
the hurned seeds were apparentK incubated at
20 (-", and unburned fresh seed would not ha\e
germinated at that temperatiu-e. We tried a
series of burning experiments on the Pitt Ri\er
bidian reservation and found burning taxored
medusahead (Young et al. 1972 1. We helped
Forest Sen ice range consenationists evaluate
burning treatment on low sagebiiish communi-
ties on the Silver Lake district of Fremont
National Forest in Oregon; the off-season burns
appeared to favor remnant perennial grasses
over medusahead.
Low sagebnish comnumities, because of
lack of herbaceous cov er, are relativelv resistant
to the spread of wildfires. Big sagebrush com-
munities, especiallv those with cheatgrass
undenstories, are ven subject to the spread of
wildfires. Invasion of medusahead into low
sagebnish communities introduces wildfires to
these communities, perhaps for the first time
since they were in pristine condition. Perennial
grass, forb, and shrub cover are all negativelv
correlated with medusahead cover in the west-
em Great Basin (Young and Evans 1970).
Grazixc Preference
It is obvious from the above discussion that
preference bv grazing animals plays an impor-
tant part in the successional dynamics of
medusahead coiiiinunities. One of the few stud-
ies of medusahead palatabilitv was conducted
on the northern coast of California using sheep
in small hurdle plots (Lusk et al. 1961).' Under
the conlinetl conditions of thc^ studv. sheep uti-
lized medusahead when it was green. When
faced with no choice, thev used some herbage
after the medusaliead matured. How nuich uti-
lization of medusahead would occur in temper-
ate desert situations is unknown.
C'heatgrass .stands [)ut a tremendous produc-
tion of grass canopses into a local eco.svstem.
\ertebrate granivores have adapted to this food
source. Savage et al. (1969) showcxl in feeding
trials that Chukar Partridges {Alcrtoris ^raeca)
could not utilize the caiyopses of medusahead
as a food source. These birds are dependent on
cheatgrass seeds in the fall and winter. We do
not know what the iulluence of medusahead
inv asion would be on other granivores. Seeds of
other recently introduced weeds in temperate
ck\sert coimnunities, such as those of barbvvire
Russian thistle {Salsola paulsvnii), are heavily
prey(xl upon by granivores. I f cheatgrass popu-
lations crash because^ of replacement bv
medusahead, what ha]-)jx'us to cheatgrass seed
predators':^
A studv c'onductetlat Washington State Uni-
versitv illustrates that granivore preference
works both ways in plant succession. Bird pop-
ulations prefer the seeds of native perennial
grass species over tho.se of clu^itgrass and
medusahead (Goebel and Bern 1976).
Utilization of medusiiliead bv large herbi-
vores of infested ranges results in increased
incidence of injun from the seeds. Data on the
level of injun' are not available for domestic
livestock and certainlv not available for wildlife.
Control of Medu.saheai:)
Kavdev eloped highlv technical and vorv suc-
cessful control and revegetation techni(jues for
the annual-dominated rangelands of cismon-
tane California using the herbicide [paraquat
( I,l'-dimethvl-4,4' bipvridinium ion) and spe-
cialized seeding equipment (Kav 1963, 1966,
Kay and McKell 1963).
This technique was not successful in the
Intermountaiu an^a because medusahead
[)lants were not susceptible to paracjuat in the
temperate desert environment antl the annuiil
legumes that proved so adapted to (ismontane
California were not adapted to the sagebmsh
environment (Young et al. 1971b'. Ilerbicidal
fallow techni(jues using atrazine (6-chIoro-N-
ethv 1-N '-[ 1 -methv letlivi 1- 1 .3,5.-tria/,ine-2,4-di
amine) or dalapon i2.2-dichl()ropropanoic
acid), and mechanical fallow techni(jues were
developed lor use in the (ireat Basin. Milken
and .Miller ( I9S()) provide a summarv of lierbi-
cidal control measures applied experimentallv
for tlie control of medusahead. A large part of
the area infested with medusahead in the west-
ern Great Basin was never adaptcnl to these
treatments because of surface rock cover that
prohibited tillage or seed-drilling techniques.
The current mass cancellation of federal regis-
tration for uses of herbicides on rangelands and
the failure of federal land management agencies
to a(k)pt the use of herbicidal revegetation tech-
ni(jues have made the use of these techniques
250
Great Basin Naturalist
[\'olunie 52
impossible. Landfornis and soils ol the sites
where niedusahead is spreading into temperate
desert rangelands are eritieal laetors in the eeo-
logical suppression ot this speeies.
Nature of Medusaiikad-infested
Landscapes
The landseape ol" the western Great Basin
where medusahead has in\aded is eomposed of
a series of fairly reeent basalt flows that eom-
prise the Modoc Plateau and the extreme south-
ern extension of the Columbia Rixer Basalts.
Superimposed on the flows are clays from a
Tertiary-age lake. This lake was much older than
pluvial Lake Lahontan, which lapped at the
lower margins of the flows. The old lake left
thick beds of cla\'-textured sediments occasion-
all)' interbedded with diatomaceous earth. The
clay minerals are predominantly double lattice
forms that expand and contract with moisture
content. This exj^iansion and shrinkage has
sorted basalt rock from the buried fk)ws into
giant polygons and pressure ridges until por-
tions of the landscape resemble arctic ice packs
that are black instead of white.
There are a host of topoedaphic situations
within this wilderness that support specific
assemblages of plants; however, the landscape is
characterized by upland areas of residual soils
with loam-textured surface soils that support big
sagebrush and clay-textured surface soils that
support low sagebrush. \'ast, nearK lexel
benches of lake sediments support swirling
mosaics of basin big sagebrush (Arfeinisid
triclentata ssp. trident ni(t) and a recentl\- discov-
ered t}pe of sagebrush, a subspecies of low
sagebnish known as Lah(jntan sagebrush. The
basin big sagebrush occurs in depressions whei'e
erosional products accumulate on .soils with
cla\-textured surface horizons, a ven unusual
occurrence for the Great Basin. The Lahontan
sagebrush communities occur on the lake bed
clay sediments that are veneered with thin
layers of subaerially deposited, coarser-textured
soil.
Wind erosion products accunuilate under
the shnib canopies and, coupled with organic
matter from leaffall, build mounds under the
shrubs while miniplayas develop in the inter-
spaces. Eckert et al. (1989) have described and
experimented with the seedbeds of these
mound interspace situations, particularly the
vesicular crust that forms in tlie interspaces and
limits establishment of perennial grass seed-
lings.
The area of medusahead inxasion in the
western Great Basin is a microcosm where
events in soil and plant ecolog\' that influence
millions of acres in the Intermountain area are
brought, b\ fortuitous combinations of ph\sical
and biological parameters, into shaip focus. In
the medusaliead in\ asion area, lake-deposited
red clay is in obxious disc()ntinuit^•\\^th the thin,
gravish surface soil. \n imdisturbed profiles of
this situation the influence of alle\iation of sub-
aerial deposited material is apparent on the
structure of the clav subsoil, indicating the
antiquity of this process (personal communica-
tion, Robert Blank, soil scientist, ARS, USDA).
Accumulations of medusahead litter change
wildfire characteristics, and the shiiib compo-
nent of the plant communitv' is eliminated. Con-
tinued grazing of medusahead-dominated
grasslands is extremely deleterious on remnant
perennial grasses because of differential grazing
preference. In contrast to medusahead,
cheatgrass is seasonalK' preferred forage spe-
cies, and even the dn' herbage of cheatgrass is
utilized bv li\estock. This dilutes the effect of
grazing as far as the native perennials are con-
cerned. Lack of preference for medusahead
concentrates the effects of herbi\'oi-y. Subaeri-
ally deposited surface soil is extremeh" erodible
once protection of the shnib canopy and its
dependent microph\tic cnist is lost. Loss of the
surface leads to exposiu'e of the cla\' sediments
that then function as Vertisols, shrinking, crack-
ing, and swallowing the surface and reexpand-
ingwith moisture. Medusahead is one of the few
plant species adapted to these Vertisols. Perhaps
some of the soils of these landscapes were
always Vertisols where, in wet \'ears, annual
sunflowers {Helianthus annuus) and turkey
mullein {Erenwcorjms seti^^erus) formed the
onl)' nati\e vegetation. Perhaps excessive graz-
ing conxerted some of these soils to \ ertisols
before medusahead arri\ed. The important
point is that medusahead is actixek attacking
assemblages of natixe vegetation and changing
the physical and biok)gical potential of the sites.
Management of Medusahead
Infestations
It is difficult to rexegetate \^ertisols in desert
enx ironments xxith both seedlings of xx'oodv and
herbaceous species, natixe and exotic. Not only
19921
Ecology and Ma.\ac;i;me\t of Mkdl saiiead
251
establishment but also subsefjuent growth are
problems on these soils despite both tremen-
dous eation exchange capaeit\ and moistnre-
holding capaeitA". The tremendous matrie
potential of these Hue cla\' soils is al\\a\ s suipris-
ing. Moisture is not axailable loi- normal plant
growth when soils still stick to \our boots.
NaTIH \l, SUCCESSION'
Dr. Mill Hironaka suggests that o\er pro-
longed periods perennial seedlings might estab-
lish in medusahead-intested sites, especialK the
short-li\ed perennial grass squirreltail (Hiro-
naka 1963). Dr. Hironaka and his students fol-
lowed this aspect of medusahead succession in
several studies. He demonstrated that squirrel-
tail can establish in medusahead communities,
but he found the perennial grass populations to
be CN'clic. When the squirreltail plants die, the\'
are replaced b\' medusiiliead, not longer-li\ed
perennial grasses (personal communication).
hi the western Great Basin, Dr. Hironaka's
work is borne out b\' gradual increases in
squirreltiiil plant densits' as grazing manage-
ment systems ha\e been implemented. This has
been especially noticeable during the past four
years of extreme drought. Densities of one
squirreltiiil plant per 10 square feet began to
change the aspect of medusahead-dominated
sites, but the fragile nature of this impro\ement
is apparent when bioassay of seed banks shows
250-500 viable medusahead seeds per square
foot (down frcjm 1 000 per square foot before the
drought) and fails to detect am viable squirrel-
tail seeds (unpublished research AHS, USDA,
Reno, Nevada).
As you look at medusahead-infested areas on
the X'ertisols of the western Great Basin, vou
have a nagging thomj-ht that something is miss-
ing. The Lahontan and big sagebmsh comnui-
nities of the ancient lake sediments have as their
most frequent perennial grass Sandberg blue-
grass. This species is completelv absent from the
medusahead stands and is missing from the
stands where scjuirreltail has begun to return.
What factors of seedbed qualitv exclude^ the
native invader Sandberg bhungrass and are the
same factors related to the failure of higher-level
perennial grasses to become established in
squirreltail/medusahead communities?
The striking difference between nativ e and
medusahead communities, other than loss of
shnib canopies, is loss of subcanopv mounds
and microphvtic cmst that covers the mounds
to extend down to mingle with vesicular crust in
the interspaces. The thalloplntic crust of
mosses, lichens, and livenvorts is obviouslv
gone, and we can onlv speculate on the fate of
tlie microscopic crust of algae, fungi, and bacte-
ria. Prolonged medusahead dominance mav
decrease populations of nncorrhizae spores in
the soil and thus inflnenc-e growth of artilicialK
established perennial seedlings (personal com-
munication, Jim Trent, soil microbiologist. .\liS,
USDA, Reno, Nevada).
Specific plant pathogens, developt^d and
marketed bv biotechnological companies, mav
have a role in range weed control. Perhaps a
Fusarittni species exists that would be highly
specific for medusahead (personal communica-
tion, Joe Antognini, National Program scientist.
Weed Science, ARS, USDA).
Taxonomists and greneticists who have
worked with medusahead have commented on
how variable individual collections mav be.
Common garden studies have shown this to be
tnie for collections from the American \\'est
(McKell, Robinson, and Major 1962, \bung et
al. 1971b). We found, in common garden stud-
ies, a collection from northern California that
matured 4 weeks earlier than the average for
other collections or on or before the maturitv for
cheatgrass. As medusahead evolves, we have vet
to see the limits of its potential on the vxesteni
range. The recent discoven' of medusahead in
Utah illustrates that portions of the eastern
Great Basin have the potential to be iii\ adcd bv
this weed (Horton 1991).
Literati HE Cited
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Poorest, \Mldlife and Range Experiment Station, Uni-
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medusahead into the Cweat Basin. Weed Science 18:
89-97.
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Research Service, USDA, Oakkuid, C';ilifornia.
Yoi Nc j. A.. R. A. Evans, and R. E. Eckeht. Jk 1968.
(Ti'rmiuation of mechisahead in response to tempera-
ture and aftenipening. Weed Science 16: 92-95.
. 1969. Population cKnamics of downx' brome. Weed
Science 17: 20-26.
YcUNG. J. A., R. A. ExANS, and B. L. Ivxv 1971a. (iermina-
tion of carxopses of annual grasses in simulated litter
Agrononix' [ounial 63: 551-555.
. 1971b. Response of niedusahead to paiaijuat, |our-
iial of Range .Management 24: 41—43.
Y(U NO |. A.. R. A. Ex'ANS. ;uid J. M.^JOK 1977. Sagebrush
ste])pe. Pages 76,3-797 in M. G. Barbour and |. Major,
eds.. Terrestrial xcgetation ol ( !aliloi iiia, John Wllex &
Sons. New York.
YoLNc; |. A. H. \, F,\ \\s and j. Hi HsisdN 1972, Influence
ol repealed animal buruiiig on a nu'dusalieatl conimn-
iiitx, |ouniai ol Range Management 24: 451-4.54.
Rrccivnl 2:Ulai/ Um
Arrrpird 22 Jinir I9h)2
(ireat Basin Naturalist .")2i.'Vi. pp. 25o-2Hl
R(X)ST SITES USED BY SANDHILL CRANE STA(;iNG
ALONG THE PLATTE HI\ EK. NEBRASKA
Bra(lle\' S. NOrliiiii; , Staiilcx 11. .VikIitsoii , and Wiune A. Iliihcrf
.•\bstiu(X — We iLssessed the influence of water depth, extent of unobstructed \ie\\\ and huiuan disturbance features on
use of roost sites h\ Sandliill Crimes along the Platte Ri\er, Nebraska, during .spring niigraton stopox er. .Xt'riai photos tiikcn
near dawn were used to determine areas of flock use and habitat a\ailabilitv in four sample reaches, and measurements
were made on the ground at flock roost areas. In general, depths of 1-13 cm were used bv s;uidhill cranes in greater
proportion than those a\ailable. Exposed sandbars ;uid depths >20 cm were a\()id(^d, wliile depths of 14-19 cm were u.sed
in proportion to their a\ailal)ilit\-. Sites 11-50 m from the nearest \isual t)bstruction were used significantly greater than
their availabilit); while sites ()-4 and >50 m from \isual obstnictions were avoided. Sandliill Cranes avoided sites near pa\ etl
roads, gra\el roads, single dwellings, and bridges wjien .selecting roost sites: howewr. thex' did not appear to be disturiied
b\ private roads, groups of residential buildings, graxel pits, railroads, or electrical trausniissioii lines.
Kc'i/ words: Sdiulliil! Crane. Cirus canadi-nsis, river rocistn. habitat .sclcclioii. water di])th. (li.slurhanrc. saiulhars. Platte
Hirer
Tlie impact of water re.sourcc^ clex'elopnieiit
on the Platte Rixer i.s well described
(Kroonemever 197S, Williams 1978, Eschneret
al. 19S]. Kii-cherand Kariinger 198L U.S. Fish
andWildlite Senice 1981. Krapn 1987, Sidle et
al. 1989). The major impact lias come from
irrigation projects along the North Platte Ri\er
(Krapn et al. 1982), which remo\e approxi-
mately 70% of the annual How of the Platte
Ri\er before reaching sonth central Nebraska
(Krooneme\er 1978). Concomitant with chan-
nel shrinkage, woocK' vegetation has encroached
on thonsands of hectares of former channel
area, contributing to further changes in channel
features and altering habitat for numerous .spe-
cies of migraton- birds in tlie Big f^end Reach of
the Platte River in Nebraska (U.S. ImsIi and
Wildlife Sei-vice 1981). The Big Bend l^eacli of
the Platte Ri\er in Nebraska is an area of
importance to numerous sjiecies of migraton
birds ol the Central I^1\\\a\ ( If.S. Fish and Wild-
life Senice 1981 ).
This area is an important stojioxcr area lor
most of the midcontin(mt population of Sandhill
Cranes (Cms i-ditddciisis^ i 4(H ).()()( )--6( )().()()()
birds), which roost in the riwr and feed in
neadn com fields (Krapn et al. 1981, Krapn
1987). The endangered Wliooping Cj-ane (C^,
(iincricdiui) also uses the area during migration,
and the tlu'eatened Bald Eagle {Haliacctus
lei(coccpJialiis) is a common winter resident
(U.S. Fish and Wildlife Senice 1981). The area
is also important habitat for the endangered
interior population of Least Tern {Sfcnia aiitil-
lantm) and the threatened Piping Ploxer
(Charadriiis niehxhis), both of which nest along
the Platte Ri\er (U.S. Fisli and Wildlife Senice
1981, Sidle etal. 1989).
Considerable attention lias been gi\en to the
impact of changing channel conditions on the
midcontinent population of Sandhill Cranes
{Gnis canadensis) that congregate along the
riverfront earl\' March to mid-,\pril during their
animal spring migration (Lewis 1977, Krapn
1978, U.S. Fish and Wildlife Senice 1981).
During this time approximatcK 4()(). ()()() Sand-
hill Cranes use tins an^a while euroiile to their
breeding grounds in (Canada, Alaska, and eastern
Siberia (U.S. F^ish imd Wildlife Senice 1981).
In Nebraska various facets of Sandhill Crane
roosting habitat re<juirements ha\e been stud-
ied (Frith 1974, Lewis 1974, U.S. Fi.sh and
W ildlife Senice 1981, Krapn etal. 1982. 1984).
I i()\\(n cr. these studies ha\(^ not considered the
infhience of habitat axailabilitA in ndation to
habitat use. The i)un)ose ot this stud\ was to
W'yoinini; (;o()ptTati\e Fish unci W'ikllirc Hescarch Liiit. Bo.\ :!166. L'niversih' Station. Laramie, Wyoming S207r
2o.i
254
Ghi:at Basin Naturalist
[X'olunie 52
Reach 1
Reach 3
â– Reach 4
Intensive Study Area ^1^*
1 2 3 4 5 km
Fiii. I. StiuK sites in the Platte Hi\'er, Nehraski]
(Ic'tcniiiiic tlic influence oF habitat axailahilitx,
as well as habitat use, on the selection of roost
sites b\' Sandhill Cranes.
This stiicK' was designed to assess the infhi-
ence of three tvpes of habitat features on roost
sites used by Sandhill Cranes: (1 ) water depth,
(2) magnitude of unobstructed \iew. and (3)
disturbance features.
STrnv Ahka
The study area is locatcxl in south central
Nebraska in Hall and l^ufTalo counties in die
eastern halfofthe Big Bend Hcnich of the Platte
Kiver. It encompasses a 36-km stretch of the
Platte Kiver beginning 4 km west of Shelton to
(^rand Island (Fig. 1). All held measurements
were in four 1 .6-km reaches along the main
channel of the Platte Ri\er.
Spiing precipitation in Nebraska contributes
to the Platte Kiver Basin flow, but most of the
flow is derixed from spring runoff that originates
as snownu'lt in tlie Kocky .Mountains (E.schner
et al. I9(S1). Spring runoff flows into both the
North and South Platte ri\ers, which flow nortli-
east and southeast, resi)(>ctivel\. across the
Cireat Plains to their confluence near North
Platte, Nebraska.
The stnd\' area is characterized b\ numerous
braided channels interspersed with imxege-
tated sandbars that fre(juentl\" shift. Most of the
land within and adjacent to the stuch' area is in
private ownership. Land use in the area is pre-
dominantly agriculture and includes approxi-
mately 60% cropland (mostly com), 5% tame
pasture, 20% nati\e grassland, and 15% riparian
woodland (Keinecke and Krapn 1979).
The riparian woodland comprises eastern
Cottonwood (Poptthis deJioidcs) forests with
(k)minant understoiA species of red cedar
(Jiinij)rrii.s lir^inidiia) and rough-leaf dogwood
(doniu-s (Innnmotidii). On low islands and \eg-
etat(^(l sandbars, peach-leaf willow {Salix
aini/i^ddloidcs). ccnote willow {S: cxig^nal), and
indigo bush (Aniorplia fnitirosa) are the domi-
nant species (U.S. Fish and Wildlife Senice
1981, Currier 1982).
MKTHODS
.Aerial photographx was used to determine
flock locations and delineate flock boundaries of
19921
CiiWE Roost Sites
roosting Saiulliill Cranes along a 36-kni stretcli
ot the Platte Hi\{M-. Photograpln" was restricted
to mornings with less than 10% cloud coxer and
ceilings abo\e 975 m. Flights were begun 30
minutes ht^fore sunrise ])ecanse ol the need to
pli()t()gra[)h Sandhill C'ranes before the\' lea\e
the roost in earl\' morning. Light was adequate
to piMinit photograph\' 10-15 minutes before
sunrise.
A Hasselblad 500 P.L, 70-mm camera was
used to photograph the stud\' area. The camera
was mounted in a standard camera hatch in a
Cessna 172 fixed-wing aircraft and was
equipped with an SO-mm focal length Zeiss lens.
Exposures were made at 1/60 and 1/125 second
at f2.8 using Kodak Tri-X 640 AFS Aerographic
film. The camera was equipped with a 70 expo-
sure back loaded with 5.5 m of film allowing <S0
ex[)<)sures.
The aircraft was flown at approxiiiiatcK 140
km/hr at an initial altitude of 790 m aboxe
ground lexel for the first two flights. During the
last two flights the altitude was increased to 910
HI al)o\e ground le\el. These altitudes provided
a 0.48-km" and 0.64-km" coxerage on each
frame, respectively. Frame rate was controlled
1)\ an intenalometer, calibrated for 309^ oxer-
lap, to pnnide continuous photographic co\er-
age of the study area.
Shortly after each flight the film was custom
pr()ces.sed by hand agitation in a single solution
tank, xaning time and (kn'eloper temperature
to obtain optimum dexelopment. Approxi-
matcK 150 frames were e.xposed from each
flight. Frames were examined under SX magni-
fication to identifx crane flocks and were
enlarged to 41 X 51 cm ( 16 X 20 in) and printed
on Kodak PoK' contract RC paper. Processed
photogiaplis wen^ stored for later anaKsis of
\isual obstructions and disturbance features.
Each of the tour 1 .6-km reaches was marked
on both sides of the rixer bank with 16. 1-nr
markers mack' of white cloth. The markers,
placed 100 m apaii at the edge of the rixerbank,
were positioned in such a wa\' that markers on
tlie opposite sides of the channel were parallel
to the channel. The markers enabled accurate
scale measurements to be taken from photos
and proxided position reference for tiansects
across the channel xx'hen sampling water depths.
Aerial photographs coxering each reach xxere
used to determine the position of transects
through flocks. Transects were positioned so
that each flock studied on a photo x\as dix ided
into general areas of ecjual size with txvo to fix-e
transects depending upon flock si/.e. A flock x\'as
(k'fined as a continuous distribution of birds or
an aggregation of birds sjxitiallx' independent of
other birds separated bx a distance >2() m.
Flocks usnallx' occurred in configurations that
a[)pear{>d distinct from other flocks in the xic initx.
After transects xx'cre located on [)hotograplis,
thex" xxere measured and laid out on the ground
in relation to marker locations using \inx 1 flag-
ging placed on each side of the channel. Water
depths xxere measured to the nearest 3 cm at
3-m inten als and plotted on acetate oxerlaid on
aerial photogra]:)hsx\ith delineated flock bound-
aries. Width and depth data xx^ere combined to
gix e mean estimates for each of the four reaches.
Each 1.6-km reach xvas sampled as soon as
possible after each flight, alxvavs xvithin three
dax's. Staff gauges xxere placed in each area to
measure anx* changes in xxater lexel between the
time each reach xx'as photographed and the time
it xvas sampled. Detectable changes in xxater
lex'el xx^ere recorded and used to con-ect dt^pth
distributions.
Discharge xvas measured on each flight dax'
in close proximity' to the study areas folloxxiug
the techni(jue of Buchanan and Somers (1969).
Contact prints xxere made from each roll of
film. Indixidual frames xx'ere cut out and glued
onto posterboard to form a mosaic, proxiding a
continuous coxerage of the rixer channel. Scale
was determined bx' comparing bridge segments
and transect locations on the contact prints xxith
measurements of these locations niadc^ on the
ground. Scale e,stimates were made along 2- to
3-km segments of rixer Photograph scales
ranged froiu 1 :8,681 to 1:1 0,334 for the first txxo
flights, and 1 : 10,595 to 1:11, 857 for the last txx'o
flights.
A binocular zoom iuicro,scope (1-4X) xx^as
used to ick^ntifs flocks and delineate flock
boundaries on the contact prints covered xxith
ac(Tatc. Flocks wcic delineated and subse-
(juentK nmubered on the acetate oxerlax'S on
contact photos. The distance from the edge of
each flock to the nearest xisnal obstniction x\as
measured to the nearest 0.5 nun on the photos
(ground distance = ^6 m) using a drafting cal-
iper \ isual obstructions inclnck'd xegetation, a
rixer bank, or anx otlier 'xisualK solid" object
>1 m in height.
Kandom points were plotted on contact
photos to (\stimate the featm-es of ax ailable hab-
itat. Ranck)m points xxere determined bx a .series
256
G H EAT B AS I N N ATU R A LI ST
[\ bluiiie 52
of random numbers identifying point coordi-
nates on gridded overlay coxering contact
prints. Points outside the rixer channel were
discarded. Onl\- random [X)ints located in water
were u.sed because points on sandbars, islands,
or the ri\er bank were not considered poten-
tiall\- usable roosting habitat. A total of 339
random points within the ri\er channel were
identified on the contact prints. Grid squares
were 1.25 mm" to ensure a representative
sample of locations on the ri\ er. As with flock
locations, the distance from each random point
to the nearest \isual f)bstruction was measured
on the photos t(; the nearest 0.5 nun using a
drafting calipei-.
For analvsis oi human disturbance features,
flock locations and random points along the
entire 36-km stud\' area were transferi-ed from
70 nun contact prints to acetate overlays of color
infrared aerial photographs (scale 1:25,595)
using a zoom transfer scope. The photographs
taken in April 1989 were obtained from the
Bureau of Reclamation in Cirand Island,
Nebraska. Distances were measured from the
edge of each flock and individual random points
selected b\' placing a card over the photograph
to the nearest human disturbance features.
These features included pa\ed roads, gravel
roads, prixate roads, urban dwellings, single
dwellings, railroads, connnercial development,
highwa\s, and bridges. Distances were mea-
sured to the nearest 0.5 mm on photos (ground
distance = 13 m) with a drafting caliper.
Data AnaK sis
FrequencN' histograms were plotted for mea-
sured distances from the edge of a flock and for
random distances to the nearest visual obstruc-
tion and disturbance features. Frequencv distri-
butions were plottc>d for axailable and used
selected water depths. Fre(jU(Mic\- distributions
of available and used selected water depths for
each 1 .6-km reach were determined bv combi u-
ing flock data for each reach for a given flight.
Available depths were defined as all depth mea-
surements taken along a transect, and used
depths were those depths where birds were
present along a tran.sect. Habitat selection v\as
computed by dividing the proportion of habitat
u.sed within a depth intenal bv the proportion
of depths available in that same intenal (Bovee
1986). Depths used less than their availabilitv'
were defined as being av oided, while those used
more than their availabilitv were defined as
being selected. Habitat avail abilitv, use, and
selection were summarized within reaches,
across flight dates, and from data pooled across
reaches and flight dates. Data were pooled to
generalize the selection of depths over the
course of the sampling period.
The chi-sqiuire of homogeneity (Marcum
and Loftsgaarden 1980) was used to test
whether differences existed between the distri-
l)utit)n of random points and those locations
used bv Sandhill Cranes relative to visual
obstructions and distiu'bauce features. It was
also used to determine if there were differences
between the proportion of used and available
water depths among and within reaches. Confi-
dence intervals were calculated using the
Bonferroni Z-statistic to test which intenals
within the distributions were used more or less
than exjDected (Byers et al. 1984). Differences
between selection functions were tested wdth a
Z-test. Analysis of variance (ANOVA) was used
to determine if visual obstructions had an effect
on the disturbance potential created by various
tvpes of disturbance features. Significance for
all statistical inferences v\'as P < .05.
Results
A total of four sampling flights were made:
one each on 21 and 31 NIarch and 4 and 10 April
1989. A total of 285 flocks were identified
during the four flights. Folkming the flights, 20
flock sites vwre selected and sampled and a total
of 5109 depth measurements were recorded in
the field.
Sampling areas. — Reaches I and II were
the narrowest, with mean channel widths of 254
m (range = 225-319 m) and 249 m (range =
241-263 m), respectivelv, while reaches III and
I\', located upstream, were wider. Reach III had
a mean channel width of 413 m (range = 387-
440 m), while reach W had a mean channel
v\idth of 357 m (range = 296-445 m).
Reaches I and II had similar discharge (17
mVs), while reaches III and l\ had greater
values (27 and 44 m Vs) on 21 March (Table 1 ).
Discliarge in rcnich III was tvpicalK tv\ice as
high as reaches I and II. Reach 1\' had the
highest discharge of the four reaches, often
three times greater than in reaches I and II
(Table 1). Reaches I, II, and III were located in
a braided portion of the surface along the south
chaimel and coutaineil onlv partial river flow.
19921
CiuxE Kous'i' Sites
25'
TaBI.F, 1. Disc-lianji;e in cubic meters per second (m ) for saiii|ile reaclu's on tlifTerent i'\\'j}]\ dates alon<4 tlie Flatte Ui\er
Nebraska, during spring 19S9.
Flight date
Reacli I
Reach II
Reach III
Reacli I\'
21 March-'
31 March
4 April
10 April
17.4
11.1
10.6
7.9
17.4
10.6
7.9
27.5
18.6
13.7
44.6
32.1
2S..S
21.7
â– 'Discliarge.s for all reaches on 21 March were nieasiircd cm 24 Maich. TIids. a lhii'i'-ila\ la;; |)rrio<! fxisted betxM'cii the lime each reach u'a.s flouTi and (he tin
each reach w;is measured for discharge.
Reach 1\ was located aloiiL:; tlic iiiaiii cliaiiuci
and contained total ri\er flow.
IlARIT\T WAILABILITY. — The distribution
of a\ailal)le water depths differed among
ivaches. On 21 March 1989, 82% of tlie avail-
able habitat in reaches I and II consisted of
depths 0-25 cm. In contrast, 53% and 66% of
the axailable habitat in reaches III and I\',
respectixelw consisted of depths 0-25 cm.
An increased freqnenc\ of shalhnv depths
(0-19 cm) and a decreased frequenc\- of deeper
depths (>20 cm) occurred o\er the stud\
period. This di\ision is made because cranes
seldom used depths greater than 20 cm. The
increase in exposed sandbars (depth = cm ) was
most pronounced in reaches I and II, which
showed increases of 13% and 11%, respecti\el\ .
Reaches II and III showed increases of 12% and
19%, respectiveK', in axailable depths of 1-4 cm
betAveen the first and last flight. Reaches III and
W showed decreases of 10% and 7%, respec-
tixcK; in depths >38 cm for the same period.
During the stiuK period a progressixe decrease
in discharge occurred (Table 1), causing more
shallow areas (0-19 cm).
HabiT.AT use. — Frequency di.stributions of
roosting habitat use hv cranes indicated the
liighest proportions of used water deptlis were
from the 1-4 and 5-7 cm increments. This range
ot water depth accounted for 65% of the mea-
sured depths. There was no discernible \'aria-
tion in the frequency of water dejiths us(^d
among the four reaches.
There was a small, but significant, difference
in tli(^ distribution of depths used bet^\•een the
beginning and end of the study period (F < .05).
Depths of cm showed a significant decrease in
use, while depths 20-22 cm sliowed a signihcant
increase in use {P < .05). The data showed a
significant difference between the distribution
of used and available water depths for all foui-
sampling periods (P < .001). Sandhill Cranes
used progressiyely deeper water depths as the
stud\ season progressed. Depths >20 cm were
used significanth- less than expected during the
first flight; but, b\- the last sune)', only depths
>29 cm were used less than expected {P < .05).
Depths of cm were generally avoided bx
Sandhill (>ranes during the last two sur\e\ s and
were used less than woidd be expected bx
chance (P < .05).
Habitat selection was assessed using both
habitat use and axailabiHtv' data for specific
water depths. The most frecjuentK occurring
depth intenals for which selection occurred
were 5-7 cm, fcjllowed by 1-4, 8-10, 1 1-13, and
14-16 cm in decreasing order of preferenc-(\
Visual (obstructions. — There was a sig-
nificant difference between the distribution of
flock locations and random points reiatixe to the
distance from the nearest \isual obstruction
(F < .001). Proportional u.se of sites 0-50 m
from the nearest \isual obstruction was signifi-
canth' greater than a\ailabilit\ (F < .05), while
sites >50 m from a \isual obstruction were
avoided (F< .05).
The 0-25 m interval was di\ided into six
increments: 0, 1— f. 5-10, 11-15. 16-20. and
21-25 m. There v\as a significant difference
lK^t^\'e(^u the distribution of flocks and random
point distances (F < .001 ). Sites as close as 5-10
m from the nearest visual obstruction were us(^d
by Sandhill Cranes. Onlv sites — f ni from a
\ isual obstruction v\ere avoided (F < .05), while
sites 1 1-25 m from a visual obstruction were
used more than expected (F < .05).
Msual obstructions v\ere divided into three
categories: (1 ) unvegetated bank. (2) \egetated
bank, and (3) xegetaled island. There v\'ere no
significant differences in the distribution of dis-
tances b(^h\een an unvegetated and vegetated
bank, but there were significant differences for
tlie distribution of distances between vegetated
banks and vegetated islands and between
unvegetated banks and vegetated islands (F <
.005). Sandhill Cranes roosted a mean distance
258
Cheat Basin Naturalist
[\'()]unie 52
of 45 Ml from umegetated banks. 50 in lioni
wgetated hanks, and 27 ni from v{>o;etat('d
ishmds.
Channel width. — ^There was a ndation-
ship between the niinimnm unob.stnicted chan-
nel width and distance to the nearest \isiial
obstrnction. The distance to the nearest \'isnal
obstrnctions was ahuiction of less than one-half
die niininnnn unobstrncted channel width.
There was a significant difference between
the distribntion of flock locations and random
points relati\e to minimnm nnobstructed chan-
nelwidth (P < .005). Sandhill Cranes used chan-
nels 100-200 ni wide in greater proportion than
tho.se generalK a\ailable. Channels narrower
than 100 m were axoided, while those >200 m
wide were used in proportion to their axailabil-
ih'. The mean minimum unobstioicted channel
wudth used by roosting flocks was 196 m (range
= 34-445 m).' Nearly 100% of the flocks we re hi
channels with a minimum unobstructed chan-
nel width of >50 m, and over 97% and 80% of
the flocks were in channels with a minimum
unobstructed width of >]()() and >150 m.
respectixeK'. The mean relative flock size (sui-
face area) was 3883 m~ (range = 19-55,354 nr).
There was no relationship between flock size
and minimum nnobstructed channel wi 
