HARVARD UNIVERSITY
Library of the
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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\ (■ 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: .■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,
■'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
■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.
Internationally Revue gesamten Ihdrobiologie 62:
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
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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
insects from a Quebec stream. Verhandlungen der
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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sand areas of a lowland river. Cixnadian Journal of
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(. ■( 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.
■^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
■ 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 (■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.
■ 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:
■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
■X
ACTUAL
■*
*
■ * +*
** *
**
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 * *
*
■
\.
i
-
*s<
* *
. ♦--<
* + ^\^
* *
* ^s,
>^* *
*
.
* *
^<
*
■
*
">
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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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
of the Grand Riyer/Cedar River, Sioux, and Ashland
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of ,^^erto,V ret <>nundcrstor^■ Lod^ plants. 1'^ treatments on regeneration of ,u.t.v.woodIanck^
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M^rt^ . c^ul s ^o icL Ji^vu-posuu^^ ..n Uu.sK. 1). W, and W W Pa.wtneh. 1985. Catde di.ts in a
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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 "■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.
Literature Cited
Ai.i.iA V. S, mul S. E. Mkvkr 1990. Temperature
n-qiiirnieiits lor seed germination of three Pcnstcmon
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mancy cycle in buried weed seeds: a continuum. Bio-
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f^RONyuisT K., K. H. Holmgren, N. H. Holmgren, J. L.
Re\ KAL and P. K. Holmgren 1984. Intermountain
flora. \'ol. 4. The New York Botanical Garden, New
York.
Fewer, M. 1985. Seed ecolog\-. Chapman and Hall,
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Franklin, B., and R. Tavlorson 1983. Light control of
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H. Mohr, eds., Photomoiphogenesis, Encyclopedia of
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Ghabe, D. F., ED 1970. Tetrazolium testing handbook for
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Cii TTEKVLAN, Y. 1982. Phenot\pic maternal effect of photo-
period on seed germination. Pages 67-79 in A. A. Khan,
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Hendricks. S. B., and R. B. T.wlorsun 1978. Depen-
dence of phvtochrome action in seeds on niembnuie
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1968. Restoring big game range in Utah. Utah State
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nation beha\ ior: the ontogen\- and e\'olution of somatic
poKnnoiphism in seeds. Americtui Naturalist 124: 1-16.
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through fifteen \ears of warehouse storage. Great
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.SiEV ENS, R., and S. B. MoNSEN 1988. "Ced;u' palmer
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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 .■)
none
—
r>averS
24,190 + 4.3(X)
2800
.A-2.373
Kaetan ( -avc
I,a\er i
1 1.220 - .■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
■'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
■"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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Bi-Twcoi HT j. L.. T R. \'a\ Devemm-h and R S.
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19921
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last 40.000 vcars of biotic change. Universitvol .Arizona
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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
■ 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-
■ 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
■ 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-
■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
(■\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(
■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.
(■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 ■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. ■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
subscribers, $15 for student and emeritus subscriptions, and $40 for institutions (outside the United
States, $30, $20, and $45, respectively). The price of single issues is $12. All back issues are in print and
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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'
1^^
^&-
.^^
\^-®
a^
.6^^.
Q
,^
%3:
9.e'
se^
^o^^
^vgS
<^':
.^^.
O.
,Q Q
^.:
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
!'))(■ 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>.^
■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
■D
>^
X
o
CD
03
X
-50
-70
-90
O -110
-130
■150
-| r
A
^'-E^'
■ 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.
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