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HARVARD    UNIVERSITY 

Library  of  the 

Museum  of 

Comparative  Zoology 


MCZ 
LIBRAR\7 


X  Jl  g  J'J'^  0  5  1992 

HARVARg 

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. 

The  Great  Basin  Naturalist,  founded  in  19.39,  is  pvil)lished  quarterly  by  Brigham  Young 
Uni\ersity.  Unpublished  manuscripts  that  further  our  biological  understanding  of  the  Great 
Basin  and  surrounding  areas  in  western  North  America  are  accepted  for  publication. 

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issues,  or  other  business  should  be  directed  to  the  Editor,  Great  Basin  Naturalist,  290  MLBM, 
Brigham  Yoiuig  University,  Provo,  UT  84602. 

Scholarly  Exchanges.  Libraries  or  other  organizations  interested  in  obtaining  the  Great 
Basin  Naluralist  through  a  continuing  exchange  of  scholarly  publications  should  contact  the 
Exchange  Librarian,  Harold  B.  Lee  Library,  Brigham  Young  Ihiiversity,  Provo,  UT  84602. 

Editorial  Production  Staff 

JoAimc  Abel Technical  Editor 

C;ar<)l\  u  Backman Assistant  to  the  Edih)r 

Natalie  Miles Production  Assistant 


Copyriylit  ©  1^W2  l)y  BriKliam  Yoiini;  Univt-rsitv 
Ofllci.il  piililication  dati;  22  Mav  1992 


ISSN  0017-3614 

.5-92  75055407 


LIBRARY 
JUN  0  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. 


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McArthur,  E.  D.,  A.  P.  Plummer.  and  J.  N,  D.wis  1978b. 
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M<  \RTiiUR.  E.  D.,  and  B.  L.  Welch.  1982.  Growth  rate 
differences  among  big  sagebnish  (Artemisia 
tridentata)  subspecies  and  accessions.  Jounnil  of 
Riuige  Management  35:  396-401. 


8 


Great  Basin  Naturalist 


[Volume 


Mc.Ahtiiuk.  E.  D.,  B.  L.  Wklcii,  and  S.  C.  Sandehson 
ly.SS.  Natnrd  and  iirtificial  liybridization  between  l)ig 
sai^ebnisli  (Artemisia  tridcntata)  subspecies.  Journal  of 
Heredity  79:  26S-276. 

McKl-.i.i..  C;.  M.  19S9.  The  biolo<f\  and  utilization  ol'slinibs. 
Academic  Press,  Inc.,  San  Diego.  California.  656  pp. 

McKkll.  C.  M.,J.  R  Bl.msdki.i..  and  J.  R.  CooiiiN.  TKCii 
Mc:.\i.  KDS  1972.  Wildland  slirubs — their  biolog\-  and 
utilization.  USDA  Forest  Ser\ice  (k'nerai  Technical 
Rej-xjrt  l.\T-l.  Interniountain  Forest  and  Range 
Exixriment  Station,  Ogden,  Utah.  494  pp. 

Mkvkk.  S.  E.,  E.  D.  McAbtiilik.  and  G.  L.  Johcensen 
1989.  \';u-iation  in  germination  response  to  tempera- 
tin-e  in  nibberrabbitbrush  (Clin/s<)thai)i)ius  nauscosiis: 
Asteraceae)  and  its  ecological  implications.  American 
journal  of  Botan\  76:  981-991. 

Meyeh.  S.  E.,andS.  B.  MoNSEN  1991.  ilahitat-correlated 
variation  in  mountain  big  sagebrush  (Artemisia 
tridentata  ssp.  vnseyana)  seed  germination  patterns. 
Ecolog)'  72:  739-742. 

MONSKN.  S.  B.  1975.  Selecting  phmts  to  rehabilitate  dis- 
tmbed  iireas.  Pages  76-90  in  R.  S.  Campbell  tuid  C.  H. 
Herbal,  eds..  Improved  range  plants.  Society  for  Rtmge 
Miuiagement,  Denver,  Colorado. 

MoNSEN.  S.  B.,  and  D.  R.  Ciikistensen  1975.  The  use  of 
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MoNSEN  S.  B.,  and  R.  Stexens  1985.  'Painte' 
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MooHE.  T.  B.,  R.  Stexens,  and  E.  D.  McMrniiK  1982. 
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cleaning,  and  storing  seed  of  western  shnibs.  Pages 
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Pum.mfh  a.  R,  et  al.  1956-1971. 

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A.  P  Plu.mmeh,  R.  L.Jensen  ;uid  H.  D.  .Staimev  19.57. 
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A.  P.  Plu.mmer  II.  D.  Staim.ev.  and  1).  H. 
ClIRISTENSEN  1959.  Job  completion  report  lor 
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A.  P  Plummek.  D.  R.  Ciiuistensen.  and  S.  B. 
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1963.  Job  completion  report  for  game  forage 


revegetation  project  W-82-R-8.  Bulletin  63-1 1.  Utah 
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1964.  job  t()ni])l(^ti()n  report  lor  game  foragi' 


rexi'getation  project  \\-82-R-9.  Bulletin  64-14.  Utah 
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pp. 

196 1.    Highlights,    results,    and   accomplish- 


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\.  P.  Plummek.  D.  R.  Ciinisi  i:\si a  and  R.  Stexens 
1970.  Highlights,  results,  and  accomplishments  of 
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RoTiL  A.  H.,  Jk  .  and  .\.  P.  Plu.mmek  1942.  Increasing 
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RuDOLE  P  o!,  K.  W.  Dorman,  R.  C;.  Hirr  ant!  A.  V. 
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10 


Great  Basin  Naturaijst 


[\blunie 


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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 
0 

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, 0 


4 1  0.3 

15.546  95.0 

769  4.7 

16.356  100. 0 


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 

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 

0.3 

This  stuck 

16.4 

0.2 

3.6 

lis 

0 

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 


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tion (Knaniics  in  a  guild  of  periph\ton-grazing  insects 
in  a  southern  Appalachian  stream.  l']colo_g\' 64:  12.36- 
124S. 

(;i;\v  L  j.  1981.  Species  comjxjsition  and  life  histories  of 
aquatic  insects  in  a  lowland  Sonoran  desert  stream. 
\merican  Midland  Naturalist  106:  229-242. 

11  win.roN,  A.  L.  1969.  On  estimating  annual  production. 
Linmol<)g\'  and  Oceanograph\'  14:  771-782. 

IIaiU'ER.  p.  p.  197.3.  Emergence,  reproduction,  and  growth 
of  setipalpian  Plecoptera  in  southern  Ontario.  Oikos 
24:94-107. 

.  1978.  Variations  in  the  production  of  emerging 

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- 
duction of  psammophilous  Chirononiidae  in  shifting 
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- 
duction and  drift  of  the  stream  amphipod  Gainiiiants 
pseiidohtnnacus  in  \'alle\  Creek,  Minnesota.  Lininol- 
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**^ 

0  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. 

Literature  Cited 

B.M.i,  (;.  H.,  and  D.  j.  Hall  1967.  A  chistering  technicjne 
for  summarizing  nuilti\ariate  data.  Beha\ionil  Science 
12: 153-155. 

BoLDT,  C.  E.,  D.  W.  Urksk.  and  K.  E.  Sknekson  1978. 
Riparian  woodlands  in  jeopiirdy  on  northern  High 
Plains.  Pages  184-189  //)  Strategies  for  protection  and 
management  of  floodplain  wetlands  and  other  riparian 
ecosNstems.  USDA  Forest  Senice  General  Technical 
Rejxirt  WO-12,  Washington,  D.C. 

Bhaov  N.  C.  1974.  The  natnre  and  properties  of  soils.  Sth 
ed.  Macmillan  Pnblication  Co.,  Inc..  New  York.  639  pp. 

Cl  I  AH  LEY,  J.  L.  1977.  Mineral  cycling  in  rangeland  ecosys- 
tems. Pages  21.5-256  in  R.  E.  Sosebee,  ed.,  Rangeland 
plant  physiolog)-.  Societ)-  for  Range  Management, 
Den\er,  Colorado. 

Dauben.mike.  R.  1959.  A  canopv-coverage  method  of  veg- 
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Donahue,  R.  L.,  R.  W.  Miller,  and  j.  C.  Shickluna. 
1977.  An  introduction  to  soils  and  pkuit  growth.  4th  ed. 
Prentice-Hall  Inc.,  Englewood  Cliffs,  New  Jersey.  626 
pp. 

Girard.  M.  M.,  H.  Goetz.  mid  A.  J.  B]uc;stad  1987. 
Factors  influencing  woodlands  of  southwestern  North 
Dakota.  Prairie  Naturalist  19:  189-198. 

Hansen,  P.  L.  1985.  An  ecoIogiciJ  study  of  the  xegetatioii 
of  the  Grand  Ri\er/Cedar  Ri\er,  Sioux,  and  Ashland 
Districts  of  the  Custer  National  Forest:  a  habitat  clas- 
sification. Unpublished  dissertation,  Soutli  Dakota 
State  Uni\'ersitv'.  Brookings.  249  pp. 

IIxN.SEN,  P  L.,  cindC.  R.  Hofeman  1988.  The  yegetation 
of  the  Grand  Riyer/Cedar  River,  Sioux,  and  Ashland 
Districts  of  the  Custer  National  Forest:  a  hal)itat  t\pe 
classification.  USDA  Forest  Service  General  Technical 
Report  RM-157.  Rock\  Mountain  Forest  and  Range 
Experiment  Station,  Fort  Collins,  Colorado.  68  pp. 

i  lANSEN,  P  L.,  G.  R.  Hofeman.  and  A.  J.  Bji cstad  1984. 
The  vegetation  of  Theodore  Roosevelt  National  Park, 
North  Dakota:  a  habitat  tvpe  classification.  USDA 
Forest  Service  General  Technical  Report  RM-113. 
Rock)-  Mountain  Forest  and  Range  Experiment  Sta- 
tion, Fort  Collins,  Colorado.  35  pp. 


R-VMSEN.  p.  L.,  G.  R.  Hoffman,  and  G.  A.  Steinauer. 
1984.  U[)land  forest  and  woodland  habitat  types  of  the 
Missouri  Plateau,  Great  Plains  Province.  In:  Great 
Plains  Agriculturd  Council,  Wildlife  Resources  Com- 
mittee, Wooded  Draws:  Cluiracteristics  and  Values  for 
the  Northern  Great  Plains — Proceedings  of  Sympo- 
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Rapid  City-,  12-13  June  1984.  Great  Plains  Agricultural 
Council  Publication  No.  111.  52  pp.  Biologv  Dep;irt- 
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Rapid  City. 

R-W  L.IN,  J.  L.,  and  P  N.  SoltanI'OUR  1981.  Evaluation  of 
the  NH4HCO.3-DTPA  .soil  test  for  iron  and  zinc.  Soil 
Science  Society  of  American  Journal  45:  70-75. 

Jones,  J.  B.,  Jr  1977.  Elemental  aniJysis  of  soil  extracts  and 
plant  tissue  asli  by  plasma  emission  spectroscopy.  Com- 
munications in  Soil  Science  iuul  Plant  .\nalysis  8:  349- 
365. 

Knudsen,  D.,  G.  a.  Peterson,  and  P.  F.  Pr.\tt  1982. 
Lithium,  sodium  and  potassium.  Pages  225-246  in 
A.  L.  Page,  ed..  Methods  of  soil  anal\sis.  Part  2:  Chem- 
ical and  micr()biok)gical  properties.  2nd  ed.  Mono- 
graph No.  9  (Pt.  2)  in  the  Agronomy  Series.  American 
Society  of  Agronomy,  Madison,  Wisconsin. 

Olsen.  S.  R., C.  V.  Cole.  F.  S.  Wvianabe.  and  L.  A.  Dean 
1954.  Estimation  of  available  phosphonis  in  soil  bv 
extraction  with  NallCO.?.  USDA  Circular  939.  U.S. 
Printing  Office,  Washington,  D.C.  19  pp. 

Page,  A.  L.,  ed.  1982.  Metliods  of  .soil  analy.sis.  Part  2.  2nd 
ed.  American  Societv^  of  Agronomy,  Madison,  Wiscon- 
sin. 1159  ])p. 

Peterson  K.  IL  1987.  Forage  (juaiit\- of  key  species  in 
northwestern  South  Dakota.  Unpublished  thesis. 
South  Dakota  State  University,  Brookings.  97  pp. 

Richards.  L.  A.,  ed.  1954.  Diagnosis  and  improvement  of 
.saline  and  alkali  soils.  USDA  Agricultural  Handbook 
No.  60.  U.S.  Government  Printing  Office.  Washington, 
D.C.  159  pp. 

Se\ehso\,  K.  E.,  and  C.  E.  Boldt  1978.  Cattle,  wildlife, 
and  rij^ariiui  habitats  in  the  western  Diikotas.  Pages 
90-103  in  J.  C.  Shaver,  ed..  Management  and  Use  of 
Northern  Plains  Rangeland — Regional  Rangeland 
Svinposium,  Bismarck,  Nortli  Dakota.  North  Dakota 
State  Universits',  F"argo. 

SoLTAM'OUR,  P  N.,  A.  E.  Ludwtck,  and  J.  O.  Reiss.  1979. 
Guide  to  fertilizer  recommendations  in  Colorado.  2nd 
ed.  Cooperative  Extension  Senice,  Colorado  State 
Universitx',  Fort  C'ollins.  45  pp. 

SoLTAN poi;  R,  P.  N.,  and  A.  P.  Schwab  1977.  A  new  .soil  test 
for  simultaneous  extraction  of  macro-  and  micronutri- 
ents  in  alkaline  soils.  Communications  in  Soil  Science 
and  Piiint  Analysis  8:  195-207. 


^^  GHKAT  Basin  NATUIULIST  [Volume  52 

UursK  D  W  19S7  Effects  dlivestock  grazing  and  tlnnning  Uhksk.  D.  W,  and  C.  E.  Bc)LDT  1986.  Effect  of  cultural 

of  ,^^erto,V  ret    <>nundcrstor^■  Lod^  plants.  1'^  treatments  on  regeneration  of  ,u.t.v.woodIanck^ 

,' ^  -,     •   V    n    Proxenza    I    T    Flii  clers.   E.   D.  northern  Great  Plains.  Praine  Naturalist  18:  193-202. 

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 

P    i^  e  bivoa'lr.teractions,  7-9  August  1985,  Snow-  ponderosa  pine  fojest  in  the  --^^^^^^  "ills, 

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 
species.  HortScience  25:  191-193. 

BasIvIN.  J.  M.,  and  C.  C.  Baskin.  1985.  The  annuiil  dor- 
mancy cycle  in  buried  weed  seeds:  a  continuum.  Bio- 
Science  35:  492-i98. 

Bew  i.KV.  [.  D.,  and  M.  Black.  1982.  Physiology-  and  bio- 
cliemistr)  of  seeds.  \'()1.  2.  Springer-\'erlag,  Berlin. 

Crksswell,  E.  G.,  and  J,  P.  Grime  1981.  Induction  of  a 
light  requirement  during  seed  development  and  its 
ecological  consequences.  Nature  291:  583^585. 

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, 
London. 

Franklin,  B.,  and  R.  Tavlorson  1983.  Light  control  of 
seed  germination.  Pages  428^56  in  W.  Shropshire  and 
H.  Mohr,  eds.,  Photomoiphogenesis,  Encyclopedia  of 
Plant  Phvsiok)gv'.  New  Series,  Vol.  16A.  Springer- 
Verlag,  Berlin. 

Ghabe,  D.  F.,  ED  1970.  Tetrazolium  testing  handbook  for 
agricultural  seeds.  Handbook  on  Seed  Testing.  Contri- 
bution 29.  Association  of  Offici;il  Seed  Analysts. 

Cii  TTEKVLAN,  Y.  1982.  Phenot\pic  maternal  effect  of  photo- 
period  on  seed  germination.  Pages  67-79  in  A.  A.  Khan, 
etl.  The  physiologv'  iuid  biochemistrv-  of  seed  develop- 
ment, dormancy,  and  germination.  Elsevier  Biomedi- 
c;J  Press,  New  York. 

Hendricks.  S.  B.,  and  R.  B.  T.wlorsun  1978.  Depen- 
dence of  phvtochrome  action  in  seeds  on  niembnuie 
orgiuiization.  Plant  Physiology  61:  17-19. 

Pl.lMMER.  A.  P.,  D.  R.  ClIRISTENSEN,  AND  S.  B.  MONSEN. 
1968.  Restoring  big  game  range  in  Utah.  Utah  State 
Dep;utment  of  Fish  and  Game  Publication  68-3. 

Sn.\  EHTOWN  |,  W.  1984,  Phenotvpic  variety  in  seed  germi- 
nation beha\  ior:  the  ontogen\-  and  e\'olution  of  somatic 
poKnnoiphism  in  seeds.  Americtui  Naturalist  124:  1-16. 

.Stevens.  R..  K.  R.  Jorgensen.  and  J.  N.  D.-vvis  1981. 
Niabilitv  of  seed  from  thiiiy-two  shnib  and  forb  species 
through  fifteen  \ears  of  warehouse  storage.  Great 
Basin  Naturalist  41:  274-277. 

.SiEV  ENS,  R.,  and  S.  B.  MoNSEN  1988.  "Ced;u'  palmer 
penstemon:  a  selected  penstemon  for  semiarid  ranges. 
Riuigehuids  10:  163-164. 

TooLK  \',  K,  1973.  Effects  of  light  and  temperature  and 
their  interactions  on  the  germination  of  seeds.  Seed 
Science  and  Techno!o<n  1:  339-.396. 


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 

0 

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 


0  


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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169-1S.5. 


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  0  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  ( +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 


0 

(  ( 

. 

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  (+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- 
ing among  five  species  of  darters  (Percidae: 
Etlicosfomii).  In:  D.  Ct.  Lindtjuist  and  L.  M.  Page,  eds.. 
Environmental  biologv  ol  darters.  \\.  Junk  Publishers, 
Netherlands. 

Winn,  H.  E.  19.58.  Comparative  reproductive  behavior  and 
ecologv  of  foiuteen  species  of  darters  (Pisces — Per- 
cidae). Ecological  Mongraphs  28:  15.5-191. 


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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Mack,  G.  D.,  and  L.  D.  Flake.  1980.  Habitat  rela- 
tionships of  waterfowl  broods  on  South  Da- 
kota stock  ponds.  Journal  of  Wildlife  Man- 
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- 
namics. Academic  Press,  New  York. 
Coulson,  R.   N.,  and  J.  A.  Witter.   1984,  Forest 
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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 
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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 


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Salt  Lake  City  ^^^^^^^^^^' 
Intl.  Airport     y////////. 


lie     f//////////////////////////////y///////y//////y/yy/y//// 


Pinecrest 


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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 


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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. 


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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 


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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 

0 

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 
0 


H 1 1 h 


H 1 1 h 


-H \ 1 1 1 h 


M     A      M     J 


A      S      0      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 
0 

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. 


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lished master's  thesis.  University  of  Utali,  Siilt  Lake 
City. 
Woodward,  L.,  J.  L.  Harney,  K.  M.  Donaldson,  J.  J. 
Shiozaki.  G.  W  Leishman,  and  J.  H.  Broderick. 
1974.  Soil  survey  of  Salt  Lake  area,  Utah.  U.S.  Soil 
Conservation  Service  in  Cooperation  with  Utah  Agri- 
cultural Experiment  Station.  132  pp. 

Received  14  November  1991 
Accepted  1  June  1992 

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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deer  and  bighorn  sheep.  Journal  of  the  Ameriean  Vet- 
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Jones,  F.  L.  1954.  The  Inyo-Sierra  deer  herds.  California 
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Received  15  November  1991 
Accepted  15  April  1992 


Great  Basin  Naturalist  52(2),  pp  131-138 


DIATOM  FLORA  OF  BEAVER  DAM  CREEK, 
WASHINGTON  COUNTY,  UTAH,  USA 


Kiiitis  II.  Yt-arsk 


■  ,  Sanmel  R.  Huslitortli  .  and  Jeffre\'  H.  Joluuisei 


Abstract — Tlie  diatom  flora  of  Beaver  Dam  Creek,  Washington  County,  Utah,  was  studied.  The  study  area  is  in  a  warm 
Mojave  Desert  en\ironment  at  an  elevation  bet\veen  810  and  850  m.  A  total  of  99  taxa  were  identified  from  composite 
samples  taken  in  the  fall,  winter,  spring,  and  summer  seasons.  These  taxa  are  all  hroadlv  distributed  and  no  endemic  species 
were  encoimtered.  Three  new  records  for  the  state  of  Utah  were  identified:  Gomphoiwis  cricnse  Sk-v.  &  Maver,  S'avicula 
el  sinensis  \ar.  lata  ( M.  Perag. )  Patr.,  and  Nitzschia  calkla  Cnin.  The  most  important  taxa  throughout  the  study  as  determined 
hv  multiplying  percent  presence  by  average  relative  density  (Important  Species  Index)  were  Cijniljella  ajfinis  Kiitz., 
Epithemia  sorex  Kiitz.,  Naviaila  vcneta  Kiitz.,  Nitzschia  palea  (Kiitz.)  W.  Sm.,  and  Nitzschia  microcephala  Grun. 


Kcti  liords:  Beaver  Dam  Creek,  diatoiris,  desert  streams. 

The  algal  flora  ot  the  Intermountaiii  West  of 
North  America  is  not  well  known  despite  the 
fact  that  numerous  studies  dealing  \\ith  algal 
systems  of  waters  in  this  region  have  been  com- 
pleted in  recent  years.  These  studies  have  exam- 
ined streams,  fresh  water  lakes,  saline  lakes, 
thermal  springs,  and  terrestrial  habitats 
(Sommerfeld  et  al.  1975,  Stewart  and  Blinn 
1976,  Czarnecki  and  Blinn  1977,  1978,  Blinn  et 
al.  1980,  Bush  and  Fisher  1981;  for  bibliogra- 
phies see  Rushforth  and  Merkley  1988,  Metting 
1991). 

Algal  floras  of  wanii  desert  systems  are  espe- 
cially poorly  known.  The  present  study  was  ini- 
tiated to  provide  additional  information  on  the 
diatom  flora  of  a  desert  stream  located  in  west- 
em  North  America.  We  examined  the  diatom 
communities  of  Beaver  Dam  Creek,  a  tributary 
of  the  Virgin  River  in  southwestern  Utah.  This 
paper  is  intended  as  a  baseline  floristic  and 
communit\'  study  of  the  diatom  communities 
present  in  this  Mojave  Desert  stream. 

We  had  three  objectives  in  this  study:  (1)  to 
identify  all  species  of  diatoms  present  in  Beaver 
Dam  Creek,  (2)  to  document  seasonal  variation 
in  the  diatom  communities  of  this  stream,  and 
(3)  to  compare  diatom  populations  according  to 
habitat  t\pe.  Our  stud\-  reports  all  diatom  taxa 
present  in  this  stream  across  four  seasons  of 
1987-88.  We  studied  populations  in  (1)  riffle 
areas  with  erosional  flow  velocities,  (2)  deposi- 


tional  areas  with  slower  flows,  and  (3)  epipln  tic 
habitats  on  the  stems  and  leaves  of  aquatic  xas- 
cular  plant  vegetation. 

Site  Description 

Beaver  Dam  Creek  at  L\tle  Ranch  Preserve 
is  located  37°10'  North  latitude  and  114°  West 
longitude  in  Washington  Countv,  Utah  (Fig.  1). 
The  stream  occurs  in  our  study  area  at  an  eleva- 
tion of  about  850  m  at  L\i:le  Ranch  dropping  to 
810  m  at  Tenys  Ranch.  Our  study  sites  are 
located  along  the  wash  near  the  ranch  house  at 
Lvtle  Ranch  Preserve  and  near  a  smaller  out- 
building at  Tenys  Ranch. 

Beaver  Dam  Creek  is  a  vigorous,  braided 
perennial  desert  stream.  It  is  important  to  the 
entire  biota  of  the  area  since  it  is  the  main  source 
of  perenniiil  water.  The  stream  through  the 
study  area  has  formed  a  broad  gravel  flood  plain 
due  to  frequent  flooding.  The  stream  occurs  in 
bajada  and  alluxial  fan  materials  derived  from 
the  Bull  N'alley,  Pine  Vallev;  and  Santa  Clara 
mountains  (Welsh  et  al.  1987). 

Beaver  Dam  Oeek  is  fed  by  seeps,  springs, 
and  snowmelt  primarily  from  the  Pine  Valley 
Mountains.  This  area  is  also  characterized  by 
flash  floods  caused  by  sevx^re  periodic  thunder- 
storms in  the  summer  and  fall  seasons.  For 
instance,  prior  to  the  April  1988  collection, 
Beaver  Dam  Wash  received  11  days  of  rtiin 


J  Department  of  Botany  and  Range  Science,  Bngliam  Yonng  lJnJ\ersit\ .  Provo.  Ltali  84602. 
Departmentof  Biology,  John  Carroll  University,  Universits  Heiglils,  bliio4411S. 


131 


132 


Great  Basin  Naturalist 


[Volume  52 


Fig.  1.    Map  of  Beaver  Dam  VVasli  .sliowing  tlie  location  of  colletting  lotalitit-s  at  Tern  s  Raiuli  and  Lxtle  Ranch  Preserve. 
Due  to  the  meandering  and  clianging  nature  of  Beaver  Dam  Creek,  the  stream  itself  is  not  sliown  on  this  map. 


1992] 


Diatoms  of  Bkankh  Dam  Chkkk 


133 


producing  moderate  to  severe  flooding  along 
the  stream  channel.  This  scoured  the  stream 
channel,  remoxing  large  amounts  of  ac^natic 
\e2etati0n  and  causing;  channel  relocation  in 
some  areas. 

The  gravel  bar  in  Beaver  Dam  Creek  is  gen- 
erallv  higher  in  the  center  than  at  the  margins, 
causing  the  stream  to  meander  over  a  wide  area 
\\1th  frequent  changes  of  channel  during  flood- 
ing (Welsh  et  al.  1987).  The  fall  in  elevation 
downstream  is  not  constant.  Gravel  tends  to  pile 
up  in  steps  that  vary'  in  length  and  height.  This 
uneven  granular  substrate  causes  the  stream  to 
meander  along  the  gravel  bar  and  eventually  to 
sink  underground  approximatel)  four  miles 
below  the  southernmost  collection  site  (Welsh 
et  al.  1987).  The  perennial  stream  reappears 
infrequenth'  as  seeps  and  springs  lower  in 
Beaver  Dam  Wash  until  merging  with  the  Virgin 
Rixer. 

Climate  in  the  stud\'  area  varies  consider- 
ably, not  only  diunially  and  seasonally,  but  over 
longer  periods  of  time.  Winters  are  generally 
cool  and  drv;  summers  hot  and  dry.  MiLximum 
summertime  temperatures  have  been  recorded 
at  45.6  C.  Rainfall  averages  less  than  15  cm  a 
year,  although  this  is  \ariable  due  to  intense 
storms  (Welsh  et  al.  1987). 

The  biota  of  our  study  area  is  exceptionally 
diverse.  Mammals,  birds,  reptiles,  amphibians, 
invertebrates,  and  a  great  variety  of  plants  occur 
in  Beaver  Dam  Wash  (Welsh  et  al.  1987).  The 
stream  supports  a  diverse  riparian  habitat  con- 
sisting of  Fremont  cottonwood  (Populus 
freinontii  Wats.),  Arizona  ash  (Fraxitms  vehitina 
Torr.),  black  willow  (Salix  oooddingii  Ball),  seep 
wiWow {Baccharis emorxji  Gray//;  Torn),  numer- 
ous torbes,  grasses,  and  grasslike  species  (Welsh 
et  al.  1987).  Silty  terraces  occur  immediately 
adjacent  to  the  wash  and  have  been  historically 
used  for  cultivation.  These  areas  are  dominated 
by  catclaw  acacia  {Acacia  greggii  Gray),  panicu- 
late rabbitbrush  {Chn/sotJiamnus  panicidatus 
[Gray]  Greene),  Ambrosia  species,  and  numer- 
ous others  (Welsh  et  al.  1987).  Adjacent  uplands 
support  Joshua  tree  forests  {Yucca  hreiifolia 
Engelm.),  creosote  bush  {Larrea  tridentata 
[DC]  Gov.),  prickly  pear  cactus  {Opuntia 
en^ehnannii  Engelm.),  cholla  cactus  {Opuntia 
hasilaris  Engelm.  and  Bigel.),  and  numerous 
other  xerophvtic  species  (Welsh  et  al.  1987). 


Methods 

Water  chemistn,'  was  sampled  at  the  collec- 
tion sites  for  Febmarv,  April,  and  July  1988 
using  a  portable  Hach  field  water  chemistry  lab. 
Air  temperature  and  water  temperature,  dis- 
solved oxygen,  hardness,  alkalinit\,  and  pH  were 
measured. 

Diatom  collections  were  taken  on  21 
November  1987,  20  February  1988,  30  April 
1988,  and  6  July  1988  to  docvmient  seasonal 
\ariations  in  diatom  populations.  (Composite 
samples  were  collected  from  three  habitat 
t)pes.  First,  riffle  areas  with  erosional  flow  rates 
were  sampled  by  scraping  algae  from  large 
stones  in  the  creek  bed.  Second,  slow  water 
areas  in  the  stream  were  sampled  by  obtiuning 
sediments,  rock  scrapings,  and  visible  attached 
algae.  Finally,  submerged  sedge  stems  and 
leaves  were  scraped  or  collected  at  selected 
localities  to  studv  epiph\'tic  assemblages. 

Due  to  seasonal  changes,  it  was  not  always 
possible  to  sample  all  three  substrate  t\pes  at 
both  locations.  A  total  of  19  samples  were  ana- 
ly7:ed  during  the  course  of  the  study.  Samples 
were  stored  at  air  temperature  and  retimied  to 
the  laboratoiy  at  Brigham  Young  University-  for 
analysis. 

Diatoms  were  cleared  by  boiling  in  nitric 
acid  and  potassium  dichromate  (St.  Clair  and 
Rushforth  1977).  After  rinsing,  cleared  fnistules 
were  suspended  in  distilled  water  and  allowed 
to  air  dry  on  cover  slips.  Strewn  mounts  were 
prepared  using  Naphrax  high-resolution  resin. 
Representative  slides  were  examined  with  Zeiss 
RA  microscopes  equipped  with  Nomarski 
optics  and  bright  field  illumination.  An  Olym- 
pus AD  photomicrographic  system  was  used  to 
record  each  taxon.  Strewn  mounts  ha\e  been 
placed  in  the  collections  at  Brigham  Young  Uni- 
versity. 

A  minimum  of  500  valves  was  counted  for 
each  sample,  and  a  percent  relati\  e  densit\-  was 
calculated  for  each  taxon  (Kaczmarska  and 
Rushforth  1983).  An  Important  Species  Index 
(ISI)  for  tcLxa  present  was  calculated  by  multi- 
plving  the  percent  frequency  of  occurrence  of  a 
taxon  in  the  samples  1)\-  its  oxerall  average  per- 
cent relative  densitv  in  all  samples  (Ross  and 
Rushforth  1980,  Kaczmarska  and  Rushforth 
1983).  This  method  is  useful  since  it  considers 
both  abundance  and  seasonal  distribution  of  a 
taxon  (Warner  and  Haqoer  1972).  Species  diver- 
sity for  each  sample  was  calculated  using  the 


134 


Great  Basin  Naturalist 


[Volume  52 


T./VBU.  1.    Mean  values  for  air  teinperatiue  iuid  water  chemical  paranieter.s  taken  from  collecting  loc;ilities  in  Beaver 
Dam  Creek,  Washington  Countv',  Utah. 


February 

April 

Ji 

ily 

L\tle               Terry's 

Lvtle 

Terrv's 

Lytle 

Terry's 

Air  temp.  (C) 

16.3                  17.3 

20.5 

20.5 

33.0 

26.0 

Water  temp.  (C) 

14.5                  17.5 

16.8 

16.8 

24.3 

22.3 

Di.ssolved  O2  (mg/1) 

9.5                  10.0 

9.0 

9.0 

( .  1 

7.0 

Hardness  (mg/1) 

247.3                276.1 

707.5 

707.5 

281.9 

362.4 

Alkalinitv  (mg/1 

195.6               207.1 

201.3 

224.3 

pH 

7.3                   7.1 

6.9 

7.0 

8.1 

7.7 

T.\BLE  2.  T;t\a  present  in  samples  collected  from  Beaver  Dam  Creek,  1987-88,  Listed  with  Important  Species  Index 
(ISI)  values.  When  ISI  is  below  0.01,  the  species  is  listed  ;is  a  trace  (T). 


Taxon 


ISI 


Lvtle 


Terry 


Achnanthes  affinis  Gnin. 

Achnanthes  exi^tia  Cnm. 

Achnanthes  hnceolata  (Breb.)  Gmn. 

Achnanthes  miniitissima  Kiitz. 

Amphora  libt/ca  Ehr. 

Amphora  pedictihis  (Kiitz. )  Gmn. 

Amphora  veneta  Kiitz. 

Cah>nei.s  bacilhim  (Cnm.)  Cl. 

Cah)neis  siliciiht  (Ehr.)  Cleve 

Cocconeis  pedicuhis  Ehr. 

Cocconeis  placentula  viu".  eti^hjpta  (Ehr.)  Cleve 

Cocconeis  placentula  v;xr.  lincata  (Ehr.)  VH. 

Cyclosteplianos  invisitattis  (H.  &  H.)  Ther.,  Stoerm.  &  Hak. 

Cyclotella  nu'nc<^hiniana  Kiitz. 

Ci/mbella  affinis  Kiitz. 

Cifiuhclla  niexicana  (Ehr.)  Cl. 

Ct/inhella  microccphala  Gmn. 

Ci/nihclla  silcsiaca  Bleisch 

Ci/nihclla  tiimida  (Breb.  ex  Kiitz.)  V.H. 

Dcnticula  dedans  Grun. 

Denticnia  clc<ians  f.  valida  Pedic. 

Diatoma  viil^arc  Bott 

Diatoma  vnl^are  var.  breve  CJrnn. 

Epilhemia  adnata  var.  proboscidea  (Kiitz.)  Hend. 

Epitheinia  sorex  Kiitz. 

Epithciiiia  tur^ida  (Ehr.)  Kiitz. 

Fra<iilaria  constniens  (Ehr.)  Gmn. 

Fraiiilaria  constniens  f.  venter  (Ehr.)  Hust. 

Fra^ilaria  piwiata  Ehr. 

Fraiiilaria  vaucheriae  (Kiitz.)  Peters. 

Gomphoneis  eriense  (Grun.)  Sk\'.  &  Meyer 

Gon\phoneis  olivacea  (Home.)  Dawson 

Goniphonema  acuminatttm  Ehr. 

Gomf)honcnui  an<i^usttim  Agardh 

Gomphonenia  clavatum  (Ehr.) 

Gomphonema  p^ninotvii  Patr. 

Gomphonenia  parvidnm  (Kiitz.)  Kiitz. 

Gomphonema  pseudoatiffir  L.-Bert. 

Gomphonema  truncatiim  Ehr. 

Gtjrosi<ima  nodulifemm  (Gnm.)  G.  West 

Hantzschia  amj>hioxys  (Ehr.)  Grun. 

Melosira  variam  Ag. 

Meridion  ciradare  (CJrev.)  Ag. 

Navicida  abiskoetisls  Hust. 

Navictda  atomus  var.  permitis  (Hust.)  L.-Bert. 

Navinila  baeilhnn  Ehr. 


1.92 

1.8 

2.6 

0.03 

0.1 

0.1 

2.51 

3.8 

1.1 

1.92 

3.4 

1.3 

0.10 

0.4 

0.1 

1.76 

2.5 

1.1 

0.13 

0.6 

0.1 

T 

T 

0.04 

0.1 

0.1 

1.07 

3.1 

0.8 

1.22 

1.4 

1.1 

T 

0.72 

1.0 

0.5 

17.57 

23.4 

13.2 

T 

0.58 

1.2 

a.5 

0.16 

0.4 

0.1 

T 

1.44 

2.5 

0.4 

T 

V 

0.84 

1.7 

0.5 

0.11 

0.5 

0.1 

0.07 

0.1 

0.3 

1.3.25 

1.8 

35.9 

T 

0.21 

0.5 

0.2 

0.50 

0.5 

0.8 

0.14 

0.2 

0.3 

2.21 

1.1 

3.0 

0.02 

0.1 

0.1 

0.27 

0.7 

0.2 

T 

0.51 

0.8 

0.4 

0.06 

0.2 

0.2 

0.08 

0.3 

0.2 

1.89 

2.1 

0.2 

1.32 

1.6 

1.5 

T 

T 

T 

0.06 

0.3 

0.1 

T 

T 

0.08 

0.2 

0.1 

0.09 

0.2 

0.2 

1992]  Diatoms  of  Bea\'Er  Dam  Creek  135 

Tablk  2.  Coutiiiut'il. 

Navicula  capitatoradiata  Germain 

Navicula  cincta  (Ehr.)  Ralfs 

Ndviaila  constans  \ar.  symmetrica  Hust. 

Navictihi  aispidata  Kiitz. 

Naviada  eli^ineusis  var.  lata  (M.  Perag.)  Patr. 

Naviada  gregaria  Donldn 

Naviada  menisctdus  Schumann 

Navicula  minu.scida  \ar.  muralis  (Gmn.)  L.-Bert 

Navicida  jutptda  Kiitz. 

Navictila  radiosa  Kiitz. 

Navicula  tripunctata  (OF.  Miill.)  Bor\' 

Navicula  tripunctata  \ar.  schiz-oneiiwidcs  (V.H.)  Patr. 

Naviada  trivialis  L.-Bert. 

Navicula  vciwta  Kiitz. 

Nridium  affinc  (Ehr.)  Pfitz. 

Ncidium  did>ium  (Ehr)  Cl. 

Nitzschia  acicularis  (Kiitz.)  W.Sm. 

Nitzschia  amphibia  Gnm. 

Nitz-schia  calida  Grun. 

Nitz.schia  communis  Rabh. 

Nitzschia  constric-ta  (Kiitz.)  Ralfs 

Nitzschia  di.ssipata  (Kiitz.)  Gnm. 

Nitzschia  fonticola  Gnm. 

Nitzschia Jnistulum  (Kiitz.)  Grun. 

Nitzschia  hantzschiana  Rabh. 

Nitzschia  inconspicua  Gnm. 

Nitzschia  linearis  (Ag.)  W.  Sm. 

Nitzschia  microce))hala  Grun. 

Nitzschia  palea  (Kiitz.)  W.  Sm. 

Nitzschia  si^moidca  (Nitz.)  W.  Sm. 

Nitz-scliia  std)tilis  Gnm. 

Pinnularia  appcndiculata  (Ag.)  CI. 

Plcurosi<ima  dclicatulum  W.  Sm. 

Plcurosira  lacvis  (Ehr)  Compere 

Rcimeria  sinuata  (CJreg.)  Kociolek  &  Stoermer 

Rlioicosplwuia  curvata  (Kiitz.)  Grun. 

Rhopalodia  hrchissonii  Krammer 

Rhopalodia  oihba  (Ehr)  O.  Miill. 

Rhopalodia  f^ihha  var.  vcntricosa  (Kiitz.)  Perag.  &  Perag. 

Rhopalodia  '^ihhcnda  (Ehr)  O.  Miill. 

Stauroncis  smithii  Gnm. 

Stcnoptcrohia  intemwdia  (Lewis)  V.H. 

Stcphanochscus  hantz-schii  Grun. 

Surirclla  an^usta  Kiitz. 

Surirclla  minuta  Breb. 

Surirella  (nalis  Breb. 

Sipu'dra  acus  Kiitz. 

Sijnedra  fasciadata  \-m:  tnincata  (Gre\-. )  Patr 

Stjncdra  radians  Kiitz. 

Syjicdra  ntmpcns  vnr.  mcnc^hiniana  (Irun. 

Sijncdra  ulna  (Nitz.)  Ehr 

Sijnedra  ulna  viir.  contractu  Oestr. 

Shannon-Wiener  clixersiU'  index  (Shannon  and  averages.  It  is  \videl\-  nsed  and  has  been  found 

Weaver  1949,  Zar  1986).  to  introduce  less  distortion  than  other  methods 

Siniilarit)-  indices  were  calculated  for  all  (Kaesler  and  Cairns  1972). 
pairs  of  samples  following  Ruzicka  ( 1958).  Clus- 
ter analyses  based  on  Ruzicka's  indices  using  RESULTS  AND  DISCUSSION 
unweighted  pair-group  technicjues  (UPCMA) 

were  then  performed  (Sneath  and  Sokal  1973).  Water  chemistiv  did  not  van-  significantly 

This  method  computes  the  average  similarit\-  of  according  to  collection  localit}'  (Table  1).  Stream 

each  site  to  e\'er\'  other  site  using  arithmetic  tem^^eratnre    increased   somewhat   during   the 


L99 

2.7 

2.0 

0.17 

0.5 

0.1 

T 

T 

0.06 

0.2 

0.1 

0.16 

0.5 

0.1 

T 

0.06 

0.3 

0.1 

0.16 

0.4 

0.2 

0.19 

0.3 

0.3 

0.12 

0.5 

0.1 

0.10 

0.4 

0.1 

0.28 

0.6 

0.2 

8.78 

9.0 

8.6 

T 

0.1 

T 

0.02 

0.1 

0.1 

1.51 

2.4 

0.4 

0.02 

2.0 

0.30 

0.9 

5.9 

0.19 

0.4 

0.3 

L90 

3.9 

0.4 

0.58 

LI 

0.4 

0.01 

0.1 

0.20 

0.2 

0.1 

0.65 

1.4 

0.2 

T 

5.44 

4.7 

6.3 

5.76 

8.7 

2.4 

0.01 

0.1 

0.1 

T 

T 

0.1 

0.02 

0.2 

T 

T 

0.1 

2.73 

1.4 

4.9 

T 

T 

0.01 

0.1 

0.03 

0.2 

0.1 

T 

0.1 

T 

0.02 

0.1 

0.06 

0.2 

0.1 

T 

T 

T 

T 

0.01 

0.1 

0.1 

0.11 

0.1 

0.3 

0.40 

0.7 

0.5 

0.20 

0.3 

0.3 

136 


Great  Basin  Naturalist 


[Volume  52 


summer  uionths,  hut  it  is  uoteworthy  that  teui- 
perature  variations  in  the  stream  were  relatively 
small.  The  stream  is  circumneutral  to  slightly 
alkaline. 

A  total  oi  99  diatom  taxa  in  24  genera  were 
obserxed  in  our  collections.  Three  new  records 
for  the  state  of  Utah  were  noted:  Gomphoneis 
eriense  (Gnui.)  Skv.  &  Meyer,  Naviaila  el^hien- 
sis  var.  lata  (M.  Perag.)  Patr.,  and  Nitzsclua 
calida  Gnm.  Taxa  are  illustrated  and  described 
in  Yearsley  (1988).  Nomenclature  followed  in 
Yearsley  (1988)  was  similar  to  that  used  histori- 
calK'  b\'  researchers  in  our  laboratoiy  for  com- 
parative puiposes  (Rush forth  and  Merkley 
1988).  Diatom  taxonomy  in  this  paper  is  based 
primarilv  on  the  recent  texts  of  Krammer  and 
Lange-Bertalot  (1986,  1988,  1991),  although 
other  references  were  consulted  and  sometimes 
followed.  We  did  not  follow  the  numerous 
generic  changes  proposed  in  Round  et  al.  (1990) 
due  to  the  controversy  over  many  of  their  rec- 
ommendations. 

Eighteen  taxa  in  Beaver  Dam  Creek  had  an 
Important  Species  Index  value  greater  than  1.0 
(Table  2).  The  most  important  taxa  in  the  overall 
study  with  ISIs  above  5.0  were  CijmhcUa  affinis 
(ISI  =  17.57),  Epitlu'inia  sorex  (13.25),  Navic- 
iila  verieta  (8.78),  Nitzschia  palea  (5.76),  and 
Nitz.schia  microcephala  (5.44).  Taxa  with  ISIs 
greater  than  1.0  included  Rlioicosphenia 
curvata  (2.73),  Achnanthes  lanceolata  (2.51), 
Frogilaria  vaucheriae  (2.21),  Navicula  capita- 
toradiata  (1.99),  Achnanthes  affinis  (1.92), 
AchnantJjcs  niinittissinia  (1.92),  Nitzschia  dis- 
sipata  (1.90),  G()nif)lioneimi  parviduni  (1.89), 
Nitzschia  anipJiihia  (1.51),  Denticiila  elegans 
(1.44),  Gomphoncnia  psendoaiigtir  (1.32), 
Cocconeis  placentida  var.  Uneata  (1.22),  and 
Cocconeis  placenfnia  \ar.  ew^Uipta  (1.07).  All  of 
these  taxa  are  cosmopolitan  and  found  in  a  vari- 
ety of  habitats. 

In  comparing  the  diatom  assemblage  from 
Beaver  Dam  (Jreck  with  the  floras  of  streams  of 
other  arid  regions,  we  noticed  a  striking  similar- 
it)'.  The  important  taxa  overlapped  in  all  of  the 
studies  even  though  the  streams  varied  in  terms 
of  their  flow  rate  and  climatic  regime.  Further- 
more, each  system  was  dominated  by  cosmopol- 
itan species.  Our  preliminary  data  indicate  that 
a  diatom  flora  unique  to  desert  streams  does  not 
exist.  Further  research  to  substantiate  this  con- 
clusion is  necessarx';  .some  evidence  is  gi\en 
below. 

Blinn  et  al.  (1980)  considered  substrate  col- 


onization in  Oak  Creek,  Arizona.  Thev  reported 
12  important  taxa  which,  in  order  of  decreasing 
abundance,  were:  Nitzschia  frustuhun,  Epithe- 
niia  sorex,  Cocconeis  placentida  var.  euglifpta, 
Achnanthes  niintitissima,  Navicida  cnjpto- 
cephala,  Navicula  veneta  (as  N.  cnjptocephala 
var.  veneta),  Nitzschia  dissipata,  Achnanthes 
lanceolata,  Ci/mbeUa  affinis,  Fragilaria  con- 
stniens,  Navicida  decussis,  and  Synedra  idna. 
These  diatoms  accoimted  for  90%  or  more  of 
the  total  algal  population  on  newly  introduced 
material  in  their  study.  Eight  of  these  taxa  were 
also  important  in  our  stream,  having  ISI  values 
above  1.0. 

Johnson  et  al.  (1975)  conducted  further 
study  on  the  diatom  flora  of  Oak  Creek,  Arizona. 
They  reported  41  diatom  taxa,  of  which  25  are 
common  to  our  study  area.  Cijnihella  affinis, 
Epithemia  sorex,  and  Nitz^schia  palea  were 
reported  as  common  or  abundant.  This  com- 
pares favorably  with  the  results  of  our  study 
since  these  three  were  among  the  most  common 
diatoms  in  Beaver  Dam  Creek. 

Rushforth  et  al.  (1976)  examined  the  algal 
flora  of  Freshwater  Wash,  Arches  NationiU 
Park,  in  southeastern  Utah.  Their  study  docu- 
mented 57  diatom  taxa,  29  of  which  were  tilso 
observed  in  Beaver  Dam  Creek.  Achnanthes 
niinittissinia,  Cijmhella  affinis,  Denticula  ele- 
gans, Goniphonenia  acuminatum,  Navicula 
radiosa,  Nitzschia  linearis,  Nitzschia  palea, 
Rhoicosphenia  curvata,  and  five  other  species 
not  present  in  Beaver  Dam  Creek  were  the  most 
abundant  taxa  in  Freshwater  Wash. 

In  their  analvsis  of  Sycamore  Creek,  Arizona, 
Fisher  et  al.  (1982)  reported  that  diatoms  made 
up  77%  of  the  total  algal  mass,  \v\i\\  Achnanthes 
exigua,  Gomphonema  parvidum,  and  Navicula 
pupula  being  the  most  important  taxa.  These 
taxa  were  present  in  Beaver  Dam  Creek  but  in 
lower  numbers.  Gomphonema  parvidum  was 
the  most  abundant  of  the  three  in  our  samples. 

The  flora  of  Beaver  Dam  Creek  is  also  sim- 
ilar to  that  of  other  streams  of  western  North 
America  draining  more  mesic  regions.  Gushing 
and  Rushforth  (1984)  in  a  study  of  the  Salmon 
River,  Ickilio,  identified  145  diatom  species,  48 
of  which  were  among  the  99  taxa  found  in 
Beaver  Dam  Creek.  Half  of  their  important 
species  (9  of  18)  were  also  among  the  important 
species  in  Beaver  Dam  Creek,  several  with  sim- 
ilar importance  values. 

Preliminar\'  research  also  indicates  that  a 
flora  similar  to  that  found  in  North  American 


1992] 


Diatoms  of  Bean-er  Dam  Cheek 


137 


hardwater  streams  exists  elsewhere.  S(juires  and 
Saoud  (1986)  reported  nine  ta\a  from  the 
Damour  Ri\er,  Lebanon,  with  Importance  Spe- 
cies Index  xiilues  above  1.0.  Six  of  these  also 
were  important  in  Beaxer  Dam  (>reek.  In  the 
Damoin-  Ri\er  stiid\'  Aclinantlics  inlnutissiina 
was  the  most  important  taxon  with  an  ISI  \alue 
of  44.4,  followed  bv  Nitzschia  dissipata  (5.12), 
Cyniljella  microcephala  (3.63),  and  CifmheUa 
affinis  (2.62). 

Shannon-Wiener  diversit)'  values  for  all  24 
samples  ranged  between  1.95  and  4.59.  Diver- 
sit)'  did  not  show  any  clear  trends  with  regard  to 
season  or  substrate  tyjoe.  The  overall  mean  for 
the  indices  was  3.42,  the  median  \alue  being 
3.57.  These  \alues  are  relati\el\'  high  and  indic- 
ative of  unpolluted  water. 

Oiu"  collections  did  not  cluster  well  on  the 
basis  of  habitat  t\pe  or  season.  However,  there 
was  a  tendency  for  stands  to  cluster  on  the  basis 
of  the  Terpy's  Ranch  versus  L)tle  Ranch  Pre- 
serve collecting  localities  (Fig.  2).  The  upper- 
most cluster  consists  of  samples  from  Terrv's 
Ranch,  while  the  second  cluster  contains  sam- 
ples from  the  Lvtle  Ranch  Preserve.  The  third 
cluster  has  a  mix  of  all  sites,  substrates,  and 
seasons.  The  fall  depositional  sample  from  the 
Lvtle  Ranch  Preserve  is  an  outlier. 

The  reasons  for  the  clustering  b)'  site  seen  in 
the  top  half  of  the  cluster  are  unclear.  Water 
chemistry'  and  temperature  did  not  var\"  greatlv 
between  the  sites  during  the  year  (Table  1). 
Likewise,  insolation  is  approximatelv  the  same 
for  both  sections  of  the  creek.  Stream  velocities, 
however,  appear  to  be  different.  The  creek  at 
Lytle  Ranch  Presene  is  generallv  slower,  shal- 
lower (<15  cm),  wider,  and  more  meandering 
than  the  stream  at  Terry's  Ranch  where  pools 
may  reach  depths  of  nearly  one  meter. 

The  cluster  shows  a  number  of  samples  that 
paired  b\-  date  of  collection  (Fig.  2).  However, 
seasonalit)-  was  ver)-  weak.  The  absence  of  sea- 
sonal changes  is  probably  attributable  to  one  or 
two  factors.  First,  temperatiu'e  changes 
tlu-oughout  the  year  are  minor,  and  changes  in 
photoperiod  alone  are  not  enough  to  drive  suc- 
cession. Second,  storm  events  scour  the  creek 
bed  occasionally  and  may  keep  the  diatom 
assemblage  in  an  early  successional  stage. 

The  habitat  t\pes  sampled  did  not  cluster 
separately,  indicating  they  are  fairly  similar. 
Because  of  scouring  events,  the  depositional 
areas  initially  sampled  often  had  all  sediments 
remoN'ed  at  later  sampling  dates  and  so  consist 


PERCENT   SIMILARITY 


100         90 
T  M  Nov. 


T  R  July 


T  D/R  July 


L  D/R  Apr. 

T  D/R  Apr. 

L  R  Feb. 

L  R  Apr. 

L  R  July 

T  M  Apr. 

L  M  July 

L  D/R  Feb. 

L  R  Apr. 

T  R  Apr. 

Fig.  2.  Cluster  diagram  of  19  samples  collected  from 
Beaver  Dam  Creek.  T  =  Terrv's  Ranch,  L  =  Lvtle 's  Ranch 
Preserxe,  M  =  macrophvtic  vegetation  (sedges),  R  =  riffle, 
D/S  =  depositional  area,  .sediments,  D/R  =  dejx)sitional 
area,  rock  scrapings. 

of  rock  scrapings,  just  as  in  the  riffle  areas.  The 
one  sample  that  consisted  of  sediment  only 
(Lylile  Ranch,  November  1987,  depositional 
area)  clustered  separately  from  all  other  sam- 
ples (see  bottom  line  of  cluster,  Fig.  2). 

In  summary,  the  diatom  assemblages 
observed  in  Beaver  Dam  Creek  consisted  of 
cosmopolitan  species  common  to  other  hard- 
water  rixers.  Seasonalit\'  was  minimal,  as  were 
the  effects  of  habitat  t\pe. 

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Colonization  rates  and  community  stnictnre  of  diatoms 

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British  Phycologicd  Jouniiil  15:  .30.3-.31(). 
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Warner,  J.  H.,  and  K.  T.  Harper  1972.  Understorv  char- 
acteristics related  to  site  quality  for  aspen  in  Uttili. 
Brigham  Young  Universitv  Science  Bulletin,  Biological 
Series  16(2):  1-20. 

Welsh,  S.  L.,  K.  W.  Pac;ker,  B.  N.  Smith,  and  S.  T. 
Taylor.  1987.  Brigham  Young  University  Lytle  Ranch 
Management  Plan.  Brigham  Young  Uni\'ersitv  Press, 
Provo,  Utah.  21  pp. 

Yearsley,  K.  H.  1988.  The  diatom  flora  of  Bea\er  Dam 
Creek,  Washington  Countv,  Utah,  USA.  Unpublished 
master's  thesis,  Brigham  Young  University',  Provo, 
Utiili.  30pp.,12pl.  ' 

Z\R,  J.  H.  1986.  Biostatisticalimalysis.2nded.  Prentice-Hall 
Inc.,  New  Jersey.  718  pp. 

Received  16  September  1991 
Accepted  31  March  1992 


{ ;ivat  Basin  Naturalist  52(2),  pp.  139-144 

STRATIFICATION  OF  HABITATS  FOR  IDENTIFYING  HABITAT  SELECTION 

BY  MERRIAM'S  TURKEYS 

Mark  A.  Riinible    and  Stanley  H.  Anderson" 

Abstract — Habitat  selection  patterns  of  Merriam's  Turkevs  were  conipiired  in  hierarchical  iuialvses  of  three  levels  of 
habitat  stratification.  Habitat  descriptions  in  first-le\el  analyses  were  based  on  dominant  species  of  vegetation.  Habitat 
descriptions  in  seconcl-le\el  anaKses  were  biised  on  dominant  species  of  vegetation  and  overstorv^  canopy  cover.  Habitat 
descriptions  in  third-level  anal\ses  were  based  on  dominant  species  of  vegetation,  o\erston'  canopy  cover,  and  stnictural 
stages  (dbh  categories).  First-level  analyses  showed  turkeys  selected  for  ponderosa  pine  and  selected  against  meadow 
habitats.  No  conclnsions  could  be  drav\Ti  regiucling  forest  management  on  habitat  selection  of  turkevs  at  this  le\el  of  habitat 
stratification.  Second-level  analyses  showed  that  selection  of  ponderosa  pine  and  aspen/birch  habitats  \aried  among  seasons. 
Implications  for  forest  management  actixities  on  turkevs  at  this  level  of  habitat  stratification  could  be  made.  Third-level 
aiuil\ses  added  little  to  conclusions  of  habitat  selection  patterns  drawn  from  second-le\el  analyses  and  increased  chances 
for  T\pe  II  errors.  Habitat  selection  patterns  of  Merriam's  Turkeys  were  best  described  when  habitats  were  stratified  by 
dominant  species  of  vegetation  and  overstorv  Ciuiopy  cover. 

Kjcij  words:  Merriam's  Wild  Turkeys.  Meleagris  gallopa\o  merriami,  hiihitat  descriptions,  forest  ntana^eiiwut,  habitat 
selection 


Habitat  use  and  management  of  Merriam's 
Turkeys  {Meleagris  gallopavo  merriami)  in 
northern  latitudes  have  been  studied  in  South 
Dakota  (Petersen  and  Richardson  1975)  and 
Montana  (Rose  1956,  Jonas  1966).  These  early 
studies  were  limited  to  direct  observation  of 
birds  when  assessing  habitat  use,  and  data  con- 
tained biases  in  the  assessment  of  the  birds' 
habitat  needs  (e.g.,  Jonas  1966,  Brvant  and  Nish 
1975,  Petersen  and  Richardson  1975,  Shaw  and 
Smith  1977).  Telemetrv'  has  allowed  collection 
of  data  on  habitat  use  patterns  in  an  unbiased 
manner,  but  few  studies  have  addressed  the 
detailed  stratification  habitats. 

Studies  of  habitat  use  and  selection  patterns 
by  Merriam's  Turkeys  have  delineated  habitats 
based  primarilv  on  the  dominant  species  of  veg- 
etation (DSV)'  (Jonas  1966,  Biyant  and  Nish 
1975,  Scott  and  Boeker  1975,  ^Mackey  1982, 
1986,  Lutz  and  Crawford  1989).  Because  timber 
management  activities  seldom  result  in  conver- 
sions of  vegetation  t)pes,  understanding  habitat 
selection  patterns  at  this  level  precludes  under- 
standing the  effects  of  forest  management  activ- 
ities such  as  logging  or  thinning  on  Merriam's 
Turkeys.   Increased  value  of  ponderosa  pine 


timber  resources,  emphasis  on  old-growth 
resource  values,  and  improved  technolog)'  for 
harvesting  timber  have  potential  to  impact 
Merriam's  Turkey  habitat  (Shaw  1986).  There- 
fore, stratification  beyond  dominant  species  of 
vegetation  is  necessary  to  elucidate  the  effects 
of  forest  management  on  turkeys.  Merriam's 
Turkeys  in  southeastern  Montana  demonstrated 
an  apparent  preference  for  pole-size  (<23  cm 
dbh)  ponderosa  pine  habitats  (Jonas  1966). 
Merriam's  Turkeys  in  Oregon  avoided  habitats 
that  had  been  logged  by  clear-cut  or  shelter- 
wood  methods  (Lutz  and  Crawford  1989).  To 
our  knowledge,  no  researchers  have  stratified 
habitats  in  terms  of  size  and  densitv  categories 
of  tree  species.  However,  on  lands  managed  by 
the  US  DA  Forest  Service  and  other  public 
agencies,  methods  of  habitat  stratification  that 
include  structural  stages  (SS)  and  overstory 
canopy  cover  categories  (OCC)  have  been 
described  (Thomas  1979)  to  further  stratifv hab- 
itats. 

The  objective^  of  tliis  stuck  was  to  determine 
the  level  of  habitat  stratification  that  best 
described  habitat  use  and  selection  patterns  of 
Merriam's  Turkeys  in  the  Black  Hills. 


, USD.-\  Forest  Sen.ice,  .501  E.  St  Joseph  St..  South  Dakota  School  of  Mines.  Rapid  Cit\.  South  Dakota .57701 . 
"USDI  Cooperative  Fisheries  and  Wildlife  Research  Unit.  University  of  Wyoming.  Laramie,  \W)ming  82071. 


139 


140 


Great  Basin  Naturalist 


[Volume  52 


Methods 

Study  Area 

This  study  was  conducted  in  the  central 
Black  Hills  of  South  Dakota,  16  km  west  of 
Rapid  Cit\'.  Most  of  the  land  is  vmder  manage- 
ment by  the  Black  Hills  National  Forest,  Pactola 
Ranger  District.  Some  private  holdings  associ- 
ated with  ranch  operations  are  present  in  the 
meadows,  and  several  private  homes  and  cabins 
are  located  in  the  study  area. 

Vegetation  of  the  study  area  is  primarily  pure 
ponderosa  pine  forest  (84%).  Meadows  and 
aspen/birch  (Popiihis  tremtiloicles/Betida  pa- 
pyrifera)  habitats  occur  in  drainages. 

This  study  was  conducted  over  a  three-year 
period  beginning  March  1986  and  ending  Janu- 
ary 1989.  Because  anahtical  methods  used  to 
make  statistical  tests  were  goodness-of-fit  tests 
and  nonsignificance  indicates  lit  by  the  pro- 
posed model,  hypotheses  tested  have  been 
stated  appropriately.  The  hvpotheses  tested  rel- 
ative to  Merriam's  Turkeys  in  the  Black  Hills  of 
South  Dakota  were  that  each  of  the  following 
habitats  depict  patterns  of  use  and  selection  by 
Merriam's  Turkeys:  (1)  habitats  stratified  by 
DSV  (2)  habitats' stratified  by  DSV  and  OCC, 
(3)  habitats  stratified  by  DSV  and  SS,  and  (4) 
habitats  stratified  by  DSV,  SS,  and  OCC. 

Trapping  and  locations. — Turkeys  were 
trapped  in  late  February  or  early  March  of  each 
year  of  the  study  with  rocket  nets  and  drop  nets 
over  com  bait.  This  study  was  primarih'  con- 
cerned with  hens  since  they  are  the  reproduc- 
tive segment  of  the  population.  Forty-four  (36 
females  and  8  males)  of  82  turkevs  trapped  were 
fitted  with  back|:)ack  radio  transmitters  weigh- 
ing approximate!)'  108  g. 

Locating  birds  began  after  a  one-week 
period  of  adjustment  to  the  radio  transmitters 
(Nenno  and  Healy  1979).  Each  bird  in  the  study 
area  was  located  three  times  each  week,  once 
during  each  of  the  following  time  periods:  sun- 
rise-1000  hr,  1001-1400  hr,  and  1401  hr- 
sunset.  Birds  that  emigrated  from  the  defined 
study  area  were  located  at  least  monthly  to  mon- 
itor their  activities  and  determine  if  they  had 
moved  back  into  the  study  area.  Locations  were 
determined  by  plotting  2+  bearings  (frequentlv 
5+)  from  known  locations  on  USGS  1:24,000 
contour  maps  in  the  field  using  a  luuul-held, 
two-element  yagi  antenna.  Bearings  were  usu- 
ally taken  from  positions  within  300  m  of  the 
estimated  location.  Each  location  was  assigned 


to  a  habitat  unit  (see  below)  based  on  maps  and 
Universal  Transverse  Mercator  coordinates 
recorded  to  the  nearest  100  m  in  the  field.  To 
achieve  independence  of  observations  (All- 
dredge  and  Ratti  1986),  only  one  location  was 
recorded  for  each  bird  on  any  given  day  and 
most  were  two  days  apart. 

Habitat  Descriptions 

Habitats  were  numerically  identified  geo- 
graphical units  approximatelv  4-32  ha  (10-80 
acres)  in  size.  Boundaries  were  usuallv  defined 
by  watershed  topography  such  as  ridges  and 
drainages.  Obvious  changes  in  vegetation  type 
also  were  used  to  define  boundaries  of  habitats. 
In  all,  513  habitat  units  were  delineated. 

Vegetative  descriptions  of  habitats  were 
determined  from  five  plots  located  within  each 
defined  habitat  unit.  These  plots  were  marked 
on  unit  1:24,000  contour  maps  in  the  lab  and 
distributed  evenly  across  each  habitat.  Some 
habitats  were  too  small  to  effectively  place  five 
plots,  so  fewer  plots  were  used.  Each  plot  was 
then  located  in  the  field  and  sampled  to  deter- 
mine tree  basal  area. 

Habitat  descriptions  were  made  based  on 
DSV,  SS,  and  OCC  according  to  criteria  devel- 
oped by  the  US  DA  Forest  Service,  Region  2 
(Buttety  and  Gillam  1983).  DSV  categories 
were  ponderosa  pine,  aspen/birch,  oak,  spruce, 
and  meadows.  SS  categories  were  pole  timber 
(trees  2.5-22.8  cm  dbh)  and  sawtimber  (trees 
greater  than  22.8  cm  dbh).  OCC  categories 
were  0-40%,  41-70%,  and  71-100%.  OCC  was 
estimated  based  on  the  following  equation: 
OCC(%)  =  0.5  r BASAL  AREA  (FT'/AC)  - 
1.94  (Bennett  1984).  Depending  on  the  level  of 
stratification  included  in  the  analyses,  .5-12  hab- 
itats were  delineated. 

Analyses 

Data  pertaining  to  use  of  habitats  described 
above  were  stratified  into  seasons:  December- 
February  (winter),  March-May  (spring),  June- 
August  (summer),  and  September-No\ ember 
(fall).  Chi-square  testof  independence  was  used 
to  test  the  lupothesis  that  habitat  use  patterns 
of  Merriam's  Turkeys  were  similar  among  sea- 
sons. Because  this  test  was  significant  (P  <  .001), 
tests  of  habitat  selection  at  different  levels  of 
habitat  stratification  were  made  within  seasons. 

Chi-square  goodness-of-fit  tests  with  correc- 
tion for  continuit)'  (Cochran  1963)  were  used  to 
test  hypotheses  regarding  the  level  of  habitat 


19921 


Turkey  Habitat  Stratification 


141 


stratification  that  best  depicted  habitat  selection 
patterns  of  Merriam's  Turkeys  in  a  hierarchical 
structure.  Bonferroni  confidence  intervals 
around  proportion  of  use  (Neu  et  al.  1974,  Byers 
et  al.  1984)  were  used  to  determine  habitat 
selection  patterns  that  deviated  from  expected 
use.  We  determined  differences  from  expected 
use  of  habitats  for  which  utilization  was  0  by 
examining  chi-square  residuals  with  G-stan- 
dardization  and  Bonferroni  correction  to  the 
Z-statistic  (Mosteller  and  Pamnak  1985).  An 
array  of  structural  stages  occurred  only  for 
ponderosa  pine  habitats.  Therefore,  the  test 
for  DSV  X  SS  level  of  habitat  stratification 
was  analyzed  using  data  from  ponderosa  pine 
habitats.' 

Initial  chi-square  tests  of  use  versus  avail- 
abilitv  for  DSV  x  SS,  DSV  x  OCC,  and  DSV  x 
SS  X  OCC  were  made  with  oak,  aspen,  and 
spruce  habitats  pooled  to  reduce  as  much  as 
possible  the  number  of  cells  with  fewer  than  five 
expected  observations.  Selection  of  these  hab- 
itats by  turkeys  was  evaluated  individually  with 
Bonferroni  confidence  intervals  for  comparison 
tests.  The  significance  of  confidence  intervals 
holds  regardless  of  the  overall  chi-square  test 
(Neu  et  al.  1974). 

Results 

Habitats  Determined  by  DSV 

The  hyjDothesis  that  habitats  stratified  by 
DSV  depict  patterns  of  habitat  use  and  selection 
by  Merriam's  Turkeys  was  rejected  (F  =  .06). 
Meadows  were  selected  less  than  expected 
across  all  seasons  (Table  1).  Ponderosa  pine 
habitats  were  selected  more  than  expected 
during  winter,  spring,  and  fall;  they  were  equal 
to  what  was  expected  during  summer.  Aspen 
habitats  were  selected  more  than  expected 
during  summer.  Oak  habitats  were  selected  less 
than  expected  during  spring,  while  spruce  hab- 
itats were  selected  less  than  expected  during 
winter  and  spring. 

Habitats  Determined  bv  DS\'  and  OCC 

The  hvpothesis  that  habitats  stratified  b\ 
DSV  and  OCC  depict  patterns  of  habitat  use 
and  selection  by  Merriam's  Turkeys  was 
rejected  for  all  seasons  (P  =  .04).  Stratifying 
habitats  by  DSV  and  OCC  did  not  alter' the 
results  for  meadow,  oak,  or  spnice  habitats 
(Table  2).  Oak  and  spruce  were  not  represented 


across  all  ov  erston  canopv  cover  categories  on 
this  study  area. 

Aspeii/birch  habitats  with  41-70%  OCC 
were  selected  more  than  expected  during  spring 
and  sunuiier  by  turkeys  in  the  Black  Hills.  Infre- 
quent use  of  aspen/birch  habitats  with  7 1-100% 
OCC  was  noted  over  all  seasons.  But  statisti- 
cally, this  was  less  than  expected  onlv  during 
spring.  Open  ponderosa  pine  habitats  (0-40% 
OCC)  were  selected  less  than  expected  during 
the  winter  and  spring.  Turkeys  selected  pon- 
derosa pine  habitats  41-70%  OCC  more  than 
expected  during  spring.  Dense  ponderosa  pine 
habitats  (71-100%  OCC)  were  selected  more 
than  expected  during  fall  and  winter  and  less 
than  expected  during  summer. 

Habitats  Determined  by  DS\'  and  SS 

The  hypothesis  that  habitats  stratified  by 
DSV  and  SS  depicted  patterns  of  habitat  use  and 
selection  by  Merriam's  Turkeys  was  not  rejected 
for  winter,  summer,  and  fall.  During  spring, 
ponderosa  pine  habitats  with  stems  greater  than 
23  cm  dbh  were  selected  more  than  expected. 
Othenvise,  no  differences  were  apparent  in  the 
habitat  selection  patterns  of  turkeys  when  pine 
habitats  were  stratified  based  on  dbh. 
Aspen/birch,  oak,  and  spruce  habitats  were  not 
adequately  represented  across  structural  stages 
to  make  comparisons. 

Habitats  Determined  by  DS\'  SS,  and  OCC 

The  hyjiothesis  that  habitats  stratified  by 
DSV,  SS,  and  OCC  depict  patterns  of  habitat  use 
and  selection  by  turkev  s  v\'as  rejected  {P  =  .03) 
during  winter,  spring,  and  summer  (Table  3). 
Data  from  fall  indicated  observed  differences 
from  expected  at  F  =  .11.  Since  several  habitat 
categories  were  pooled  to  achieve  minimum 
sample  .size  in  the  overall  chi  s(juare  test,  F  =  .11 
was  considered  sufficient  indication  of  differ- 
ence from  expected  to  proceed  with  the 
Bonferroni  confidence  intervals. 

Use  patt(M-ns  of  meadov\',  oak,  and  spruce 
habitats  bv  Merriam's  Turkeys  v\'ere  unchanged 
from  previous  levels  of  habitat  stratification. 
However,  because  more  habitats  were  included 
in  the  analyses,  selection  of  spruce  during 
winter  and  aspen/birch  habitats  with  41-70% 
overstory  canopy  cover  during  summer  no 
longer  differed  from  expected. 

Turkev  s  selected  open  ponderosa  pine  habi- 
tats in  both  structural  stages  less  than  expected 
durine  winter,  and  the  2..5-22.8  cm  dbh  stnictnral 


142 


Great  Basin  Naturalist 


[\blume  52 


Table  1.   Seasooiil  utilization  by  Merriam's  Turkeys  of  habitats  described  by  dominant  species  of  vegetation  in  the  Black 
HiUs  of  South  Dakota/'-^' 


Habitat 


Proportional 
area 


Winter 

(205) 


Spring 

(S78)" 


Summer 

(126) 


FaU 

(218) 


Aspen 

Meadow 

Pine 

Oak 

Spnice 


0.0516 

4— 

61 

17+  + 

14 

0.1016 

11— 

9— 

5 — 

/  — 

0.8371 

186+ + 

807+ + 

100 

195+ + 

0.0044 

4 

0— 

1 

1 

0.0056 

0— 

1— 

3 

1 

"Sample  sizes  (teleinetn'  fixes)  are  in  parentheses.  Expected  use  can  he  calculated  from  proportional  use  X  sample  size. 
Differences  [P  <  .101  among  hahitats  selected  versus  aviulahle  are  indicated  hy  —  it  used  less  than  exjiected  and  ++  if  used  more  than  ex]«?cted. 


Table  2.  Seasonal  utiUzation  bv  Merriam's  Turkevs  of  habitats  described  bv  dominant  species  and  overstoiy  c;xnopy 
cover  of  vegetation  in  the  Black  Hills  of  South  Dakota.'  ' 


Habitat 

Percent 

Proportional 

Winter 

Spring 

Summer 

FaU 

canopy  cover 

area 

(205) 

(878) 

(126) 

(218) 

Aspen/birch 

0-iO 

0.0148 

2 

14 

4 

1 

Aspen/birch 

41-70 

0.0191 

0 

46+  + 

11  +  + 

12 

Aspen/birch 

71-100 

0.0177 

2 

1— 

2 

1 

Ponderosa  pine 

0-^0 

0.1199 

.3— 

6.3— 

26 

29 

Ponderosa  pine 

41-70 

0.3760 

65 

430+ + 

45 

71 

Ponderosa  pine 

71-100 

0.3412 

118+  + 

314 

29— 

95+  + 

Meadows 

0.1016 

11— 

9— 

.5— 

7 — 

Oak 

0-100 

0.0044 

4 

0 

1 

1 

Spnice 

0-100 

0.0056 

0 

1— 

3 

1 

''Sample  sizes  (telemetr\'  fixes)  are  in  parentheses.  Expected  use  can  be  calculated  from  proportiona!  use  X  sa 
'Differences  (P  <  .10)  among  habitats  selected  versus  available  are  indicated  by  —  if  used  less  than  expected  ; 


nple  size 
nd++if 


used  more  than  expected. 


Table  3.   Seiisonal  utibzation  by  Merriam's  Turkeys  of  habitats  determined  bv  dominant  .species,  overstor\'  canopv  cover, 
and  structural  stage  in  the  Black  Hills  of  South  Dakota."' 


Habitat 

Structuriil 

Percent 

Proportional 

Winter 

Spring 

Summer 

Fall 

stage 

canopv  cover 

area 

(205) 

(878) 

(126) 

(218) 

Aspen/birch 

2.,5-22.8  cm 

0-40 

0.0148 

2 

14 

4 

1 

Aspen/birch 

2.5-22.8  cm 

41-70 

0.0191 

0 

46+  + 

11 

12 

Aspen/birch 

2..5-22.S  cm 

71-100 

0.0177 

2 

1— 

2 

1 

Ponderosa  pine 

2.,5-22.8  cm 

0^0 

0.0701 

1— 

9— 

20+  + 

18 

Ponderosa  pine 

2.5-22.8  cm 

41-70 

0.1677 

32 

143 

2.5 

.'3.3 

Ponderosa  pine 

2.5-22.8  cm 

71-1(K) 

0.2173 

85+  + 

222 

16— 

62 

Ponderosa  pine 

>22.8  cm 

0-40 

0.0498 

2 

54 

6 

11 

Ponderosa  pine 

>22.8  cm 

41-70 

0.2083 

33 

287+  + 

20 

38 

Ponderosa  pine 

>22.8  cm 

71-KK) 

0.1239 

33 

92 

13 

33 

Meadows 

0.1016 

11— 

9— 

5— 

7 

Oak 

0-100 

().(K)44 

4 

0 

1 

1 

Spruce 

0-100 

0.0056 

0 

1— 

3 

1 

J^Sample  sizes  (telemetry  fi,\es)  are  in  parentheses.  Expected  use  can  be  calculated  from  proportional  use  X  sample  size. 
Differences  (P  <  .10)  among  habitats  selected  versus  aviiilable  are  indicated  l)y  —  if  used  less  than  expected  and  ++  if  used  more  tli.u 


stage  was  selected  less  than  expected  during 
spring.  No  differences  were  noted  for  pon- 
derosa pine  with  41-70%  overstory  canopv 
cover  and  2.5-22.8  cm  dbh  across  seasons. 
However,  the  structural  stage  greater  than  22.8 
cm  dbh  and  41-70%  overstory  canopy  cover  was 


selected  more  than  expected  during  spring. 
Dense  ponderosa  pine  ( >71%  overstorx'  canopv 
cover)  2.5-22.8  cm  dbh  was  selected  more  than 
expected  during  winter  and  less  than  expected 
during  summer.  No  differences  were  noted  for 
dense  ponderosa  pine  >22.8  cm  dbh. 


1992] 


TuHivt:Y  Habitat  Stratification 


143 


Discussion 

The  highest  level  of  stratification  of  habitats 
that  added  new  information  to  use  and  selection 
patterns  of  Merriam's  Turkeys  in  this  study  area 
was  b\-  DS\' and OCC.  Despite  statistical  signif- 
icance of  differences  when  habitats  were  strati- 
fied by  DS\',  SS,  and  OCC,  trends  in  habitat 
selection  were  similar  to  analyses  for  which  data 
were  pooled  across  SS  categories.  Shaw  and 
Smith  (1977)  noted  apparent  habitat  selection 
b\-  Merriam's  Turkevs  in  Arizona  when  pon- 
derosa  pine  habitats  based  on  diameter  classes 
were  ignored.  However,  pole-size  ponderosa 
pine  habitats  were  used  more  than  other  size 
classes  b\'  turkevs  in  Montana  (Jonas  1966). 
Within  our  studv  area,  12  ot  the  372  ponderosa 
pine  habitats  had  an  average  dbh  of  less  than  15 
cm  (6  in);  the  lowest  average  dbh  was  10.7  cm 
(4.2  in).  Thirt\-se\en  of  the  ponderosa  pine 
habitats  in  the  stud\  area  had  dbh  greater  than 
30  cm  (12  in),  of  which  the  majoritv"  were  in  the 
0-40%  OCC  category  indicative  of  large  over- 
mature trees.  Most  of  the  study  area  had  been 
logged  in  the  past  one  hundred  \ears.  Because 
excellent  germination  conditions  for  ponderosa 
pine  in  the  Black  Hills  result  in  overstocked 
stands  with  reduced  growth  rates  (Boldt  and 
\'an  Duesen  1974),  ponderosa  pine  habitats 
larger  than  30  cm  dbh  were  rare.  Ponderosa 
pine  habitats  in  this  study  were  representative 
of  a  narrow  range  of  the  potential  tree  dbh 
classes  for  ponderosa  pine.  However,  they  did 
represent  the  size  classes  of  ponderosa  pine 
throughout  the  Black  Hills. 

The  tests  of  the  model  for  DSV  x  SS  sug- 
gested good  agreement  between  the  model  and 
observed  use  bv  turkevs  from  a  statistical  point 
of  \iew.  These  results  suggest  random  selection 
of  habitats  when  stratified  by  DSV  x  SS.  Non- 
random  selection  of  habitats  had  already  been 
demonstrated.  We  also  beliexe  that  stratifica- 
tion of  habitats  bv  DSV  x  SS  obscured  biologi- 
cal patterns  alreacK'  demonstrated  In  the  test  of 
DSV  X  OCC.  Many  of  the  relationships  of  OCC 
were  contrasted  between  high  and  low  OCC. 
These  results  were  pooled,  resulting  in  the 
apparentlv  good  fit  of  the  DS\'  X  SS  UKxlel. 

Our  approach  to  these  anahses  was  hierar- 
chical in  nature;  and  since  patterns  of  habitat 
selection  by  turkeys  had  been  demonstrated  at 
higher  le\els,  it  would  not  be  prudent  to  ignore 
those  biological  patterns.  Howexer,  to  ensure 
that  no  oversights  were  made,  we  made  tests  of 


hal)itat  selection  based  on  habitats  stratified  b\- 
SS,  OCC,  and  SS  x  OCC.  The  test  of  the  model 
for  SS  was  not  rejected.  Tests  of  the  model  for 
OCC  and  SS  x  OCJC  were  rejected,  but  were 
influenced  b\'  the  preponderance  of  the  studv 
occupied  b\  ponderosa  pine  (84%)  and  the 
range  of  dbh  classes  in  the  Black  Hills.  Interpre- 
tations of  results  from  these  latter  tests  were 
similar  to  tests  of  DSV  X  SS  and  DSV  X  OCC. 
Stratification  of  habitats  bcNond  that  neces- 
sary' to  depict  the  dispersion  patterns  of  the 
animal  decreases  the  sensitivit)-  of  tests  and 
increases  the  probabilits  of  T\pe  H  error  in  the 
anahses  (Alldredge  and  Ratti  1986).  The  effect 
of  adding  stratification  factors  is  to  dilute  the 
sample  sizes  in  indixidual  cells,  thus  increasing 
the  chance  of  Type  H  error.  Apparent  T\pe  II 
errors  occurred  in  the  determination  of  habitat 
selection  patterns  when  habitats  were  stratified 
b\  DS\'  X  SS  X  OCC.  At  the  highest  level  of 
habitat  stratification,  apparent  differences  from 
expected  use  for  three  habitat  categories  disap- 
peared from  the  analyses. 

Acknowledgments 

This  research  was  supported  b\'  the  USDA 
Forest  Ser\ice,  Rocky  Mountain  Forest  and 
Range  Experiment  Station;  National  Wild 
Turkev  Federation;  Black  Hills  National  Forest; 
and  South  Dakota  Came,  Fish  and  Parks.  We 
extend  special  thanks  for  the  support  and 
encouragement  of  Dr.  A.  J.  Bjugstad 
(deceased).  Technical  assistance  of  R.  Hodorff, 
T.  Mills,  C.  Oswald,  K.  Thorstenson,  K.  Jacob- 
son,  and  L.  Harris  was  appreciated.  M.  Green 
\'olunteered  his  time  throughout  this  study,  and 
R.  Taylor  allowed  access  to  his  property-  for 
trapping  and  data  collection.  Dr.  G.  Hur.st, 
Dr.  R.  fonas,  and  H.  Shaw  reviewed  earlier 
drafts  of  tliis  manuscript. 

LiTER.ATURE  CiTED 

Alldrf.dgf..  J.  R..  and  J.  T  Rvni  1986.  Comparison  of 
some  statistical  techuiejiii's  for  iuialysis  of  resource 
selection,  jounial  of  Wildlife  Management  .50:  157- 
16.5. 

Bf.wktt  D.  L.  I9S4.  Criizinn  potential  of  major  soils 
within  the  Black  Hills  of  South  Dakota.  Unpublished 
master's  thesis.  South  Dakota  State  Uni\ersit\,  Brook- 
ings. 199  pp. 

Boldt,  C.  E.,  and  J.  L.  Van  Duf.sf.n.  1974.  Sikiculture  of 
ponderosa  pine  in  the  Black  Hills:  the  status  of  our 
knowledge.  USDA  Forest  Ser\ice  Research  Paper 
RM-124.  Fort  Collins.  Colorado.  45  pp. 


144 


Great  Basin  Naturalist 


[Volume  52 


Bkvant.  F.  C].,  and  D.  Nisii  1975.  Habitat  use  by  Merriains 
Turkey  in  southwestern  Utdi.  In:  L.  K.  Halls,  ed., 
Proceedings  of  the  Third  National  Wild  Turkey  Sym- 
posium 3:6-13.  Texas  Parks  and  Wildlife  Department, 
Austin. 

Buttery.  R.  F.,  and  B.  C.  Gillam.  1983.  Forest  ecosys- 
tems. Pages  43-71  in  R.  L.  Hoover  and  D.  L.  Wills, 
eds..  Managing  forested  lands  for  wildlife.  Colorado 
Division  of  Wildlife,  in  cooperation  with  US  DA  Forest 
Service,  Rock-v  Mountain  Region,  Denver,  Colorado. 
459  pp. 

Byeks,  C.  R.,  R.  K.  Steinhokst.  and  P.  R.  Kjuu.sman 
1984.  Clarification  of  a  technique  for  analysis  of  utili- 
zation-a\ailability  data.  Jouniiil  of  Wildlife  Manage- 
ment 48:  1050-1053. 

Cochran,  W.  G.  1963.  Sampling  techniques.  John  Wiley 
and  Sons,  Inc.,  New  York.  413  pp. 

Jonas.  R.  1966.  Merriam's  Turkeys  in  southeastern  Mon- 
tana. Techniciil  Bulletin  3.  Montana  Game  and  Fish 
Depiu^tment,  Helena.  36  pp. 

LUTZ,  R.  S.,  and  J.  A.  Crawford  1989.  Habitat  use  and 
selection  of  home  nuiges  of  Merriam's  Turkey  in 
Oregon.  Great  Basin  Naturdist  49:  252-258. 

Mackey,  D.  L.  1982.  EcologyofMerriam's  Turkeys  in  south 
central  Washington  with  special  reference  to  habitat 
utilization.  Unpublished  m;ister's  thesis,  Washington 
State  University  Pullman.  87  pp. 

.  1986.  Brood  habitat  of  Merriam's  Turkeys  in  south- 
central  Washington.  Northwest  Science  60:  108-112. 

MOSTELLER,  F.,  and  A.  Parunak.  1985.  Identifying 
extreme  cells  in  a  sizeable  contingency  table:  probabi- 
listic and  exploratory  approaches.  Pages  189-224  in 
D.  C.  Hoaglin,  F.  Mosteller,  and  J.  W.  Tukey,  eds.. 


Exploring  data  tables,  trends,  and  shapes.  John  Wiley 
and  Sons,  Inc.,  New  York.  527  pp. 

Nenno,  E.  S.,  and  W  M.  Healy.  1979.  Effects  of  radio 
packages  on  behavior  of  wild  turkey  hens.  Journal  of 
Wildlife  Management  43:  460^65. 

Neu.  C.  W.,  C.  R.  Byers.  and  J.  M.  Peek  1974.  A  tech- 
nique for  analysis  of  utilization-availability  diita.  Jour- 
nal of  Wildlife  Management  38:  .541-545. 

Petersen.  L.  E.,  and  A.  H.  Richardson  1975.  The  wild 
turkey  in  the  Black  Hills.  Bulletin  No.  6.  South  Diikota 
Game,  Fish  ;uid  Parks,  Pierre.  51  pp. 

Rose,  B.  J.  1956.  An  evaluation  of  two  introductions  of 
Merriam's  Wild  Turkey  to  Montiina.  Unpublished 
master's  thesis,  Montana  State  College,  Bozemtui.  37 
pp. 

Scott,  V.  E.,  iuid  E.  L.  Boeker  1975.  Ecology  of 
Merriam's  Wild  Turkey  on  the  Fort  Apache  Indian 
Reservation.  In:  L.  K.  Halls,  ed..  Proceedings  of  the 
Third  National  Wild  Turkey  Symposium  3:141-158. 
Texas  Parks  and  Wildlife  Department,  Austin. 

Shaw,  H.  G.  1986.  Impacts  of  timber  harvest  on  Merriam's 
Turkey  populations.  Problem  analysis  report.  Arizona 
Depiirtment  of  Game  and  Fish,  Tucson.  44  pp. 

Shaw,  H.  G.,  and  R.  H.  Smith  1977.  Habitat  use  patterns 
of  Merriam's  Turkey  in  Arizona.  Federal  Aid  Wildlife 
Restoration  Project  W-78-R.  Arizona  Department  of 
Game  and  Fish,  Tucson.  33  pp. 

Thomas,  J.  W.  1979.  Wildlife  habitats  in  managed  forests: 
the  Blue  Mountains  ofOregon  and  Washington.  USDA 
Forest  Service  Handbook  553.  U.S.  Government  Print- 
ing Office,  Washington,  D.C.  512  pp. 

Received  24  June  1991 
Accepted  15  March  1992 


Great  Basin  Naturalist  52(2),  pp.  145-148 

POLLINATOR  PREFERENCES  FOR  YELLOW,  ORANGE,  AND  RED  FLOWERS  OF 
MIMULUS  VERBENACEUS  AND  M.  CARDINALIS 


Robert  K.  Vickerv,  |r. 

Abstract. — Red,  orange,  and  yellow  niorphs  of  Mimitltis  verhetuwens  and  M.  cardiimlis  were  field  tested  for  pollinator 
preferences.  The  species  are  closely  similar  except  that  M.  vcrhoiaccus  flowers  ha\e  partiallv  refle.xed  corolla  lobes,  whereas 
M.  ccirdinalis  flowers  ha\e  fullv  reflexed  corolla  lobes.  On  the  basis  of  oxer  6()(X)  bumblebee  and  hummingbird  visits,  highly 
significant  (/;  <  .001)  patterns  emerged.  Yellow,  which  is  the  mutant  color  morph  in  both  species  and  is  determined  by  a 
single  p;ur  of  genes,  was  strongly  preferred  bv  bumblebees  ;uid  strongly  eskewed  by  Innnmingbirds  in  both  species.  Orange 
and,  to  a  lesser  extent,  red  were  strongK  preferred  b\  hummingbirds  but  eskewed  by  bumblebees  in  both  sjiecies.  Thus, 
strong,  but  partial,  reproductive  isolation  was  observed  between  the  yellow  mutants  and  the  orange-  to  red-flowered 
populations  from  which  they  were  derived.  Color — yellow  versus  orange  iind  red — appeared  more  iniportant  than 
shape — piirtiallv  reflexed  versus  fullv  reflexed  corolla  lobes — in  determining  the  preferences  of  tlu-  guild  of  pollinators  in 
tliis  particular  test  environment  for  Mimulus  vcrhenaceiis  and  M.  cardinalis. 

Kl'ij  words:  Mimulus,  spcciation,  flower  colors,  pollinator  preferences,  bun^lAcbees,  luaninin^hirds. 


How  mucli  of  a  change  in  flower  color  ancl/or 
shape  is  enough  to  lead  to  a  change  in  pollinators 
and  hence  to  reproductive  isolation  and  poten- 
tially to  speciation?  The  flower  color  and  shape 
nioq^hs  o^ Mimulus  verbenaceus  Greene  and  M. 
cardinalis  Douglas  provide  an  excellent  system 
for  addressing  this  intriguing  question. 

Materials 

Mimulus  verbenaceus  and  M.  cardinalis  are 
tspicallv  bright  red  flowered  and  hiuiuningbird 
pollinated.  However,  yellow-flowered  morphs 
occur  in  M.  verbenaceus,  e.g.,  in  a  population  at 
N'assey's  Paradise,  Grand  Canyon,  Arizona,  and 
in  M.  cardinalis  populations,  e.g.,  on  Cedros 
Island,  Baja  California,  Mexico,  and  in  the 
Siskyou  Mountains,  Oregon.  My  experimental 
hybridizations  show  that  yellow  is  due  to  a  single 
pair  of  recessive  genes  that  limit  the  floral 
anthocyanins  to  small  dots  in  the  corolla  throat. 
Intermediate,  orange-flowered  forms  are 
known  in  M.  verbenaceus,  specificallv  the  pop- 
ulation at  Yecora,  Sonora,  Mexico.  And,  an 
intermediate,  orange-flowered  form  of  M.  car- 
dinalis was  obtained  bv  repeated  cycles  of  selec- 
tion. In  both  cases  orange  is  due  to  a  single  pair 
of  quantitative  genes  that  reduce  the  amount  of 


anthocyanin  pigments  in  the  corolla  lobes. 
Thus,  parallel  series  of  red,  orange,  and  vellow 
color  forms  are  available  for  both  M.  ver- 
benaceus andM.  cardinalis  (Table  1). 

Mimulus  verbenaceus  and  M.  cardinalis  are 
similar,  closely  related  species  in  section 
Enjthranthe  (Grant  1924);  however,  their  flow- 
ers differ  in  shape.  Those  of  M.  verbenaceus 
have  only  the  upper  pair  of  corolla  lobes  sharplv 
reflexed,  giving  the  flowers  a  partiallv  tubular 
aspect.  The  side  pair  of  lobes  and  the  labellum 
curve  gently  forward  forming  a  bee  landing  [)lat- 
form.  Flowers  of  M.  cardinalis  have  both  the 
upper  and  side  corolla  lobes  shaq)K'  reflexed, 
giving  the  flowers  a  fully  tubular  shape.  The 
labellum  is  thrust  fonvard  and  is  fokk-tl  on  itself 
forming  a  ridge  instead  of  a  landing  platform. 
Shapes  of  the  flowers  of  both  species  would 
seem  to  invite  hummingbirds.  Flowers  of  M. 
verbenaceus  but  not  those  of  M.  cardinalis 
would  appear  adaj)ted  for  bees  as  well.  How- 
ever, flowers  of  all  three  color  moq^hs  of  both 
species  showed  no  reflectance  patterns  in  the 
ultraviolet,  that  is,  no  putative  bee  nectar 
guides.  Thus,  flower  shapes  of  A/,  verbenaceus 
and  M.  cardinalis  are  similar  in  some  respects 
but  differ  in  others  of  potential  significance  to 
pollinators. 


Department  of  Biolog\',  University  of  Utah,  Salt  Lake  CAW.  Utah  841  IS 


145 


146 


Great  Basin  Naturalist 


[Volume  52 


Plan 

The  effect  of  flower  color  and  flower  shape 
on  pollinator  preferences  will  be  addressed 
stepwise.  First,  pollinator  preferences  for 
color — red,  orange,  and  yellow — will  be  ascer- 
tained for  M.  verhenaceus  plants  onl)',  holding 
flower  shape  constant.  Second,  red-,  orange-, 
and  yellow-flowered  M.  cardinalis  plants  will  be 
added  to  the  experiment.  Are  pollinator  prefer- 
ences for  red,  orange,  and  yellow  flowers  of  M. 
cardinalis  the  same  as  for  those  of  the  M.  ver- 
henaceus series?  Note  that  the  pigments  are 
identical  (Vickery  1978).  Or,  does  the  difference 
in  corolla  shape  between  the  two  species  lead  to 
a  difference  in  pollinator  preferences? 

Methods 

Seeds  for  each  of  the  six  populations  of  the 
study  (Table  1)  were  collected  in  the  wild  or 
harvested  from  transplants  brought  into  the 
greenhouse  except  those  of  orange  M.  car- 
dinalis, which  were  obtained  by  selection.  A 
large  population  of  red  M.  cardinalis  from 
Cedros  Island  was  grown  and  the  most  orange- 
red  flowered  plant  self-poUinated.  Its  progeny 
included  several  orange-flowered  plants.  Prog- 
eny of  these  plants  were  grown  and  found  to 
breed  true  for  orange  and  were  used  as  the 
source  of  seeds  for  the  orange  M.  cardinalis 
moiph. 

Seeds  of  the  sLx  populations  were  sown  in 
early  April  1988  in  the  University  of  Utah  green- 
house, following  which  seedUngs  were  trans- 
planted into  4"  plastic  pots  and  grown  to 
flowering.  Pots  were  placed  in  plastic  travs  to 
facilitate  bottom-watering,  plants  being  ran- 
domly arranged  as  to  flower  color. 

When  plants  began  flowering,  they  were 
moved  outdoors  to  test  pollinator  preferences. 
Instead  of  using  Red  Butte  Canyon  Natural 
Research  Area  as  before  (Vickery  1990),  with  its 
relative  paucity  of  pollinators,  I  scattered  the 
plants  on  a  lawn  adjacent  to  native  gambel  oak 
clumps  at  the  mouth  of  Parley  s  Canyon  of  the 
Wasatch  Mountains  in  an  area  rich  in  pollina- 
tors. Some  50  to  100  plants  of  each  color  morph 
of  M.  verhenaceus  made  up  the  artificial  popu- 
lation of  the  first  part  of  the  experiment.  Some 
50  to  100  plants  of  red  and  of  orange  M.  car- 
dinalis plus  20  plants  of  yellow  A/,  cardinalis  (all 
that  were  available)  were  added  to  the  M.  ver- 


Tablf,  L  Origin  of  populations  studied. 

Mimulus  verhenaceus  Greene 

Vasscij's  Paradise,  Grtuid  Caiivon,  Arizona,  USA,  elev. 

-650  m 

Red  moq^h  =  culture  number  14,088 
Yellow  morph  =  culture  number  14,089 

Yrcora,  Sonora,  Mexico,  elev.  —1,550  m 
Orange  =  culture  number  13,256 

Mimulus  cardinalis  Douglas 

Isia  Cedros.  Baja  California,  Mexico,  elev.  -100  m 
Red  moq^h  =  culture  number  13,106 
Yellow  morph  =  culture  number  13,2.50 
Orange  =  culture  number  13,249 

(obtained  by  selection  from  the  red  moiph) 


henacens  plants  for  the  second  part  of  the  exper- 
iment. 

Pollinator  visits  to  the  flowers  were  observed 
and  recorded  for  an  average  of  \Vi  hours  per 
observation  period  for  15  periods  for  each  of  the 
two  parts  of  the  experiment  (Tables  2,  3).  Time 
of  day  of  the  observations  was  varied  to  be  sure 
of  noting  all  the  different  kinds  of  \isitors.  To 
count  as  a  visit,  a  hummingbird  had  to  thrust  its 
beak  into  a  flower.  A  bee  had  to  land  on  the 
flower  and  crawl  into  the  flower  far  enough  to 
brush  the  stigma  and  anthers.  A  fly,  butterfly, 
etc.,  had  to  walk  on  the  reproductive  structures. 
The  numbers  of  flowers  rather  than  plants  of 
each  color  of  each  species  were  recorded  for 
each  observation  period. 

For  analvsis  of  visits,  chi-square  tests  were 
Rm  for  each  obseivation  period  for  each  part  of 
the  experiment.  The  null  hyj^othesis  was  that 
hummingbirds  or  bumblebees  (very  few  flies, 
butterflies,  etc.,  visited  the  flowers  and  were  not 
listed)  would  visit  the  three  colors  of  flowers  of 
M.  verhenaceus  in  the  first  part  of  the  experi- 
ment and  the  three  colors  of  M.  verhenaceus 
and  M.  cardinalis  in  the  second  part  of  the 
experiment  in  proportion  to  the  mmibers  of 
those  flowers  in  the  experimental  population 
(Tables  2,  3).  If  the  overall  chi-square  value  for 
a  period  of,  for  example,  bee  visits  to  M.  ver- 
henaceus or  hummingbird  visits  to  M.  cardinalis 
indicated  a  significant  deviation  from  expected 
values,  then  the  frecjuencv  of  \isits  to  each  color 
was  inspected.  Those  high  or  low  enough  that 
their  term  in  the  chi-square  equation  was  large 
enough  b\'  itself  to  produce  a  significant  devia- 
tion at  the  5%  level  were  considered  to  be 
significant  (Tables  2,  3). 


19921 


MiMUIA'S  FOLIJNATOH  PREFERENCES 


14' 


TablK  2.  Pollinator  pifffrencL'S  for  rt-d.  orange,  or  \x'llow  noweis  ol  Mimulus  rcrhciuiccii.s  in  19S8. 


Numbers  of  flowers 

Bumblebee  visits 

Hummingbi 

ird  visits 

Month:cla\:time 

Red 

Orange 

Yellow 

Red 

Orange 

Yellow     P 

Red 

Orange 

Yellow     P 

7:26:1630 

48 

56 

70 

28i" 

523T 

1984- 

<.001 

0 

3 

0 

<.100 

7:29:0745 

56 

91 

74 

30 

50 

58 

<.200 

0 

SIT 

29 

<.010 

7:30:0710 

46 

79 

114 

24 

67 

67 

<.010 

55T 

66 

70 

<.010 

8:02:1640 

85 

77 

74 

3i 

8ST 

53 

<.001 

27 

49 

27 

<.010 

8:03:0630 

92 

101 

133 

53 

99T 

81 

<.()01 

33 

79T 

36i 

<.001 

8:03:1.540 

120 

117 

172 

3U 

74 

209T 

<.()01 

lOOi 

24  IT 

183 

<.001 

8:04:0640 

S6 

73 

178 

(U 

5 

52T 

<.0()] 

S3 

145T 

170 

<.001 

8:05:0715 

120 

UK) 

169 

33i 

71 

125 

<.()01 

9i 

77T 

28i 

<.001 

8:05:1645 

126 

104 

174 

12i 

22i 

126T 

<.001 

36i 

149T 

92i 

<.001 

8:05:1830 

126 

104 

174 

5i 

4i 

73T 

<.001 

75 

150T 

82i 

<.001 

8:06:0<S40 

126 

88 

151 

74i 

159T 

291T 

<.001 

66 

lOOT 

26i 

<.001 

8:06:1445 

126 

9S 

150 

6i 

61 

60T 

<.001 

49 

94T 

24i 

<.001 

8:06:1810 

130 

117 

142 

50 

105 

257T 

<.001 

31 

4ST 

U 

<.001 

8:07:1515 

130 

119 

142 

Oi 

4i 

68T 

<.001 

52 

125T 

5i 

<.001 

8:08:0725 

118 

91 

124 

12i 

32 

131 T 

<.001 

32 

67T 

5i 

<.(X)1 

■"T  or  J.  =  significantly  high  or  low;  see  text. 


Table  3.  Pollinator  preferences  for  red,  orange,  or  yellow  flowers  of  M.  verhenaceus  and  M.  canliiuilis  in  1988. 


Number  of  flowers 


Month:day:time  Red      Orange      Yellov 


Bumblebee  visits 


Hummingbird  visits 


Red 

Orange 

Yellow     P 

Red 

Orange 

Yellow     P 

Mil 

mtiltis  vc 

rhenacens 

17i'' 

29 

62T 

<.(X)1 

23 

40T 

34 

<.001 

16i 

36 

131T 

<.001 

70 

70T 

184 

<.()01 

2i 

2U 

124T 

<.(K)1 

171 

167T 

1.354 

<.001 

841 

129T 

190T 

<.001 

13 

10 

3 

<.001 

13i 

56 

202T 

<.(K)1 

404 

80T 

384 

<.001 

.844- 

1()6T 

237T 

<.(M)1 

60 

.5()T 

04 

<.001 

mi 

97 

160 

<.0Ol 

196 

166T 

994 

<.001 

5i 

94 

120T 

<.(K)1 

168 

147T 

163 

<.001 

54i 

66 

172T 

<.(K)1 

115 

63T 

.564 

<.001 

3i 

4i 

160T 

<.0()1 

44 

31 T 

24 

<.001 

27i 

37 

162T 

<.(X)1 

71 

WT 

244 

<.()01 

39i 

36 

167T 

<.001 

54 

37 

38 

<.010 

2i 

3 

50T 

<.(K)1 

7 

2 

2 

<.300 

14i 

84, 

174T 

<.CX)1 

21 

.3()T 

04 

<.001 

3i 

24 

128T 

<.(M)1 

66 

72T 

264 

<.0()1 

Mil 

<nultis  CO 

rdinalis 

36-L 

59 

89T 

<.001 

28 

25 

14 

<.001 

2U 

37 

34T 

<.001 

61 

36 

04 

<.001 

18 

8 

22T 

<.(X)1 

137 

117T 

244 

<.001 

49 

59 

26 

<.010 

4 

6 

0 

<.1(X) 

4S 

104 

48T 

<.(X)1 

81 

.53 

124 

<.010 

274 

62 

49T 

<.0()1 

.34 

20 

0 

<.020 

33 

26 

34T 

<.(X)1 

6,3 

95T 

13 

<.001 

18 

S 

10 

<.().50 

1.30 

88 

21 

<.100 

35 

40 

17 

<.010 

59 

41 

04 

<.010 

28 

15 

16T 

<.0Ol 

91 

92 

14 

<.300 

39 

24 

19T 

<.(X)I 

77 

52 

34 

<.020 

20 

33 

26T 

<.(X)1 

.55 

58 

04 

<.001 

13 

4 

6T 

<.010 

11 

24 

0 

<.010 

18i 

21 

34 

<.(X)1 

.34 

74T 

04 

<.001 

19i 

47 

12 

<.(X)1 

204 

115T 

44 

<.001 

8:08: 16(M) 
8:09:07.50 
8:09:1705 
8:10:0815 
8:10:1640 
8:11:0810 
8:12:0805 
8:12: 17(X) 
8:13:08.55 
8:1.3:1800 
8:14:0815 
8:15:0740 
8:1.5:17(X) 
8:16:08.30 
8:17:06.30 

8:08: 16(X) 
8:09:0750 
8:09:1705 
8:10:0815 
8:10:1640 
8:11:0810 
8:12:0805 
8:12: 17(X) 
8:13:08.55 
8:1.3: 18(X) 
8:14:0815 
8:15:0740 
8:1.5:1700 
8:16:08.30 
8:17:06.30 


117 

92 

132 

115 

73 

116 

115 

73 

116 

145 

90 

143 

145 

90 

143 

175 

83 

177 

200 

111 

198 

200 

HI 

198 

180 

83 

175 

180 

87 

175 

212 

81 

165 

184 

94 

183 

184 

94 

183 

206 

112 

1.53 

214 

S6 

177 

79 

47 

61 

69 

45 

32 

69 

45 

32 

61 

39 

23 

61 

39 

23 

61 

55 

12 

65 

51 

18 

65 

51 

18 

64 

42 

14 

89 

81 

14 

83 

69 

15 

53 

71 

15 

53 

71 

15 

79 

78 

21 

89 

79 

18 

'T  or  i  =  significantly  liigh  or  low;  see  text. 


148 


Great  Basin  Naturalist 


[Volume  52 


Results 

Pollinators  showed  clear,  veiy  highly  signifi- 
cant if)  <  .001)  preference  for  or  avoidance  of 
yellow  flower  color,  but  less  clear  preferences 
for  or  avoidance  of  orange  or  red  flower  colors. 
Bunil)lehees — ^principally  Bomlms  appositiis  and 
B.  huntii — strongly  preferred  vellow  in  both  M. 
verhenaceus  and  M.  cardinalis.  Difference  in 
shape  did  not  appear  to  matter.  Humming- 
birds— principally  Selasphonis  plati/cenis — 
strongly  eskewed  yellow  in  both  species  (Tables 
2,  3).  Agiiin,  difference  in  shape  did  not  appear 
to  matter. 

Hummingbirds  significantly  {p  <  .001)  pre- 
ferred orange  M.  verhenaceus  flowers  and 
showed  a  tendency  to  prefer  orange  M.  car- 
dinalis  flowers  as  well  (Tables  2,  3).  This  prefer- 
ence for  orange  over  red  flowers  should  not  have 
been  surprising  in  view  of  the  fact  that  orange 
and  red  are  equally  conspicuous  to  humming- 
birds (Grant  and  Grant  1968,  Raven  1972). 

Strong  preferences  and  aversions  for  yellow 
are  particularly  interesting  because  yellow  is  the 
mutant  color  in  both  species.  So,  a  new  yellow 


mutant  of  either  species  would  be  preferentially 
visited  by  bumblebees  and  preferentially 
avoided  bv  hummingbirds,  but  not  in  all-or- 
none  reactions.  Apparently  then,  with  the  spe- 
cies of  pollinators  tested,  we  are  seeing  the 
establishment  of  real,  but  partial,  reproductive 
isolation  due  to  the  mutation  of  a  single  pair  of 
color  senes. 


Literature  Cited 


Grant.  A.  L.  1924.  A  monograph  of  the  genus  Mimiilus. 

Annals  of  the  Missouri  Botanical  Garden  IL  99-.3S9. 
Grant,  K.,  and  V.  Grant.  1968.  Hummingbirds  and  their 

flowers.  Columbia  University  Press,  New  York.  11.5  pp. 
Raven,  R  H.  1972.  Why  are  bird-\'isited  flowers  predomi- 
nantly red?  Evolution  26:  674. 
ViCKERY,  R.  K.,  Jr    1978.  Case  studies  in  the  evolution  of 

species  complexes  in  Mimulus.  Evolutionary  Biology 

11:404^506. 
.  1990.  Pollination  experiments  in  the  Mimulus 

cardinalis-M.  lewisii  complex.  Great  Basin  NatunJist 

.50:  1.55-159. 


Received  21  October  1991 
Accepted  1  May  1992 


Crrat  Basin  Xatiinilist  52(2),  pp.  149-154 

SOIL  LOOSENING  PROCESSES  FOLLOWING  THE  ABANDONMENT 
OF  T\VO  ARID  WESTERN  NEVADA  TOWNSITES 

Paul  A.  Kiiapp 

ABSTRACrr. — Soil  compaction  was  measured  at  four  sites  within  two  abandoned  mining  camps  in  the  western  CIreat  Basin 
Desert,  Ne\ada.  Bulk  densitv*  and  macroporositv  xiilues  were  generated  from  soil  samples  collected  in  areas  ol  different 
liuid  use  intensities  in  camps  that  had  been  abandoned  for  approximatek-  70  \e;irs.  Results  show  that  significant  differences 
remain  in  bulk  density  values  bet^\'een  abandoned  roads  and  undisturbed  areas  in  both  towiis,  and  that  the  areas  around 
foundation  peripheries  ;u"e  still  signifkiuitK'  more  compacted  in  one  towai.  There  were  no  significant  differences  between 
liuid  use  groups  as  measured  bv  macroporositv.  Estimated  soil  recoven',  based  on  a  linear  model  using  bulk  densitv  v;iJues, 
suggests  that  appro.ximatek-  KX)  to  1.30  ve;u-s  are  necessary  for  complete  loosening  to  occur  for  abandoned  roads,  and  that 
100  or  fewer  \ears  are  necessar\-  for  complete  amelioration  of  the  foundation  peripher\-  iireas.  The  wetter  towaisite,  with 
more  freeze-thaw  davs,  finer-grained  soils,  and  greater  plant  cover,  had  shorter  recoverx  estimates.  These  findings  suggest 
that  die  results  of  human-use  impacts  in  arid  areas  may  still  be  apparent  long  ;ilter  disturbances  cease. 

Ki-i/  tcords:  soil  rccovrn/.  soil  roinpnrfioii.  arid  hnuh.  Great  Basin  Desert,  g/iosf  toiins. 


Arid  lands  are  undergoing  enxironmental 
degradation  processes  at  a  rapid  rate  worldwide 
and  are  being  severely  disturbed  by  excessive 
soil  erosion  and  salinization  (Allen  1988,  Goudie 
1990).  The  explosion  in  human  population 
levels  in  the  last  sexeral  decades  in  arid  regions 
has  been  a  major  cause  for  land  degradation, 
especially  considering  that  arid  regions  are 
particularly  sensitive  to  anthropogenic  land  use 
impacts  (Goudie  1990).  While  the  greatest 
extent  of  soil  degradation  has  occurred  in  StJiel- 
ian  Africa,  other  arid  zones  of  the  world  are  also 
\ulnerable  (Goudie  1990). 

The  arid  American  West  is  one  such  region 
where  human  use  impacts  liave  risen  dramati- 
cally in  the  last  sexeral  decades  (Francis  and 
Ganzel  1984).  The  increased  popularits'  of  back- 
country  visits  by  off-road  vehicles,  mountain 
bikes,  backpackers,  or  horseback  riders  has  had 
a  considerable  impact  on  the  surrounding  envi- 
ronment, either  damaging  or  altering  both  the 
flora  and  soils  of  affected  areas  (Cole  1983, 
1987,  1990,  Lathrop  1983,  Webb  1983,  Prose 
andMetzger  1985). 

Compaction  of  desert  soils  caused  bv  back- 
country  activities  can  decrea.se  infiltration  rates, 
increase  nmoff,  and  impede  plant  root  growth, 
which  favors  further  soil  degradation  processes 
(Vollmeretal.  1976,  Rowlands  and  Adams  1980, 


Hincklev  et  al.  1983,  Lathrop  1983,  Prose  et  al. 

1987,  Goudie  1990).  While  the  impacts  of  back- 
countiv  activities  have  been  documented  over 
short  time  spans  (often  less  than  30  years),  little 
is  known  about  long-term  consequences  of 
these  activities  (Knapp  1991).  Few  studies  exist 
that  document  how  well  a  disturbed  area  recov- 
ers following  cessation  of  disturbances,  particu- 
larly in  areas  traditionally  considered  to  have 
little  economic  value,  such  as  arid  lands. 

Recovery  processes  of  compacted  soils  are 
not  well  understood  (Webb  et  al.  1983,  1986) 
and  liave  been  conducted  primarily  in  more 
mesic  environments  (Webb  et  al.  1983,  Knajip 
1989).  Recovery  estimates  varv'  considerably, 
ranging  from  less  than  10  years  on  Minnesota 
forest  soils  (Thorud  and  Fris,sell  1976),  to  23 
years  on  Idaho  forest  .soils  (Froehlich  et  al. 
1985),  to  50  years  on  forest  soils  in  South  Aus- 
tralia (Greacen  and  Sands  1980),  and  up  to  200 
years  on  soils  in  southwestern  Montana  (Knap[) 
1989). 

The  few  studies  that  have  examined  .soil 
recoven  rates  in  the  arid  American  West  have 
been  confuuHl  to  the  Mojave  Desert  (i.e.,  Webb 
and  Wilshire   1980,  Webb  et  al.   1983,  1986, 

1988,  Prose  and  Metzger  1985).  Rates  of  .soil 
recoven  from  the.se  studies  of  abandoned 
mining  camps  ranged  from  80  to  140  years  and 


Department  of  GedKrapliy.  Uiii\ersit\  of  Nevada.  Keiio,  Nevada  S9.557-0()4S. 


149 


150 


Great  Basin  Naturalist 


[Volume  52 


-^■^^j^!^f:^i'^4^Mtijik 


Fig.  la.  Terrill,  ca  1920,  looking  northwest.  Photo  by  Roly  Ham,  courtesy  Special  Collections,  University  of  Nevada, 
Reno,  Librarv. 


■-"NIC^, 


V^ 


^*^*    «dlirHi- 


Fig.  lb.  Terrill,  1990.  Photograph  I )\  author. 


averaged  100  years.  Comparable  studies  have 
yet  to  be  conducted  in  the  Great  Basin  Desert. 
Ghost  towns  abandoned  in  the  earlv  twenti- 
eth century  in  the  western  Great  Basin  Desert 
showcase  the  long-term  effects  of  soil  compac- 


tion. Built  because  of  the  discovery  of  valuable 
ores  such  as  gold  and  silver,  these  towns  were 
short-lived  as  the  ores  became  too  scarce  to 
extract  profitably  (Palier  1970,  Carlson  1974, 
Shamberger   1974).   These  towais  have  been 


1992] 


Great  Basin  Soil  Kkcon  ehy 


151 


Table  1.  Cliinatic  ;uicl  soils  data  lor  tlie  hvo  selected  Great  Basin  Desert  townisites. 


Est. 

Est." 

p:st.° 

annual 

mean 

mean 

Sand,  silt 

Townsite 

Ele\atioii 

(m) 

precipitation 

(mm) 

Jan.  temp. 

{°C) 

JiiK  temp. 

■(°c) 

.Soil 
type 

and  cla\ 

(%)  ' 

Terrill 
Wonder 


1305 
1740 


125 
2.50 


-0.8 
-3.9 


22.8 
20.5 


loamy  sand 
sandy  loam 


S4/12/4 
46/50/4 


SoiiR-e  of  estimate:  HouglUon  et  al.  1975. 

ex|X).sed  to  a  xarietv  of  envdronmental  impacts, 
including  trampling  by  livestock,  humans,  and 
\ehicles,  and  lia\e  shown  a  \ariet\'  of  vegetation 
reco\er\'  responses  (Knapp  1992).  The  piupose 
of  this  paper  is  to  examine  the  effects  of  soil 
reco\"er\-  in  two  abandoned  mining  towns  in  the 
Great  Basin  De.sert  in  similar  fashion  to  those 
studies  conducted  in  the  Mojave  Desert. 

Study  Areas 

Two  measures  of  soil  compaction,  bulk  den- 
sity' and  percentage  macroporositv;  were  gath- 
ered from  Terrill  and  Wonder.  Terrill  (39°05'N, 
11S°46'\V)  and  Wonder  (39°35'N,  118°04'W) 
were  abandoned  in  ca  1915  and  ca  1925,  respec- 
tivelv  (Figs,  la,  lb).  Both  sites  lie  at  the  base 
of  north-south  trending  fault-block  mountiiin 
ranges  in  central  western  Nevada,  although 
Terrill's  elevation  ( 1305  m)  is  substantially  lower 
than  Wonder's  (1740  m).  Terrill  is  the  drier  site, 
receiving  approximatelv  130  mm  of  precipita- 
tion annuallv  with  the  estimated  mean  January 
and  July  temperatures  being  -0.8  G  and  22.8  G, 
respectively  (Houghton  et  al.  1975;  Table  1). 
The  vegetation  in  Terrill  is  a  salt  desert  scrub 
habitat  txpe  (Tueller  1989),  and  common  spe- 
cies are  the  shnibs  Sarcobatus  bailey i,  Atriplex 
confertifolia,  and  Tetradijmio  spp.;  the  grasses 
Onjzop.sis  Ju/men()i(les  and  Bromus  tectoniin; 
and  the  forb  Spliaeralcea  ainlji^iia.  (iroimd 
cover  in  Terrill  is  approximatelv  2()9f  (Knapp 
1992).  Wonder  receives  appro.ximateK'  250  mm 
of  annual  precipitation,  has  mean  January-  and 
July  temperatures  of  -3.9  G  and  20.5  G,  respec- 
tively (Houghton  et  al.  1975;  Table  1),  and  sup- 
ports a  sagebnish/grass  habitat  t\pe  (Tueller 
1989)  with  appro.ximatelv  35%  ground  cover 
(Knapp  1992).  Gommon  species  in  Wonder  are 
the  shrub  Artemisia  tridentata  and  the  grasses 
6.  tectonim  and  Sitatiiou  hifstrix. 

Both  towiisites  have  alluviallv  deposited,  vol- 
canic sandv-loam  to  loam\'  sand  soils  (Stewart 
and  Garlson  1978;  Table  1).  The  soils  in  Terrill 


are  sandy,  mixed,  T\pic  Galciorthids,  while 
Wonders  soils  are  fine-loamv;  mixed,  T\pic  C Galci- 
orthids (USDA-SGS  1975).  Organic  matter  was 
estimated  to  be  less  than  1%  at  both  towiisites. 
Terrill  and  Wonder  have  been  subjected  to 
minimal  human -caused  impacts  since  abandon- 
ment because  of  their  remote  locations.  Little 
grazing  by  domestic  animals  has  occurred  in 
Terrill  because  of  the  lack  of  a  nearby  water 
source.  Wonder  has  experienced  greater  graz- 
ing pressiues  by  sheep,  cattle,  and  feral  horses. 
Neither  sheep  nor  cattle  have  grtized  the 
Wonder  area  since  1980  (A.  Anderson,  District 
Range  Gonservationist,  BLM,  personal  com- 
munication, 1990). 

Methods 

Soil  samples  for  bulk  densitv  and  macro- 
porosit)'  measurements  were  gathered  at  foiu" 
different  land  use  categories  at  each  town.  Data 
were  collected  from  active  roads  (to  get  a  theo- 
retical upper  limit  of  compaction),  abandoned 
roads  (representing  prior  high-intensit)'  land 
use),  areas  within  5  m  of  foundation  peripheries 
(representing  prior  moderate-intensit\'  land 
use),  and  contrf)l  plots  (an^is  of  minimal  distur- 
bance located  near  [<2  km]  the  townsite).  .All 
efforts  were  made  to  ensure  that  the  four  differ- 
ent land  use  groups  within  each  towaisite  were 
similar  to  each  other  in  terms  of  slope,  aspect, 
.soil  texture,  elevation,  and  parent  material  so 
that  accurate  coiuparisons  could  be  mack-.  Trails 
caused  bv  either  (era!  liorses  or  small  iiianmials 
were  avoided. 

Soil  data  from  the  controls,  active  roads,  and 
abandoned  roads  were  gathered  using  a  strati- 
fied, imaligned  sampHng  method.  Thirt}'  5-m 
line  transects  were  set  parallel  to  both  the  active 
and  abandoned  roads,  and  one  soil  core  was 
gathered  at  a  random  point  along  each  line 
transect.  Soil  cores  from  control  plots  were 
gathered  at  a  random  point  on  each  of  fort}-  5-m 
line  transects.  Soil  cores  were  also  gathered  at  a 


152 


Gre.at  Basin  Natuh.m.isi 


[X'olume  52 


T.ABLE 

2. 

Bulk  del 

Lsity 

and  macroporo.sity  \ 

iilues. 

and 

recovei^'  period  estim 

ate.s 

for 

abandoned  townsites. 

Bulk  Den.sity"' 

(g/cm^) 

Macroporosity 
(%byvol.) 

Recovery  period  (years) 

Site 

Bulk  densit\'       Macroporositv 

Terrill 


Active  road 

1.65  ±  0.04'' 

19.7  ±  2.4'' 

Abandoned  road 

1.51*    ±  0.05 

21.6*  ±  2.3 

Foundation 

peripheries 

1.47*  ±0.07 

21.9*  ±2.7 

Control  plot 

1.41*'^  ±  0.03 

22.7*  ±  2.4 

1.59  ±  0.08 

Wonder 

Active  road 

17.3  ±  2.0 

Abandoned  road 

1.48*   ±  0.07 

20..3*  ±  1.9 

Foundation 

peripheries 

1.46*  ±  0.07 

20.8*  ±  0.8 

Control  plot 

1.42*  ±0.06 

21.1*  ±  1.1 

130 
100 


100 

85 


120 
100 


80 
70 


■"Bulk  density  data  with  exception  of  active  roads  and  standard  deviation  vahies  are  trom  Knapp  1992. 
One  standard  deviation. 

=  Significantly  different  {p  =  .0.5)  from  active  road  based  on  Tukey  test. 
=;  Significantlv  different  ip  =  .05)  from  control  plot  based  on  Tiikev  test. 
=  Significantlv  different  (/)  =   (15)  from  fbnndation  peripheries  based  on  Tukey  test. 


random  point  on  each  of  forty  5-ni  line  transects 
that  were  set  perpendicular  to  the  foundation 
periphery  sides.  The  cores  were  oven-dried 
overnight  and  then  weighed  for  bulk  densitv 
(Blake  1965).  One-fourth  of  the  cores  also  were 
kept  intact  for  macroporosity  readings  that  were 
measured  under  30  cm  of  tension  using  a  ten- 
sion-table (Orr  1960).  Soil  te.xture  was  mea- 
sured using  the  micro-pipette  method  (Miller 
and  Miller  1987). 

Analysis  of  variance  (ANOVA)  was  used  to 
examine  whether  differences  in  either  bulk  den- 
sity or  macroporosity  values  existed  between 
land  use  categories  (abandoned  roads,  founda- 
tion peripheries,  and  control  plots)  for  each 
town  (Zar  1984,  SAS  1985).  Where  significant 
overall  differences  existed,  Tukey  multiple  com- 
parison tests  were  used  to  determine  between 
which  groups  these  differences  occurred  (Zar 
1984).  Soil  recovery  was  considered  complete 
when  no  significant  differences  existed  between 
disturbed  sites  and  their  respective  control 
plots. 

Soil  recover)'  estimates  were  based  on  the 
equation  (corrected  from  Webb  et  al.  1986): 

T=[(Ia-Iu)/(Ia-It)]'TA 
where     It  =  townsite  (either  abandoned  road  or 
foundation  periphery) 

In  -  undisturbed  soils  (control  plots) 

la  =  active  road 

Ta  =  time  since  abaudomnent  of 
towTisite 


The  data  collected  from  active  roads  were  used 
only  for  estimates  generated  by  this  equation. 
This  equation  generates  rough  estimates  of  soil 
recoven.'  times.  Webb  et  al.  (1986)  state  that  an 
exponential  deca\'  model  might  gi\e  more  real- 
istic soil  reco\eiy  estimates,  although  onlv  one 
abandonment  time  per  site  excludes  the  use  of 
the  exponential  decay  model. 

Results 

Bulk  densit\'  measurements  for  the  aban- 
doned road  (1.51  g/cm^)  and  foundation  periph- 
eries (1.47  g/cm')  were  significantl)-  greater  dian 
for  the  control  plot  (1.41  g/cm^)  in  Terrill,  but  in 
Wonder  onlv  the  abiuidoned  road  (1.48  g/cm^) 
had  significanth'  greater  bulk  densits'  values 
than  tiie  control  plot  (1.42  g/cm^)  CRible  2). 
Macroporositv'  measurements  in  botli  Terrill  and 
Wonder  were  not  significantK  different  between 
land  use  categories. 

Estimated  recoven'  times  ranged  from  85  to 
1 30  years  when  based  on  bulk  density  measure- 
ments, and  from  70  to  120  years  when  based  on 
macroporositv  measurements  (Table  2).  All 
measiuvments  were  greater  on  abandoned 
roads  than  on  foimdation  peripheries  and  were 
comparatively  longer  in  Terrill  than  in  Wonder. 
While  these  values  are  derixed  b\  a  linear  recov- 
eiv  model,  it  is  most  likeh  that  soil  recovery 
follows  more  of  a  nonhnear  path  with  rapid 
reco\eiy  early,   then  recover)'  rates   slowing. 


1992] 


Great  Basin  Soil  Hkcon  ehv 


153 


Heinonen  (1977)  has  suggested  that  the  l)ulk 
densitv'  of  soils  may  decrease  to  a  certain  point, 
then  le\el  off  without  reaching  predisturbance 
conditions. 

Discussion 

Soil  loosening  is  dependent  upon  shrink- 
swell,  freeze-thaw,  and  biological  acti\it}-  pro- 
cesses (Larson  and  Alhnaras  1971,  Akrani  and 
Kemper  1979,  Webb  1983,  Webb  et  al.  1986, 
Knapp  1989).  These  processes  in  turn  may  be  a 
function  of  soil  type,  climate,  and  biological 
acti\it\.  The  recover)'  times  for  Terrill  and 
Wonder  show  a  relationship  to  all  three  ot  these 
processes,  with  recovery  times  in  Terrill  being 
longer  than  those  in  Wonder. 

Soil  texture  is  important  because  finer- 
grained  soils  are  more  prone  to  freeze-thaw  and 
shrink-swell  loosening  processes  than  are 
coarser-grained  soils  (Webb  et  al.  1986).  Fine- 
textured  soils  have  more  total  pore  space  and 
have  a  higher  water-holding  capacit\',  thereby 
pro\'iding  the  soils  of  Wonder,  that  are  more 
fine-grained  than  Terrill,  more  opportunities 
for  expansion-contraction  processes  to  occur 
(Millar  et  al.  1958).  While  percentages  of  clay 
mav  also  be  important,  particularly  if  the  clay 
has  a  high  shrink-swell  ratio,  total  amounts  of 
clay  at  the  two  towns  were  the  same  and  should 
not  have  a  greater  effect  at  one  place  than  at  the 
other. 

Climate  plaws  an  important  rc^le  in  soil  loos- 
ening processes,  particularlv  where  there  is  a 
high  frequency  of  wetting  and  drying,  freezing 
and  thawing,  or  heating  and  cooling  processes. 
Three  climatic  features  favor  faster  soil  loosen- 
ing processes  in  Wonder  than  in  Terrill.  First, 
\V  onder  is  435  m  liigher  than  Terrill  and  Wonder 
has  a  shorter  freeze-free  period  by  approxi- 
mately a  month  to  a  month  and  a  half  (J.  James, 
Nevada  State  Climatologist,  personal  communi- 
cation 1991).  Second,  Wonder  lies  at  the  base  of 
a  bowl-shaped  depression  and  receives  maxi- 
mum cold-air  drainage.  Typical  diurnal  temper- 
ature contrasts  for  Wonder  range  from  22  to  28 
C,  with  the  greatest  contrasts  occurring  in  the 
summer  and  the  least  contrast  in  the  winter 
(Houghton  et  al.  1975,  J.  James,  personal  com- 
munication, 1991).  Terrill,  on  the  other  hand, 
experiences  a  16.5  to  22  C  diurnal  temperature 
range  (Houghton  et  al.  1975,  J.  James,  personal 
communication,  1991).  These  differences  in 
diurnal   temperature   range   suggest   that   the 


heating-cooling  and  expansion-contraction  pro- 
cesses are  more  pronounced  for  Wonder  Third, 
Wonder  receiws  approximateK  twice  as  much 
annual  precipitation  as  Terrill;  therefore,  the 
freezing-thawing  and  wetting-diving  processes 
should  occur  more  often  in  Wonder,  facilitating 
the  soil  loosening  processes. 

Biological  acti\it\'  through  plant  root  growth 
can  also  ameliorate  soil  compaction.  Cxrasses 
and  forbs  are  particularly  effective  for  loosening 
of  topsoil  because  they  have  manv  diffuse,  shal- 
low roots  that  penetrate  the  topsoil  with  subse- 
quent minimal  increases  in  soil  strength,  but 
leave  behind  small  channels  after  the  roots  die 
(Webb  et  al.  1983).  Plants  such  as  shrubs,  with 
a  central  taproot,  however,  cause  localized  com- 
paction around  the  root,  yet  have  fewer  roots 
per  unit  volume  and  are  less  effectixe  for  soil 
loosening  (Webb  et  al.  1983).  Total  plant  cover 
in  Wonder  was  substantiallv  (approximately 
20%)  greater  than  in  Terrill,  especialK*  with  the 
grasses  Bromns  tectontm  and  Sitanion  Jii/strix, 
which  both  have  extensive,  shallow  root  sys- 
tems. Therefore,  it  appears  that  if  soil  loosening 
can  be  attributed  to  biological  activity;  it  would 
be  more  pronounced  in  Wonder  than  in  Terrill, 
although  controlled,  detailed  experiments  are 
necessai"v  for  confirmation. 

Conclusions 

After  75  years  of  recoveiy,  significant  dilfer- 
ences  remain  between  disturbed  and  undis- 
turbed sites  in  Terrill  as  measured  by  bulk 
densitv.  Estimates  for  recoverx  based  on  bulk 
densitv'  are  from  100  to  130  \ears.  In  Wonder, 
after  65  years  of  recover) ,  significant  differences 
remain  only  between  abandoned  roads  and  con- 
trol plots.  Estimated  reco\en-  for  the  aban- 
doned road  is  100  \ears.  These  results  an^  in 
close  agreement  with  similar,  previous  studies 
that  examined  soil  reco\en'  times  in  the  Mojave 
Desert  (e.g.,  Webb  and  Wilshire  1980,  Wehh  et 
al.  1986)  and  suggest  that  the  results  of  soil 
compaction  processes  that  occur  in  arid  en\i- 
ronments  are  long-li\('d.  but  aiv  not  irreversible. 

Ac  K  N  OW  L  E  D  C  M  E  N  TS 

I  wish  to  thank  the  Universit)-  of  Nevada 
Graduate  School  for  funding,  Louis  R.  Loftin 
for  field  and  laboratorv  assistance,  and  Diana  F. 
Thran,  Can  J.  Hausladen,  and  Chris  R.  Ryan 
for  comments  and  suggestions. 


154 


Great  Basin  Naturalist 


[Volume  52 


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Received  5  Ati^ust  1991 
Accepted  16  April  1992 


Great  Basin  NatiinJist  52(2).  pp.  155-159 

BIOCHEMICAL  DIFFERENTIATION  IN  THE  IDAHO  GROUND  SQUIRREL, 
SPERMOPHILUS  BRUNNEUS  (RODENTIA:  SCIURIDAE) 

Ayesha  E.  Gill   and  Eric  Yensen""' 

Abstract — SpenmypJdltis  hnmncus  is  restricted  to  a  90  x  125-km  area  of  west  central  Idalio,  with  two  distinct  (northern 
and  southern)  groups  of  populations  \sithin  this  limited  range.  Morphological  differences  in  pelage  length  and  coloration, 
e.xtemal  and  cr;uii;il  measurements,  imd  hacula  suggest  that  these  groups  are  either  \er\'  distinct  subspecies  or  species.  We 
used  starch-gel  electrophoresis  to  estimate  the  tmiount  of  genetic  differentiation  accompan\ing  these  morphological 
differences  bv  assaying  genetic  \ariation  at  31  loci  in  the  two  geographic  groups.  Fifteen  lcx.i  were  poKuioiphic  (13  in  the 
northern  group,  12  in  the  southern),  and  mean  heterozNgosits'  (H)  was  high  (12.3%  northern  and  lO.S'^  southern).  Nei's 
genetic  distance  (0.057)  is  in  the  range  usualK'  associated  with  subspecific  differences.  However,  Jaccards  association 
coefficient  (0.893)  is  about  the  same  as  that  found  between  se\eral  ground  squirrel  taxa  currently  recognized  as  species. 
The  high  levels  of  heterozygosity  suggest  that  S.  hntntwtis  is  a  neoendemic  rather  than  a  paleoendemic  species. 

Ki'ij  words:  .SjTermophilus  bnumeus,  Spermophilus,  Iddlio  <^nniiul  s(jiiinvl.  <^r(>uiul  s<juinTl.s.  electrophoresis,  taxonoiiiij. 
biochemical  differentiation. 


Spermophilus  hnmneus  is  one  of  the  rarest, 
least  studied,  and  most  geographically  restricted 
of  the  North  American  ground  squirrels.  Within 
its  restricted  range  of  ca  90  X  125  km  in  west 
central  Idaho  there  tire  two  well-differentiated 
subspecies,  S.  /;.  hninneus  and  S.  b.  endemicus 
(Yensen  1991).  Significant  differences  in  pelage 
length  and  color,  e.xtemal  and  cranial  measure- 
ments, and  bacular  moq^holog)'  suggest  that  the 
two  taxa  ma)'  be  close  to  species-level  separation 
(Yensen  1991).  The  northern  Spernwphihis 
h.  hninneus  is  known  from  onl\'  ca  20  isolated 
sites  in  mountain  meadows  in  Adams  and  \^alley 
counties.  These  denies  consist  of  <200  individ- 
uals and  are  separated  from  each  other  b\-  dis- 
tances of  1-70  km.  In  contrast,  the  southern  S. 
/;.  endemicus  is  patchily  distributed  over  a  con- 
tiguous area  70  km  long  and  up  to  20  km  wide 
in  the  lower-elevation  foothills  of  Gem,  Payette, 
and  Washington  counties  (Yensen  1991). 

Da\is  (1939)  divided  the  North  American 
species  of  subgenus  Spermophilus  into  "small- 
eared"  and  "large-eared"  groups  and  placed  S. 
hninneus  within  the  large-eared  group.  Nadler 
et  al.  ( 1973)  found,  however,  that  the  karyotyjoes 
of  S.  hninneus  and  S.  townsendii  mollis  (small- 
eared  group)  differed  onlv  in  the  presence  or 
staining  intensity'  of  minor  bands  on  six  chromo- 


somes, indicating  a  close  affinity  behveen  S. 
hninneus  and  the  small-eared  group.  Nadler  et 
al.  (1974)  analyzed  serum  transferrins  of  S. 
hninneus  using  starch-gel  electrophoresis  and 
concluded  that  it  was  biochemicallv  "intermedi- 
ate" and  possiblv  ancestral  to  both  the  Nearctic 
"small-eared"  and  "big-eared"  species  groups  of 
subgenus  Spennophilus.  Nadler  et  al.  (1982) 
extended  their  analysis  to  21  Holarctic  species 
using  18  loci  and  concluded  that  S.  hninneus 
was  a  paleoendemic  species  most  closeK'  related 
to  the  Eurasian  S.  dmiricus.  Nadler  et  al.  (1984) 
revised  their  phvlogeny  to  incoq3orate  chromo- 
somal data  and  placed  the  e\'()lutionaril\'  con.ser- 
vatixe  S.  hninneus  within  the  S.  townsendii  group. 
The  present  study  was  conducted  to  estimate 
the  genetic  differentiation  accompaming  the 
substantial  moiphological  differences  behveen 
the  two  geographic  groups  of  S.  hninneus  and 
to  assess  the  hypothesis  that  S.  hninneus  is  a 
paleoendemic  species  with  small,  rclictual 
populations. 

Material. s  and  Methods 

Specimens  AnaK/.ed 

A  total  of  82  specimens  were  analyzed  from 
the  following  localities:  Spermophilus  hninneus 


^  University  of  Nevada,  Reno,  Nevada  89557,  Present  address:  Institute  of  I  leallli  Policv  Studii 
'Museum  of  Natural  History',  Albertson  College,  Caldwell.  Idaho  83605. 
.\ddress  for  reprint  requests. 


versitv  of  California.  San  Francisco.  California  94143. 


155 


156 


Great  Basin  Naturalist 


[Volume  52 


bniniu'iis — Adams  Co.:  1  mi  NE  Bear  Guard 
Station,  3;  Bear  Cemetery,  2;  Cold  Springs  Cr., 
1;  Little  Mud  Cr.,  5;  Mill  Cr.  3  mi  N  Hornet 
Guard  Station,  2;  New  Meadows,  12;  Price 
Vall(n,  2;  Lick  Cr.,  6;  Summit  Cr.,  9.  Sper- 
mophiliis  hninneus  endemicxis — Gem  Co.: 
Sucker  Cr.  11  mi  N  Emmett,  20;  12.6  mi  N 
Emmett,  1;  Payette  Co.:  Big  Willow  Cr.,  1;  Dr)/ 
Cr.  Road,  3;  Washington  Co.:  Lower  Mann  Cr., 
10;  Weiser  Cove,  5.  These  specimens  have  been 
deposited  as  vouchers  in  the  Albertson  College 
Museum  of  Natural  History. 

Laboratory'  Methods 

Blood  was  collected  from  the  suborbital 
sinus  of  living  animals  (samples  sizes  were  21  S. 
h.  hnmneiis,  9  S.  b.  endemicus).  Liver  and 
kidney  tissues  were  from  sacrificed  animals  ( 10 
S.  b.  bniniietis,  6  S.  b.  endemicus)  or  frozen 
carcasses  collected  for  other  purposes  (IS  S.  b. 
hninneus,  31  S.  b.  endemicus).  Carcasses  were 
stored  at  -20  C  for  1-6  months. 

Tissue  sample  preparation  and  horizontal 
starch-gel  electrophoresis  follow  Selander  et  al. 
( 1971 )  with  slight  modifications.  We  used  1 1 .0% 
electrostarch  for  lithium  hydro.xide  gels  and 
12.4%  for  all  other  gels.  Enzyme  locus  designa- 
tions follow  standardized  Enzyme  Commission 
(E.G.)  nomenclature  (Harris  and  Hopkinson 
1976).  The  enzymes  and  nonenzymatic  proteins 
screened  in  this  studv,  with  tissue  and  buffer 
svstems  used,  were:  alcohol  dehydrogenase, 
E.G.  No.  1.1.1.1  (ADH),  liver,  tris'-citrate,  pH 
8.0;  glycerol-3-phosphate  dehydrogenase,  E.G. 
No.  1.1.1.8  (GPD),  liver,  tris-citrate,  pH  8.0; 
L-iditol  dehydrogenase,  E.G.  No.  1.1.1.14 
(IDDH),  liver,  tris-citrate,  pH  8.0;  lactate  de- 
hydrogenase, E.G.  No.  1.1.1.27  (LDH),  kidney, 
tris-citrate,  pH  8.0;  malate  dehydrogenase,  E.G. 
No.  1.1.1.37  (MDH),  liver,  tris-citrate,  pH  6.3; 
isocitrate  dehydrogenase,  E.G.  No.  1.1.1.42 
(ICD),  kidney,  tris-citrate,  pH  8.0;  superoxide 
dismutase,  E.G.  No.  1.15.1.1  (SOD),  kidney, 
tris-maleate  or  tris-citrate,  pH  8.0;  aspartate 
aminotransferase,  E.G.  No.  2.6.1.1  (AAT),  liver, 
lithium  hvdroxide;  hexokinase,  E.G.  No.  2.7.1.1 
(HK),  kidney,  tris-citrate,  pH  8.0;  phosphoglu- 
comutase,  E.G.  No.  2.7.5.1  (PGM),  kidney,  tris- 
citrate,  pH  8.0;  esterase,  E.G.  No.  3.1.1.1  (ES), 
hemolvsate,  tris-hvdrochloric  acid;  peptidase, 
E.G.  No.  3.4.1 1  orl3.''  (PEP),  liver,  tris-citrate, 
pH  6.3;  hemoglobin  (HGB),  hemolysate,  tris- 
hydrochloric  acid;  albumin  (ALB),  plasma,  lith- 
ium   hydroxide;    transferrin    (TRF),    plasma, 


litliium  hydroxide;  general  proteins  (GPl  and 
GP2),  hemolysate,  tris-hydrochloric  acid;  and 
general  proteins  (GP3  and  GP4),  plasma,  tris- 
hvdrochloric  acid.  The  proteins  were  numbered 
in  order  of  decreasing  mobilitv,  with  the  most 
anodal  labeled  1. 

The  buffer  and  stain  systems  for  the  proteins 
screened  in  this  study  were  described  by  Selan- 
der et  al.  (1971),  except  for  stains  for'iDDH, 
HK,  and  PEP  (Gill  et  al.  1987).  Of  the  esterases, 
only  acetylesterases  were  stained  and  were 
numbered  1  (most  anodal)  to  5.  PEP-G  was 
detected  with  L-leucyl-L-alanine.  ADH  does 
not  have  to  be  stiiined  specifically  and  is  seen  on 
many  dehydrogenase  gels.  It  was  read  on  gels 
stained  for  GPD. 

Computational  Methods 

Gene  frequencies,  measures  of  genetic  vari- 
ation, Nei's  (1978)  unbiased  genetic  distance 
and  unbiased  genetic  identity,  and  the  average 
inbreeding  coefficient  (Est)  were  derived  from 
input  on  single  individual  genotypes  (elec- 
tromoiphs)  using  the  computer  program 
BIOSYS-1  (Swofford  and  Selander  1981). 
Jaccard's  association  coefficient,  S,  -  a/(a+u), 
where  a  =  the  number  of  matched  elec- 
tromoq3hs  (1:1)  and  u  =  the  number  mis- 
matched (1:0  or  0:1)  (Sneath  and  Sokal  1973), 
was  also  calculated  for  the  two  groups.  Sj 
depends  only  upon  the  presence  ( 1 )  or  absence 
(0)  of  alleles,  as  indicated  b\'  bands  on  the  starch 
gels  (electromoq^hs),  not  on  cillehc  frequencies 
as  do  measures  of  genetic  distance.  Negative 
matches  were  excluded. 

Results  and  Discussion 

SpermopJidus  b.  bninneus  was  polymorphic 
at  13  loci  (42%),  whereas  S.  b.  endemicus  was 
polyinoq:)hic  at  12  loci  (39%).  If  esterases  are 
excluded,  polvmoq^hism  is  reduced  to  31%, 
which  is  similar  to  the  29%  reported  for  Mus 
rnusculus  and  Homo  sapiens  (Lewontin  1974). 
Average  number  of  alleles  per  locus  (A)  was  1.48 
±0.11  (X  ±  SE)  in  S.  b.  bninneus  and  1.48  ± 
0. 12  in  S.  b.  endemicus.  All  polymoqjhic  loci  had 
two  alleles,  except  for  peptidase  and  hvo  of  the 
esterases,  which  had  three. 

Mean  heterozvgositv  per  individual  per 
locus  in  our  sample  was  12.3  ±  3.7%  in  S.  b. 
bninneus  and  10.8  ±  3.9%  in  S.  h.  endemicus. 
These  values  are  much  higher  than  the  2.7% 
heteroz)'gosit)'  reported  b)  Nadler  et  al.  (1982) 


1992] 


Sfehmoi'hiia'sbri'nneus  Electropiiokksis 


157 


for  S.  h.  hniniu'us.  The  loci  coininoii  to  both 
studies,  however,  were  less  variable  than  some 
of  our  18  additioHcil  loci.  Even  if  esterases  are 
excluded  from  the  anahsis,  our  measures  of 
genetic  variabilis  (S.  b.  bniiineus,  H  =  8.2%,  A 
=  1.35;  S.  b.  endemicus,  H  =  7.4%,  A  =  1.38)  are 
still  much  higher  than  theirs.  They  found  H 
values  of  0.0-10.4%  (X  =  3.5%)  for  other  species 
o{  SpcrniopJiiliis.  Cothran  et  al.  (1977)  found 
high  heteroz\gosit\'  (9.3%)  in  the  ground  squir- 
rel subgenus  Ictidomys.  The  average  hetero- 
z\'gosity  for  26  taxa  of  rodents  was  5.4% 
(Selander  1975),  so  Idalio  ground  scjuirrels  ha\e 
relativelv  high  le\els  of  hetero/Agositv  Thus, 
the  levels  of  genetic  \ariabilit\'  are  high  for  a 
species  postulated  to  be  a  paleoendemic 
(Nadler  et  al.  1974,  Cothran  et  al.  1977,  Nadler 
et  al.  1982)  with  small  isolated  denies  and  con- 
fined to  a  small  geographic  area  (Yensen  1991). 

Sixteen  of  31  protein  systems  scored  for  S. 
bniiuu'iis  were  monomoiphic  (GPD,  LDH-A, 
K:D-2,  HK-1,2,  PGM-1,2,  AAT-1,2,  iddh, 
SOD-B,  ADH,  ALB,  TRF,  GP-1,2).  Frequen- 
cies of  alleles  in  the  pol)TOoq3hic  systems  (the 
most  common  allele  <0.99)  are  shown  in  Table 
1.  As  in  other  species  (Kojima  et  al.  1970, 
Lewontin  1974),  non -glucose-metabolizing 
enzymes  were  more  polymorphic  than  glucose- 
metabolizing  enzMiies  wdth  five  monomoq)hic 
(AAT-1,2,  IDDH,  SOD-B,  and  ADH),  while 
PEP-C,  S()D-A  and  all  fixe  esterases  were  poK  - 
moiphic  (Table  1).  The  two  taxa  of  S.  bniniietis 
did  not  differ  substantially  in  glucose- metabo- 
lizing enz\  nies,  with  the  majoritv  of  loci  mono- 
moiphic, and  the  sairie  allele  conuuon  in  the 
poKuioiphic  loci. 

Nadler  et  al.  ( 1982)  found  LDH  to  be  mono- 
moq:)hic  in  all  21  North  American  and  Eurasian 
Sj)i'nu()j)liilus  species  examined.  Howexer,  we 
found  t\v()  indi\iduals  of  S.  b.  bntmu'tis  that 
were  homoz)gous  for  a  fast  iillele  at  the  LDH-B 
locus.  Nadler  et  al.  (1982)  assayed  from  LDH  in 
red  blood  cells  while  we  used  kidney  extracts,  so 
the  difference  ma\- be  between  the  two  tissues. 
Both  groups  of  S.  bninneus  were  pol\  nioqihic 
for  ICD-1  and  HK-3,  while  onl\\S.  b.  endemicus 
was  poKmorphic  for  MDH-1. 

Of  the  enz\nies  not  invoked  in  glucose 
metabolism,  the  esterases  were  the  most  \ari- 
able  (Table  1).  We  also  found  considerable  dif- 
ferences between  S.  b.  bninneus  and  S.  b. 
endemicus  in  the  other  non-glucose-metaboliz- 
ing enzymes.  Different  alleles  were  conunon  for 
PEP-C    and    ES-4    in    the    two    groups    of 


Tahlk    I.    Allelic   IrcijiR'iicit's   oi    [Xilvinoipliif 
Speniiuphilii.s  h  ni  n  iwtis. 


Locus" 

Allele- " 

bninneus 

endemicus 

Glucose 

-METABOLIZINC; 

ENZYMES; 

LDH-B 

a 

0.929 

L(K)0 

b 

0.071 

0.(X)0 

MDH-1 

a 

0.(X)() 

0.018 

b 

1.000 

0.911 

c 

().(X)0 

0,071 

ICD-1 

a 

0.926 

0.986 

b 

0.074 

0.014 

HK-3 

a 

0.132 

0.097 

b 

0.868 

0.903 

NON-GLUCOSE-METABOLIZINC;  ENZYMES: 

SOD-A 

a 

0.786 

0.957 

h        -.. 

0.214 

0.043 

PEP-C 

a 

0.365 

0.329 

b 

0.13.5 

0..343 

c 

0.500 

0.329 

ES-1 

a 

0.179 

0.056    . 

b 

0.107 

0.167       . 

c 

0.714 

0.778 

ES-2 

a 

0.969 

l.(X)0 

b 

0.031 

0.000 

ES-3 

a 

0.971 

0.944 

b 

0.000 

0.056 

c 

0.029 

0.000 

ES-4 

a 

0.714 

0.389 

b 

0.286 

0.611 

ES-5 

a 

0.656 

0.944 

b 

0.344 

0.056 

Non  ENZYMATIC  I'KOTEINS 

HGB-1 

a 

0.233 

0.667 

b 

0.767 

()..333 

HGB-2 

a 

0.100 

0.500 

b 

0.900 

()..500 

GP-3 

a 

0.000 

1.000 

b 

1.000 

O.CXK) 

GP-4 

a 

0.962 

0.750 

b 

0.038 

0.250 

"See  text  for  iicronvm.s  of  loci. 

""Alleles  are  listed  in  order  of  increasing  mobilit)';  a  is  slowest. 


S.  bninneus.  In  both  cases  the  differences  were 
in  allelic  frequenc)-  rather  than  in  the  presence 
or  absence  of  alleles. 

Nonenz\inatic  proteins  were  .scored  in  both 
hemoKsate  and  plasma,  .-\lbumin  and  transfer- 
lin  in  plasma  and  hvo  general  proteins  in 
hemolysate  were  monomorphic.  We  found  vari- 
ability- at  the  two  hemoglobin  loci  and  at  two 
general  protein  loci  in  plasma  (Table  1). 
Heterozygosit\  of  hemoglobins  has  been  found 
in  the  closely  related  TowTisends  ground  s(juir- 
rel  (S.  fownsendii),  in  which  the  hvo  hemoglo- 
bins ha\e  identical  a-chiiins  and  differ  by  only 
one  amino  acid  in  the  sequence  of  their  p-chains 


158 


Great  Basin  Naturalist 


[Volume  52 


(Kleinschniidt  et  al.  1985).  They  found  no 
difference  in  the  oxvgen  affinity  of  the  two 
lienio2lol)ins. 

A  general  protein  in  plasma  (GP-3)  repre- 
sented by  a  band  just  anodal  to  albumin  distin- 
guished the  tu'o  S.  hninneus.  A  fast  allele 
apparently  has  reached  fixation  in  S.  h. 
hrnnneiis,  whereas  a  slow  allele  appears  fixed  in 
S.  h.  cndemiais  (Table  1).  This  is  the  only  locus 
that  can  serxe  as  a  marker  gene  among  the  31 
loci  scored,  although  LDH-B  and  MDH-1  had 
alleles  that  were  fixed  in  one  taxon  and  polymor- 
phic in  the  other.  The  other  presumed  loci  dif- 
fered in  allelic  frequency  only. 

Nei's  (1978)  genetic  distance  is  a  measure  of 
the  accumulated  number  of  gene  differences 
per  locus  between  populations.  The  genetic  dis- 
tance of  0.057  found  between  the  two  S. 
hninneus  was  within  the  range  associated  with 
subspecific  differentiation  (Avise  1974).  The 
average  inbreeding  coefficient  (Fsr  =  0.167) 
indicated  moderately  high  genetic  differentia- 
tion. The  two  S.  hninneus  have  a  genetic  iden- 
tity of  0.944.  By  comparison,  Cothran  et  al. 
(1977)  found  genetic  identities  of  0.808 
between  S.  spilosonia  and  S.  niexicanus,  0.835 
between  S.  spilosonui  and  S.  tridecemlineatus, 
and  0.965  between  S.  tridecemlineatus  and  S. 
mexicanus  in  the  subgenus  Ictidonu/s. 

To  compare  our  results  with  other  results 
from  the  subgenus  Spenriophihis  (Nadler  et  al. 
1982),  we  also  calculated  Jaccard's  association 
coefficient.  This  measure  is  less  sensitive  to 
sample  size  and  depends  on  presence  or 
absence  of  an  allele,  rather  than  on  allelic  fre- 
quencies. Jaccard's  coefficient  of  similarit)' 
between  the  two  groups  of  S.  hninneus  was 
0.893.  Judging  from  Figure  2  in  Nadler  et  al. 
(1982:206),  the  similarity  between  the  two 
groups  of  S.  hninneus  is  about  the  same  as  the 
similarit)'  between  S.  anmitus  and  S.  heldin^i, 
or  between  some  of  the  putative  semispecies  in 
the  S.  townsendii  complex,  the  Eurasian  S.  sus- 
licus  and  S.  citeUus,  or  S.  major  and  S. 
enjthro^enijs.  SpenmyphUus  richanlsoni  and  S. 
elegans  are  more  similar  electrophoretically 
than  the  two  Idaho  ground  scjuirrels.  However, 
direct  comparisons  are  difficult  since  the  simi- 
larity coefficients  computed  by  Nadler  et  al. 
(1982)  were  based  on  a  different,  and  appar- 
ently less  variable,  set  of  loci. 

The  electrophoretic  data  confirm  that  the 
two  Idaho  ground  s(|uirrels  are  genetically  as 
well  as  moqohologicall\-  differentiated  taxa.  The 


evidence  does  not  clearly  resolve  the  question 
of  whether  the  two  are  separated  at  the  subspe- 
cies or  species  level.  The  presence  of  one 
marker  gene  and  the  observed  frequency  differ- 
erices  at  others  could  be  consistent  with  either 
inteq3retation.  The  high  levels  of  heterozygos- 
ity, however,  do  not  support  the  paleoendemic 
hvpothesis. 

Acknowledgments 

We  thank  D.  B.  Hammond,  W.  F.  Laurance, 
and  D.  A.  Stephens  for  field  assistance;  P.  L. 
Packard  for  specimen  shipment;  and  R.  S.  Hoff- 
man, W.  F  Laurance,  C.  F.  Nadler,  E.  A.  Rick- 
art,  P.  W.  Sherman,  O.  G.  Ward,  and  two 
anonymous  reviewers  for  comments  on  the 
manuscript. 

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Received  8  JaniKinj  1991 
Accepted  18  April  1992 


Great  Basin  Naturalist  52(2),  pp.  160-165 

NEW  GENUS,  APLANUSIELLA,  AND  TWO  NEW  SPECIES  OF      - 
LEAFHOPPERS  FROM  SOUTHWESTERN  UNITED  STATES 
(HOMOPTERA:  CICADELLIDAE:  DELTOCEPHALINAE) 

M.  W.  Nielson'  luid  B.  A.  Haws" 

Abstract. — A  new  genns,  Aplaimsk'lla  (type-species,  Aplanusidla  utahensis,  n.  sp.)  and  bA'o  new  species,  A.  ittahemis 
and  A.  calif onriensis ,  are  described  antl  illustrated.  The  two  species  are  allopatric  and  coexist  on  the  same  host  genus, 
(Atriplex)  with  members  of  a  closely  allied  leafliopper  genus,  Aplamis.  Notes  on  distribution  of  hosts  and  leaflioppers  as 
well  as  leafliopper  intergeneric  relationships  are  also  given. 

Ki'ij  words:  leaflioppers.  new  species,  new  gentis,  Cicadcllidae,  Aplanusiella,  distrihntion.  hosts. 


In  a  1986-89  suney  of  rangeland  leafliop- 
pers of  Utah  (Haws  et  al.  1989),  two  populations 
were  taken  from  Atriplex  spp.  and  tentatively 
identified  as  members  of  the  genus  Aplamis. 
One  population  was  later  identified  as  Aplamis 
alhidus  (Ball)  from  shadscale,  Atriplex  con- 
ferfifolia  (Torn  &  Frem.)  Wats.  The  other  pop- 
ulation was  collected  from  four- winged 
saltbush,  Atriplex  canescens  (Pursh)  Nutt.  and 
is  described  herein  as  a  new  genus  and  new 
species  closely  allied  to  Aplamis.  An  additional 
new  species  is  also  described  from  specimens 
collected  in  California  on  Atriplex  sp.  Notes  are 
given  on  the  ph\1:ogeography  of  the  host  genus, 
Atriplex,  the  distribution  of  the  two  genera,  and 
their  taxonomic  and  host  relationships. 

The  general  habitus  (form  and  color  pattern) 
of  the  component  populations  are  so  remark- 
ably similar  that  it  is  likelv  that  additional  mate- 
rial of  the  new  taxa  will  be  found  in  other 
repositories.  Only  after  dissection  and  examina- 
tion of  the  male  genital  structures  will  their  tnie 
identity  be  revealed.  Moreover,  it  is  probable 
that  additional  new  species  will  come  to  light 
after  more  thorough  collecting  is  done  on 
Atriplex  spp.  in  southwestern  United  States  and 
northeni  Mexico.  This  assimiption  is  based  on 
two  additional  populations  of  female  specimens 
in  hand  from  Nexada  and  California  for  which 
males  are  presently  unknown  and  are  required 
for  definitive  generic  placement.  The  female 
seventh  sternal  characters  appear  to  place  these 
populations  in  the  new  genus  (sensu  .stricto). 


Populations  of  these  groups  are  rather  rare  in 
Atriplex  host  areas  of  the  high-  to  low-desert 
regions  of  western  North  America. 

Aplanusiella,  new  genus 

T\TE SPECIES. —  Aplanusiella  utahensis,  n.  sp. 

Small,  rather  slender  species.  Related  to 
Aplamis  Oman  but  smaller  and  with  distinctive 
nicile  genital  characters.  General  color  light 
yellow  to  ivory  with  numerous,  nearly  concen- 
tric, tiny  rufous  spots  on  forewings,  spots  not 
usually  forming  lines  as  typically  present  in 
Aplamis,  large  spots  in  claviis  and  in  apical 
crossveins  of  costa  fonned  by  aggregation  of 
smaller  spots,  pronotum  and  scutellum  some- 
times with  tiny  spots. 

Head  narrower  than  pronotum,  anterior 
margin  obtusely  angled  and  rounded  to  front, 
crowii  produced  mediallv  to  about  one  and  one- 
half  times  length  next  to  inner  margin  of  eve, 
disk  somewhat  depressed  in  middle  but  lacks 
transverse  depression  before  apex  as  in  Aplamis; 
pronotum  and  scutellum  as  in  Aplamis;  fore- 
wings  with  imier  anteapical  cell  open  basally, 
appendix  well  de\  eloped;  cKpeus  and  cKpellus 
as  in  Aplamis. 

Male  pvgofer  with  macrosetae  in  apical  half 
and  with  prominent  caudoventral  spine,  some- 
times crossing  over  in  caudal  view;  aedeagus 
small,  base  large  in  lateral  \iew,  apical  htilf 
narrow,  tubular,  sometimes  with  smdl  angulate 
protrusion  at  base  of  shaft  on  dorsal  margin, 
gonopore  subapical  on  ventral  margin;  connectiv  e 


Monte  L.  Bean  Museum.  Brigham  Young  University.  Provo.  Utah  S4602. 
Department  of  Biolog)-,  Utah  State  Universitv-,  Logan,  Utah  84322. 


160 


1992] 


New  Genus,  Apianusieija 


161 


short,  Y-shaped,  articuhitrd  with  acdeagus;  st\ie 
hroad,  apophxsis  short:  plate  trianpilate  with 
row  of  niacrosetae  subinarginalK  and  row  of 
microsetae  marginally,  female  seventh  sternum 
with  short  projection  medially  on  caudtJ  margin. 
Two  aliopatric  species  are  known  that  occur 
in  the  southwestern  states  of  Utah  and  Califor- 
nia on  desert  shrubs  of  the  genus  Atriplex. 
Aplanusiella  can  be  distinguished  from  Aplantis 
b\  the  sniiiller  size,  by  the  absence  of  a  preapical 
depression  on  the  crown,  by  the  presence  of  a 
prominent  caudoventral  pvgofer  spine,  by  the 
smaller  aedeagus  that  lacks  apical  processes, 
and  by  the  female  seventh  sternum  that  has  a 
more  distinctive  spatulate  process  on  the 
middle  of  the  caudal  margin. 

Aplanusiella  utahensis,  n.  sp. 

Figs,  la-ll 

Length. — Male  3..5-3.75  mm.  female  4.00- 
4.20  mm. 

General  color  pale  yellow  to  ivor)'  with 
numerous,  nearly  concentric,  tiny  nifous  spots 
on  forewdngs,  large  aggregate  spots  on  apex  of 
clavus  and  in  apical  crossveins  of  costa,  some- 
times with  few  similar  spots  on  pronotum  and 
scutellum.  Related  to  AplamisieUa  californien- 
sis,  n.  sp.,  but  with  distinctive  male  genital  and 
female  seventh  sternal  characters. 

Male. — Pygofer  in  lateriil  view  with  long, 
stout  caudoventral  process  that  sometimes 
crosses  its  counteipart  in  caudal  view,  but  usu- 
ally closely  appressed  to  caudal  margin  of  pygo- 
fer (Fig.  lb);  plate  long,  triangulate  with 
uniserate  niacrosetae  submarginallv  and  uniser- 
ate  microsetae  marginallv  on  outer  margin  (Fig. 
Ic);  st)'le  in  dorsal  view  long,  broad  in  basal  2/3, 
apophysis  short,  curved  and  pointed  apically 
(Fig.  Id);  connective  short,  Y-shaped  (Fig.  le); 
aedeagus  in  lateral  view  short,  ventral  margin 
abruptly  angled  near  middle,  broad  basallw 
shaft  narrow,  tubular  with  basal  triangulate  pro- 
jection on  either  side  of  dorsal  margin,  gono- 
pore  subapical  on  ventral  margin  (Figs.  If-lk). 

Ft:MALE. — Seventh  sternum  broadly  exca- 
vated on  caudal  margin,  with  prominent  median 
spatulate  process  (Fig.  U). 

HOLOTYPE  (male).— UTAH:  Daggett  Co., 
Browns  park,  Pyke  plots,  roadside,  12.\T.1987, 
four-winged  saltbush,  Atriplex  canescens,  B.  A. 
Haws  (CAS).  Paratvpes,  1  male,  Daggett  Co., 
Brown's  park,  3.5  mi  E  Jams  ranch,  26.\T.19S7, 
on  four- winged  saltbush,  Atriplex  canescens. 
Haws,   Nelson  (authors  collection);  2   males. 


1  female,  San  Juan  Co.,  Div  \alley,  8.IX.1987, 
four-winged  saltbush,  Atriplex  canescens.  B. 
Haws,  A.  Issa  (USU);  2  males,  2  females,  Uintah 
Co.,  Bonanza,  14.VII.  1975-3.IX.  1976,  Afn>/ex 
canescens,  G.  E.  Bohart  (USU);  1  male.  Grand 
Co.,  Jughandle  Potash  Rd.,  19.Vni.l987,  four- 
winged  saltbush,  Atriplex  canescens,  B.  A. 
Haws,  C.  R.  Nelson  (BYU);  1  male.  Grand  Co., 
Colorado  River,  Hwy  128, 6  mi  NE  jet.  Uwy  191, 
26.V.19H7,  Atriplex  canescens,  B.  A.  Haws.'C.  R. 
Nelson  (USU). 

Remarks, — This  species  can  be  distin- 
guished from  calif orniensis,  n.  sp.,  by  the  longer 
caudoventral  pygofer  process,  by  the  abruptly 
angled  ventral  margin  of  the  aedeagus,  b\'  the 
presence  of  a  small  ba.sal  triangulate  process  on 
the  dorsal  margin  of  the  aedeagal  shaft,  and  by 
the  prominent  spatulate  process  on  the  middle 
of  the  female  seventh  sternum. 

The  species  is  known  from  the  eastern  coim- 
ties  of  Utah  bordering  Colorado  and  is  likely 
present  in  the  western  part  of  that  state  and  in 
northern  Arizona  where  the  host  occurs.  Collec- 
tion dates  suggest  that  the  species  has  two  gen- 
erations per  \'ear  and  presuiuabK'  cnerwinters  as 
eggs  on  its  host. 

Aplanusiella  californiensis,  n.  sp. 

Figs,  liii-ls 

Length. — Male  3.30-3.50  mm,  female 
3.60^3.80  nmi. 

General  color  as  in  A.  titahensis,  n.  sp.,  and 
related  to  that  species  but  with  distinctive  male 
genital  and  female  seventh  sternal  characters. 

Head  similar  to  utahensis  except  not  as 
pointed  apicallv 

Male. 

ately  long  caudoventral  process  that  usualK 
crosses  its  counteqoart  in  caudal  \iew,  not 
closely  appres.sed  to  margin  of  pygofer  (P'ig. 
Im);  plate  long,  triangulate,  with  row  of  mar- 
ginal micr().setae  and  submarginal  niacrosetae 
(Fig.  In);  .style  in  dorsal  \iew  long,  narrow, 
apophysis  sliort,  ol)li(juel\'  tnmcate  apicalK 
(Fig.  lo);  aedeagus  in  lateral  view  short,  \entral 
margin  gradualK'  curved,  apical  third  tubular 
broad  basalK  in  ventral  view,  tapered  toward 
apex,  gonopore  subapical  on  ventral  margin 
(Figs.  lj>-lr). 

Fe.\L\LE. — Seventh  sternum  with  truncate 
caudal  margin  except  for  sliort,  median  process 
(Fig.  Is). 

HOLOTYPE  (miile). — CaLIFORNL\:  Riverside 
Co.,    Indio,     12.1.1988,    Atriplex   sp.,    G.    N. 


-Pv  gofer  in  lateral  view  with  nioder- 


162 


Great  Basin  Naturalist 


[Volume  52 


Figs,  la-ll.  Aplfinusk'Ua  utahensis,  n.  sp.:  la,  head  pronotum,  and  scutellnm,  dorsal  view;  lb,  male  pygofer,  lateral  view; 
Ic,  right  plate,  ventral  view;  Id,  right  style,  dorsal  view;  le,  connective,  dorsal  view;  If,  aedeagus,  dorsal  view;  Ig,  same, 
lateral  view;  Ih,  same  (enlarged),  .showing  triangulate  process,  lateral  view;  li,  same  (showing  variation),  lateral  \ie\v;  Ik, 
same  (enlarged),  showing  apex  of  aedeagus,  ventral  view;  11,  female  seventh  sternum,  ventral  view. 

Figs.  Im-ls.  Aplanmiella  calif omiensis,  n.  sp.:  Im,  male  pygofer,  lateral  view;  In,  right  plate,  ventral  view;  lo,  right 
style,  dorsal  view;  Ip,  aedeagus,  lateral  view;  Iq,  same,  ventral  view;  Ir,  same  (enlarged),  showing  apex  of  aedeagus,  ventral 
view;  Is,  female  seventh  sternum,  ventral  view. 


1992] 


New  Genus,  Aplanusieija 


163 


Figs.  2a-2f,  2in.  Aplanus  pauperciilus  (Ball):  2a,  nude  pvgofer,  lateral  \iew;  21).  right  plate,  ventral  view;  2c,  aedeagus, 
dorsal  view;  2d,  same,  lateral  view;  2e,  right  sUle,  dorsal  \iew;  2f,  connectixe,  dorsal  \iew;  2m,  female  seventh  sternum' 
ventral  view. 


Figs.  2g-21,  2n.  Aplanus  albidns  (B;ill):  2g,  male  pvgofer,  lateral  view;  2h.  right  plate,  ventral  view;  2i,  aedeagus.  dorsal 
\iew;  2j,  same,  lateral  view;  2k,  right  style,  dorsal  view;  21,  connective,  dorsal  view;  2n,  female  seventh  sternum,  ventral  view. 


164 


Great  Basin  Naturalist 


[Volume  52 


Oldfield  (CAS).  Parat)pes,  2  males,  6  females, 
same  data  as  holotype  (OSU);  5  males,  16 
females.  Imperial  Co.,  Brawlev,  23.VIII.1983, 
Atriplex  .sp.,  J.  Williams  (OSU,  BYU). 

Remarks. — This  species  can  be  separated 
from  utahensis  by  the  shorter  caudoventral 
pvgofer  spine,  by  the  smoothly  curved  ventral 
margin  of  the  aedeagus,  by  the  lack  of  a  basal 
process  on  the  aedeagal  shaft,  by  the  broader 
base  of  the  aedeagus  in  ventral  view,  and  by  the 
truncate  caudal  margin  and  shorter  median  pro- 
cess of  the  female  seventh  sternum. 

This  species  is  knowTi  from  southern  Califor- 
nia on  Atriplex  (species  unknown)  at  elevations 
below  sea  level.  Collection  dates  suggest  that 
the  species  overwinters  in  the  adult  stage  and 
ma\'  have  as  many  as  three  generations  per  year. 

Aplanus  Oman 

Aplanus     Onuui,      1949:138.     Tvpe     species,     Eutctfix 
paiipcrctihis  Ball. 

Only  two  species  are  known  in  the  genus, 
both  assigned  by  Oman  (1949).  Crowder  (1952) 
treated  the  group  with  a  key  to  species,  rede- 
scriptions,  and  illustrations  of  the  genital  char- 
acters. The  range  of^ Aplanus  is  much  broader  in 
western  United  States  than  the  presently  known 
range  o{ Aplonusiella. 

Characters  are  given  for  Aplanus 
pauperculns  (Figs.  2a-2f,  2m)  and  Aplanus 
aUndus  (Ball)  (Figs.  2g-2l,  2n)  to  show  generic 
relationships  between  them  and  species  of 
AplanusieUa.  In  Aplanus  the  pygofer  lacks  the 
caudal  spine,  and  the  aedeagus  is  alxnit  twice  as 
long  with  distinctive  terminal  processes.  The 
female  seventh  sternum  lacks  the  obvious 
median  caudal  process  that  is  present  in 
AplanusieUa.  Ball  (1900)  reported  that  shad- 
scale,  Atriplex  eon feiii folia  (Torn  &  Frem.) 
Wats.,  was  the  host  of  Aplanus  alhidus.  The 
specific  host  of  A.  pauperculns  is  yet  unknown. 

Phytogeographv  o{  Atriplex 

Four-winged saltbush  (Atriplex  eanescens)  is 
endemic  to  western  North  America.  Its  range 
extends  from  southern  Canada  to  northern 
Mexico.  Shadscale  (Atriplex conferiifolia)  is  also 
endemic,  but  its  range  is  more  restrictive  within 
western  United  States  (Stutz  and  Sanderson 
1979, 1983,  Sanderson  et  al.  1990).  Both  species 
produce  hybrids  between  themselves  and  other 
species  of  Atriplex.  However,  autopk)idy  is  the 
most  common  genetic  mechanism  in  both  spe- 
cies, which  have  produced  a  number  of  races 


throughout  their  range.  These  races  and  other 
ecotypes  have  been  identified  and  mapped  by 
these  workers. 

The  biogeographical  relationships  between 
Aplanus  and  AplanusieUa  species  and  their  host 
species  are  poorly  knouai.  Although  hosts  have 
been  identified  for  two  leaflioppers  (Aplanus 
albiclus  and  AplantisieUa  utahensis)  of  the  four 
known  species,  nothing  is  known  about  the 
others  nor  has  preference,  if  any,  of  these  leaf- 
hoppers  for  races  or  ecotypes  been  studied  in 
Atriplex.  The  role  of  Atriplex  in  the  evolutionary 
development  and  speciation  of  the  group  is  like- 
wise unknown. 

Deposition  of  type  specimens 

The  holotvpe  specimens  of  AplanusieUa 
utahensis  and  AplanusieUa  califomiensis  are 
deposited  in  the  California  Academv  of  Sci- 
ences, San  Francisco  (CAS);  parat\pes  are  in 
Oregon  State  University,  Corvallis  (OSU),  Utah 
State  University,  Logan  (USU),  and  Monte  L. 
Bean  Museum,  Brigham  Young  University, 
Provo,  Utah  (BYU). 

Acknowledgments 

We  thank  Paul  W.  Oman,  Oregon  State  Uni- 
versit\',  Corvallis,  for  loan  of  material  of  Aplanus 
and  C.  Rilev  Nelson,  Universitv  of  Texas,  Austin, 
for  his  assistance  in  collecting  material  in  Utah. 
We  iilso  appreciate  helpful  comments  by  H. 
Derrick  Blocker,  Kansas  State  University,  Man- 
hattan, and  Paul  H.  Frevtag,  University  of  Ken- 
tuck"\',  Lexington,  who  reviewed  the  paper.  This 
studv  was  supported  in  part  by  endowanent 
funds  from  the  Monte  L.  Bean  Life  Science 
Museum,  Brigham  Young  University,  Provo, 
Utah,  for  which  we  are  grateful. 

Literature  Cited 

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C:anadi;in  Entomologist  32:  200-20.5. 

(^ROWDKR,  II.  \V.  1952.  A  re\ision  of  some  phlepsiuslike 
genera  of  the  tribe  Delttxephdini  (Homoptera, 
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versitv  Science  Bulletin  35:  .309^541. 

H.\ws,  B.'a.,  G.  E.  Boh.'KRT.  C.  R.  Nelson,  ;uid  D,  L. 
Nelson  1990.  Insects  and  shnib  die-off  in  western 
states:  1986-1989  survey  results.  Pages  127-151  in 
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Report  INT-276.  351  pp.  tional  Arid   Land  Conference  on   Plant   Resonrccs. 

Oman,  P.  W.  1949.  The  Nearctie  leiiflioppers.  A  generic  International  Center  for  Arid  and  Seini-iirid  Land 

classification  and  check-list.  Memoirs  of  the  Entonio-  Studies,  Texas  Tech  Uni\ersitv.  Liihhock.  621  pp. 

logical  Society  of  Washington.  .  1983.  EvoKitionarv  studies  oi'Atriplcx:  chromo- 

Sanukkson,  S.  C'.,  H.  C.  Silt/,  and  E.  1).  McArtihh  some  races  of  A,  confertifolid  (Shad.scale),  American 

1990.  Geographical  differentiaHon   in  Atriplcx  am-  Journal  of  Botany  70:' 1536-1547. 

fertifolia.  American  Journal  of  Botany  77:  490-498. 
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Creiit  Basin  Natunilist  52(2),  pp.  166-173 

SUMMER  HABITAT  USE  BY  COLUMBIAN  SHARP-TAILED  GROUSE 
IN  WESTERN  IDAHO 

Victoria  Ann  Saab    and  Jeffrey  Sliavv  Marks" 

Abstra(ti" — We  shidiecl  smnnier  habitat  use  by  Columbian  Shaip-tailed  Grouse  {Tytnpuuuchus  pJuisiancllus  co- 
hunhiaims)  in  western  Idaho  during  198.3-S5.  Vegetative  and  topographic  measurements  were  recorded  at  716  locations 
of  15  radio-tagged  grouse  and  at  180  random  sites  within  the  major  vegetation/cover  types  in  the  study  area.  The  mean  size 
of  summer  home  ranges  was  1.87  ±  1.14  km".  Of  eight  cover  types  identified  in  the  study  area,  individual  grouse  used  the 
big  sagebrush  {Artcviisia  tridentata)  cover  type  more  than  or  in  proportion  to  availability,  the  low  sagebmsh  [A.  arhusaila) 
type  in  proportion  to  availability,  and  avoided  the  shrubby  eriogonum  (Eriooonum  spp.)  tyjie.  Characteristics  of  the  big 
sagebmsh  cover  tyj^e  that  Sharp-tailed  Cirouse  preferred  include  moderate  vegetative  cover,  high  plant  species  diversity, 
and  high  stnictural  dixersitv.  Grouse  used  areas  of  dense  cover  (i.e.,  mountiiin  shrub  and  riparian  cover  tyjjes)  primarily  for 
escape  cover.  Compared  with  random  sites,  grouse  selected  areas  with  (1)  greater  horizontal  ;uid  vertical  cover,  (2)  greater 
canopv  coverage  of  forbs  tyj^ically  decreased  by  livestock  grazing,  (3)  greater  density  and  canopy  coverage  of  arrowleaf 
balsamroot  (Balsamorhiza  sagittata),  and  (4)  greater  canopy  coverage  of  bluebunch  wheatgrass  (Agroptjron  spicatum)  in 
the  big  sagebmsh  cover  type  in  1984  ;uid  the  low  sagebrush  cover  type  in  1985.  The  importance  of  the  native  perenniiils 
arrowleaf  biilsamroot  and  bluebunch  wheatgrass  became  apparent  chiring  a  drought  year  when  many  exotic  annuals  dried 
up  and  provided  no  cover.  Overall,  grou.se  selected  vegetative  communities  that  were  least  modified  bv  lixestock  grazing. 

Key  words:  Tympanuchus  phasi;uiellus  columbianus,  C(>luml>itin  SJiaiy-taiJcd  Gnnise.  Idaho,  stnnuwr  habitat  charac- 
teristics, nmnaocment . 


Coliinibian  Shaip-tmled  Grouse  {Tynipa- 
nuchiis  phasianellus  columbianus)  have 
declined  in  both  numbers  and  distribution  since 
European  settlement,  currently  occupying 
<10%  of  their  former  range  (Miller  and  Graul 
1980).  Degradation  of  native  habitat  by  live- 
stock grazmg  and  agriculture  are  thought  to  be 
major  factors  in  this  decline  (Yocom  1952, 
Aldrich  1963,  Zeigler  1979).  Overgrazing 
reduced  bunchgrasses  and  perennial  forbs  that 
are  important  components  of  nesting  and 
brood-rearing  habitat  (Yocom  1952,  Jewett  et  al. 
1953,  Klott  and  Lindzey  1990).  Conversion  of 
range  to  cropland  destroyed  nesting  and  brood- 
rearing  habitat  and  deciduous  shrubs  that  are 
critical  for  winter  food  and  escape  cover  (Zeigler 
1979,  Giesen  1987,  Marks  and  Marks  1988).  As 
a  result,  Columbian  Sharp-tiiiled  Grouse  were 
designated  as  a  candidate  species  for  listing  as 
federally  threatened/endangered  (Federal  Reg- 
ister 1989). 

Quantitative  information  on  home  range  size 
and  habitat  preferences  of  (Columbian  Shaip- 
tailed  Grouse  throughout  their  range  is  lacking. 


especially  data  based  on  radio-tagged  individu- 
als during  the  summer  reproductive  period  (see 
Klott  and  Lindzey  1990).  We  studied  Colum- 
bian sharjDtails  in  areas  with  eight  vegeta- 
tion/cover types.  The  primary  objective  of  our 
study  was  to  provide  information  on  summer 
habitat  preferences  by  Columbian  Sharp-ttiiled 
Grouse. 

Study  Area 

The  2000-ha  study  area  is  23  km  north  of 
Weiser  in  Washington  Countv;  Idaho.  Elevation 
ranges  from  970  to  1188  m.  Annual  precipita- 
tion averages  39  cm.  The  springs  and  summers 
of  1983  and  1984  were  relatively  cool  and  wet, 
whereas  those  of  1985  were  unusually  hot  and 
dry.  Sharp-tiiiled  Grouse  had  not  been  himted 
in  the  study  area  since  1974. 

Vegetation  is  characteristic  of  a  sliRibsteppe 
communitv  (Marks  and  Marks  1987a).  The 
greatest  proportion  of  the  studv  area  (40%)  was 
occupied  bv  the  big  sagebmsh  (Artemesia 
trident  (it  a)  cover  t\pe;  low  sagebmsh  (A. 
arhusndo)  and  shmbby  eriogonum  {Eriogonum 


Biology  Department,  Montana  .Slate  Univt-nsitv.  Bo/cmaii,  Montana  59717.  Present  address:  USDA  Forest  Ser^nce,  Intermonntain  Ke 
M)Ttle  Street,  Boise,  Idaho  8.3702. 

Di\nsion  of  Biological  Sciences,  University  of  Montana,  Missonla.  Montana  .'59812, 


•arcli  Station.  .316  E, 


166 


19921 


Sharp-tailed  Grouse  Summer  Habitat 


16- 


sphacrocephaluiii  and  E.  thijiiioidca)  Upes 
occupied  21  and  20%,  respectively.  The  remain- 
ing 19%  of  the  stud\-  area  was  occupied  bv'  five 
other  cover  tvpes  (see  below). 

The  big  sagebnish  cover  t\pe  was  dominated 
bv  big  sagebrush,  with  lesser  amounts  of 
bitterbrush  {Purshia  tridentata)  and  low  sage- 
brush. The  greatest  canopv  coxerage  of  blue- 
bunch  wheatgrass  {Agropyron  spicatuin)  was 
found  in  this  cover  type;  arrowleaf  btilsamroot 
(BaJsanwrhiza  sogittata)wds  the  dominant  forb. 
Bulbous  bluegrass  was  the  most  common  her- 
baceous plant  in  the  understor\'  of  the  low  sage- 
brush co\er  t\pe  with  lesser  amoimts  of 
willoweed  {Epilobium  paniadatum),  blue- 
bunch  wheatgrass,  and  Sandberg's  bluegrass 
{Poa  sandhergii).  The  herbaceous  layer  of  the 
shnibbv  eiiogonum  cover  t\pe  was  relativelv 
sparse  and  dominated  bv  Sandberg's  bluegrass. 
The  mountain  shrub  cover  type  occurred  in 
dense  patches  on  hillsides;  common  species 
were  bittercherr\'  {Pnimis  emarginatus), 
common  chokecherr}'  [P.  virginiana),  snow- 
brush  ceanothus  {Ceanothus  vehitimis),  and 
Saskatoon  serviceberry  {Amelanchier alnifolia) . 
The  shrub  layer  of  the  bitterbrush  (Purshia 
tridentata)  cover  type  was  almost  exclusively 
bitterbnish,  while  the  herbaceous  layer  was  sim- 
ilar to  that  found  in  the  big  sagebrush  t)pe. 
Riparian  vegetation  was  dominated  by  Douglas 
hawthorn  (Crataegus  douglasii),  with  lesser 
amounts  of  wallow  (Salix  spp.)  and  Woods  rose 
(Rosa  woodsii).  Bulbous  bluegrass  (Poa 
btdhosa),  an  exotic  grass,  was  widespread 
throughout  the  study  area.  Plant  nomenclature 
follows  Hitchcock  and  Cronquist  (1976). 

Two  vegetation  t\pes  were  almost  exclu- 
sively comprised  of  nonnative  vegetation.  A 
small  portion  of  the  study  area  contained  agri- 
culture, composed  of  dryland  wheat  and  barley, 
and  monocultures  of  intermediate  wheatgrass 
(Agropyron  interniedinni)  seedings. 

The  study  area  was  grazed  by  livestock  since 
at  least  1900.  Before  about  1940,  large  bands  of 
sheep  were  driven  through  the  area.  Since  then, 
the  major  land  use  in  the  studv  area  has  been 
cattle  grazing.  No  livestock  grazing  occurred 
during  this  study. 

Methods 

Trapping  and  Monitoring 

Grouse  were  captured  on  dancing  grounds 
using  funnel  traps,  mist  nets,  and  drop  nets.  Sex 


was  determined  1)\'  examination  of  crown  feath- 
ers (Henderson  et  al.  1967)  and  age  by  exami- 
nation of  outer  primaries  (Ammann  1944). 
ThirtA-eight  grouse  (28  males  and  10  females) 
of  46  captured  were  fitted  with  solar-powered 
radio  transmitters  attached  to  Herculite  pon- 
chos (Marks  and  Marks  1987b).  Radios  weighed 
between  13.5  and  14.5  g.  Fifteen  (13  males  and 
2  females)  grouse  provided  data  for  home  range 
and  microhabitat  analyses.  The  other  23  grou.se 
with  radios  were  relocated  for  two  months  or 
less  as  a  result  of  mortality  (Marks  and  Marks 
1987b)  or  dispersal  from  the  stud\'  area.  Data 
from  these  birds  were  used  in  the  microhabitat 
analyses  but  not  in  the  calculation  of  home 
range  size.  Sample  sizes  were  not  large  enough 
to  compare  habitat  use  or  home  range  size 
between  male  and  female  grouse. 

Radio-tagged  grouse  were  monitored  from 
May  to  September  1983—85.  Each  time  a  grouse 
was  located,  it  was  flushed  (hereafter  these  loca- 
tions are  called  flush  sites).  Flush  sites  sened  as 
focal  points  for  habitat  sampling  and  for  calcu- 
lation of  home  ranges.  Grouse  were  located 
throughout  the  day  and  locations  were  stratified 
into  four  time  intervals:  sunrise  to  0800,  0801  to 
1100,  1101  to  1700,  and  1701  to  sunset.  On 
average,  each  radio-tagged  bird  was  flushed 
four  days  a  week,  once  in  each  of  the  four  time 
intervals. 

Habitat  Sampling 

The  stud\'  area  boundaiv  was  determined  by 
grouse  movements  during  1983.  C^cn-er  tvpes 
were  digitized  and  areas  calculated  for  each  t) pe 
using  GEOSCAN  (Software  Designs  1984),  a 
geographic  information  program.  Flush  sites 
were  plotted  and  home  range  sizes  (Mohr  1947) 
were  calculated  using  the  compute^-  program 
TELDAY  (Lonner  and  Burklialter  1986).  ' 

Use  vs.  availabilih'  of  cover  t\pes  (i.e., 
macrohabitat)  was  assessed  by  (1)  using  the 
proportion  of  cover  t\pes  within  each  bird's 
home  range,  and  (2)  using  the  proportion  of 
cover  tyj:)es  within  a  1.2-km  radius  of  the  danc- 
ing ground  at  which  each  bird  was  captured. 
The  1.2-km  radius  around  each  of  three  dancing 
grounds  (upper,  middle,  and  lower)  encom- 
passed 90%  of  all  grouse  locations.  Flush  sites 
within  50  m  of  a  dancing  ground  during  the 
spring  and  autumn  display  periods  were  omitted 
from  macrohabitat  analyses. 

We  measured  \egetation  at  each  flush  site 
(i.e.,  microhabitat)  to  estimate  plant  .species 


168 


Great  Basin  Naturalist 


[Volume  52 


composition,  frequency,  percent  canopy  cover- 
age, and  bare  ground  using  a  20  X  50-cm  frame 
(Daubenniire  1959).  Five  frames  were  read  at 
each  flush  site:  one  at  the  approximate  center 
and  one  in  each  of  the  four  compass  directions 
at  randomly  chosen  distances  of  2,  4,  6,  or  8  m 
from  the  center  location.  Vertical  structure  of 
the  vegetation  was  evaluated  by  a  coxer  board 
that  was  a  16.5  x  49.5-cm  rectangle.  The  cover 
board  was  placed  at  the  center  of  the  flush  site 
and  read  twice  from  5  m  away  in  each  of  the  four 
compass  directions  while  the  observer  was 
prone  and  standing,  respectively.  A  total  reading 
of  150  squares  was  possible  from  each  compass 
direction.  In  total,  five  canopy  coverage  and  four 
cover  board  measures  were  taken  at  each  site. 
Other  variables  recorded  at  flush  sites  included 
(1)  cover  type,  (2)  distance  to  water,  (3)  percent- 
age of  slope,  (4)  distance  to  nearest  riparian  or 
mountain  shrub  cover  type,  and  (5)  cover  type 
where  flushed  grouse  landed  (landing  site). 

We  recorded  vegetative  and  topographic 
measurements  at  randomly  located  sites  to 
assess  microhabitat  avmlability  in  the  cover 
t\pes  used  most  bv  grouse.  Habitat  characteris- 
tics were  sampled  with  similar  methods  as 
described  at  flush  sites.  A  total  of  180  random 
sites  were  sampled  during  the  study,  30  each 
month  during  Mav  through  Julv  in  1984  and 
1985.  The  number  of  random  sites  located  in 
each  cover  type  was  based  on  the  percentage  of 
area  occupied  by  that  cover  type  in  the  study 
area.  Canopy  coverage  and  cover  board  read- 
ings were  recorded  at  the  origin  and  at  points 
every  10  paces  along  a  straight  line  until  20 
readings  were  completed.  Slope  and  distance  to 
the  nearest  mountain  shiub  or  riparian  cover 
type  were  recorded  onlv  at  the  first,  tenth,  and 
twentieth  frames  of  each  random  site. 

Data  AnaK'sis 

Data  were  anab'zed  with  the  Statistical  Anal- 
ysis System  (SAS  Institute,  Inc.  1982).  Use- 
availabilit)'  analyses  of  cover  types  were 
conducted  with  chi-s(juare  goodness  of  fit  tests 
(Neu  et  al.  1974)  and  Bonferroni  z-tests  (B\ers 
et  al.  1984).  Data  were  analyzed  separately  for 
each  year  and  pooled  when  differences  were  not 
significant.  For  analyses  of  canopy  coverage, 
each  plant  species  was  placed  into  one  of  10 
categories:  ( 1)  big  sagebrush,  (2)  low  sagebrush, 
(3)  bitterbrush,  (4)  other  shrubs,  (5)  arrowleaf 
balsamroot,  (6)  other  composites,  (7)  non- 
composite  forbs,  (8)  bluebunch  wheatgrass,  (9) 


bulbous  bluegrass,  and  (10)  other  grasses.  Non- 
parametric  statistics  (Mann-Whitney  U-  and 
Kruskal-Wallis  tests)  were  used  to  anaK'ze 
canopy  coverage  and  vertical  stmcture  because 
these  data  were  not  nonnally  distributed  (Con- 
over  1980).  Vegetative  measurements  at  flush 
sites  from  May  through  July  were  combined  by 
cover  tvjje  and  month  for  comparisons  with  data 
collected  at  random  sites  for  the  same  period. 
All  multiple  comparisons  were  computed  with 
Tukey  tests  (Zar  1974).  The  Shannon-Wiener 
index  was  used  to  calculate  plant  species  diver- 
sity (Hill  1973).  Proportions  entered  into  the 
diversit)'  formula  were  derived  from  the  total 
number  of  plant  species  occurrences  within  the 
frames  used  to  estimate  canop\'  coverage.  The 
significance  level  for  all  tests  was  P  <  .05,  and  all 
tests  of  means  were  two-tiiiled.  Means  are  fol- 
lowed by  ±  one  standard  deviation. 

Results 

Home  Ranges  and  Macrohabitat  Selection 

The  mean  size  of  summer  home  ranges  was 
1 .87  ±  1 .  14  km-  (N  =  15,  range  =  36-68  locations 
per  grouse).  Based  on  habitats  within  home 
ranges,  three  trends  emerged  from  the  use- 
availabilit)'  analysis  of  coxer  t\pes:  (1)  grouse 
used  the  big  sagebrush  cover  t}pe  more  than  or 
in  proportion  to  availabilitv;  (2)  the  low  sage- 
brush cover  t\pe  was  used  in  proportion  to 
a\ailabilif\',  and  (3)  the  shrubby  eriogonum  and 
intermediate  wheatgrass  coxer  txpes  xvere 
avoided  (Table  1).  These  trends  xvere  similar 
xvhether  use-aviulabilit\'  xvas  assessed  xxithin 
estimated  home  ranges  or  xxithin  a  fixed  radius 
aroimd  the  upper  and  lower  dancing  grounds 
(Table  1).  In  addition,  a  single  grouse  from  the 
middle  dancing  ground  used  the  big  sagebiTish 
coxer  t\pe  more  than  that  expected  bx'  chance 
xxithin  its  home  range  and  the  fixed  radius. 
Grouse  were  seldom  found  in  the  denser  cover 
txpes,  i.e.,  riparian  and  mountain  shrub  habitats. 
Hoxx'exer,  thex  used  these  coxer  txpes  as  escape 
coxer  in  77%  of  the  cases  xxhere  the  landing  site 
of  a  flushed  radioed  bird  xx'as  obsened  (N  = 
338). 

Microhabitat  Selection 

Mean  distance  to  xvater  did  not  differ  signif- 
icantly bet\x'een  flush  (.V  =  297.6  ±  183.3  m)  and 
random  (.v  =  295.9  ±  211.7  m)  sites  (F  <  .40), 
and  no  evidence  xvas  found  that  Shaip-tiiiled 
Grouse  sought  free  xvater.  The  range  of  slopes 


1992] 


Sharp-tailed  Grousk  Summer  Habitat 


169 


Table  1.  Sunnner  hahitat  usf-a\ailal)ilih  analysis  showing  tli(-  iniinl)er  oi  raclio-taggml  (Columbian  Sliarp-tailed  Crouse 
using  the  major  cover  types  more  than  ( + ),  less  than  ( — ),  or  in  projxjrtion  to  (NS)''  that  expected  by  chance ',  1983-85. 


Cover  types 


Home  range 


NS 


1.2-km  fixed  radius 


NS 


Upper  dancing  ground 

Big  sagebnish 

Low  sagebrush 

Shnibbv  eriogonuni 

Mountain  shrub 
Number  of  grouse 
Lower  dancing  ground 

Big  sagebrush 

Low  sagebrush 

Intermediate  wheatgrass 
Number  ot  grouse 
Total  number  ot  grouse 


2 

0 

3 

0 

1 

4 

0 

5 

0 

1 

0 

4 

N  =  5 

7 

0 

2 

0 

3 

6 

0 

2 

' 

N  =  9 

N  =  14 

0 

0 

5 

0 

0 

5 

0 

5 

0 

1 

0 

4 

8 

0 

I 

0 

1 

8 

0 

6 

3 

■'Not  sigiiitkant, 
'T  <  .05. 


used  by  grouse  was  0-47%.  Grouse  used  three 
classes^  of  slope  intervals  (0-9%,  10-29%, 
>30%)  in  proportion  to  their  availabilit\;  with 
>95%  of  the  use  occurring  on  slopes  <30% 
(Marks  and  Marks  1987a). 

Grouse  did  not  show  a  strong  preference  for 
sites  that  were  close  to  nioinitiiin  shmb  or  ripar- 
ian \egetation  except  in  1985,  the  drought  year. 
The  mean  distance  to  mountain  shrub  and  ripar- 
ian habitats  measured  at  flush  sites  {x  =  151.5  ± 
156.5  m)  was  farther  than  that  measured  at 
random  sites  (.v  =  120  ±  99.7  m)  in  1983  and 

1984  (Mann-Whitney  L'-test  P  <  .04)  but  signif- 
icantly closer  (flush  sites,  x  -  84.4  ±  90.9  m)  in 

1985  (F<  .0001). 

Vertical  cover  measured  at  random  sites  dif- 
fered significantly  among  cover  types  (Kruskal- 
Wallis  P  <  .001).  Mean  cover  board  readings 
indicated  that  the  bitterbrush  cover  tyjie  pro- 
vided the  greatest  cover;  big  sagebrush,  inter- 
mediate wheatgrass,  and  low  sagebrush  tyjoes 
])ro\ided  intermediate  cover;  and  eriogonmn 
sites  had  verv  little  cover  (Fig.  1).  A  drought 
during  1985  resulted  in  significantly  less  vertical 
cover  in  1985  than  in  1984  (Mann-Whitney  li- 
test P  <  .01 ).  However,  the  rank  order  of  cover 
availabilitv  was  the  same  among  all  cover  t\pes 
except  intermediate  wheatgrass,  which  de- 
creased substantiiillv  in  1985. 

Eight\'-three  percent  of  the  flush  sites  for 
which  microhabitat  measurements  were  taken 
occurred  in  big  and  low  sagebrush  cover  t\pes. 
\'egetative  data  on  microsite  use  vs.  availability- 
were  evaluated  onlv  for  big  and  low  sagebrush 


cover  types  because  sample  sizes  were  too  small 
for  the  other  types. 

Vertical  cover  measured  at  flush  sites  dif- 
fered among  years  within  big  and  low  sagebrush 
cover  types  (Kmskal-Wallis  P  <  .05).  As  noted 
at  random  sites,  there  was  significantly  less 
cover  in  1985  than  in  1984.  A  comparison  of 
grouse  flush  sites  with  random  sites  revealed 
that  grouse  selected  denser  cover  than  that  mea- 
sured at  random  sites  (Fig.  1). 

The  cover  types  used  most  by  grouse,  big  and 
low  sagebnish,  had  a  higher  diversitv  of  shrub, 
forb,  and  grass  species  than  the  otlier  cover 
tvpes  (Fig.  2).  The  big  sagebrush  cover  type  had 
the  highest  diversit)'  of  shnibs  and  grasses,  and 
the  low  sagebrush  cover  tvpe  had  the  highest 
diversity  of  forbs.  Overall,  the  big  sagebnish 
cover  tyjDe  had  the  highest  stnictural  heteroge- 
neity (measured  as  the  coefficient  of  variation  of 
canopv  coverage  and  cover  board  readings). 

During  198.3-85,  canopv  coverage  oi  shnibs 
at  grouse  flush  sites  averaged  about  9%  in  both 
big  and  low  sagebnish  cover  types.  Forb  cover- 
age averaged  about  30%,  and  grasses  ranged 
from  28%  to  32%  canopv  coverage  in  low  sage- 
brush and  big  sagebrush  cover  tvpes,  respec- 
tively. Overall,  canopy  coverage  at  flush  sites 
was  significantlv  greater  than  at  random  sites 
due  largelv  to  greater  total  forb  coverage  at  flush 
sites  (Table  2).  (^onverselv,  percentage  of  bare 
ground  was  less  at  flush  sites  than  random  sites 
in  all  cases  (Table  2).  Sites  chosen  by  grouse  in 
1984  and  1985  had  significantlv-  higher 
arrowleaf  balsamroot  cover  than  did  random 
sites.   There  was   significantly  higher  canopy 


170 


Great  Basin  Naturalist 


[Volume  52 


O     RANDOM 
D     FLUSH 


t 


6 


^ 


ARAR        ERIO 


COVER  TYPES 


Fig.  L  Mean  (±  SD)  cover  board  readings  at  random 
sites  and  Sharp-tailed  Grouse  flush  sites  in  the  major  cover 
types  (big  sagebnish  [ARTR],  low  sagebnish  [ARAR], 
shrubby  eriogonum  [ERIO],  intermediate  wheatgrass 
[AGIN],  bitterbrush  [PUTR],  1984-85  (°  =  F  <  .001). 
Vertical  tixis  represents  the  number  of  boxes  visible  on  the 
cover  board  (see  Methods). 


26- 

24- 

22 
X 
3.  20 

>- 

(/)         ' 

EC        , 
liJ 

I     1^ 

LLI 
CL 

cn     6 


(66) 


(67) 


(45)^5 


.{24) 


COVER  TYPES 

Fig.  2.  Plant  species  diversit)  (e"  )  at  random  sites  for 
shrubs,  forbs,  and  grasses  in  the  major  cover  tyjies  (big 
sagebnish  [ARTR],  low  sagebnish  [ARAR],  shrubby 
eriogonum  [ERIO],  intermediate  wheatgrass  [AGIN]), 
1984-85.  The  total  number  of  plant  species  sampled  in  each 
cover  type  is  in  parentheses. 


coverage  of  bhiebunch  wheatgrass  at  grouse 
flush  sites  than  at  random  sites  in  the  big  sage- 
brush cover  ty|3e  in  1984  and  in  the  low  sage- 
brush cover  ty|)e  in  1985. 

Canopy  coverage  at  grouse  flush  sites  in  the 
big  sagebrush  type  differed  among  years  in  five 
of  six  vegetative  categories  (Fig.  3).  Bare  ground 
increased  while  bulbous  bluegrass,  other  forbs, 
and  other  composites  decreased  during  the 
drought  of  1985  as  compared  to  1983  and  1984. 
However,  bluebunch  wheatgrass  increased  in 
1985,  while  the  cover  of  arrowleaf  balsamroot 
was  not  significantly  different  among  years. 

Bluebunch  wheatgrass  and  arrowleaf 
balsamroot  are  native  perennials  that  are  con- 
sidered decreaser  species  (Bhiisdell  and 
Pechanec  1949,  Evans  and  Tisdale  1972);  i.e., 
they  tvpicallv  decrease  or  are  eliminated  under 
heaxy  livestock  grazing  (Dyksterhuis  1949). 
Canopy  coverage  of  decreaser  forbs  was  signif- 
icantly greater  at  flush  sites  than  at  random  sites 
in  the  big  and  low  sagebrush  cover  types  ( IVIarks 
and  Marks  1987a). 

Discussion 

Summer  home  ranges  for  this  subspecies  in 
Colorado  (Giesen  1987)  and  for  other  subspe- 
cies (Artman  1970,  Christenson  1970,  Ramhar- 
ter  1976,  Gratson  1983)  were  sniiiller  than  we 
observed  in  this  study.  Differences  in  home 
range  size  were  probably  a  reflection  of  habitat 
condition;  larger  home  ranges  were  obsened  in 
western  Idaho,  where  decreaser  forbs  were  lim- 
ited and  historic  livestock  grazing  apparently 
had  a  greater  influence  on  the  vegetation. 

From  spring  to  fall,  >90%  of  all  grouse  loca- 
tions were  within  1.2  km  of  a  dancing  ground. 
Similarly,  locations  of  Sharp-tailed  Grouse  in 
other  studies  were  within  1.0  and  2.5  km  of 
dancing  grounds  (Pepper  1972,  Oedekoven 
1985,  Giesen  1987,  Nielsen  and  Yde  1982). 
These  results  suggest  that  maintiiining  habitats 
within  2.5  km  of  dancing  grounds  will  provide 
summer  habitat  recjuirements  for  Shaip-tiiiled 
Grouse. 

Compared  with  other  cover  tyjoes,  big  sage- 
brush sites  had  a  high  diversit)' of  shrubs,  forbs, 
and  grasses;  the  highest  structural  diversity;  and 
the  greatest  canopx'  coverage  of  perennial  bimch- 
grasses.  The  sharptails"  overall  preference  for 
the  big  sagebrush  cover  type  indicated  that  they 
likely  selected  for  habitat  diversitv  relative  to 
surrounding  areas. 


1992] 


Sh.\hp-tailed  Grouse  Summer  Habitat 


171 


T.^BLK  2.  Mean  can<)p\- coverage  (%)  of\'cgetativecateg()rie.siiil)igsagebnish  (ARTR)  and  low  sage! )ni.sli  (ARAR)  cover 
tvpes  at  Columbian  Sliarp-tailed  Grouse  flush  sites  vs.  random  sites. 


Year 

1984 

1985 

ARTR 

ARAR 

ARTR 

ARAR 

Vegetative 

Flush 

Random 

Flush 

Random 

Flush 

Random 

Flush 

Random 

category' 

(io7r 

(42) 

(21) 

(24) 

(107) 

(42) 

(21) 

(24) 

Big  sagebrush 

3.43 

4.03'' 

0.02 

0.07 

4.97 

6.. -32 

0.22 

0.33'' 

Low  sagebnish 

0.21 

0.49'' 

5.45 

7.. 84 

0.55 

0.79'' 

7.03 

7.88 

Bitterlmish 

1.52 

1.02 

0.86 

0.17 

2.76 

l.w'' 

1.15 

0.88 

Otlier  shmbs 

1.73 

0.89 

0.14 

0.59'' 

2.21 

2.69'' 

1.36 

0.40 

Total  shrubs 

6.89 

6.43 

6.47 

8.67 

10.49 

11.84 

9.76 

9.49 

Arrowleaf  balsamroot 

13.60 

6.55'' 

12.21 

3.91'' 

13.06 

7.40'' 

11.91 

5.28 

Other  composites 

7.05 

3.78'- 

5.14 

2.95'' 

2.90 

3.33 

3.02 

3.19 

Otlier  forbs 

12.76 

15.3l'' 

12.83 

14.24 

9.70 

7.87 

14.97 

7.22 

Total  forbs 

33.40 

25.64'' 

30.18 

21.10'' 

25.66 

I8.6O'' 

29.90 

15.69 

Bluebnnch  wheatgrass 

2.93 

2.56'' 

1.02 

0.85 

5.18 

2.91 

4.72 

0.46'' 

Bulbous  bluegrass 

35.87 

24.59'' 

36.83 

23.09 

15.97 

16.52 

13.20 

22.  a3'' 

Other  grasses 

3.76 

4.32 

2.52 

3.32 

3.01 

2.02 

3.33 

3.29 

Tcjtal  grasses 

42.56 

31.47 

40.37 

27.26 

24.16 

21.45 

21.2,5 

26.08'' 

Bare  ground 

23.93 

35.93'' 

28.05 

42. 30'' 

40.23 

48.62'' 

39.31 

48.94'' 

■'Sample  .size  (number  ot  trausecLs  conducted  in  each  tspe). 

'Indicates  significant  difference  (P  <  .05)  in  mean  canopy  coverage  behveen  flush  and  random  sites  withi 


ver  tjpes. 


Slinibb\'  eriogonum  sites,  which  were 
strongK'  axoided  by  grouse,  contained  a  low 
di\ersit\  of  forbs,  and  even  in  the  absence  of 
grazing  proNided  Rttle  cover.  Exchiding  dancing 
grounds,  Shaip-tailed  Grouse  studied  else- 
where have  exliibited  similar  selection  against 
areas  of  sparse  cover  (Pepper  1972,  Ziegler 
1979,  Klott  and  Lindzey  1990).  The  intermedi- 
ate wheatgrass  cover  type  also  was  avoided  by 
grouse.  Grouse  were  particularly  absent  from 
intermediate  wheatgrass  during  years  with  rela- 
ti\elv  low  numbers  of  grasshoppers. 

Mountain  shrub,  riparian,  and  bitterbnish 
habitats  were  used  primariK  as  escape  cover 
during  spring  and  summer.  Beginning  in  late 
siunmer,  moimtaiu  shrub  and  riparian  plant  spe- 
cies produced  fniits  that  became  an  important 
part  of  the  grouse  diet  (Marks  and  Marks 
19S7a).  Proximity  to  this  shrubby  vegetation 
ma\-  not  have  been  critical  during  earK"  to  mid- 
summer when  the  cover  types  preferred  by 
grouse  were  providing  adequate  food  and  cover. 
Grouse  were  found  closer  to  mountain  shrub 
and  riparian  habitat  than  expected  l)y  chance 
only  in  the  drought  year  (1985),  when  xertical 
cover  decreased  significantK'  in  all  cover  t\pes 
that  were  measured. 

Shaq^tails  apparentK'  selected  areas  least 


modified  by  lixestock  grazing.  Grouse  locations 
were  characterized  by  greater  herbaceous  cover 
and  less  bare  ground  than  random  sites.  Studies 
of  plant  communities  with  and  without  gnizing 
indicate  that  areas  with  relati\eK-  little  bare 
ground  are  least  modified  b\-  li\estock  (.see 


40   - 

a 

e 

/ 

S.BAGR 
/ 

30   ^ 

y 

^^ 

y?\ 

\ 

20  - 

\ 

\ 

>POBU 

10    - 

'^""'g'"--.,^ 

. 

~-^OTFO 
-AGSP 

~"OTCO 

^' 

_^^Z-=-' 

"^ 

- 

k 

k 

1 

1 

1 

1984 
YEARS 


Fig.  3.  Comparison  of  canopy  coverage  at  Sharp-tailed 
(Jrouse  flush  sites  in  tlie  big sagel)nish  co\er  t%pe  in  western 
IiliJio.  198.3-85.  On  each  line  different  letters  indicate  that 
corresponding  means  are  significantK'  different  at  P  =  .05. 
(BAGR  =  bare  ground.  POBU  =  bulbous  bluegrass,  BASA 
=  arrowleaf  b;i]samroot,  OTFO  =  other  forbs,  AGSP  = 
bluebiuich  wheatgrass,  OTCO  =  otlier  composite  forbs.) 


172 


Great  Basin  Naturalist 


[Volume  52 


Holechek  et  al.  1989).  When  eoinpared  with 
random  sites,  grouse  locations  had  significantly 
higher  proportions  of  forb  species  that  decrease 
from  overgrazing  (Dyksterhuis  1949).  In  partic- 
ular, grouse  preferred  microhabitats  with 
greater  abundances  of  arrowleaf  btilsamroot 
and  bluebunch  wheatgrass,  two  plant  species 
that  ty}3icallv  decline  with  overuse  by  livestock 
grazing  (Blaisdell  and  Pechanec  1949,  Evans 
andTisdale  1972,  Muegglerand  Stewart  1980). 
These  native  perennials  are  major  components 
of  later  serai  stages  (Hironaka  et  al.  1983). 

The  presence  of  arrowleaf  balsamroot  and 
bluebunch  wheatgrass  as  cover  plants  during  a 
drought  year  is  especially  noteworthy.  These 
plants  are  particularlv  drought  resistant  (Tisdale 
and  Hironaka  1981,  \Vasser  1982).  Bulbous  blue- 
grass,  the  most  abundant  and  widespread  grass 
in  the  study  area,  is  an  introduced  perennial 
with  root  systems  that  die  each  year;  it  is  virtu- 
ally nonexistent  during  years  of  low  moisture 
(Monsen  and  Stevens,  in  preparation).  Indeed 
bulbous  bluegrass  contributed  lower  cover 
values  in  1985  than  in  1983  and  1984  (years  with 
average  moistiu-e)  (Table  2).  In  contrast,  cover 
of  bluebunch  wheatgrass  was  similar  among 
those  years.  In  the  absence  of  nati\e  perennials, 
grouse  would  not  have  had  as  much  cover  dining 
drought  years.  The  loss  of  these  important  cover 
plants  may  have  contributed  to  the  disappear- 
ance of  Columbian  Shaq:)-tailed  Grouse  from 
large  portions  of  their  historic  range. 

CONCLUSlOxNS  AND  MANAGEMENT 

Implications 

Ciiven  the  widespread  decline  of  the  Colum- 
bian Sharp-tailed  Grouse  and  the  fragmented 
nature  of  extant  populations,  consenation  of  all 
potential  sources  of  genetic  variation  should  be 
a  critical  concern  to  managers.  Maintenance  of 
shniljsteppe  coiumunities  in  advanced  serai 
stages  is  especially  important  for  con.servation  of 
summer  habitat  in  the  Intermountain  region. 

Habitat  features  that  characterize  occupied 
habitats  in  western  Idaho  are  flat  to  rolling 
rangeland  in  relatively  good  condition  with  a 
diversity  of  native  shmbs,  forbs,  and  grasses. 
Native  perenniiils  arrowleaf  balsamroot  and 
bluebunch  wheatgrass  are  critical  for  cover 
during  a  drought  year.  Also  important  is  riparian 
vegetation  and  numerous  patches  of  mountiun 
shrubs  for  escape  cover  and  late  summer  food. 
These  habitat  characteristics  suggest  that 
Columbian  Sharp-tailed  Grouse  are  an  indica- 


tor of  good  range  condition  in  the  mesic 
shnibsteppe  of  the  Intermountain  region. 

Federal  land  management  agencies  are 
directed  to  conserve  candidate  species  and  their 
habitats  and  to  avoid  actions  that  mav  cause  the 
species  to  become  listed  as  federally  threat- 
ened/endangered. Conservation  efforts  for 
Columbian  Shaqo-tailed  Grouse,  a  candidate 
species,  must  include  protection  and  enhance- 
ment of  habitats  that  are  occupied  by  the  sub- 
species throughout  their  range,  especially 
disjunct  populations  in  jeopardy  of  extirpation. 
The  success  of  attempts  to  improve  their  cur- 
rent status  is  dependent  on  reducing  distur- 
bances that  may  damage  the  natural  diversity  of 
shrubsteppe  habitat  (e.g.,  overgrazing  by  live- 
stock and  agricultural  development). 

Protecting  habitats  within  2.5  km  of  dancing 
grounds  is  critical  for  mmntainence  of  summer 
habitat.  Suitable  habitats  for  reestablishment 
within  their  historic  range  need  to  be  identified. 
However,  reestabHshment  efforts  for  this  native 
species  should  not  take  precedence  over  pres- 
ervation and  restoration  of  habitats  that  cur- 
rently support  sharptails  (cf.  Griffith  et  al.  1989). 

Acknowledgments 

We  thank  A.  Sands,  L.  Nelson,  S.  Mattise,  R. 
Eng,  T  Lonner,  R.  Autenrieth,  R.  Nelson,  and 
J.  Connelly  for  their  contributions  to  the  study, 
and  the  G.  Tarter  and  T  Nelson  families  for 
granting  access  to  their  lands.  We  are  also  grate- 
ful for  the  field  assistance  provided  by  B.  Czech, 
J.  Berr)'hill,  S.  Lisle,  and  R.  Morales.  J.  Craw- 
ford, C.  Groves,  K.  Giesen,  and  T.  Martin  pro- 
vided useful  suggestions  for  manuscript 
improvement.  The  research  was  funded  bv  the 
U.S.  Bureau  of  Land  Management;  additional 
support  was  provided  bv  the  Idalio  Department 
of  Fish  and  Game  and  Montiuia  State  Universit)'. 

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Received  25  September  1991 
Accepted  16  March  1992 


Great  Basin  NatiinJist  52(2),  pp.  174-178 

CHARACTERISTICS  OF  SITES  OCCUPIED  BY  SUBSPECIES  OF  ARTEMISIA 
TRIDENTATA  IN  THE  PICEANCE  BASIN,  COLORADO 

Thomas  R.  Cottrell    and  Chiules  D.  Bonham" 
Kit/  words:  Artemisia  tridentata.  Colorado,  sagehnisli,  clirointifoij^raplu/,  factor  analysis,  sod. 


Artemisia  tridentata,  big  sagebiiish,  is  the 
dominant  plant  species  in  the  Piceance  Basin  of 
western  Colorado  and  displays  great  morpho- 
logical variabilitv  between  sites.  The  existence 
of  at  least  three  subspecies  is  widely  accepted 
(McArthur  et  al.  1981,  1988).  These  are  A. 
tridentata  spp.  tridentata  Beetle,  A.  tridentata 
spp.  ivi/oming^ensi.s  Beetle  and  Young,  and  A. 
tridentata  spp.  vaseyana  Beetle. 

Despite  extensive  research  in  the  Piceance 
Basin  (Redente  and  Cook  1986),  we  have  found 
only  one  study  referring  to  intraspecific  taxa  of 
sagebrush  (Ward  et  al.  1985).  This  work 
referred  to  subspecies  tridentata  but  did  not 
indicate  where  this  taxon  was  found.  Because 
the  taxa  are  known  to  respond  differentiiilly  to 
soil  and  climate  factors  (Hironoka  1978,  Sturges 
1978)  their  existence  in  the  basin  should  be 
recognized.  The  present  study  was  designed  to 
identify  the  subspecies  o{  Artetnisia  tridentata 
present  in  the  Piceance  Basin  and  to  describe 
soil  characteristics  of  sites  occupied  by  sub- 
species. 


Study  Site 

The  Piceance  Basin  comprises  about  3()()() 
km"  in  Garfield  and  Rio  Blanco  counties  of 
northwest  Colorado  (Fig.  1).  The  cUmate  of  the 
Piceance  Basin  is  semiarid  and  shows  extreme 
variability  in  monthlv  precipitation  (Wymore 
1974).  Consecutive  months  often  receive  little 
precipitation.  The  mean  annual  precipitation 
for  eight  weather  stations  in  the  region  for  the 
period  1951-70  was  35.3  cm,  with  a  95%  confi- 
dence intei-val  of  ±18.7  cm.  About  one-half  of 
the  total  precipitation  falls  as  snow.  The  average 


annual  temperature  ranges  from  7  C  at  1800  m 
to  - 1  C  at  2700  m. 

The  strong  influence  of  topography  on  tem- 
peratiu'e  and  precipitation  results  in  a  complex 
of  habitats  in  the  basin  (Tiedeman  and  Terwilli- 
ger  1978).  Generally,  soil  development  is  corre- 
lated to  elevation.  At  higher  elevations,  except 
ridge  tops,  soils  are  dark  brown,  shallow 
mollisols.  At  mid-elevations,  aridisols  are 
common  on  deep  loess.  The  lowest  elevations 
are  characterized  by  entisols  developed  on 
heavy  clays  and  deep,  sandy  alluvial  soils. 

Methods 

Six  sites  dominated  by  sagebrush  were 
selected  for  this  study  (Table  1).  These  sites 
spanned  the  environmental  extremes  of  sage- 
brush habitat  in  the  Piceance  Basin.  Two  sites 
were  selected  from  each  of  three  broad  topo- 
graphic regions.  High  mounttiin  sites  were 
about  2000  ni;  upland  terraces  and  valley 
bottom  sites  were  below  2000  m. 

Sagebrush  subspecies  were  identified  bv  the 
combined  information  of  three  techniques  and 
verified  by  A.  H.  Winward,  regional  ecologist 
for  Range  and  Watershed  Management,  USFS 
Intermountain  Region,  Ogden,  Utah.  The  first 
technique  involved  field  identification  using 
moqihological  characteristics  based  on  keys  by 
A.  H.  Winward  and  Tisdale  (1977).  Leaf  sam- 
ples were  taken  for  the  other  two  procedin-es. 
Two-dimensional  chromatography,  as  described 
by  Hanks  et  al.  (1973),  was  done  on  persistent 
overwintering  leaves  from  three  plants  at  each 
site  except  site  5,  where  the  moqihologiciil  vari- 
abilit)'  of  the  sagebnish  plants  was  greater  than 
at  the  other  sites.  At  this  site  five  plants  were 


^Department  of  Biology,  Colorado  State  University,  Fort  C^ollins.  Colorado  80523. 
Range  Science  Department,  (Colorado  State  University,  Fort  Collins,  Colorado  80.52,3. 


174 


1992] 


Notes 


175 


m. 


''830 


WYOMING 


PICEANCE 
BASIN 


C     OLORADO 


NEW         MEXICO 


2130- 
24  30- 


Meeker 


P'il 


e^^ 


40°N 


/ 


PICEANCE    BASIN 
COLORADO 


SCALE    IN    MILES 


SCALE    IN    KILOMETERS 
CONTOUR   INTERVAL    300m 


Fig.  1.   The  shulv  iirea  of  the  distribuHcMi  ,A  Anemisia  trklcutata  subspecies  i„  northuest  Colorado. 

tested  by  chromatography.  Results  were  com-  matelv  IS  other  plants  in  the  stucK-  sites  Leaves 

pared  with  representative  chromatograms.  The  were  crushed  In  hand  and  placed  in  glass  con- 

hird  procedure  was  a  leaf  extract  in  uater.  This  tainers  for  four  hours.  These  were  viewed  under 

Uitter  method  was  performed  on  all  plants  tested  long-wave  ultraviolet  light  and  compared  to 

by  chromatography  and  on  a  total  of  approxi-  descriptions  by  Stevens  and  McArthur  (1974) 


176 


Great  Basin  Naturalist 


[Volume  52 


Tabi.K  1.  Location,  elevation,  ;uid  sagebnisli  subspecies  of  stutly  sites.  VAS  —  ssp.  id.sci/ana:  TRI  —  ssp.  tridoitata:  WTO 
=  ssp.  wuoimngensis.  Selected  soil  characteristics  are  listed  for  ()-L5  cm  and  16^30  cm  soil  samples  for  each  site. 


L()cation 

Site 

Elev. 

ssp. 

Depth 
in  cm 

%  sand 

%  silt 

pH 

C:aCO^ 
est. 

llji^h  mountain 

I 

2.365 

VAS 

0-15 

54 

26 

6.9 

low 

1.5-30 

52 

26 

6.8 

low 

2 

2585 

VAS 

0-15 

40 

33 

6.8 

low 

1.5-.30 

36 

33 

6.5 

low 

Viilley  bottom 

3 

1987 

TRI 

0-15 

74 

13 

8.2 

med 

1.5-.30 

67 

20 

8.1 

med 

4 

2057 

TRI 

0-15 

56 

30 

8.1 

med 

1.5-.30 

52 

32 

8.2 

med 

Upland  terrace 

5 

1920 

WYO 

0-15 

46 

27 

8.2 

med 

15-30 

51 

27 

8.3 

med 

6 

2070 

WTO 

0-15 

32 

45 

7.7 

med 

1.^.30 

28 

44 

8.2 

med 

In  each  site,  soil  samples  were  collected  at 
two  random  locations  from  two  depths,  0-15  cm 
and  16-30  cm.  These  were  analyzed  for  pH, 
organic  matter,  electrical  conductivity,  esti- 
mated CaCOa,  sand,  silt,  clay,  K,  Mn,  Zn,  Cu,  P, 
and  Fe.  These  data  were  used  in  a  factor  analysis 
as  described  by  Affifi  and  Clark  (1984).  the 
factor  scores  for  each  site  and  depth  were  then 
graphed.  This  graph  was  used  to  inteqDret  the 
axes  that  usually  represent  some  environmental 
characteristic  associated  with  plant  species. 

Results 

Sites  1  and  2  were  high  mountain  sites.  Sage- 
brush plants  averaged  less  than  50  cm  in  height. 
(Common  associated  plants  were  Liipimis  .sp., 
ChnjsotJiamnus  viscidiflonis,  Erio<s<^iuiin  iimlwl- 
latum,  Stipa  letfentuinii,  and  Si/nifjlioricaiyo.s 
oreophilus.  Soils  were  all  deeper  than  40  cm  and 
dark  in  color.  Near  these  sites  Populus 
tremuloides  stands  were  c-onunon  in  favorable 
microenvironments. 

Sites  3  and  4  were  in  a  valley  bottom  in  the 
Yellow  Creek  area.  The  sagebrush  in  these  sites 
commonly  reached  heights  greater  than  2  m. 
Associated  vegetation  included  the  moss  Tor- 
tula  niralis,  CJwnopodium  prdtcricohh  and 
Lepidhun  laiifolhnn.  Soils  were  light  in  color, 
and  depths  greater  than  40  cm  were  common. 

Sites  5  and  6  were  at  similar  elevations  to  3 
and  4,  but  away  from  streams.  Sagebrush  plants 
averaged  40  cm  in  height.  Site  5  soils  were 
approximately  10  cm  in  depth.  Bromiis  fec- 
tonim,  Gutierrezia sarodirac,  Ah/ssuni  ahfssoidcs. 


and  Ort/zopsis  Ju/moioides  were  the  common 
plant  species.  Site  6  soils  averaged  20  cm  deep. 
Common  understoiy  species  were  Koeleria 
crisfafa,  A^ropijwn  smitliii,  and  PJilox  hoodii. 
Site  6  was  surrounded  by  forests  ofPintis  ednlis 
and  Jiinipenis  osteospemw. 

Locations  of  A.  fridentata  subspecies  in  the 
study  area  relate  generalK'  to  elexation.  The 
lowest  elevations  supported  both  ssp. 
wijominfj^ensis  and  ssp.  fridentata.  Factor  anal- 
ysis results  indicate  that  soil  texture  and  chem- 
istiy  differences  existed  between  the  sites  (Fig. 
2).  Subspecies  tiidentata  was  found  in  sandier 
soils  and  wijomin^ensis  in  siltier  soils.  The  tex- 
ture differences  were  generally  related  to  topo- 
graphic position.  Subspecies  tiidentata  was 
most  common  in  xallev  bottoms,  and  ssp. 
wyoniin^ensis  was  tvpicallv  dominant  awaN' 
from  streams,  at  the  lowest  elexations  to  approx- 
imately 2100  m.  Sites  above  2100  m  supported 
ssp.  vasei/ana.  Soil  te.xtures  in  vaseijana  site  1 
were  similar  to  those  in  tridentata  sites,  while 
vaseijana  site  2  textures  more  closely  resembled 
those  of  the  wt/oinin^iensis  sites.  Soil  pH  was 
lower  in  the  vasetjana  sites  than  sites  with  the 
other  siibspecies. 

Moiphologiciil  identification  of  ssp.  vasei/ana 
and  tridentata  generalK-  agreed  with  the  results 
from  two-dimensional  chromatograph\  and  the 
leaf  extracts.  Subspecies  wyomingensis  chro- 
matograms  were  not  consistently  separable 
from  those  of  subspecies  tridentata.  None  of 
the  wi/oniin(^en.si.s  chromatograms  closely 
matched  published  chromatograms.  Leaf  ex- 
tracts from  ssp.  ivt/innin^ien.si.s  showed  almost  no 


1992] 


Notes 


177 


LJ 

< 

Ld 

u 


O 
< 


Q 

< 


1.5 


0.5  - 


-  -0.5 


ct: 
O 
I— 
u 
< 


-1  - 


-1.5 
-1.5 


3s 

tridentata 

3d 

4d 

^=d 

5s 

vaseyana 

6s 

2d 

wyomlngensis 

2s 

6d 

1                1 

1                 1                 1 

-1     -0.5     0      0.5      1      1.5 
FACTOR  2:  ELEV.  INCREASES,  pH  DECREASES 


Fig.  2.  Results  of  factor  analysis  on  all  soil  data.  Y  axis  corresponds  to  increasing  sand:silt  ratio;  X  axis  corresponds  to 
decreasing  pH.  Stand  numbers  are  as  in  Table  1;  s  indicates  sample  from  0-15  cm;  d  indicates  sample  from  15-30  cm. 
Names  show  approximate  region  of  ordination  occupied  bv  each  subspecies. 


fluorescence  and  were  not  separable  from  ssp. 
tridentata.  Moqohologicallv,  howev'er,  this  sub- 
species was  separable  from  vasci/ana  and 
tridentata  bv  the  ke\s  of  Winward  and  Tisdale 

(1977). 

Discussion 

Three  subspecies  of  A.  tridentata  were  iden- 
tified in  the  Piceance  Basin  b\  reference  to 
moiphologv;  chromatography,  and  leaf  extracts. 
The  subspecies  identified  were  wyomingensis, 
tridentata,  and  vasei/ana.  Two-dimensional 
cliromatograpliN  and  leaf  extracts  \ielded  pre- 
Hininary  evidence  to  suggest  that  ssp. 
wyominiiensis  in  the  Piceance  Basin  is  chemi- 
cally different  from  those  previously  identified. 

The  distributions  of  A.  tridentata  subspecies 
are  generaliv  related  to  soil  moisture,  tempera- 
ture, depth,  and  parent  material  (Hironaka 
1978).  The  overall  tendency  seems  to  be  for  ssp. 
tridentata  to  occupy  deep,  somewhat  sandv 
soils.  Although  subspecies  wijcnningensis  occurs 


in  an  overlapping  zone  \\A{\\  tridentata,  it  is  more 
common  in  shallow,  silt\'  soils  where  moisture 
stress  is  greater.  Subspecies  lasei/ana  occurs  in 
cool,  moist  sites,  usutilly  above  2100  m,  but 
lower  elevations  have  been  documented  (Good- 
rich et  al.  1985). 

Each  subspecies  was  found  at  eJexations  and 
in  soil  textures  similar  to  those  reportcnl  in  the 
literature.  Soil  texture,  expressed  as  a  ratio  of 
sand  to  silt,  explains  the  first  factor  in  the  factor 
analvsis  and  distinguishes  i)etween  sites  of  ssp. 
tridentata  and  wyomingensis  (Fig.  2).  That  is, 
the  vertical  axis  in  Figure  2  corresponds  to  this 
ratio.  It  appears  that  the  relative  proportion  of 
sand  and  silt  determines  whether  ssji.  tridentata 
or  ssp.  wt/oniingensis  will  be  dominant.  Barker 
and  McKell  ( 1983)  reported  similar  results  and 
suggest  that  the  characteristics  of  soils  associ- 
ated with  these  subspecies  are  different.  Fine- 
textured  soils  ha\e  been  implicated  in  increased 
water  stress  in  ssp.  wyoniingensis  sites  (Shumar 
and  Anderson  1986).  This  might  indicate  a 
differential    adaptation    to   water   stress    and. 


178 


Great  Basin  Naturalist 


[Volume  52 


consequently,  different  life  history'  strategies  in 
the  subspecies  (Bonham  et  al.  1991). 

Soils  at  sites  with  ssp.  vaseyana  are  distin- 
guished from  the  other  sites  by  factor  2  of  the 
factor  analysis.  This  axis  represents  both  an  ele- 
vational  and  soil  pH  gradient  (Table  1,  Fig.  2). 
Sites  with  ssp.  vaseyana  were  at  a  higher  eleva- 
tion, and  soils  were  lower  in  pH  and  CaCO^ 
values.  The  textures  at  these  sites  did  not  differ 
substantiiillv  from  the  other  sites. 

No  previous  study  in  the  area  has  identified 
these  taxa  or  characterized  their  habitats.  The 
great  differences  in  habitat  preference  among 
these  subspecies  suggest  this  is  a  major  over- 
sight. 

Acknowledgments 

The  research  was  supported  jointly  by  the 
U.S.  Department  of  Energy,  Contract  No.  DE- 
AS02-76EV04018  to  Colorado  State  Universit)- 
and  the  Agricultural  E.xjoeriment  Station,  Colo- 
rado State  University,  Project  660(4242). 

Literature  Cited 


Affifi.  a.  a.,  and  Clark.  V.  1984.  Computer-aided  multi- 
variate analy.sis.  Lifetime  Learning  Publications,  Bel- 
mont, California.  458  pp. 

Bakker,  J.  R.,  and  C,  M.  McKell  1983.  Habitat  differ- 
ences between  basin  and  Wyoming  big  sagebrusb  in 
contiguous  populations.  Journal  of  Range  Manage- 
ment 36:  450-454. 

BoNiiAM.  C.  D.,  T.  R.  CoTTHELL.  and  J.  E.  Mitchell. 
1991.  Inferences  for  life  history  strategies  of  Artemisia 
tridentata  subspecies.  Journal  of  Vegetation  Science  2: 
339-;344. 

GooDRiCM,  S.,  E.  D.  McArthur,  and  A.  H,  Winward 
1985.  A  new  combination  and  a  new  variety  oi  Artemi- 
sia tridentata.  Great  Basin  Naturalist  45:  99-104. 

Hanks,  D.  L.,  E.  D.  McArthuk.  R.  Steven.s,  and  A.  R 
PiAJMMER.  1973.  Chromatographic characteri.stics  and 
phylogeiietic  relationships  of  Artemisia  section 
Tridentatae.  USDA  Forest  Service  Research  Paper 
INT-141.24pp. 


HiRONAKA.  M.  1978.  Basic  svnecological  relationships  of 
the  Columbia  River  sagebrush  type.  Pages  27-.32  in 
The  sagebnish  ecosystem:  a  symposium.  Utah  State 
University,  LogiUi. 

McARTHUR,  E.  D.,  C.  L.  Pope,  and  D.  C.  Freeman  1981. 
Chromosomal  studies  of  subgenus  Tridentatae  ofArte- 
misia:  evidence  for  autopoKploidv.  American  Journal 
of  Botany  68:  589-605. 

McArthur,  E.  D.,  B.  L.  Welch,  and  S.  C.  Sanderson. 
1988.  Naturd  and  artificial  hybridization  between  big 
sagebnish  {Artemisia  tridentata)  subspecies.  Journal  of 
Heredit)'  79:  268-276. 

Redente,  E.  F.,  andC.  W.  Cook,  eds  1986.  Structural  and 
functional  changes  in  early  successional  stages  of  a 
semiarid  ecosystem.  Progress  Report  to  U.S.  Depju't- 
ment  of  Energy.  Depiutment  of  Range  Science,  Colo- 
rado State  University,  Fort  Collins.  67  pp. 

Shumar,  M.  L.,  and  J.  E.  Anderson  1986.  Gradient  anal- 
ysis of  vegetation  dominated  by  two  subspecies  of  big 
sagebnish.  Journal  of  Range  Miuiagement  39:  156- 
159. 

Stevens,  R.,  iuid  E.  D.  McArthur  1974.  A  simple  field 
technique  for  identification  of  some  sagebnish  taxa. 
Journal  of  Range  Management  27:  .325-.326. 

Sturc:es.  D.  L.  1978.  Hydrologic  relations  of  sagebrush 
lands.  Pages  86-100  in  The  sagebnish  ecosystem:  a 
symposium.  Utah  State  Universitv',  Logan.  251  pp. 

TiEDEMAN.  J.  A.,  and  C.  Terwilliger.  Jr  1978.  A  phy- 
toedaphic  classification  of  the  Piceance  Basin.  Colo- 
rado State  University,  Department  of  Range  Science, 
Science  Series  31.  265  pp. 

Ward.  R.  T,  W  L.  Slauson,  and  C.  W.  Weldon  1985. 
Response  of  shnib  ecotvpes  to  mining  waste  material 
in  soil  profiles  and  competitive  interactions  of  woodv' 
species  under  experimental  and  naturiJ  conditions. 
Pages  87-94  in  E.  ¥.  Redente,  C.  W.  Cook,  J.  M.  Stark, 
and  C.  L.  Simmons,  eds.,  Semiarid  ecosvstem  de\eIop- 
ment  as  a  function  of  resource  processing  and  alloca- 
tion. Progress  Report  to  U.S.  Department  of  Energ\-. 
Colorado  State  University,  Department  of  Range  Sci- 
ence, Fort  Collins. 

Winward,  A.  H.,  and  E.  W.  Tisdale  1977.  Titxonomv  of 
the  Artemisia  tridentata  complex  in  Idiilio.  Uni\ersit\' 
of  Idiilio,  Forest,  Wildlife  and  Range  Experiment  Sta- 
tion Bulletin  No.  19.  15  pp. 

WvMORE,  I.  F.  1974.  Estimated  average  annual  water  bal- 
ance for  Piceance  and  Yellow  Creek  watersheds.  En\a- 
ronmentiJ  Resources  Center,  Colorado  State 
University,  Fort  Collins.  Technical  Report  Series  No. 
2.  60  pp.  ■ 

Received  20  November  1991 
Accepted  4  May  1992 


Great  Basin  Natunilist  52(2).  pp.  179-184 

USE  OF  LAKES  AND  RESERVOIRS  BY  MIGRATING  SHOREBIRDS  IN  IDAHO 


)aiiiil  \l.Ta\I()r    and  (  iharlfs  il.lVo.st 


Kit/  uonh:  shorrhird.s.  habitat  ii\c.  iiiiiilfldts.  uaU'r  (Irandoini.  irri<^atiiiii  rcsi'troirs.  ini^ratin^  liinl.s 


ShorebircLs  migrating  long  cli.stances  are  \iil- 
lu^able  because  their  wetland  .stopover  sites  are 
limited  in  number  and  susceptible  to  distur- 
bance or  destniction  b\'  humans  (Senner  and 
Howe  1984,  Myers  et  al.  1987).  It  is  therefore 
critical  to  know  which  wetland  areas  migrating 
shorebirds  use,  and  the  factors  making  the.se 
sites  attracti\e  to  shorebirds. 

\\  e  conducted  shorebird  censuses  at  numer- 
ous wetland  sites  in  Idaho  with  these  objectives: 
(1)  to  identify  t>pes  of  lakes  and  reservoirs  that 
are  important  for  migrating  shorebirds,  (2)  to 
identih'  habitat  characteristics  at  these  wetlands 
used  b\  shorebirds,  (3)  to  determine  the  inilu- 
ence  of  mudflat  exposure  and  water  le\el 
cliang(^s  on  shorc^bird  use. 

Study  Ahk.\s  and  Methods 

A  total  of  19  lakes  and  resenoirs  were  cen- 
sused  at  least  once  in  1989  (Table  1).  Nine 
high-ele\'ation  lakes  were  visited  in  the  Saw- 
tooth Wilderness  in  earlv  September  1976,  and 
three  high -elevation  lakes  in  the  Seafoam  area 
of  the  Frank  Church  River  of  No  Return  Wil- 
derness in  earlv  .August  1990.  Additional  obser- 
vations from  Lake  Lowell  were  made  in  1986, 
1987,  and  1990.  All  shorebirds  were  censused 
within  100  m  of  the  shoreline  in  and  out  of  the 
w  ater  at  all  sites;  thus,  evei-v  500  m  of  transect 
censused  was  equal  to  0.1  km".  We  estimated 
birds  per  500  m  of  shoreline  for  our  densitv 
estimates.  The  Springfield  area  of  American 
Falls  Reservoir  had  over  15  km  of  mudflat 
exposed  by  drawdown  during  the  study  period 
and  also  included  numerous  seep  areas  awav 
from  the  main  shoreline;  because  of  this,  it  was 
not  possible  to  make  density-  estimates  from  this 
site.  Four  of  the  lakes  and  reservoirs  visited  in 
1989  had  mudflat  areas  that  were  censused  at 


least  six  times  at  roughly  weekly  inteivals  from 
mid-Julv  to  earlv  September,  the  time  of  peak 
shorebird  abundance  in  Idaho  (Tavlor  et  al. 
1992).  We  used  ANOVA  and  Newman-Keuls 
tests  (Zar  1974)  to  compare  differences  in 
shorebird  numbers  at  these  four  sites.  Birds 
were  censused  bv  walking  from  10  to  1 00  m  back 
from  the  shoreline  and  using  binoculars  and  a 
25X  spotting  scope.  Care  was  taken  not  to  dis- 
turb birds.  If  birds  moved,  their  numbers  were 
kept  track  of,  or  the  entire  coimt  was  restartc^d 
to  avoid  counting  birds  more  than  once. 

Results 

The  natural  lakes  at  high  elevations  we  cen- 
sused in  1989  (Table  2)  had  onfy  0-2  Spotted 
Sandpipers  (see  Table  3  for  cill  scientific  names). 
Only  a  single  Spotted  Sandpiper  was  found  at 
nine  high-elevation  lakes  visited  in  the  Sawtooth 
Wilderness  in  September  1976.  No  shorebirds 
were  found  at  three  high-elevation  lakes  in  the 
Seafoam  area  in  early  August  1990. 

At  the  Lowell,  Walcott,  American  Falls,  and 
Carey  areas  we  found  significant  differences  in 
the  densities  of  total  shorebirds  (ANO\'A,  F2(3) 
26  =  88.76,  P  <  .001).  Lake  Lowell  had  signifi- 
cantly the  most  shorebirds,  American  Falls  had 
significantlv  more  than  Carey  Lake,  but  Carey 
Lake's  higher  mean  was  not  significantlv  more 
than  Lake  Walcott  s  (Newnian-Keuls,  q  =  29.89 
to  7.47,  for  significant  differences  P  <  .05  or 
greater;  (j  =  2.04,  P  =  :2  for  Carey  Lake-Lake 
Walcott).  These  differences  in  shorebird  num- 
bers reflect  the  amount  of  mudflat  available  at 
the  different  sites;  the  larger  the  mudflats,  the 
greater  the  number  of  shorebirds. 

The  pattern  of  more  shorebirds  being 
attracted  to  larger  mudflats  is  further  supported 
In  shorebird  numbers  at  different  Lowell  sites 


Department  of  Biolopcal  Sciences.  Idaho  State  University,  Pocatello.  Idalio  83209 


179 


180 


Great  Basin  Naturalist 


[Volume  52 


TaBI.f:  1.  Characteristics  of  hkilio  lakes  ;uicl  reservoirs  sur\'eyecl  for  shorehirds  in  19S9. 


Transect 

Elevation 

length 

Name 

County 

(m) 

(m) 

Habitat 

Reservoirs  and  lakes  with  mudflats 

American  Falls 

Power 

1321 

900 

500  m  mudflat 

Lowell 

C;inyon 

757 

4600 

1200  m  mudflat 

Walcott 

Minidoka 

1279 

1500 

20  m  mudflat 

Carey 

Blaine 

1453 

2200 

200  m  mudflat 

Little  Camas 

Elmore 

1502 

800 

120  m  mudflat 

Dry 

Ciuiyon 

818 

15(X) 

50  m  mudflat/700  m  grass 

Mackay 

Custer 

1849 

1400 

200  m  mudflat 

Palisades 

Bonneville 

1708 

1600 

1000  m  mudflat 

Reservoirs  and  lakes 

without  mudflats 

Cascade 

ViJley 

1472 

2600 

1-2  m  sandy  or  muddy  shore 

Wilson 

Jerome 

1224 

1800 

dirt  or  grass  shore 

Boulder 

Valley 

2127 

900 

2  m  mud  or  rocky  shore 

Bnineaii 

Owyhee 

763 

23(:K) 

1  m  mud  or  sandy  shore 

High-elevation  lakes 

Alice 

Blaine 

2622 

1000 

herb  or  rocky  shore 

Toxaway 

Custer 

2539 

9(K) 

herb  or  rockv  shore 

Edith 

Custer 

2611 

6(X) 

herb  or  rocky  shore 

East 

Valley 

2373 

1100 

herb  or  rocky  shore 

West 

Valley 

2361 

900 

herb  or  rocky  shore 

North 

Valley 

2367 

7(X) 

herb  or  rocky  shore 

Payette 

Valley 

1522 

700 

herb  or  rocky  shore 

responding  to  changes  in  mudflat  conditions  in 
1989  (Fig.  1).  In  July  Public  Access  No.  1  had 
verv'  few  shorebirds,  and  nearly  all  of  its 
mudflats  were  submerged  by  water  (Fig.  lb). 
The  New  York  Canal  site  was  submerged  at  this 
time  and  had  no  birds  (Fig.  la).  When  the  large 
mudflats  of  the  New  York  Canal  site  became 
exposed  in  August,  thousands  of  shorebirds 
appeared  there  (Fig.  la).  Numbers  of  shore- 
birds  at  some  of  the  other  sites  declined  (Fig. 
lb),  which  may  have  been  due  in  part  to  birds 
shifting  to  the  New  York  Canal  site.  The  reflood- 
ing  of  Lowell  in  late  September  1989  com- 
pletely eliminated  shorebirds  from  census  areas 
by  27  September  (Fig.  1),  although  American 
Falls  Reservoir  had  over  500  shorebirds  at  this 
time.  On  27  September  1990,  uith  Lake  Lowell 
very  low  due  to  dam  reconstniction,  there  were 
extensive  mudflats  at  the  New  York  Canal  site, 
and  926  individuals  of  10  species  of  shorebirds 
were  present.  In  earlv  Julv  1986  there  were 
hundreds  of  shorebirds  on  the  exposed  mudflats 
at  Public  Access  No.  1,  but  in  early  July  1987, 
with  high  water  flooding  into  riparian  vegetation 
at  this  site,  there  were  no  shorebirds. 

The  reservoirs  we  counted  once  or  a  few 
times  in  1989  usually  supported  the  pattern  of 
total  shorebird  numbers  declining  with  decreas- 


ing mudflat  size,  but  there  were  some  excep- 
tions (Table  2).  Wilson,  Boulder,  and  Cascade 
reservoirs  all  had  zero  or  onh'  a  few  meters  of 
exposed  shoreline,  and  thev  had  only  1  or  2 
shorebirds.  Mackay  Reservoir  had  onlv  2  shore- 
birds  on  3  July  when  no  mudflats  were  exposed, 
but  351  two  weeks  later  when  there  was  200  m 
of  mudflat.  The  Drv^  and  Little  Camas  reservoirs 
supported  hundreds  of  shorebirds  (Table  2), 
and  these  sites  had  mudflats  of  50-120  m.  How- 
ever, Bruneau  had  onlv  1-2  m  of  mud  or  sandy 
beach,  and  it  had  79  individual  shorebirds.  An 
even  stronger  anomalv  was  Palisades,  a  reservoir 
which  had  exposed  mudflats  of  about  1000  m 
and  water  drawdown  continually  exposing  new 
areas,  but  practicallv  no  birds  (Table  2). 

Black-bellied  Plovers,  Lesser  Golden-Plo- 
vers, Sanderlings,  Pectoral  Sandpipers,  and  Stilt 
Sandpipers  were  found  only  on  mudflats  with 
>500  m  of  exposed  mud  (Table  3).  Ten  other 
shorebirds  species  were  most  abundant  at  sites 
with  >500  m  of  exposed  mudflat.  Eight  shore- 
bird  species  had  similar-sized  peaks  at  sites  with 
>500  m  or  between  20  and  200  m  of  ex'posed 
mudflat.  The  onlv  species  with  a  maximmn  peak 
on  mudflats  between  20  and  200  m  was  the 
uncommon  Long-billed  Curlew.  No  individual 
shorebird  species  had  maximum  numbers  at 


1992] 


Notes 


181 


Table  2.  Total  numher  and,  in  parentheses,  densit\  per  0.5  km  of  transect  ol  sliorehirds  counted  at  lakes  and  reservoirs 
in  Idaho  in  1989. 


Count  area 


Mean 


SD 


Range 


Springfield 
American  Falls 

9 
9 

Lowell 

8 

Wiilcott 

9 

Carey 

6 

Little  Camas 

4 

Da' 

4 

Macka\- 

2 

Palisades 

4 

Cascade 
Boulder 

2 
1 

Wilson 

Bnmeau 

Alice 

Payette 

Edith 

Toxawav 

West 
East 
North 

2296 

578.1 

1698-3252 

209 

87.2 

92-337 

(105) 

(43.6) 

(46-168.5) 

3061 

1839.6 

752^5739 

(323) 

(230.6) 

(7^717) 

54 

40.6 

17-153 

(18) 

(13.4) 

(6-.50) 

254 

111.9 

80-393 

(58) 

(25.4) 

(1^89) 

294 

161.5 

117^46 

(184) 

(101.0) 

(7;3-279) 

132 

28 

93-158 

(44) 

(9.3) 

(31-53) 

177 

2^51 

(62) 

(1-125) 

18 

23.6 

0-70 

(6) 

(8.3) 

(0-18) 

0 

1 

(0.6) 

0 

79 

(17) 

1 

(1) 

0 

0 

1 

(0.7) 

0 

0 

0 

sites  with  <5  m  of  mudflats  or  rocWlierh  shore- 
lines. 

Discussion 

Tlie  \irtiial  absence  of  shorebirds  from  the 
19  hi<rh-ele\ation  lakes  we  \isited  in  1976,  1989, 
and  1990  is  similar  to  the  findings  of  the  only 
previous  study  of  a  high-elevation  lake  in  Idaho. 
Visits  annually  to  Fish  Lake,  Idaho  Co.,  from 
1923  to  1929  found  only  a  few  Solitary  Sandpip- 
ers and  Spotted  Sandpipers,  and  one  or  two 
indi\iduals  of  four  other  species  (Hand  1932). 
Burleigh  (1972)  reported  no  large  numbers  of 
shorebirds  at  any  high-elevation  lakes  in  Idaho. 
Further  investigation  may  reveal  some  high-ele- 
vation lakes  to  be  important  for  migrating  shore- 
birds,  but  the  lack  of  mudflats  at  most  of  these 
lakes  probabK'  limits  their  use  by  most  shorebird 
species. 

The  concentration  of  most  shorebirds  at 
large  mudflats  is  consistent  with  our  previous 


findings  at  American  Falls  Resenoir,  where  we 
foimd  verv  few  shorebirds  on  sand\',  clay,  or 
boulder  beaches  or  bedrock  (Taylor  et  al., 
unpublished  data).  Shorebirds  also  concen- 
trated on  mudflats  at  inland  studies  done  in 
Nevada  (Hainline  1974),  Missouri  (Rundle  and 
Fredrickson  1981),  and  Saskatchewan  (Colwell 
and  Oring  1988),  although  the  latter  study  also 
had  some  shorebird  species  associated  with  dif- 
ferent habitats.  Our  stud)  also  shows  that  small 
and  moderate-sized  mudflats  of  both  natunil 
lakes  and  reservoirs  mav  attract  some  shore- 
birds,  especially  those  that  often  feed  in  water. 
Shorebird  species  that  primarily  or  com- 
pletel)  feed  b)-  probing  in  or  gleaning  off  land 
surfaces  or  very  shallow  water  almost  always  had 
higher  peaks  on  the  larger  mudflats,  or  were 
foiuid  there  exclusi\ely.  An  exception  was 
Baird's  Sandpiper,  which  had  a  similar  peak 
between  large  and  moderate  mudflats.  Five  of 
the  shorebird  species  with  equal-sized  peaks  on 
large  and  moderate  mudflats,  the  Black-necked 


182  Great  Basin  Naturalist  [Volume  52 

Table  3.  Slioiehird  sjx-cies  (ouiid  at  19  tfsenoiis  and  hikes  in  Idalio  in  1989. 
Species  Abundance''  and  habitat  use ' 

Black-bellied  Plover  Uncommon  on  large  mudflats. 

Pluiialis  scjiiatawla 
Lesser  Colden-Plover  Rare  on  large  mudflats. 

Fluvialis  dominica 
Semipalmated  Plover  Uncommon  on  large  mudflats;  rare  on  moderate  nmdflats  luid  muddy  shores. 

Cluirad lilts  scmipaliiuitiis 
Killdeer  Common  on  large  and  moderate  mudflats;  uncommon  on  muddv  shores;  occasional  on 

Charadriiis  vocifcnis  rocky/lierb  shoreline. 

Black-necked  Stilt  Uncommon  on  kri'ge  and  moderate  nmdflats;  rare  on  muddv  shores. 

Himantopus  mexkamis 
American  Avocet  Abundant  on  Luge  mudflats;  uncommon  on  moderate  mudflats  and  muddv  shores. 

Real m irostra  anie rica 1 1 a 
Greater  Yellowlegs  Uncommon  on  laige  and  moderate  mudflats;  occasion^  on  muddy  shores. 

Tringa  melanoleuca 
Lesser  Yellowlegs  Conuiion  on  large  nmdflats;  uncommon  on  moderate  mudflats;  occasional  on  muddy 

Tringa  flavipes  shorelines. 

Solitary  Siuidpiper  Occasional  to  rare  on  all  shore  t\pes. 

Tringa  solitaria 
Willet  Unconunon  on  Uirge  mudflats;  occasional  on  moderate  nmdflats;  rare  on  muddv  shorelines. 

Catoptwphonis  scmipahnatus 
Spotted  Sandpiper  Uncommon  on  large  and  moderate  mudflats,  muddy  shorelines;  occasional  on  rock)'/herb 

Actitis  maadaria  shorelines. 

Long-billed  Curlew  Occasional  on  moderate  mudflats;  rare  on  large  mudflats. 

Numeii iu.s  aincricanus 
Marbled  Godwit  Common  on  large  mudflats;  occasional  on  moderate  mudflats. 

Lhnosa  fedoa 
Sanderling  Uncommon  on  huge  mudflats. 

Calidris  alba 
Semipalmated  Sandpiper      Uncommon  on  large  mudflats;  occasional  on  moderate  nuidflats. 

Calidris  pusilla 

Western  Sandpiper  Abundant  on  huge  mudflats;  common  on  moderate  mudflats;  uncommon  on  mudd\'  shores. 

Calidris  iiuiuri 

LeiLst  Sandpiper  Unconunon  on  hu^ge  muiiflats;  occasional  on  moderate  mudflats. 

Calidris  minittilla 

Baird's  Sandpiper  Conunon  on  large  and  moderate  mudflats;  occasional  on  nnidd\'  shores. 

Calidris  hairdii 

Pectoral  Sandpiper  Uncommon  on  large  mudflats. 

Calidris  mclaiiotus 

Stilt  Sandpiper  Riu-e  on  large  mudflats. 

Calidris  hiiiuiiitopus 

Short-billed  Dowitcher  Occasional  on  large  and  moderate  mudflats. 

Limnodromus  grisciis 

Long-billed  Dowitcher  Conunon  on  large  uuuinats;  uncommon  on  moderate  nuidflats  and  nuidd\  shores. 

Limnodromus  scolopaccns 

Common  Snipe  Uncoiumon  on  huge  nuidflats;  occasional  on  moderate  nmdflats  and  uurIcIv  shores. 

Gallinago  gallinago 

Wilson's  Phiilarope  Conunon  on  large  and  moderate  mudflats;  uncommon  on  uuuldv  shores. 

Phalaropiis  tricolor 

Red-necked  Pluilarope  Conunon  on  large  and  moderate  nuidflats;  occasional  on  imidd)  shores. 

Phakiropxis  lobatiis 

■'A  species  was  considered  abundant  if  it  had  a  single  peak  count  over  10(K)  at  a  siiecific  site,  cuinnion  u-itli  a  peak  o\er  100,  iiiiconinion  wUh  a  peak  <)\er  10,  ocxiisional 
with  a  peak  under  10,  and  rare  if  only  one  or  two  individuals  were  found 

Uirge  mudflats  include  American  Falls,  Springfield,  Palisades,  and  Lowell,  and  all  had  water  drawdnwii  exposing  mndllats  of  distances  >.5tK)  in.  Moderate  mudflats 
include  Carey,  Little  Camas,  Di\'  (in  part),  Mackay,  and  V\'alcott.  and  had  water  drawdouii  exposing  20  2IKI  ni  ol  mudflat.  Muddy  shores  included  Dry  (in  part), 
Bnmeau.  Oiscade,  Boulder,  and  Payette  (in  part),  and  tliese  included  small  muddv  shorelines  or  iiiudflais  of  .5  m  widtli  or  less  and  also  sandy  or  dirt  shorelines. 
Rocky/herb  shorelines  included  Alice.  Dry(in])art),  Kitst.  F.ditli.  North.  Pavette  l  in  parti,  Toxawav,  and  Wil.son. 


19921 


Notes 


183 


shorebirds 


1500  - 

Public  #1                            0 
Public  #2                          /\ 

1000  - 

Public  #3                           /    \ 

B 

Q                 /       1 

500  - 

\ 

j\l          \ 

• 

^ 

:^^^*=*-fcJl 

Q' '   T  '^  T 

■    -r    ■     1     ■      1      '     1      ■     1     '    T     '^P    • 

isades  Reservoir  in  tliis  stiicK,  indicates  there  are 
additional  factors  inflnencing  shorebird  use. 
This  could  include  food  abundance  (Harrison 
1982,  Myers  et  al.  1987),  which  is  important  at 
American  Falls  Resenoir  (Mihuc  1991),  tradi- 
tional use  (Myers  et  al.  1987),  and  in  the  case  of 
Palisades  Reservoir  possible  difficulty  of  shore- 
birds  locating  it  because  it  is  enclosed  by  high 
mountains  in  all  directions  (personal  observa- 
tion). Steep-sided  resenoirs,  such  as  C.  J. 
Strike,  Hells  Canyon  (personal  observation), 
and  Lower  Granite  Creek  (Monda  and  Reichel 
1989)  on  the  Snake  River,  and  stretches  of  the 
Columbia  River  subject  to  water  level  fluctua- 
tions (Books  1985),  supported  few  shorebirds 
even  with  water  drawdown  in  summer  and  tail. 
The  absence  of  shorebirds  at  Lake  Lowell 
and  Mackay  Reservoir  from  sites  when  high 
water  covered  mudflats  shows  the  importance 
of  water  drawdown  exposing  these  areas  during 
migration.  At  American  Falls  Resenoir  we  have 
previously  found  shorebird  numbers  to  be  cor- 
related with  rate  of  drawdown  (Tavlor  et  al., 
unpublished  data).  Water  levels  at  reservoirs  in 
this  region  are  usually  determined  by  irrigation, 
power  generation,  recreational  activities  such  as 
boating,  or  waterfowl  management.  It  is  impor- 
tant that  controllers  of  water  levels  at  reservoirs 
and  lakes  (1)  become  aware  of  the  potential  or 
real  use  of  shorebirds  in  their  area  and  (2) 
manage  water  levels  for  shorebirds  whenever 
feasible. 


Fig.  1.  W'eeklv  counts  of  the  total  number  of  shorebirds 
at  four  sites  at  Lake  Lowell,  Canvon  Co.,  Idaho,  in  1989.  (A) 
New  York  Canal  Mouth  site,  with  both  total  number  of 
shorebirds  and  the  amount  of  mudflat  exposed.  (B)  Open 
circle  is  Public  Access  No.  1  site;  open  triangle  is  Public 
Access  No.  2  site;  vertical  line  is  Public  Access  No.  3. 


Stilt,  Greater  Yellowlegs,  Short-billed  Dow- 
itcher,  Wilsons  Phalarope,  and  Red-necked 
Phalarope,  along  with  the  Long-billed  Curlew, 
all  often  feed  in  water.  The  two  remaining  spe- 
cies with  similar-sized  peaks  between  large  and 
moderate  mudflats,  the  Killdeer  and  Spotted 
Sandpiper,  were  the  most  widespread. 

This  study  indicates  that  most  reservoirs  and 
lakes  in  Idaho  and  the  Intermountain  West  can 
provide  habitat  for  shorebirds  in  fall  migration 
if  they  have  moderate  to  large  mudflats  that  can 
be  exposed  by  water  drawdown  during  summer 
and  fall.  The  absence  of  shorebirds  at  some 
reservoirs  with  large  mudflats,  in  particular  Pal- 


ACKNOWLEDCMENTS 

We  would  like  to  thank  E.  Stone,  S.  Bailey, 
S.  Hart,  and  two  anonvmous  reviewers  for  their 
comments  on  earlier  drafts  of  this  paper  This 
studv  was  funded  in  part  by  the  Department  of 
Biological  Sciences,  Idalio  State  Universit}'. 

Literature  Cited 

BooK.s,  G.  G.  1985.  Avian  interactions  with  mid-Columbia 
River  level  fluctuations.  Northwest  Science  59:  304- 
312. 

Bl  KLF.ICH,  T.  D.  1972.  Birdsof  Idaho.  The  Caxton  Printers, 
Ltd.,  CiJdwell,  Idaho.  467  pp. 

CoLWELL,  M.  A.,  and  L.  W.  Ohinc;  1988.  Habitat  use  by 
breeding  and  migrating  shorebirds  in  soudicentral  Sas- 
katchewan. Wilson  Bulletin  100:  .554-566. 

H.MNLINE.  J.  L.  1974.  The  distribution,  migration,  and 
breeding  of  shorebirds  in  western  Nevada.  Unpub- 
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lt\ND,  R.  L.  19.32.  Notes  on  the  occurrence  of  water  and 
shorebirds  in  the  Lochsa  region  of  Idaho.  Condor  34: 
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184 


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H\RRisoN,  B.  A.  1982.  Untviiig  the  enigma  of"  the  Red 
Knot.  Living  Bird  Quarterly  1:  4-7. 

MlHUC,  J.  1991.  An  experimental  study  of  tlie  impact  ot 
shorebird  predators  on  benthic  invertebrates  in  Amer- 
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MONDA,  M.  J.,  iuid  J.  D.  Reichel  1989.  Aviiui  connnunitv 
changes  following  Lower  Granite  Dam  construction  on 
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Myers.  J.  R,  R.  I.  G.  Morrison.  R  Z.  Antus.  B.  A.  Har- 
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for  migratorv  shorebird  species.  American  Scientist  75: 
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RUNDLK  VV.  D.,  and  L.  H.  Fredrickscjn  1981.  Miuiaging 
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Received  15  September  1991 
Accepted  1  May  1992 


Great  Basin  Natuidist  52(2),  pp.  185-188 

DISPERSAL  OF  SQUARROSE  KNAPWEED 

{CENTAUREA  VIRGATA  SSP  SQUARROSA) 

CAPITULA  BY  SHEEP  ON  RANGELAND  IN  JUAB  COUNT\',  UTAH 

Cind\  Talbott  Roc-he  ',  Ben  F.  Roche,  Jr.  ,  and  G.  Allen  Rasniu.s.sen 
Key  words:  Centaurea  \irgata  ssp.  s(|uarrosa,  sciiiarrosc  knapweed,  weed  dispersal.  ranf^eUnid  weeds,  wool,  sheep. 


Among  Centaurea  species  naturalized  in 
western  North  America,  squarrose  knapweed 
(Centaurea  virgata  Lam.  ssp.  sqiiarrosa  Gugl.) 
has  a  unique  dispersal  mechanism.  The  seeds 
(achenes)  of  other  CentourtY/  species  (C.  diffusa 
Lam.,  C.  maculosa  Lam.,  C.  solstitialis  L.,  C. 
jacea  L.  x  C  nigra  L.)  disperse  either  as  indi- 
viduals with  crop  seed,  vehicles,  and  gravel,  or 
as  branches  or  entire  plants  moved  by  wind  or 
vehicles,  or  in  hay.  Squarrose  knapweed  involu- 
cral  bracts  recurve  or  spread  outward  with  a 
short  tenninal  spine  about  1-3  mm  long.  The 
entire  head  (capitulum)  is  deciduous  via  an 
abscisson  laver  at  the  base  of  the  capitulum. 
Thus,  the  capitula  of  squarrose  knapweed  func- 
tion like  burs  clinging  to  passing  animals  as 
l)urdock  {Arctium  minus  (Hill)  Bemh.),  cockle- 
bur  {Xantliium  strumarium  L.),  or  buffalobur 
{Solanum  rostratnm  Dunal).  Soon  after  the  dis- 
covery of  squarrose  knapweed  in  California 
(1950)  and  in  Utah  (1954),  its  occurrence  was 
linked  to  the  practice  of  trailing  rangeland  sheep 
from  one  area  to  another  (Bellue  1954,  Tingey 
1960).  On  Utah  rangeland  squarrose  knapweed 
is  more  abundant  along  sheep  trails  and  on 
bedgrounds  than  in  other  areas  (H.  Gates  and 
T  Roberts,  personal  communication).  Wool  is 
idealK  suited  to  catching  and  holding  the 
burlike  capitula,  but  squarrose  knapweed  along 
trails  and  in  sheep  bedgrounds  may  have  been 
carried  by  vehicles  or  other  means  and  estab- 
lished in  soil  disturbed  b\'  sheep.  The  objective 
of  this  study  was  to  determine  if  the  distribution 
of  squarrose  knapweed  in  Utah  is  due  to  seed 
carried  in  the  wool  of  rangeland  sheep. 


Methods  and  Materials 


In  mid-April  1990,  sheep  examined  in  this 
study  were  trailed  from  winter  range  west  of 
Tintic  Junction,  Juab  Comity,  Utali,  and  sheared 
before  being  mo\'ed  to  spring  range.  We 
received  permission  from  the  owoiers  to  collect 
wool  samples  during  shearing  of  a  band  that  had 
wintered  on  rangeland  known  to  have  squarrose 
knapweed.  We  had  predicted  that  sheep  would 
pick  up  the  "burs'"  by  lying  on  or  brushing 
against  knapweed  plants  growing  on  their 
bedgrounds.  However,  we  saw  no  obvious  knap- 
weed capitula  in  bellv  wool  or  on  the  sides  of 
sheep  being  sheared.  One  shearer  pointed  out 
several  ewes  with  a  profusion  of  kiiapweed 
capitula  around  their  faces  and  on  top  of  their 
heads  (Fig.  1).  We  then  collected  samples  of 
topknot  wool  (that  shorn  from  the  top  of  the 
head)  from  458  randomly  selected  white  ewes 
from  a  band  of  approximately  2500  ewes  at  the 
Jericho  shearing  station  in  Juab  Count);  Utah. 
Black  ewes  were  not  sampled.  Samples  from 
individual  ewes,  averaging  10  g,  were  kept  sep- 
arate in  small  plastic  bags.  Squarrose  knapweed 
capitula  were  sorted  bv  hand  from  each  sample, 
and  the  number  of  achenes  per  capitulum  was 
recorded.  Filled  achenes  (hard,  plump,  dark  tan 
or  browni  achenes)  and  light  aclienes  (.softer, 
flatter,  pale  tan  or  whitish  achenes)  were 
recorded  separately.  Presence  or  absence  of 
insect  o^AhiUropJiora  ajfinis  Frauenfeld  and  U. 
(juadrifasciata  [Meigen])  in  the  knapweed 
capitula  was  noted. 

Achene  viabilit)- was  determined  with  germi- 
nation trials  nm  for  10  da\s  at  20  C,  12  hours 


^Department  of  Natural  Resource  Sciences.  Washington  State  University.  Pullman,  Washington  99164-6410. 
"Present  address:  Department  of  Plant.  Soil,  and  Entomological  Sciences,  University  of  Idaho,  Moscow.  Idaho  8.3843. 
'  Department  of  Range  Science.  Utah  State  University,  Logan.  Utah  84.322-5230. 


185 


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Great  Basin  Naturalist 


[Volume  52 


Fig.  1.  Numerous  squarrose  knapweed  capitula  were  caught  as  burs  in  the  topknot  wool  of  sheep  that  Iiad  wintered 
where  squarrose  k-napweed  occurred  on  rangekind  in  Juab  Count\',  Utah. 


T.^BLE  1.  Proportion  ol  capituki  containing  0-6  aclienes 
per  capituhnn,  comparing  all  capitula  from  iui  intact  plant 
with  sheep-gathered  capitula  removed  from  topknot  wool, 
in  Juab  Count)',  Utah. 


Achenes/capituluni      Intact  pkuit 

'7c 


Extracted  from  wool 


0 

14 

75 

1 

12 

IS 

2 

19 

6 

3 

35 

1 

4 

17 

trace 

5 

3 

0 

6 

trace 

0 

light  alternating  with  12  hours  dark.  Seeds  were 
placed  in  germination  bo.xes  on  wetted  blotter 
paper.  Filled  and  light  achenes  were  tested  sep- 
arately. We  germinat(Hl  30  filled  achenes  in  four 
replications  in  each  of  two  trials.  Two  trials  of 
light  achenes  were  run  with  20  seeds  in  each  of 
two  replications. 

In  August  19<S9,  a  scjuarrose  knapw  eed  plant 
with  all  ol  its  capitula  was  collected  in  a  bag.  We 
dissected  the  capitula  and  recorded  the  number 
of  achenes  per  capitulum.  These  \alues  were 


compared  to  capitula  and  achenes  found  on 
sheep. 

Results 

We  determined  that  sheep  on  rangeland 
infested  with  squarrose  knapweed  picked  up 
and  carried  its  burhke  capitula.  Squarrose  knap- 
weed capitula  were  present  in  topknot  wool 
samples  from  73%  of  the  ewes.  A  total  of  2469 
knapweed  capitula  were  reco\ered  from  the  458 
ewes,  an  average  of  5.5  capitula  per  10  g  wool. 
Most  capitula  were  on  the  wool  surface, 
although  a  few  were  embedded  deeply  and 
appeared  to  have  been  there  longer  as  the\"  were 
satmated  with  lanolin  and  spines  had  worn  off 
the  in\()lucral  bracts. 

Sevent)'-five  percent  of  the  sheep-gathered 
capitula  were  barren,  compared  with  14%  of  the 
capitula  produced  on  a  whole  plant  (Table  1). 
()iil\-  49%  of  the  wool  samples  that  contained 
capitula  had  one  or  more  achenes.  Barren  capit- 
ula in  this  study  were  not  the  result  of  biocontrol 
insects  because  we  foinid  no  insect  galls. 

The  nimiber  of  knapweed  capitula  on  sheep 


1992] 


Notes 


18' 


<  V' 


•/>" 


Fig.  2.  Squarrose  knapweed  phuits  along  the  sheep  trails 
west  ot  the  Jericho  she;iring  station  were  grazed  in  mid-April 
1990.  A  few  capitula  remain  on  the  npper  right  side  of  the 
plant. 

heads  would  lead  a  casual  obsener  to  couclude 
that  the  sheep  carty  more  achenes  than  we 
found  by  dissecting  the  capitula.  Among  all  ewes 
sampled,  only  36%  carried  achenes  in  the  sam- 
pled topknot  wool.  These  seed-carriers  aver- 
aged 4.5  filled  achenes  per  10  g  wool.  Those 
filled  achenes  averaged  69%  germination.  In 
addition  to  the  filled  achenes,  5%  of  the  light 
achenes  germinated.  Light  achenes  composed 
only  23%  of  the  total  numbc^r  of  achenes. 


Discussion 

Sheep  carried  squarrose  knapweed  capitula 
but  not  as  many  achenes  as  the  ninnber  of 
capitula  woidd  indicate  if  the  proportion  were 
the  same  as  that  estimated  in  August.  This  find- 
ing could  indicate  one  of  two  conditions:  ( 1 )  the 
capitula  were  picked  up  in  late  winter  or  early 
spring,  when  only  the  lighter  capitula  remained 
on  the  plants,  or  (2)  some  achenes  were  lost 
from  capitula  lodged  in  the  wool  during  late 
summer  or  fall.  In  late  summer  heavier  capitula 
are  more  easil\-  dislodged  from  plants  than  are 
the  lighter  capitula.  Capitula  do  not  open  wideK' 
at  maturity-;  instead,  achenes  sift  out  throush  a 


small  opening  created  as  the  dried  flowers  fall 
from  the  capitulum.  The  proportion  of  empt)' 
capitula  increases  with  time  following  maturity 
as  plants  are  shaken  b\-  wind,  animals,  or  \ehicles. 

Sheep  acquired  knapweed  capitula  in  a 
manner  different  from  what  we  had  predicted. 
Although  some  capitula  clung  to  sheep  brushing 
against  plants  or  King  upon  them,  the  numerous 
knapweed  capitula  in  the  wool  aroiuid  their 
faces  suggest  that  ewes  searched  out  squarrose 
knapweed  as  a  food  source.  We  observed  that 
scjuarrose  knapweed  plants  along  the  sheep 
trails  had  been  grazed  (Fig.  2).  This  relationship 
was  nuitually  beneficial  for  knapweed  and 
sheep,  providing  propagule  dispersal  for  the 
knapweed  and  nourishment  for  the  sheep. 

Previousl)  reported  to  be  poor  forage 
(Tingev  1960),  squarrose  knapweed  rosette 
leaves  may  be  an  excellent  source  of  protein  in 
late  winter  and  early  spring.  Nutrient  content  of 
spotted  knapweed  rosette  leaves  is  comparable 
to  native  forage  plants  with  9-18%  crude  pro- 
tein (Kelsey  and  Mihalovich  1987).  Similar 
values  have  been  obtained  for  diffuse  knapweed 
and  yellow  starthistle  rosette  leaves  (Roche, 
unpublished  data).  In  the  stud\'  area,  Septem- 
ber 1989  through  Mav  1990  was  unusualK'  dry 
(Utah  State  University'  Tintic  research  site 
weather  station,  unpublished  data),  and  the 
normal  growth  of  cheatgrass  {Bromus  tectorum 
L.)  was  not  present  on  the  winter  range. 
Squarrose  knapweed,  a  deep-rooted  perennial 
forb,  was  one  of  the  few  plants  exhibiting  new 
growth  at  the  time  sheep  would  normalK  forage 
on  cheatgrass. 

Although  we  found  that  sheep  carr>- 
squarrc:>se  knapweed  seeds  as  they  move  across 
rangeland,  they  are  by  no  means  the  only  dis- 
persal mechanism  for  squarrose  knapweed. 
Other  animals,  both  domestic  and  wild,  may 
carry  knapweed  seeds.  In  addition,  these 
rangelands  are  hea\il\-  used  b\-  off-road  \ehicle 
recreationists.  Mining  traffic,  railroad  acti\it\'. 
and  militar\'  maneuxers  are  important  in  certain 
areas.  Hunters,  rockhounds,  and  other 
recreationists  also  frequent  the  area. 

Shearing  limits  the  dispersion  of  scjuarrose 
knapweed  b\- sheep.  It  is  unlikeK  that  knapweed 
achenes  remained  on  sheep  after  shearing. 
These  ewes  had  not  yet  lambed,  and  so  all  sheep 
in  this  band  left  the  knapweed-infested  winter 
range  shorn  of  seeds.  Seeds  in  the  wool  are 
remox  ed  at  the  woolen  mill,  which  has  been  one 
of  the  fates  of  squarrose  knapweed  seed  for 


188 


Great  Basin  Naturalist 


[Volume  52 


centuries,  as  evidenced  by  squarrose  knapweed 
found  at  Juvenal  Gate,  a  woolen  mill  in  France 
where  imported  wool  was  washed  for  200  years, 
beginning  in  1686  (TheUung  1912). 

Acknowledgments 

This  study  was  made  possible  by  the  cooper- 
ation of  H.  Gates  and  T.  Roberts  (Bureau  of 
Land  Management),  S.  Dewey  (Utah  State  Uni- 
versity), and  the  ranchers  who  permitted  us  to 
sample  wool  during  their  shearing  operation.  J. 
Miller,  Universit)'  of  Idaho,  was  consulted  con- 
cerning vegetable  matter  in  wool.  E.  Evans  and 
D.  Scamecchia  reviewed  the  manuscript  and 
provided  valuable  suggestions. 

The  project  was  fimded  in  part  by  the 
Renewable  Resources  Extension  Act  through 
Washington  State  University  Cooperative 
Extension. 


Literature  Cited 

Bkllue,  M.  K.  1952.  Virgate  stiir  thistle,  Ccntaurca  virgata 
vai".  squarrosa  (Willd.)  Boiss.  in  California.  California 
Dep;xrtnient  of  Agriculture  BuOetin  41:  61-63. 

Kelsey,  R.  C,  and  R.  D.  Mhialovich.  1987.  Nutrient 
composition  of  spotted  knapweed  {Centatirea 
maculosa).  Journal  of  Range  Management  40:  277- 
281. 

Roche,  C.  T,  and  B.  F.  Roche,  Jr  1989.  Introductory- 
notes  on  squarrose  knapweed  (Centatirea  virgata  Lam. 
ssp.  squarrosa  Gugl.).  Northwest  Science  63: 246-2.52. 

Thellung,  a.  1912.  La  flore  adventice  de  MontpelLier. 
Memoires  de  la  Societe  Naturelles  et  Mathematiques 
de  Cherbourg  38:  57-728. 

TiNGEY,  D.  C.  1960.  Control  of  squarrose  knapweed.  Utixli 
State  University  Experiment  Station  Bulletin  No.  432. 
11  pp. 

Received  11  March  1991 
Accepted  31  March  1992 


Great  Basin  Natunilist  52(2).  1992.  pp.  189-193 

VEGETATION  ASSOCIATED  WITH  TWO  ALIEN  PLANT  SPECIES  IN  A  FESCUE 
GRASSLAND  IN  GLACIER  NATIONAL  PARK,  MONTANA 

R()l)in  W.  Tyser 


Ki'i/  icord.s:  alini  flora.  CJacicr  F<nk.  FesiwcA grasslands. 

The  presence  of  alien  flora  in  natunil  area 
grasslands  of  the  Great  Basin  and  surrounding 
areas  is  of  considerable  concern,  given  the  wide- 
spread success  of  alien  flora  and  associated 
decline  of  nati\e  species  in  this  region  (Young et 
al.  1972,  Mack  1986,  1989).  Suiveys  of  indige- 
nous bunchgrass  communities  in  northern 
Roclcs  Mountain  national  parks  have  detected 
the  occurrence  of  several  alien  plant  species 
(Koterba  and  Ilabeck  1971,  Stringer  1973, 
Weaver  and  Woods  1985,  1986,  Tvser  and 
Worley  1992).  In  addition,  alien  species  have 
commonK'  been  used  to  revegetate  human-dis- 
turbed sites  such  as  roadsides  and  housing  areas 
in  national  parks.  Livestock-related  introduc- 
tion of  Eurasian  pasture  grasses  by  private  out- 
titters  is  also  known  to  have  occurred  (Glacier 
National  Park  Records  1939).  Ho\ve\er,  in  spite 
of  these  observations  and  practices,  the  effects 
of  alien  vegetation  in  natural  area  grasslands  of 
this  region  remain  poorly  studied. 

This  study  compares  the  indigenous  \ascular 
flora  and  crvptoganiic  ground  cover  associated 
with  two  cdien  species,  Centoiirea  ituicidosa 
Lam.  (spotted  knapweed)  and  Phleiim  pratense 
L.  (common  timothv),  that  ha\'e  in\'aded  a 
fescue  grassland  in  Glacier  National  Park,  Mon- 
tana. C.  nmcitlosa,  now  a  noxious  rangeland 
imader  throughout  the  Pacific  Northwest 
(Watson  and  Renney  1974,  Lacey  1989),  was 
first  detected  in  the  park  in  the  mid- 1960s  (R. 
Wassem,  personal  communication).  Earlier 
obserxations  have  shown  that  this  species  is 
expanding  into  grasslands  adjacent  to  roadsides 
in  the  park  and  reducing  species  richness  (Tyser 
and  Key  1988).  The  impact  of  C.  nuiculoso  on 
the  cr\ptogamic  ground  crust — of  potential 
importance  in  soil  stabilization,  moisture  reten- 
tion, and  nitrogen  fixation  (Rvchert  and  Skujins 


1974,  Anderson  et  al.  1982,  Brotherson  and 
Rushforth  1983) — has  not  yet  been  considered, 
nor  has  the  impact  of  C.  maculosa  been  com- 
pared to  that  of  other  alien  species.  P.  pratense 
is  widelv  distributed  throughut  the  park  s  grass- 
lands. Unlike  C.  maculosa,  this  species  appears 
to  have  been  intentionally  seeded  in  grasslands 
by  outfitters  before  the  1940s  and  along  road- 
sides by  park  personnel  before  the  198()s  (D. 
Lange  and  J.  Potter,  personal  comnumication). 

Study  Site  and  Methods 

The  ca  10-ha  stud\-  area,  located  adjacent  to 
Going-to-the-Sun  Highway  in  the  St.  Mary 
drainage  of  Glacier  National  Park,  Montana,  is 
fairly  topographically  homogeneous,  i.e.,  clearK' 
defined  drainage  channels  are  absent,  and 
slope,  exposure,  and  substrate  texture  are  rela- 
tively uniform.  The  study  area  includes  a  large 
(ca  5  ha),  irregularly  shaped  stand  dominated  bv 
PJileum  pratense  and  a  small  (ca  0.1  ha)  stand 
adjacent  to  the  roadside  ditch  dominated  b\ 
Centaurea  maculosa.  The  Centaurea  stand 
extends  >20  m  away  from  the  ditch  and  is  not 
part  of  the  road-cut  corridor.  The  remaining 
portion  of  the  studv  site  is  composed  of  a  stand 
of  natixe  Festuca  grasses  and  associated  \egeta- 
tion  in  which  inxasion  by  alien  species  has  been 
minimal.  Though  no  homesteading  is  known  to 
ha\  e  occurred  in  the  .studv  area  before  establish- 
ment of  the  park  in  1910,  this  area  was  likel\ 
used  as  summer  pasture  for  concession  trail 
horses  from  approximately  1915  to  1940  (B. 
Fladmark,  personal  communication).  The  study 
area  has  not  been  used  for  stock  grazing  since 
that  time.  Othenvise,  there  is  no  indication  that 
any  of  the  areas  sampled  in  the  three  stands  have 
been  subjected  to  anthropogenic  disturbance 


Department  of  Biolog\  and  MicTol)iolog\,  I'niversitv  of  Wisconsin-La  Oosse,  La  Crosse.  Wisconsin  546()L 


189 


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Great  Basin  Naturalist 


[Volume  52 


since  the  park  was  established.  In  addition,  no 
fires  have  been  recorded  in  or  near  the  study 
area  since  1910.  A  cnptogani  ground  layer  com- 
posed of  small  lichens  and  mosses  covering 
undisturbed  soil  surfaces  is  commonly  present. 
Qualitative  observation  suggests  that  mosses  are 
the  dominant  element  in  this  layer.  Mean  annual 
precipitation  in  the  study  area  is  ca  65  cm 
(Finklin  1986). 

In  each  stand,  vegetation  was  sampled  using 
20  X  50-cm  quadrat  frames  along  two  transects 
placed  in  representative  areas.  Within  each 
quadrat,  presence  of  all  vascular  species  was 
determined,  and  the  canopy  cover  of  each  vas- 
cular species  and  the  surface  cover  of  the  cry|3- 
togamic  ground  crust  were  estimated  to  the 
nearest  percentage.  A  stratified  random  sam- 
pling procedure  was  used  in  which  quadrats 
were  randomly  placed  along  transect  segments 
of  fixed  length.  For  the  Centaurea  stand,  tran- 
sects were  20  m  long,  and  one  quadrat  was 
randomly  placed  within  each  2-m  segment  (N  = 
20  quadrats).  For  the  Plilcmn  and  Festuca 
stands,  transects  were  100  m  long,  and  one 
quadrat  was  randomly  placed  within  each  5-m 
segment  (N  =  40  quadrats  per  stand). 

Five  vegetation  measures  were  calculated 
for  each  individual  quadrat:  (1)  vascular  species 
cover  diversity  using  the  Shannon -Wiener  index 
(H'  =  -S  Pi  log  p,),  (2)  vascular  species  richness, 
(3)  cumulative  canopy  cover  of  native  forb  spe- 
cies, (4)  cumulative  canopy  cover  of  native  grass 
species,  and  (5)  surface  cryptogam  cover.  One- 
way ANOVAs  were  used  to  assess  among-stand 
differences  for  each  of  these  quadrat  measures. 
With  the  exception  of  the  diversity  measures, 
data  did  not  meet  parametric  assumptions 
(normal  distributions,  homogeneous  variances) 
and  could  not  be  transformed  using  standard 
logarithmic,  arcsine,  or  square  root  transforma- 
tions. Therefore,  data  were  analyzed  by  the 
Kruskal-Wallis  nonparametric  one-way 
ANOVA  procedure  as  described  by  Conover 
and  Iman  (1983).  The  Tukey  multiple  compari- 
son procedure,  applicable  to  both  parametric 
and  nonparametric  ANOVAs  (Conover  and 
Iman  1981),  was  used  to  make  pair-wise  among- 
stand  comparisons.  Species  nomenclature  fol- 
lows that  of  Hitchcock  and  C^rontjuist  (1973). 

Results  and  Discussion 

Prominent  graminoid  and  forb  species  in  the 
Festuca  stand  included  Achillea   millefolium. 


Carex  spp.,  Festuca  idahoensis,  F.  scabrella, 
Gaillardia  aristata,  and  Lupinus  sericeus  (Table 
1).  Species  composition  of  this  stand  was  similar 
to  prairie  communities  described  elsewhere  in 
the  Pacific  Northwest,  e.g.,  the  Festuca 
scabrella/F.  idahoensis  association  of  western 
Montana  (Mueggler  and  Stewart  1980),  the 
Festuca/Danthonia  prairie  of  Alberta  (Stringer 
1973),  and  the  Washington  Palouse  prairie 
(Daubenmire  1970).  Estimated  surface  cover  of 
the  cryjDtogam  layer  in  this  stand  was  relatively 
high,  characteristic  of  western  bunchgrass  prai- 
ries (Daubenmire  1970,  Mack  and  Thompson 
1982).  Three  alien  species  were  sampled  within 
the  Festuca  stand,  though  total  cover  of  these 
species  was  <1.0%. 

Significant  among-stand  variation  occurred 
for  all  community  measures  (Table  2).  Each  of 
these  measures  was  lowest  in  the  Centaurea 
stand.  Canopy  cover  of  native  forbs  and  crypto- 
gam ground  cover  in  this  stand  were  particularly 
low,  only  ca  18%  and  4%,  respectively,  of  the 
corresponding  Festuca  stand  measures.  Thus, 
effects  of  the  Centaurea  macidosa  invasion  on 
the  native  flora  in  the  study  site  appear  to  be 
relatively  severe.  The  impact  of  this  species  is 
even  more  impressive  considering  its  relatively 
recent  entry  into  the  park. 

All  but  one  of  the  Phleum  stand  measures 
were  significantly  lower  than  those  of  the 
Festuca  stand  (Table  2).  Canopy  cover  by  native 
graminoids  in  the  Phleum  stand  was  reduced  to 
about  50%  of  its  level  in  the  Festuca  stand. 
However,  forb  cover  differences  between  these 
two  stands  were  not  statisticiilly  significant 
(Table  2).  Three  species  {Achillea  millefolium, 
Agoseris  glauca,  and  Lupinus  sericeus)  were 
among  the  four  forb  species  with  highest  canopy 
cover  in  each  stand,  suggesting  that  the  forb 
components  of  these  two  stands  were  relatively 
similar.  These  observations  suggest  that  Phleum 
invasion  has  affected  natixe  graminoids  more 
than  native  forbs.  It  should  also  be  noted  that 
while  mean  quadrat  richness  was  lower  in  the 
Phleum  stand  (Table  2),  eight  more  species  were 
recorded  in  this  stand  than  in  the  Festuca  stand 
(N  =  59  vs.  N  =  51;  see  Table  1).  Thus,  different 
Phleum  vs.  Festuca  richness  patterns  may  occur 
if  comparisons  are  derived  from  sampling  units 
larger  than  the  0.1-m~  quadrats  used  in  this 
study. 

Cryptogam  cover  in  the  Phleum  stand  was 
approximately  50%  lower  than  in  the  Festuca 
stand  (Table  2).  Mack  and  Thompson  (1982) 


1992] 


Notes 


191 


Table  1.  Canopv  cover  (%)  estimates  of  six^cies  within  the  Fcstuca,  Phlcitin,  and  Ccnidurcd  stands.  °  =  iJien  sjieeies. 


Species 

Festuca 

Phleuni 

Centaurea 

GlUMINOIDS 

Agropijron  caninum 

0.4 

0.6 

Agroptjwn  spicatum 

0.3 

0.3 

Carex  spp. 

12.3 

5.6 

9.3 

Danthonia  intcmivdia 

4.2 

0.9 

Fcstuca  iclaliocnsi.s 

9.2 

4.3 

0.2 

Fcstuca  saibrclla 

7.1 

4.1 

2.1 

Hclictotrichon  hookcri 

0.9 

<0.1 

Kitclcria  crlstata 

1.4 

0.4 

<0.1 

Flileum  pratense' 

0.2 

38.4 

0.7 

Fodjuitcifolui 

<0.1 

Poa  pratcnsis' 

<0.1 

0.9 

1.0 

Stipa  occidentalis 

3.7 

2.1 

Stipa  rirhnrrlxonii 

0.1 

0.8 

FOKBS 

Achillea  millefoliuiu 

11.7 

8.6 

0.8 

Agoscri.'i  glaiica 

4.0 

4.3 

Allium  cenmiim 

0.1 

<0.1 

Atnehmchicr  alnifolid 

0.3 

0.9 

0.5 

And  rosace  septenthoiidli.s 

1.0 

0.3 

Anemone  midtifida 

1.4 

1.0 

<0.1 

Antennaha  inicroplujlld 

O.S 

0.3 

1.7 

Arahis  <^lahra 

<0.1 

Arahis  nuttallii 

0.2 

0.1 

<0.1 

Arctostaplujlos  u vd-tt rsi 

0.4 

0.2 

Aster  laevis 

1.8 

0.9 

Berheris  repens 

0.1 

0.6 

0.3 

Campanula  rotundifolid 

0.5 

1.0 

<0.1 

Castilleja  aisickii 

0.3 

<().! 

Centaurea  nmculosa' 

62.0 

Cerastiu m  arven.se 

4.0 

3.1 

0.7 

Colloniia  linearis 

<0. 1 

Comandra  nmtjeltata 

0.5 

0.3 

Species 

Festuca 

Phleum 

Centaurea 

Epilohium  angustifoliu m 

0.5 

F,rigeron  suhtrinervi.s 

1.5 

Erysimum  inconspicuum 

0.3 

Fru'^a ri a  v i r^i niana 

<0.1 

0.7 

Gaillardia  aristata 

1.9 

0.6 

<0.1 

Galium  horeale 

0.6 

1.8 

0.2 

Gentiana  amarella 

1.3 

0.7 

Geran iu m  viscosissimum 

<0.1 

1.2 

Hedijsanun  horeale 

0.5 

Heuehera  cijlindhca 

0.1 

0.2 

0.2 

Hieracium  umhellatum 

0.2 

Jiinctts  haltieus 

1.0 

Latlujriis  oehroleucus 

0.2 

Lithospermu  m  niderale 

1.9 

3.9 

0.7 

Lonmtium  tritematum 

1.0 

2.4 

0.3 

Ltipinus  serieeus 

5.6 

6.0 

<0.1 

Monarda  fistulosa 

0.6 

Orthocarpus  tenuifolius 

1.2 

<().! 

Oxijt  n  )p  is  cam  pest  ris 

2.8 

0.9 

Oxtjtropi.s  splendens 

0.3 

Penstenum  confertus 

0.8 

1.9 

0.7 

Potentilla  arguta 

<0.1 

1.1 

Potentilla  is^racilis 

<0.1 

0.4 

0.3 

Potentilla  hippiana 

0.5 

Pninus  vir<i^iniana 

0.1 

Fdnnanthus  cri.Hta-<icdli 

0.9 

0.4 

Rosa  woodsii 

1.3 

2.3 

0.2 

Silene  pamji 

0.4 

0.1 

<0.1 

Sisijrinchium  an^iiistifolium   0.2 

0.4 

Soliddfio  missouriensis 

1.6 

1.8 

0.2 

Taraxacum  officinale ° 

0.2 

1.4 

0.3 

Traoopogon  duhius  ° 

<0.1 

0.4 

Vicia  americana 

1.6 

1.0 

Zioadcnus  venenosus 

<0.1 

Tablk  2.  Among-stand  compari.sons  of  quadrat  means  for  five  vegetation  measures.  N  =  40,  40,  antl  20  cjnadrats, 
resjDectively,  for  the  Festuca,  Phleum,  and  Centaurea  stands. 


H' 


Richness 


Native 
graminoids 


Native 
forbs 


Cryptogam 
crust 


Festuca " 

Phleum 

Centaurea 

F2.97 

P 


0.966" 
0.872'' 
0.385'' 
90.084 
<.001 


14.8-' 
12.9'' 

7.2' 
41.1.50 

<.001 


.39.. 5^' 
19.2*' 
11.6' 
.53.807 
<.001 


48.4" 

55.  r' 

8.8'' 
40.896 
<.001 


28.^ 
15.1'' 

1.3'' 
31.835 
<.001 


°\\  ithin  each  vegetation  me;Lsurt-.  means  with  clitferf  nt  letters  differ  significantl\'  Fn 


otlier  (P  <  ,05,  Tiikey  multiple  conipi 


suggest  that  the  extensive  rhizome-tiller  mats  of 
Eurasian  grasses  limit  cryptogam  colonization 
sites,  which  may  account  for  the  reduced  cryp- 
togam cover  observed  in  the  Phleum  stand.  A 
large  elk  herd  overwinters  in  the  St.  Mar\' \alley 
grasslands  in  which  the  study  area  was  located 
(Martinka  1983).  Thus,  it  is  possible  that  elk 
trampling/grazing  may  reduce  cryjotogam  cover 
and  facilitate  Phleum  invasion. 


The  role  plaved  bv  pre- 1 940  horse  grtizing  in 
the  occurrence  of  Phleum  in  the  study  site 
cannot  be  assessed.  However,  the  prominence 
of  this  species  some  50  years  after  the  cessation 
of  horse  grazing  ck)es  indicate  that  ongoing  live- 
stock griizing  is  not  necessarv  for  its  persistence. 
The  more  recent  Centaurea  maadosa  invasion 
in  the  study  site  and  in  other  fescue  grasslands 
in  the  park  (Tyser  and  Key  1988,  Tyser  and 


192 


Great  Basin  Naturalist 


[Volume  52 


Worley  1992)  suggests  that  livestock  grazing  is 
not  a  prerequisite  for  successful  invasion  of  nat- 
ural areas  by  this  species.  The  success  of  both  P. 
pratense  and  C.  maculosa  in  the  study  site  sug- 
gests that  mechanisms  proposed  for  the  success 
of  alien  flora  in  agro-systems,  e.g.,  rapid  coloni- 
zation of  disturbed  sites,  structural  and  physio- 
logical adaptations  to  grazing  and  trampling, 
and  low  piilataliilit}'  (Mack  and  Thompson  1982 
and  references  therein,  Lacey  et  al.  1986, 
Locken  and  Kelsey  1987,  Kelsey  and  Bedunah 
1989),  may  also  operate  in  natural  area  systems. 
In  addition  to  overwintering  elk,  diggings  of 
other  native  herbivores,  especially  ground 
squirrels  (Sperniopliilus  Columbia  mis),  were 
common  throughout  the  study  area.  At  any  rate, 
though  the  status  and  impacts  of  C.  maculosa 
and  P.  pratense  require  additional  research,  this 
study  shows  that  the  potential  effects  of  these 
species — particularlv  that  of  C.  maculosa — in 
natural  area  bunchgrass  prairies  need  to  be  seri- 
ously contemplated. 

Reduction  of  Plileum  pratense  is  not  a  real- 
istic option  in  Glacier  National  Park  or  other 
natural  areas  in  which  this  species  is  now  widely 
established.  Perhaps  the  most  reasonable  rec- 
ommendation for  this  species  and  other  Eura- 
sian grasses  is  simply  that  resource  managers  not 
use  these  species  for  revegetation  (see  also 
Wilson  1989).  Centaurea  maculosa,  though 
potentially  more  ecologically  disruptive  than  P. 
pratense,  is  at  an  earlier  stage  of  invasion  in  the 
park  and  probabK'  in  other  natural  areas  in  this 
region  as  well.  Thus,  the  fate  of  this  species  may 
yet  be  influenced  by  appropriate  resource  man- 
agement actions. 

Acknowledgments 

I  thank  Tom  Jacobsen  and  Andy  Tyser  for 
their  assistance  with  fieldwork,  Andy  Matchett 
for  statistical  advice,  and  Glaciers  research  and 
resource  management  staff  for  support  and 
assistance  with  this  stud\'.  The  study  was  funded 
by  a  grant  from  the  Universit\'  of  Wyomincj- 
National  Park  Service  Research  Center. 

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Notes 


193 


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194-201. 

Received  26  April  1991 
Accepted  16  April  1992 


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Mack,  G.  D.,  and  L.  D.  Flake.  1980.  Habitat  rela- 
tionships of  waterfowl  broods  on  South  Da- 
kota stock  ponds.  Journal  of  Wildlife  Man- 
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 
ecology  of  natural  disturbance  and  patch  dy- 
namics. Academic  Press,  New  York. 
Coulson,  R.   N.,  and  J.  A.  Witter.   1984,  Forest 
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Wiley  and  Sons,  Inc.,  New  York.  669  pp. 
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(ISSN  0017-3614) 

GREAT  BASIN  NATURALIST    voi  52  no  2  June  1992 

CONTENTS 

Articles 

Red  Butte  Canyon  Research  Natural  Area:  history,  flora,  geology,  climate,  and 

ecology James  R.  Ehleringer,  Lois  A.  Arnow,  Ted  Arnow, 

Irving  R.  McNulty,  and  Norman  C.  Negus        95 

Influences  of  sex  and  weather  on  migration  of  mule  deer  in  California 

Thomas  E.  Kucera      1 22 

Diatom  flora  of  Beaver  Dam  Creek,  Washington  County,  Utah,  USA 

Kurtis  H.  Yearsley,  Samuel  R.  Rushforth,  and  Jeffrey  R.  Johansen      131 

Stratification  of  habitats  for  identifying  habitat  selection  by  Merriam's  Turkeys 

Mark  A.  Rumble  and  Stanley  H.  Anderson      139 

Pollinator    preferences    for    yellow,    orange,    and    red    flowers    of   Mimulus 

verbenaceus  and  M.  cardinalis Paul  K.  Vickery,  Jr.      145 

Soil  loosening  processes  following  the  abandonment  of  two  arid  western  Nev- 
ada townsites Paul  A.  Knapp      149 

Biochemical  differentiation  in  the  Idaho  ground  squirrel,  Spennophilus  brun- 

neus  (Rodentia:  Scuridae) Ayesha  E.  Gill  and  Eric  Yensen      155 

New  genus,  Aplanusiella,  and  two  new  species  of  leafhoppers  from  south- 
western United  States  (Homoptera;  Cicadellidae:  Deltocephalinae) 

M.  W.  Nielson  and  B.  A.  Haws      160 

Summer  habitat  use  by  Columbian  Sharp-tailed  Grouse  in  western  Idaho  .  .  . 

Victoria  Ann  Saab  and  Jeffrey  Shaw  Marks      166 

Notes 

Characteristics  of  sites  occupied  by  subspecies  of  Artemisia  tridentata  in  the 

Piceance  Basin,  Colorado  .  .  Thomas  R.  Cottrell  and  Charles  D.  Bonham      1 74 

Use  of  lakes  and  reservoirs  by  migrating  shorebirds  in  Idaho 

Daniel  M.  Taylor  and  Charles  H.  Trost      1 79 

Dispersal  of  squarrose  knapweed  {Centaurea  virgata  ssp.  squarrosa)  capitula  by 

sheep  on  rangeland  in  Juab  County,  Utah Cindy  Talbott  Roche, 

Ben  E  Roche,  Jr.,  and  G.  Allen  Rasmussen      185 

Vegetation  associated  with  two  alien  plant  species  in  a  fescue  grassland  in  Gla- 
cier National  Park,  Montana Robin  W.  Tyser      189 


H       E 


MCZ 
LiijRARY 

V  mo 

HARVARD 
UNIVhHSlTY 


GREAT  BASIN 


MURAUST 


VOLUME  52  NO  3  -  SEPTEMBER  1992 


BRIGHAM    YOUNG    UNIVERSITY 


GREAT  BASIN  NATURALIST 

Editor 

James  R.  Barnes 

290  MLBM 

Brigham  Young  UniversiU' 

Provo,  Utah  84602 

Associate  Editors 


Michael  A.  Bovvers 

Blandy     Experimental     Fann,     University    of 

Virginia,  Box  175,  Boyce,  Virginia  22620 

J.  R.  Callxhan 

Museum  of  Southwestern  Biolog)',  Universit)'  of 

New  Mexieo,  Albuquerque,  New  Mexico 

Mailing   address:    Box   3140,    Hemet,    California 

92546 

Jeanne  C.  Chambers 

USDA   Forest    Ser\ice    Research,    University   of 

Ne\ada-Reno,  920  Valley  Road,  Reno,  Nevada 89512 

Jeffre\'  R.  Johansen 

Depiirtment  of  Biology,  John  Carroll  University, 

Universit)'  Heights,  Ohio  44118 

Paul  C.  Marsh 

Center  for  En\ironmental  Studies,  Arizona  State 

University;  Tempe,  Arizona  85287 


Brian  A.  Maurer 

Department  of  Zoolog);  Brigham  Young  Uni\ersity, 

Provo,  Utah  84602 

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 

Robert  C.  Whitmore 

Division  of  Forestry',  Box  6125,  West  \'irginia  Uni- 

versits',  Morgiuitowai,  West  Virginia  26506-6125 


Editorial  Board.  Richard \V.  Bauniann,  Ch;xirman,  Zoolog)';  H.  Duane  Sniitli,  Zoolog);  Cla\ton  M. 
White,  Zoologv;  Jerran  T.  Flinders,  Botany  and  Range  Science;  William  Hess,  Botany  anc{  Range 
Science.  AJl  are  at  Brigham  Young  University.  Ex  Officio  Echtorial  Board  members  include  Clayton  S. 
Huber,  Dean,  College  of  BiologiccJ  and  Agricultural  Sciences;  Norman  A.  Darais,  Director,  University 
Publications;  James  R.  Banies,  Editor,  Great  Basin  Naturalist. 

The  Great  Basin  Naturalist,  founded  in  1939,  is  published  quarterl\-  b\  Brigham  Young  Uni\ersit\'. 
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Copyright  ©  1992  b\  Brigluuii  Young  University 
Official  publication  date:  18  December  1992 


ISSN  0017-3614 
12-92  750  2473 


The  Great  Basin  Naturalist 


Plblishkd  atPhono,  Utah,  by 
Bricham  Young  Um\ersi'it 

ISSN  0017-3614 


Volume  52  September  1992  No.  3 

Great  Basin  Naturalist  52(3),  pp.  195-215 

PLANT  ADAPTATION  IN  THE  GREAT  BASIN  AND  COLORADO  PLATEAU 

Jonathan  P.  Comstock    antl  James  R.  Elileringer 

Al5STlU(X — Adapti\e  features  of  plants  of  tlie  Great  Basin  are  reviewed.  The  combination  of  cold  winters  and  an  arid 
to  semiarid  precipitation  regime  results  in  the  distinguishing  features  of  the  vegetation  in  the  Great  B;isin  and  Golorado 
Plateau.  The  priniaiy  effects  of  these  climatic  features  luise  from  how  the\  structure  the  hvdrologic  regime.  Water  is  die 
most  limiting  factor  to  plant  growth,  and  water  is  most  reliabK  axailahle  in  the  earl\-  spring  after  winter  recharge  of  soil 
moisture.  This  factor  determines  main  characteristics  of  root  moipholog\,  growth  phenolog\-  of  roots  and  slKX)ts,  and 
photos\ndietic  physiolog):  Since  winters  are  hpicallv  cold  enough  to  suppress  growth,  and  drought  limits  growth  during 
the  summer,  the  cool  temperatures  characteristic  of  the  peak  growing  .season  are  the  second  most  importiuit  climatic  factor 
influencing  plant  habit  luid  perform;uice.  The  combination  of  several  distinct  stress  periods,  including  low-temperature 
stress  in  winter  and  spring  and  high-temperature  stress  combined  with  drought  in  summer,  appears  to  have  limited  plant 
habit  to  a  greater  degree  thiui  found  in  the  warm  de.serts  to  the  south.  Nonetheless,  cool  growing  conditions  and  a  more 
reliable  spring  growing  season  result  in  higher  water-u.se  eiiiciencv  and  productiv  ih"  in  the  vegetation  of  the  cold  de.sert 
than  in  warm  deserts  with  equiv;ilent  total  rainfall  amounts.  Edapliic  factors  are  also  importimt  in  structuring  communities 
in  these  regions,  and  halophvtic  connnunities  dominate  main  landscapes.  These  haloph\-tic  communities  of  the  cold  desert 
share  more  sj^ecies  in  common  with  warm  deserts  than  do  the  nonsdine  communities.  The  Golorado  Plateau  differs  from 
the  Great  Basin  in  having  greater  amounts  of  smniner  rainfall,  in  some  regions  less  predictable  riiinfall,  sandier  soils,  and 
streams  which  drain  into  river  .systems  rather  than  closed  basins  and  salt  plavas.  One  result  ofthe.se  climatic  and  edapliic 
differences  is  a  more  important  summer  grov\ing  seasf)n  on  the  (Colorado  j'lati'au  and  a  sonu-wliat  <ii"eater  di\c'rsilication 
of  plant  habit,  phenolog),  and  physiolog)'. 

Key  icords:  cold  desert,  plant  adaptation,  water  stress,  phenalo'^ij.  salinitt/.  Great  Basin.  Colorado  Plateau. 

Several  features  arising  from  climate  and  Nevada  and  increase  both  to  the  north  and  ea,st, 

geolog)'    impose   severe   limitations   on   plant  and  to  the  southeast  moving  into  the  Colorado 

gro\\i:h  and  activit)^-  in  the  Great  Basin  and  Col-  Plateau  (Fig.  1,  Table  1).  The  fraction  of  annual 

orado  Plateau.  The  climate  is  distinctlv  conti-  precipitation  during  the  hot  sununer  months 

nental  with  cold  winters  and  warm,  often  dn  (|une-Se[)tember)    varies   considerabh;    from 

sununers.  Annual  precipitation  levels  are  low  in  l()-2()9f  in  northern  Nevada  to  30-40%  along 

the  basins,  ranging  from  100  to  300  mm  (4-12  the  boundaiAof  theCold  and  .\Iojave  deserts  in 

inches),  and  t)piciilly  increasing  with  elevation  southwestern  Nevada  and  southern  Utah,  and 

to  500  mm  (20  inches)  or  more  in  the  montane  35-50%  throughout  much  of  the  Colorado  Pla- 

zones.  Precipitation  levels  are  lowest  along  tlu^  t(^au.    Winter   jirecipitation    falls   primariK'   as 

southwestern  boundan'  of  the  Great  Basin  in  snow  in  the  Great  Basin  and  liii£her  elevations 


Department  of  Biologv-.  Universit)-  of  t'tali.  Salt  Lake  Cit^■.  Utiih  84 1 12. 


195 


196 


Great  Basin  Naturalist 


[\blume  52 


TaHI.K  1.  Srlec'tec!  climatic  data  tor  l(m-elc'\ation  sites  in  different  regions  of  the  Great  B;Lsin,  Moja\e  Desert,  and 
Colorado  Platean.  Viilues  are  based  on  a\erages  for  the  U.S.  Weather  Bureau  stations  indicated.  The  tliree  dixisions  of  the 
year  presented  here  reflect  ecologically  relevant  units,  but  are  unequal  in  length.  The  fixe  months  of  October-Februan 
represent  a  period  of  temperature-imposed  plant  dorm;uicy  and  winter  i-echarge  of  soil  moisture.  The  spring  mondis  of 
March-Mav  represent  the  potential  growing  period  at  cool  temperatures  immediately  follcnving  winter  recharge.  The 
summer  and  ei\r\\  fall  from  Jiuie  through  September  represent  a  potential  warm  growing  season  in  areas  with  sufficient 
summer  rain  or  access  to  other  moisture  sources. 


Map  # 

Weather        E 

levation 

Total  precipitation 

Mean 

temperature 

Region 

Annua 

1  Oct-Feb 

Mar-Ma\  J 

ini-Sep 

Annual  Oct-Fe 

'b  Mar-Ma\ 

Jiui-Sep 

(Fig.  1) 

station 

(m) 

(mm) 

(%) 

(%) 

(%) 

(°C) 

(°C) 

(°C)   ' 

(°C) 

Northern 

I 

Fort  Bidwell 

1370 

402 

63 

24 

13 

9.0 

3.0 

8.0 

17.3 

Great  Basin 

r> 

l^eno 

1340 

1S2 

61 

24 

15 

9.5 

3.3 

8.4 

18.0 

3 

Elko 

1547 

230 

52 

29 

19 

7.6 

0.1 

7.1 

17.5 

4 

Snowxille 

1390 

300 

43 

33 

24 

7.4 

0.7 

6.2 

18.4 

Southern 

5 

Sarcobatus 

1225 

85 

45 

22 

33 

13.5 

6.4 

12.5 

23.1 

Great  Basin 

rs 

C;iliente 

1342 

226 

47 

24 

29 

11.7 

4.1 

11.2 

21.5 

' 

Fillmore 

1573 

369 

44 

34 

•1-1 

11.0 

3.0 

10.0 

21.7 

Moja\e  Desfc 

'rt    S 

Trona 

517 

102 

70 

19 

11 

19.0 

11.3 

18.4 

29.0 

9 

Bea\erdam 

570 

169 

50 

23 

28 

18.3 

11.0 

16.9 

28.6 

Colorado 

10 

Hanksxille 

1313 

132 

36 

19 

45 

11.4 

2.1 

11.5 

22.8 

Plateau 

11 

Clrantl  Junction 

147S 

211 

39 

25 

36 

11.3 

2.4 

10.9 

22.9 

12 

Blanding 

1841 

336 

48 

19 

.33 

9.7 

2.1 

8.7 

19.9 

13 

TnbaCit\' 

1504 

157 

38 

21 

41 

12.6 

4.8 

12.0 

22.8 

14 

(]haco  Canvon 

1S67 

220 

35 

20 

45 

10.3 

2.6 

9.4 

20.6 

of  the  Colorado  Plateau,  which  is  thought  to  be 
a  critical  feature  ensiuins  soil  moisture  recharge 
and  a  reliable  spring  growing  season  (West 
1983,  CakKvell  1985,  Dobrowolski  et  al.  1990). 
During  the  winter  period,  precipitation  is  gen- 
eralK'  in  excess  of  potential  exaporation,  but  low 
temperatures  do  not  permit  gro\\1:h  or  photo- 
sxnthesis,  and  exposed  plants  may  experience 
shoot  desiccation  due  to  dry  winds  and  frozen 
soils  (Nelson  and  Tienian  1983).  Strong  winds 
can  also  cau.se  major  redistributions  of  the  snow- 
pack,  sometimes  rexersing  the  expected 
increase  in  [)recipitation  with  ele\ation  and 
having  important  consecjuences  to  plant  distri- 
butions (Branson  et  al.  1976,  Sturges  1977,  W'e.st 
and  Caldwell  1983).  The  important  growing 
season  is  the  cool  spring  when  the  soil  profile  is 
recharged  from  winter  precipitation;  growth  is 
usualK'  curtailed  b\-  dning  soils  coupled  with 
high  temperatures  in  earl\-  or  mid-smnmer.  A 
clear  pictiu-e  o(  this  climatic  regime  is  essential 
to  an\-  chscussiou  of  plant  adaiitations  in  the 
region. 

A  second  major  feature  affecting  plant  per- 
formance is  the  prevalence  of  saline  soils  caused 
l)\'  the  C()ml)ination  of  low  precipitation  and  the 


internal  drainage  txpical  of  the  Great  Basin.  In 
this  paper  we  address  the  salient  morphological, 
physiological,  and  phenological  specializations 
of  nati\ e  plant  species  as  the\'  relate  to  siua i\al 
and  growth  tmder  the  constraints  of  these 
potentialK  stressful  limitations.  We  emphasize 
(1)  edaphic  factors,  particularK  soil  salinit\  and 
texture,  and  (2)  the  climatic  regime  ensuring  a 
fairlv  dependable,  deep  spring  recharge  of  soil 
moisture  despite  the  overall  ariditv;  as  factors 
molding  plant  adaptations  and  producing  the 
uni(jue  aspects  of  the  regional  plants  and  vege- 
tation. The  majoritv  of  the  Great  Basin  lies  at 
moderatelv  high  elevations  (4000  ft  and  aboxe), 
and  it  is  occupied  bv  cold  desert  plant  comnm- 
nities.  Reference  to  "the  Great  Basin"  and  its 
environment  in  this  paper  will  refer  to  this  high- 
{^]e\ation  region  as  distinct  from  that  corner  of 
the  Mojave  Desert  that  occupies  the  southwest- 
em  corner  of  the  Cireat  l^asin  geographic  unit 
(Fig.  1 ).  Our  emphasis  will  be  placed  on  these 
cold  desert  shnib  communities  in  both  the 
Great  Basin  and  the  Colorado  Plateau  ranging 
from  the  topographic  low  points  of  the  saline 
plavas  or  cauNon  bottoms  up  to  the  pinvon-juni- 
per  woodland.  The  lower-elevation,  warmer. 


19921 


Plant  Adaptation 


197 


Great  Basin 


Mojave 
Colorado  Plateau 


Fig.  1.  Distribution  of  the  major  desert  vegetation  zones 
ill  tlie  Great  Bitsin  and  Colorado  Plateau.  Numbers  indicate 
l(K'atioiis  of  climate  stations  for  which  data  are  presented  in 
Table  1.  Most  of  the  Mojave  Desert  indicated  is  geologically 
part  of  the  Great  Basin,  but,  due  to  its  lower  elevation  and 
warmer  temperatures,  it  is  climaticallv  distinct  from  the  rest 
of  the  region. 


antl  drier  Mojax'e  De.sert  portion  of  the  Cjreat 
Basin  will  he  considered  primariK  as  a  point  of 
e()inj)arison,  and  for  more  tlioronii;h  coxerage  of 
that  region  we  recommend  the  reviews  h\ 
Ehleringer  (1985),  MacMahon  (1988),  and 
Smith  and  Nowak  (1990).  For  the  higher  mon- 
tane and  alpine  zones  of  the  desert  mountain 
ranges,  the  reader  is  referred  to  rexiews  l)\ 
\'asek  and  Thome  (1977)  and  Smith  and  Knapp 
( 1990).  We  are  indebted  in  onr  own  c()\erag(^  of 
the  cold  desert  to  other  rec-ent  rexic^ws.  includ- 
ing Caldwell  (1974,  19S5).  West  (19SS). 
Dobrowolski  et  al.  (1990),  and  Smith  and 
Nowak  (1990). 

The  Great  Basin  and  the  Colorado  Plateau 
share  important  climatic  features  such  as  overall 
ariditv;  frequent  summer  droughts,  and  conti- 
nental winters;  yet  the\^  differ  in  other  ecjualK 
important  features.  Temperatures  on  the  Colo- 
rado Plateau  are  similar  to  ecjuixalent  elexatioiis 


in  the  southern  (Ircat  liasin.  hut  suiiiiiici'  pre- 
cipitation is  suhstantially  greater  on  the  Colo- 
rado I'lateau  (Tahk"  \).  Soils  and  drainage 
patterns  also  differ  in  crucial  wa\s.  The  high- 
lands of  the  Colorado  Plateau  generally  drain 
into  the  Colorado  Hixer  sv'stem.  In  manv  areas 
extensive  exposure  of  marine  shales  from  the 
Chinl(\  \hmcos,  and  Morrison  Brnshv-Basin 
formations  wc^ithcr  into  soils  that  restrict  plant 
diversitv  and  total  cover  due  to  high  concentra- 
tions of  NaSOa,  and  the  formation  of  clavs  that 
do  not  allow  water  infiltration  (Potter  et  al. 
1985).  In  other  areas  massive  sandstone  out- 
crops often  dominate  the  landscape.  Shrubs  and 
trees  mav  root  through  ven  shallow  rock"v  soils 
into  natural  joints  and  cracks  in  tlie  sub.stratum. 
Deeper  soils  are  generallv  aeolian  deposits 
forniinti  sands  or  sandv  loams.  In  contrast,  high 
elevations  of  the  Cireat  Basin  drain  into  closed 
\alleys  and  evaporative  sinks.  This  results  in 
greater  average  salinitA'  in  the  lowland  soils  of 
the  Great  Basin,  with  NaCd  being  the  most 
common  salt  (Flovxers  1934 ).  and  a  more  exten- 
sive development  of  haU)ph\  tic  or  salt-tolerant 
vegetation.  Soils  tend  to  be  deep,  especialK  at 
lower  elevations,  and  van'  in  texture  from  clav 
to  sandv  loams.  Summer-active  species  with 
different  photosvnithetic  pathwavs,  such  as  C4 
grasses  and  CAM  succulents,  are  poorlv  repre- 
sented in  nuich  of  the  Crreat  Basin,  but  the 
combination  of  increased  summer  rain,  sandier 
soils,  and  milder  winters  at  the  lower  elevations 
of  the  Colorado  Plateau  has  resulted  in  a  greater 
expression  of  phenological  diversit\. 

The  interactions  of  edaphic  factors  and  cli- 
mate are  complex  and  often  subtle  in  their 
effects  on  plant  performance.  Furthermore, 
[)Iant  distributions  are  rarelv  determined  bv  a 
single  factor  throughout  th(ir  geographic  range. 
e\en  though  a  single  factor  mav  appear  to  con- 
trol distribution  in  the  context  of  a  local  ecosvs- 
tcMii.  Species-spcxific  characteristics  generally 
do  not  inqxirt  a  narrow  re(|uirement  for  a  spe- 
cific environment,  but  rather  a  unique  set  of 
"ranges  of  tolerance"  to  a  large  arrav  of  enxiron- 
mental  j)arameters.  In  different  enviromncntal 
contexts,  different  tolerances  mav  be  more  lim- 
iting, both  abiotic  and  biotic  interactions  may  be 
altered,  and  the  same  set  of  species  may  .sort  out 
in  different  spacial  ])attenis.  A  further  conse- 
(juence  of  this  is  that  a  local  combination  of 
species,  whicli  we  might  refer  to  as  a  Great 
Basin  plant  communitv.  represents  a  region  of 
oxerlap  in  the  geograpln'calK    more  extensive 


198 


Great  Basin  Naturalist 


[N'oluiiie  52 


and  treiieralK  miicjue  tlistrihutioiis  ot  each  coni- 
ponenf  species.  In  fact,  the  distributions  of  spe- 
cies commonly  associated  in  the  same  Great 
Basin  connnunitv'  may  be  strongly  contrasting 
outside  the  Great  Basin.  This  is  an  essential 
point  in  attempting  to  discuss  plant  adaptations 
with  the  implication  oi  cause  and  effect, 
because  few  species  discussed  will  have  a  strict 
and exclusixe  relationship  with  the  environment 
of  interest.  Unless  we  can  show  local,  ecot\pic 
differentiation  in  the  traits  discussed,  we  need 
to  take  a  broad  view  of  the  relationship  between 
environment  and  plant  characters.  In  a  few 
instances,  a  small  number  of  edaphic  factors  and 
plant  characters,  such  as  tolerance  of  veiy  high 
salinity  in  soil  wdth  shallow  groundwater,  seem 
to  be  of  overriding  importance.  In  most  cases  we 
need  to  ask,  what  are  the  common  elements  of 
climate  over  the  diverse  ranges  of  all  these  spe- 
cies? One  such  common  element,  which  will  be 
emphasized  throughout  this  re\iew,  is  the 
importance  of  reliable  winter  recharge  of  soil 
moisture  in  an  arid  to  semiarid  climate.  B\-  iden- 
tifying such  common  elements  and  focusing  on 
them,  we  do  not  fully  describe  the  autecologv  of 
an\'  species,  but  we  attempt  a  cogent  treatment 
of  plant  adaptations  to  the  Great  Basin  environ- 
ment, and  an  explanation  of  the  unicjue  features 
of  its  plant  connnunities. 

Climate,  Edaphic  Factors,  and  Plant 
Distribution  Patterns 

Typical  zonation  patterns  observed  in  spe- 
cies distributions  around  playas  (the  saline  flats 
at  the  bottom  of  closed-drainage  basins)  are 
quite  dramatic,  refl(^cting  an  o\  erriding  effect  of 
salinit)'  on  plant  distribution  in  the  Cireat  Basin. 
Moving  out  from  the  basin  center  is  a  gradient 
of  decreasing  soil  salinity  often  correlated  with 
progressively  coarser-textured  soils.  Along  this 
gradient  there  are  conspicuous  species  replace- 
ments, often  resulting  in  concentric  zones  of 
contrasting  vegetation  (Flowers  1934,  Vest 
1962).  In  the  lowest  topographic  zone,  saline 
groundwater  may  be  very  neav  the  surface.  Soils 
are  ven'  saline,  fine  textured,  and  subject  to 
occasional  flooding  and  anoxic  conditions,  in 
this  enxiromnent  the  combination  of  available 
moisture  with  other  poteutiallv  stressful  soil 
characteristics  seems  to  be  more  important  than 
climatic  factors  of  temperatiu'e  or  seasonal  rain- 
fall patterns.  Speci(>s  found  here,  such  as  desert 
saltgrass     {Distichlis     spic<il(i),     pickleweeds 


(Allciirolfia  occich'ittdlis  and  Salicontia  spp.), 
and  greasewood  (Sarcobatiis  vcniiicitkitiis), 
may  themselves  show  zonation  due  to  degrees 
of  tolerance.  They  tend  to  occur  in  close  prox- 
imitv,  however,  on  the  edges  of  salt  plavas,  saline 
flats  with  shallow  water  tables,  and  near  saline 
seeps  over  a  wide  range  of  elevations,  tempera- 
tures, and  seasonal  rainfall  patterns  in  both  the 
Great  Basin  and  southern  warm  deserts 
(MacMahon  198S).  This  relative  independence 
of  distribution  from  prevailing  climate  is  a  spe- 
cial characteristic  of  strongly  halophytic  plant 
communities  throughout  the  world  and  reflects 
the  overriding  importance  of  such  extreme 
edaphic  conditions.  Species  found  on  better- 
drained,  moderately  saline  soils,  where  groimd- 
water  is  not  found  within  the  rooting  zone, 
include  winterfat  {Ccratoidcs  laiuita)  and 
shadscale  {Atiiplcx  confeififolia).  These  species 
are,  in  turn,  replaced  at  higher  elevations  and  on 
moister,  nonsaline  soils  bv  big  sagebnish  iAiic- 
inisia  tridcntatd),  rabbitbrush  [Chnjsoiluntinus 
sp.),  bitterbnish  {Piirsliia  sp.),  and  spinv  hop- 
sage  {GiYHfia  spinosa).  Shadscale  is  often  fcnmd 
in  areas  where  soils  are  notably  saline  in  the 
lower  half  of  the  rooting  zone,  but  not  in  the 
upper  soil  lavers.  Such  a  tolerance  of  mt)der- 
ately  saline  soils  seems  to  control  its  distribution 
around  playas,  especially  in  the  wetter,  eastern 
portion  of  the  Great  Basin  (western  Utah)  and 
lower  elevations  in  the  warm  Mojave  Desert.  In 
the  more  arid  regions  of  western  and  central 
Nevada,  however,  the  shadscale  connnunitv 
occurs  widely  on  nonsaline  slopes  lower  in  ele- 
vation, warmer,  and  drier  than  those  dominated 
by  big  sagebrush.  These  higher  bands  of 
shadscale  have  been  variously  inteipreted  in 
terms  of  ariditv  tolerance  and  climate  (Billings 
1949)  or  an  association  with  limestone-derived 
calcareous  soils  (Beatlev  1975).  The  latter 
author  points  out  that  even  on  nonsaline  soils 
percent  cover  in  the  shadscale  connnunitv  is 
lower  than  expected  for  the  level  of  precipita- 
tion, and  argut^s  that  this  indicates  stress  from 
ecUiphic  factors.  Thus,  shadscale  distribution  is 
most  correlated  with  salinitv  tolerance  in  some 
regions  and  other  eckiphic  and  climatic  tolcMan- 
ces  in  other  regions. 

Where  the  higher  elevations  of  thc>  Cyreat 
Basin  conu^  in  contact  with  the  lower-elevation, 
generallv  drier,  and  warmer  Mojave  Desert 
region,  comminn'ties  ck)minated  by  creosote 
(Larrca  tridfufafa)  replace  sagebrush  commu- 
nities    on     nonsaline     slopes     and     bajadas. 


19921 


Plant  Adaitxtion 


199 


Shadscak'  can  ht'  toiiiul  liotli  on  saline  soils  at 
\en  low  t'k'\ ations  in  tlu^  Mojaw  and  as  a  tran- 
sitional band  at  liigli  ekn  ations  l)et\\een  creo- 
sote and  sagebmsh.  Elements  of  the  cold  desert 
shnib  conimnnities,  adapted  to  continental  win- 
ter's and  a  cool  s[)ring  growing  season,  can  be 
tonnd  throughout  the  winter-rain-doniinated 
\h)ja\"e  Desert  region  as  a  high-elexation  band 
on  arid  mountain  ranges.  They  also  extend  to  the 
southeast  at  high  ele\ations  into  the  strongK- 
bimodal  precipitation  regime  of  the  Colorado 
i'latean,  and  northward  at  low  elexations  into 
Idaho.  Washington,  andexen  British  (-oluinbia. 
Nhning  up  from  bajadas  of  the  southern  warm 
deserts,  there  appears  to  be  no  suitable  combi- 
nation of  temperature  and  precipitation  at  an\' 
elevation  to  support  floristic  elements  of  the 
cold  desert.  As  precipitation  increases  with  alti- 
tude, zones  with  equivalent  total  precipitation 
do  not  \et  ha\e  cold  winters  and  are  occupied 
In  warm  desert  shnib  connnunities  grading  into 
chaparral  composed  of  unrelated  ta.xa.  Higher 
ele\  ations  with  cold  winters  have  sufficient  pre- 
cipitation to  support  forests.  Other  elements 
coimnon  in  shadscale  and  mixed-shrub  connnu- 
nities of  the  Great  Basin,  such  as  winterfat  and 
budsage  (Ai-tcmisia  spiiiosa),  are  often  found 
outside  the  Great  Basin  in  cold-winter  but 
largel\-  summer-rainfall  grasslands. 

f^rom  these  patterns  of  communitv-  distribu- 
tion within  the  Great  Basin  and  Colorado  Pla- 
teau, and  also  from  a  consideration  of  the  more 
extensive  ranges  and  affinities  of  the  component 
species,  we  can  isolate  a  few  ke\-  features  of  the 
environment  that  are  largely  responsible  for  the 
unique  plant  features  seen  in  the  Great  Basin. 
The  most  obvious  features  are  related  to  the 
patterns  of  soil  salinitv  and  texture  generated  bv 
the  (Aerall  ariditv  combined  with  either  internal 
drainasie  basins  or  tlie  in  situ  weathering  of 
specific  rock  tvpes.  The  most  important  climatic 
variables  are  slightlv  more  subtle.  There  is 
cknulv  not  a  requirement  for  exclusivelv  winter 
rainfall,  but  there  is  a  re(|uirement  for  at  least  a 
substantial  portion  ol  the  annual  rainfall  to  come 
dniing  a  continental  winter  This  permits  v\inter 
(iccitninlatioit  of  precipitation  iod 'greater  depth 
in  the  soil  profile  than  w  ill  occur  in  response  to 
less  predictable  sunnner  replenishment  of 
dning  soil  moisture  reserves.  Unck'r  an  overall 
arid  climate,  winter  n^charge  maintains  a  pre- 
ilictablv  favorable  and  ck)minant  spring  growing 
season  even  in  manv  areas  of  strongly  bimodal 
rainfiill.  Most  of  the  phvsiological.  moqihologi- 


cal.  and  plieiiological  traits  lonnd  in  llie  (k)mi- 
nant  shrubs  rell(^ct  the  [)riman  importance  of 
such  a  cool  spring  growing  .season. 

PlI()T(lSY\'THKSIS 

Piiotosyxtiiktk;  I'ATIIWAVS. — The  pro- 
cess of  photosvnthesis  in  plants  relies  on  the 
acquisition  of  CO2  from  the  atmo.sphere,  which, 
when  coupled  with  solar  energ\',  is  transformed 
into  organic  molecules  to  make  sugars  and  pro- 
vide  for  plant  growth.  In  moist  climates  plant 
communities  often  achieve  clo.sed  canopies  and 
1(){)%  cover  of  the  ground  surface.  Under  these 
conditions  competition  for  light  may  be  among 
the  most  important  plant-plant  interactions.  In 
the  deserts  total  plant  cover  is  much  less  than 
100%,  and  in  the  Great  Basin  closer  to  259f. 
Photosviithesisisnotgreatlvlimitcxlbv  available 
light,  but  rather  bv  water,  mineral  nutrients 
needed  to  .synthesize  enzAines  and  maintain 
metabolism,  and  maximum  canopv  leaf-area 
development. 

Three  biochemical  pathwavs  of  photosvii- 
thesis  have  been  described  in  plants  that  differ 
in  the  first  chemical  reactions  associated  with 
the  capture  of  CO2  from  the  atmosphere.  The 
most  common  and  most  fundamental  of  these 
pathways  is  referred  to  as  the  C3  pathway 
because  the  first  product  of  photosynthesis  is  a 
3-carbou  molecule.  The  other  two  pathways,  C4 
and  CAM,  are  basically  modifications  of  the 
primaiy  C3  pathway  (Osmond  et  al.  1982).  The 
C4  pathwav  (first  product  is  a  4-carbon  mole- 
cule) is  a  morphological  and  biochemical 
arrangement  that  overcomes  photorespiration, 
a  process  that  results  in  reduced  photosviithetic 
rates  in  C3  plants.  The  C.i  pathway  can  confer  a 
much  higher  temperature  optimum  for  photo- 
.synthesis  and  a  greater  water-use  efficiency.  As 
water-use  efficiencv  is  the  ratio  of  photosvn- 
thetic  carbon  gain  to  transpirational  water  loss, 
C4  plants  have  a  metabolic  advantage  in  hot,  dn^ 
environments  w4ien  soil  moisture  is  available.  In 
grasslands  C4  grasses  become  dominant  at  low 
elevations  and  low  latitudes  where  animal  tem- 
]x^ratur(\s  are  warmest.  In  interpreting  })lant 
distribution  in  deserts,  the  .seasonal  pattern  of 
rainfall  usuallv  restricts  the  periods  of  plant 
growth,  and  the  temperature  during  the  rainy 
season  is  thus  more  important  than  m(\ui  annual 
temperature.  The  C4  pathwav  is  ofti'u  associated 
with  smnmei-active  species  and  also  with  plants 
of  saline  soils.  While  C3  grasses  pre(k)minate  in 


200 


Great  Basin  Naturalist 


[\'olunie  52 


most  of  the  Cireat  Basin,  C4  grasses  beeonie 
iiicreasinglv  important  as  summer  rain  increases 
to  the  south,  and  especiaHv  on  the  Colorado 
Plateau.  Halophvtic  plants  are  often  C4,  such  as 
saltbush  iAfrij)Icx  spp.)  and  saltgrass  (Disticlilis 
spp.),  and  tliis  mav  gixe  the  plants  a  competitixe 
advantage  from  increased  water-use  efficienc\- 
on  saline  soils. 

The  third  photo.sMithetic  pathway  is  CAM 
photosMithesis  (Crassulacean  Acid  Metabolism). 
CAM  plants  open  their  stomata  at  night,  capture 
COo  and  store  it  as  malate  in  the  cell  \acuole, 
and  keep  theii  stomata  closed  dining  the  dav 
(Osmond  et  al.  1982).  The  CO2  is  then  released 
from  the  vacuole  and  used  for  photos)aithesis 
the  folloxxing  da^'.  Because  the  stomata  are  open 
onl\  at  night  when  it  is  c()t)l,  water  loss  associ- 
ated with  CAM  photosNuthesis  is  greatlv 
reduced.  This  pathwa\'  is  often  found  in  succu- 
lents such  as  cacti  and  agaxe,  and,  although 
C^AM  plants  are  present  in  the  Great  Basin,  they 
are  a  i-elati\eK-  minor  component  of  tlie  vegetation. 

Photosxnthetic  rates  of  plants,  like  most  met- 
abolic processes,  sho\\'  a  strong  temperatm-e 
dependence.  UsualK,  photosvnthetic  rates  are 
reduced  at  low  temperatures  because  of  the 
temperature  dependence  of  enz^'uie-catah'zed 
reaction  rates,  increase  with  temperature  mitil 
some  maximum  \alue  (which  defines  the  "tem- 
perature optimum"),  and  then  decrease  again  at 
higher  temperatures.  The  temperature  optima 
and  niiuimum  photosxnthetic  rates  of  plants 
show  considerable  variation,  and  the\'  generalK 
reflect  the  growing  conditions  of  their  natural 
environments. 

PHOTOSYNTHETIC  adaptation. — In  the 
spring  the  photosynthetic  temperature  optima 
of  the  dominant  shrub  species  are  tvpicalK'  as 
low  as  15  C  (40  F)  (Caldwell  1985),  correspcMid- 
ing  to  the  prevailing  en\ironmental  tempera- 
tures (mi(kla\-  ma.xima  generally  less  than  20  C). 
As  ambient  temperatures  rise  10-15  C  in  the 
summer,  there  is  an  upward  shift  of  only  ,5-10  C 
in  the  photos\iithetic  temperature  optima  of 
most  shrubs,  coupled  with  a  slower  decline  of 
photosynthesis  at  high  temperatures.  The  max- 
imimi  ph()t()s\nithetic  rates  measunxl  in  most 
Great  Basin  shrubs  under  either  natural  or  irri- 
gated conditions  range  from  14  to  19  jjluioI  ClO^ 
m-  s'  (DePuit  and  Caldwell  1975,  Caldwell  et 
al.  1977,  Evans  1990).  These  rates  are  (|uite 
mode.st  compared  to  t\ie  high  maxima  of  25  to 
45  jjLmol  CO2  m  "  s  '  ob.sened  in  man\-  warm- 
dc^sett  species  adapted  to  rapid  growth  at  higher 


temperatures  (Ehleringer  and  Bjorkman  1978, 
Mooney  et  al.  1978,  Comstock  and  Ehleringer 
1984,  1988,  Ehleringer  1985).  This  presumably 
reflects  the  specialization  of  these  Great  Basin 
shiiibs  towards  utilization  of  the  cool  spring 
growing  season.  Positive  photosynthetic  rates 
are  maintained  even  when  leal  temperatures 
are  near  freezing,  which  permits  photosvnthetic 
activity  to  begin  in  the  very  early  spring  (DePuit 
and  Caldwell  1973,  Caldwell  1985). 

Unusuallv  high  maximmn  photosvnthetic 
rates  of  46  ixmol  CO2  m  ~  s  '  have  been  reported 
for  one  irrigated  Great  Basin  shnib,  rabbitbrush 
{Chnjsothamnus  nauseosus)  (Da\is  et  al.  1985). 
This  species  is  also  unusual  in  having  a  deep  tap 
root  that  often  taps  groundwater,  unusuallv  high 
levels  of  summer  leaf  retention  (Branson  et  al. 
1976),  and  continued  photo.sxnthetic  activitx* 
throughout  the  summer  drought  ( Donoxan  and 
Ehleringer  1991).  All  of  these  characters  sug- 
gest greater  photosvnthetic  activity  during  the 
warm  summer  months  than  is  found  in  most 
Great  Basin  shrub  species. 

Shoot  ACTTIMTY'  and  phenology. — Gener- 
ally speaking,  there  is  a  distinct  drought  in  early 
summer  (June-|ulv)  in  the  Great  Basin  Cold 
Desert,  the  Mojave  Desert,  and  the  Sonoran 
Desert.  All  of  these  deserts  ha\e  a  substantial 
winter  precipitation  season,  but  they  differ  in 
the  importance  of  the  summer  and  early  fall 
rainy  seas(jn  (|ul\-October)  in  supporting  a  dis- 
tinctive period  of  plant  growth  and  acti\itv 
(MacMahon  1988).  The  relationship  between 
climate  and  plant  growing  season  is  complex  and 
includes  total  rainfall,  seasonal  distribution  of 
rainfall,  and  predictabilitv  of  rainfall  in  different 
seasons  as  important \ariables.  Fmthermore,  in 
\en  arid  areas  the  seasonalih'  of  temperatures 
may  be  as  important  as  that  of  rainfall.  In  the 
Great  Basin,  cold  winters  allow  the  moisture 
from  low  lexels  of  precipitation  to  accumulate 
in  the  soil  for  spring  use,  while  hot  summer 
temperatiu'es  cause  rapid  evaporation  from 
plants  and  soil.  High  temperatures  can  prevent 
wetting  of  the  soil  profile  bevond  a  few  centime- 
t(Ms  depth  in  response  to  sununer  rain,  even 
when  sununer  rain  accounts  for  a  large  fraction 
of  the  animal  total  (Caldwell  etal.  1977).  As  total 
annual  rainfall  decreases,  the  relative  impor- 
tance of  the  cool  spring  growing  season 
i  I  icreases  as  the  oiiK  potential  growing  period  in 
which  available  soil  moisture  approaches  the 
evaporative  demand  (Thornthwaite  1948,  Com- 
stock and  Ehleiintier  1992).  Finally,  reliabilih 


19921 


Plant  Adaitation 


201 


of  nioisturc  is  important  to  [XTcnnials,  and  as 
axerage  total  precipitation  decreases,  the  \ari- 
ance  bet\veen  \ears  increases  (Ehleringer 
1985);  \ariabilit\'  of  annuiil  precipitation  is  dis- 
cussed in  more  detail  later  in  the  section  on 
annuals  and  life-histor\'  dixersitv.  Summer  rain 
is  more  likel\-  to  be  concentrated  in  a  few  high- 
intensit\  storms  that  max  not  happen  e\eiA' \ear 
at  a  gi\en  site  and  ma\'  cause  more  nmoit  when 
the\  do  occur.  The  abilits'  of  moisture  from 
w  inter  rain  to  accumidate  o\er  several  months 
greatly  enhances  its  reliabilits'  as  a  moisture 
resource.  Thus,  most  plants  in  the  Great  Basin 
have  their  priniar\-  growing  season  in  the  spring 
and  earl\-  summer.  Some  species  achie\e  an 
e\ergreen  canop\'  b\'  rooting  deepK;  but  few 
species  occur  that  specialize  on  growth  in  the 
hot  summer  season  (Branson  et  al.  1976,  Cald- 
well et  al.  1977,  Everett  et  al.  1980).  Ehleringer 
et  al.  (1991)  measured  the  abilitv  of  common 
perennial  species  in  the  Colorado  Plateau  to  use 
moisture  from  summer  convection  storms. 
The\-  obserxed  that  less  than  half  of  the  water 
uptake  b\-  wood\'  perennial  species  was  from 
suriace  soil  laxers  saturated  b\'  summer  rains. 
Prexalence  of  summer-active  species  increases 
along  the  border  betxveen  higher  basins  and  the 
southeast  Mojaxe  Desert  xvhere  summer  rain  is 
more  extensixe,  and  especialK'  on  the  Colorado 
Plateau  xx'here  summer  rain  is  greatest.  Summer 
temperatures  are  also  lower  on  the  Colorado 
Plateau  than  in  the  eastern  Mojaxe  (Table  1), 
alloxxing  more  effectixe  use  of  the  moisture. 

Most  phenolog)-  studies,  especiallx'  from  the 
more  northern  areas,  emphasize  the  directional, 
sequential  nature  of  the  phenological  phases 
(Branson  et  al.  1976,  Saner  and  Uresk  1976, 
Cambell  and  Harris  1977,  West  and  Gastro 
1978,  Pitt  and  W'ikeem  1990).  A  single  period  of 
spring  vegetative  groxvth  is  usually  folloxved  by 
a  single  period  of  floxxering  and  reproductix'e 
groxx'th.  Manx-  species  produce  a  distinct  cohort 
of  ephemeral  spring  leaves  and  a  later  cohort  of 
exergreen  leaxes  (Daubenmire  1975,  Miller  and 
Schultz  1987).  For  most  species,  multiple 
groxxth  episod(\s  associated  xxith  intermittent 
spring  and  summer  rainfall  exents  do  not  occur. 
In  xears  xxith  unusually  heavy  August  and  Sep- 
tember rains,  a  distinct  second  period  of  xegeta- 
tixe  growth  may  occur  in  some  species  (West 
and  Gastro  1978).  Climatic  xariations  from  xear 
to  xear  do  not  change  the  primaty  importance 
of  spring  gro\xi:h  or  the  order  of  phenological 
exents.   In  particular  \ears,   thex'  ma\-  cause 


expansion  or  contraction  of  xc^gt^tatixc  pluuses 
and  exen  the  omission  of  reproductix-e  pha.ses. 

Most  species  initiate  grox\th  in  earlx'  spring 
(March)  xvhen  maximum  da\time  temperatures 
are  5-15  C  and  xx'hile  nighttime  temperatures 
are  still  freezing.  Delaxed  initiation  of  spring 
groxxth  is  generally  associated  xxith  greater 
summer  actixit\-  and  max-  be  related  to  an  exer- 
green habit,  a  phreatophxtic  habit,  or  occupa- 
tion of  habitats  xxith  greater  sununer  moisture 
axailabilitx  from  regional  rainfall  patterns, 
nmoff,  or  tirovmdxx'ater.  Higher-than-ax-erase 
xxinter  and  spring  precipitation  tends  to  prolong 
vegetatixe  growth  and  delax-  reproductive 
groxx'th  till  later  in  the  sununer  ( Saner  and  U  re.sk 
1976,  Cambell  and  Harris  1977).  Strong xegeta- 
tive  dormancy  ma\'  be  displayed  in  mid-summer 
(Everett  et  al'.  1980,  Evans  1990),  although  root 
groxx'th  (Hodgkinson  et  al.  1978)  and  increased 
reproduction  (W'est  and  Gastro  1978,  Exans, 
Black,  and  Link  1991)  max'  still  occur  in 
response  to  rain  at  that  time.  In  xears  with 
beloxx'-axerage  spring  and  svunmer  precipita- 
tion, leaf  senescence  is  accelerated  and  floxx'er- 
ing  may  not  occur  in  man\-  species. 

The  time  taken  to  complete  the  full  annual 
groxxth  cxcle  including  both  xegetatixe  and 
reproductixe  stages  is  stronglx  related  to  rooting 
depth  in  most  of  these  conmumities,  xxith  deep- 
rooted  species  prolonging  actixit\'  further  into 
the  summer  drought  (Pitt  and  Wikeem  1990). 
The  exact  timing  of  floxx'ering  and  fniit  set  shoxvs 
considerable  xariation  among  different  shrub 
species.  Some,  especiallx  those  that  are 
drought-deciduous,  lloxxer  in  late  sprin>j;  and 
earlx  summer  just  prior  to  or  concurrent  xxith 
the  onset  of  summer  drought.  Manx-  exergreen 
shRib  species  begin  floxxering  in  midsummer 
(Artonisia)  or  in  the  fall  {Gutierrczia  and 
Chn/sothainntts).  These  late-flowering  species 
generallxdo  not  aj)pear  to  utilize"  stored  reserx'es 
for  floxx'ering.  but  relx  on  current  photo.sxnthe- 
sis  during  this  least  fax-orabk"  period.  In  the  case 
(){  Aticmisia  fridoitafa.  it  has  been  shoxxn  that 
earlx )lix-drates  used  to  fill  fruits  arc  dcrixcd 
exclnsixi'lx  from  the  inflorescences  theniselxes, 
xxhile  photosxnthate  from  xegetatixe  l)ranches 
presumablx  continues  to  support  root  groxx'th. 
Summer  rain  during  this  time  period  does  not 
promote  xegetatixe  shoot  groxxth  but  does 
increase  xvater  use  by  and  the  ultimate  size  of 
inflorescences  (Exans  1990).  Exans,  Black,  and 
Link  (1991)  haxe  argued  that  this  pattern  of 
floxx'ering,  ba.sed  on  residual  deep  soil  moisture 


202 


Great  Basin  Naturalist 


[Volume  52 


and  the  unreliable  summer  rains,  ma)'  contrib- 
ute to  competitixe  dominance  within  these 
comnumities.  The  more  favorable  and  much 
more  reliable  spring  growing  season  is  used  for 
\egetative  growth  and  coiupetitive  exploitation 
of  the  soil  \olume,  while  reproductive  gro\\i:h  is 
delayed  until  the  less  favorable  season,  and  may 
be  successful  only  in  years  with  adequate 
su mmer  precipitation . 

Most  grasses  in  the  northern  part  of  the 
Great  Basin  utilize  the  G,5  pathway  and  begin 
growth  very  early  in  the  spring.  These  species 
complete  fruit  maturation  by  early  or  mid- 
sunnner,  often  becoming  at  least  partially  dor- 
mant thereafter.  On  the  Colorado  Plateau,  with 
much  greater  amounts  of  summer  precipitation, 
there  is  a  large  increase  in  species  number  and 
cover  by  C4  grasses  such  as  bluestem 
(Andropogon)  and  grama  {Bouteloua),  espe- 
cial K  at  warmer,  lower  elevations  and  on  deep 
sandy  soils.  Many  of  these  species  occur  in 
mixed  stands  with  the  C3  species  but  delay  ini- 
tiation of  growth  until  May  or  Jime;  they  can  be 
considered  suiumer-active  rather  than  spring- 
actix'e.  In  contrast,  some  C4  grasses  such  as  sand 
dropseed  {Sporoholii.s  cri/ptcmdnis),  galleta 
grass  (Hilaria  jainesiii),  and  three-awn  {Arisfkla 
purpurea)  are  widespread  in  the  Great  Basin 
where  sunuuer  rain  is  only  moderate  in  long- 
term  averages  and  veiy  inconsistent  year  to  year. 
Spring  growth  of  these  widespread  species 
tends  to  be  slighth'  or  moderately  delayed  com- 
paied  to  co-occurring  C5  grasses,  but  they  are 
still  able  to  complete  all  phenological  stages 
based  on  the  spring  moisture  recliarge.  The\' 
show  a  greater  abilit)'  than  the  G,;  species  to 
respond  to  late  spring  and  simuiier  rain  witli 
renewed  growth  (Everett  et  al.  1980),  however, 
which  compensates  in  some  years  for  their  later 
developuKMit.  Other  C4  grasses  in  the  Great 
Basin  may  be  associated  with  seeps, 
streamsides,  or  salt-marshes,  and  show  a 
summer  activity'  pattern.  G4  shrubs  such  as  salt- 
bush  (Atriplex)  show  similar,  spring-actixe 
growth  patterns  to  the  (v;  shrubs,  but  may  show 
slightly  greater  tolerance  of  sunuuer  drouglit 
(Caldwell  et  al.  1977). 

Phenolog)'  in  the  Mojave  Desert  shows  both 
similarities  and  strong  contrasts  to  the  Great 
Basin.  Although  rainfall  is  largeK  biiuodal  in  the 
eastern  Mojave,  absolute  amoimts  are  vvw  low. 
The  sunuuer  is  so  hot  that  little  growth  normally 
occurs  at  that  time  unless  more  than  25  nun  (1 
inch)  comes  in  a  single  storm  (Ackerman  et  al. 


1980).  Fall  and  winter  precipitation  is  the  mo.st 
important  in  promoting  good  spring  growth  of 
perennials  (Beatley  1974).  Comstock  et  al. 
(1988),  looking  at  one  years  growth  in  19 
Mojave  species,  described  an  important  cohort 
of  twigs  initiated  during  the  winter  period  that 
accounted  for  most  vegetative  growth  during 
the  following  spring.  Although  new  leaves  were 
produced  in  response  to  summer  rain,  summer 
growth  in  many  of  the  species  was  largeK'  a 
further  ramification  of  spring-initiated  floral 
branches.  In  most  species  summer  growth  made 
little  contribution  to  perennial  stems.  Despite 
the  preferred  winter-spring  orientation  of  many 
shmbs,  winter  recharge  is  much  less  effective 
and  reliable  in  the  Mojave  Desert  than  in  the 
Great  Basin.  Due  to  warmer  temperatures, 
winter  dormancy  may  be  less  complete,  but 
vigorous  growth  rarely  occurs  until  tempera- 
tures rise  further  in  the  early  spring.  Rapid 
growth  luay  be  triggered  by  rising  spring  tem- 
peratures or  may  be  delayed  until  major  spring 
raius  provide  sufficient  moisture  (Beatley  1974, 
Ackenuan  et  al.  1980).  Furthermore,  a  shal- 
lower soil  moisture  recharge  often  results  in 
fluctuating  plant  water  status  and  multiple 
episodes  of  growth  and  flowering  during  the 
spring  and  early  fall.  Some  Great  Basin  species 
that  also  occur  in  the  Mojave,  such  as  winterfat 
and  shadscale,  commonly  show  multiple  growth 
and  reproductive  episodes  per  year  under  that 
climate  (Ackennan  et  al.  1980)  but  not  in  the 
Great  Basin  (West  and  Gastro  1978).  The 
degree  to  which  this  difference  is  due  entirely 
to  environmental  differences  as  opposed  to  eco- 
t\pic  differentiation  does  not  appear  to  have 
been  studied. 

Water  Relations 

Ai:)APTATION  TO  LIMITED  W.ATER. — Stoma- 
tal  pores  provide  the  pathvx'av  by  which  atmo- 
spheric COo  diffuses  into  the  leaf  replacing  the 
CO2  incorporated  into  sugar  molecules  during 
photosynthesis.  Because  water  vapor  is  present 
at  \eiy  high  concentrations  inside  the  leaf, 
opening  stomata  to  capture  COo  inevitably 
results  in  trauspi rational  water  loss  from  the 
plant;  thus,  leaf  water  content  is  decreased, 
resulting  in  a  decrease  in  plant  water  potential 
(^).  Thus,  plant  water  status,  transpiration,  and 
ac(juisiti()n  of  water  from  the  soils  have  a  tre- 
mendous impact  on  photosynthetic  rates  and 
overall  plant  grovxth. 


1992] 


Plant  AnAPTYnox 


203 


Main  soils  in  the  (Ticat  Basin  arc  liiu^  t(^\- 
tured,  which  has  botli  atKantagcs  and  disadxan- 
tages  for  plant  growth.  Infiltration  of  water  is 
slower  in  fine-textured  soils,  increasing  the  like- 
lihood of  runoff  and  reduced  spring  recharge, 
especialK'  on  steeper  slopes.  They  are  also  more 
prone  to  water-logging  and  anoxic  c-onditions. 
The  deep  root  systems  of  Cireat  Basin  sluMihs  are 
ver\'  sensitive  to  anoxia,  and  this  can  be  a  strong 
determining  factor  in  species  distributions 
(Limt  et  al.  1973,  CiroeneN'eld  and  Crowley 
1 9S8).  Unnsualh'  wet \ears  ma\' e\en cause  root 
dieback,  especially  at  depth.  Once  water  enters 
the  soil  profile,  the  extremely  high  surface  areas 
of  fine-textured  soils  with  high  clav  and  silt 
content  gi\e  them  a  much  higher  water-holding 
capacit\'  than  that  foimd  in  sandy,  coarse-tex- 
tured soils.  Much  of  this  water  is  tighth'  bound 
to  the  enormous  surface  area  of  the  small 
particles,  howe\er,  and  is  released  onl\  at  \en' 
negatixe  matric  potentials.  Plants  nuist  be  able 
to  tolerate  low  tissue  water  potentials  to  make 
use  of  it. 

As  soil  water  is  depleted  during  sunuuer, 
plants  reduce  water  consumption  b\  closing  sto- 
mata  (DePuit  and  Caldwell  1975,  CambeJl  and 
Harris  1977,  Caldwell  1985,  Miller  1988)  and 
reducing  total  canop\'  leaf  area  to  a  minimum 
(Bran.son  et  al.  1976).  Evergreen  species  shed 
only  a  portion  of  the  total  canop\,  however, 
maintaining  the  youngest  leaf  cohorts  through- 
out the  drought  (Miller  and  Schulz  1987). 
Although  plnsiological  actixit)'  is  greatK' 
reduced  b\'  water  stress,  exergreens  such  as 
sagebnish  can  still  have  positive  photosviithetic 
rates  at  leaf  water  potentials  as  low  as  —50  bars 
(Exans  1990)  and  may  surxive  even  greater 
](nels  of  stress.  In  contrast,  crop  plants  can 
rareK"  sunixe  prolonged  M^  of  less  than  - 15  bars. 
Remaining  functional  at  loxx'  xx'ater  potentials 
requires  the  concentration  of  solutes  in  the  cell 
sap,  and  this  appears  to  be  accomplished  b\ 
several  mechanisms.  In  manx  mesic  species, 
increases  in  organic  solutes  may  account  for 
most  of  the  change  in  osmotic  potential.  In  other 
species,  and  especialK'  tho.se  that  experience 
xeiy  loxv  leaf  xvater  potentials,  a  large  fraction  of 
the  solutes  is  acquired  by  the  uptake  of  inor- 
ganic ions  such  as  K+  (Wvii  fones  and  (^orhani 
1986).  High  concentrations  of  inorganic  ions 
may  l)e  toxic  to  enzx'me  metabolism,  hoxxexer. 
and  they  are  xxidely  thought  to  be  se(juestered 
largely  in  the  central  vacuole,  xvhich  accoimts 
for  90%  of  the  total  cell  xolume.  exen  thoush 


much  of  the  exick^ice  for  this  is  (|uite  indirect. 
Nonetheless,  the  osmotic  potential  of  the  cxto- 
plasm  irnist  also  be  balanced  or  suffer  dehxdra- 
tion.  The  cytoplasmic  .solutes  must  haxe  the 
special  propeitx  of  lowering  the  osmotic  poten- 
tial of  the  cell  sap  xxathout  dismpting  enz\nne 
function  or  cellular  metabolism,  and  are  hence 
termed  "compatible"  solutes  (W'xii  Jones  and 
Gorham  1986).  The  use  of  specific  molecules 
shows  considerable^  xariation  across  species 
apparentlx'  due  to  both  species-specific  xaria- 
tions  in  cell  metabolism  and  taxonomic  relation- 
ships. Some  frecjuentlx  encountered  molecules 
thought  to  function  in  this  manner  include 
amino  acids  such  as  proline  and  also  special 
sugar-alcohols.  Soiue  plants  appear  to  generate 
low  osmotic  potentials  bx'  actixeK"  manufactur- 
ing larger  quantities  of  dissolxed  organic  mole- 
cules per  cell  in  response  to  water  st^^ss.  a 
process  referred  to  as  "osmotic  adjustment."' 
This  process  ma\'  be  costh;  hoxx'exer,  recjuiring 
the  inxestment  of  large  amovmts  of  materials 
(nexv  solutes)  at  a  time  xx'hen  xx'ater  stress  is 
largely  inhibiting  photosvnthetic  activitv'.  An 
alternatixe  method  seems  to  inxolve  dramatic 
changes  in  cell  xx'ater  xolume.  Sexeral  desert 
species  haxe  been  obserx'ed  to  reduce  cell  xx'ater 
xolume  bx'  as  much  as  80%  xx'ithoutxxiltingxx'hen 
subjected  to  extremelx'  loxx'  soil  xxater  potentials 
(Moore  et  al.  1972,  Meinzer  et  al.  1988,  Evans 
et  al.  1991).  This  alloxx'ed  the  leaf  cells  to  have 
sufficiently  loxv  osmotic  potentials  for  xx'ater 
uptake  exen  though  solute  content  })er  cell  xx'as 
actually  reduced.  Although  total  solutes  per  leaf 
(and  presumablx  per  cell)  decreased,  the  rela- 
tix'e  concentrations  of  specific  solutes  changed 
(Evans  et  al.  1991)  such  that  "compatible" 
solutes  made  larger  contributions  to  the  osmotic 
potential  untk'r  stress.  Thus,  tolerance  of  loxv 
leaf  xxater  potentials  was  achieved  bv  a  combi- 
nation of  anatonncal  and  phxsiological  special- 
izations. The  anatomical  mechanisms  inxolxed 
in  this  magnitude  of  xolume  reduction  and  the 
im]ilied  cell  elasticitx'  haxe  not  been  studied,  but 
tlie  process  has  been  shown  to  be  fnllx  rexcrsible. 
Soil  texture^  is  also  an  important  factor  in 
determining  the  abilitx'  of  plant  connnunities  in 
a  coId-x\int('r  climate  to  respond  to  summer 
rain.  In  areas  xxith  moderate  lexels  of  precipita- 
tion, sandx'  soils,  because  of  their  loxx-  xxater- 
holding  capacitx.  nsuallx'  hold  a  sparser,  more 
drought-adapted  x  (^getation  than  finer-textured 
ones.  In  xvarm,  arid  areas,  however,  what  has 
been  called  the  "rexerse  texture"  effect  results 


204 


Great  Basin  Naturalist 


[\'oliime  52 


ill  the  more  liisli  xegetation  oceiirrintj;  in  tlie 
coarse-textured  soils.  This  occurs  because  the 
high  water-holding  capacit)'  of  fine-textured 
soils  allows  them  to  hold  all  the  moisture 
deri\'ed  from  a  single  rainfall  event  in  the  upper- 
most layers  of  the  soil  profile,  where  it  is  liigliK 
subject  to  direct  e\aporation  from  the  soil.  The 
same  amount  of  rainfall  entering  a  sandv  soil, 
precisely  because  of  its  low  \\'ater-h()lding 
capacity',  will  penetrate  to  a  much  greater  depth. 
It  is  also  less  likeK'  to  return  to  the  dning  surface 
b\'  capillaiv  action.  Less  of  the  rain  will  exapo- 
rate  from  the  soil  surface,  and  a  greater  fraction 
of  it  will  await  extraction  and  use  by  plants.  This 
inverse-textiu-e  effect  further  restricts  the  effec- 
tiveness of  summer  rains  in  the  fine  soils  of  the 
Great  Basin.  The  effect  is  less  operative  in 
respect  to  winter  precipitation  in  the  Great 
Basin,  however,  because  of  the  gradual  recharge 
of  the  soil  profile  to  considerable  depth  under 
conditions  where  surface  e\aporation  is  mini- 
mized by  cold  temperatures.  The  combination 
of  much  sandier  soils  and  greater  amounts  of 
summer  rainfall  in  the  region  of  the  Colorado 
Plateau  is  largely  responsible  for  the  major  flo- 
ristic  and  ecological  differences  bet\\'een  the 
two  regions.  At  lower  elevations  on  the  south- 
east edge  of  the  plateau,  shiid^-dominated 
desert  scnib  mav  be  replaced  by  grassland  dom- 
inated by  a  mix  of  spring-active  C5  and  summer- 
active  C4  species. 

ROOTINC;  DEPTH,  MORPHOLOGY,  AND  PHE- 
NOLOGY.— One  of  the  unique  and  ecologicallv 
most  important  features  of  the  Great  Basin 
shmb  communities  is  not  apparent  to  abo\e- 
ground  obseners.  This  is  the  investment  of  the 
vast  inajorit\-  of  plant  resources  in  the  growth, 
maintenance,  and  tunioxer  of  an  enormous  root 
system.  The  dominant  slinibs  of  the  Great  Basin 
usually  root  to  the  full  depth  of  the  winter-spring 
soil  moisture  recharge.  Depending  on  soil  tex- 
ture, slope  aspect,  and  elevation,  this  is  gener- 
ally between  1.0  and  3.0  m  (Dobrowolski  et  al. 
1990).  Although  this  represents  a  wide  range  of 
absolute  ck^pths,  nianv  of  the  ([ualitatixe  pat- 
terns of  rooting  behaxior  are  similar  on  most  of 
these  sites.  Ratios  of  rootishoot  standing  bio- 
mass  iang(^  from  4  to  7,  and  estimates  of 
root:shoot  annual  carbon  inxe.stment  are  as  high 
as  3.5.  Most  of  the  shrubs  ha\e  a  flexible,  gen- 
eralized root  system  with  dexelopment  of  both 
deep  taproots  and  laterals  near  the  surface. 
Moreover,  it  is  the  categon  of  fine  roots  <  3.0 
mm  in  diameter  that  constitutes  50-95%  (Cald- 


well et  al.  1977,  Sturges  1977)  of  the  total  root 
biomass.  The  veiy  large  annual  root  inxest- 
ments,  therefore,  are  not  primariK-  related  to 
large  storage  compartments,  but  to  the  tunioxer 
of  fine  roots  and  root  respiration  necessan-  for 
the  acquisition  of  water  and  mineral  nutrients. 

The  fine  root  network  thoroughK' permeates 
the  soil  x'olume.  Root  densities  are  grenerallv 
quite  high  throughout  the  upper  0.5  meters  of 
the  profile,  but  depth  of  maximum  root  devel- 
opment \aries  with  depth  of  spring  soil-mois- 
ture recharge,  species,  and  lateral  distance  from 
the  trunk  or  crowai.  A  particularly  high  densit)' 
in  the  first  0.1  m  ma\'  often  occur,  especially 
immediateh  under  the  shmb  canopx  (Branson 
1976,  Caldwell  et  al.  1977,  Dobrowolski  et  al. 
1990).  AlternatixeK',  maximal  densit)  mav  occur 
at  depths  between  0.2  m  and  0.5  m  (Sturges 
1980),  and  sometimes  a  second  zone  of  high  root 
densit}'  is  reported  at  depths  of  0.8  m 
(Daubenmire  1975)  to  1.5  ni  (Reynolds  and 
Fralev  1989).  Regardless  of  the  precise  depth  of 
maximum  rooting,  sliRib  root  densit\'  is  usualK' 
high  throughout  the  upper  0.5  m  and  then 
tapers  off,  but  max  continue  at  reduced  densi- 
ties to  much  greater  depth.  The  radius  of  lateral 
spread  is  usuallx'  much  greater  for  roots  ( 1-2  m) 
than  for  canopies  (0.1-0.5  m).  Percent  plant 
coxer  abox'eground  is  often  in  the  neighborhood 
of  25%  xxdth  75%  bare  ground,  but  beloxvground 
the  interspaces  are  filled  xvith  roots  throughout 
the  profile,  and  root  sxstems  of  adjacent  plants 
xxdll  overlap.  The  perennial  grasses  that  are 
potentiallv  co-dominant  xxith  shnibs  in  manx  of 
these  communities,  such  as  xxheatgrass 
{A^ropi/roii  sp.),  xx'iklne  (Eh/nui.s  sp.), 
squirreltail  {Sitaiiioii  liisti-ix).  Indian  ricegrass 
(On/zopsis  lu/i)icii()i(h:s).  and  galleta  grass 
{Hilaiia iainesii),  generallx  haxe  somexxhat  shal- 
loxxer  root  .sxstems  than  the  shrubs  (Branson  et 
al.  1976,  Rexiiolds  and  Fralex-  1989,  Dobro- 
xvolski  et  al.  1990).  Root  densities  of  grasses  are 
often  as  high  as  or  higher  than  those  of  shrubs 
in  the  upper  0.5  m  but  taper  off  more  rapidlx 
such  that  shnibs  usuallx  haxe  greater  densitx  at 
depth  and  greater  maximum  penetratit)n. 

The  moisture  resource  supporting  the  great- 
est amount  of  plant  groxx'th  is  usuallx-  the  xx'ater 
ston^l  in  the  soil  profile  during  the  xxinter.  This 
j)r()(ile  usuallx  has  a  positixe  balance,  xxith  more 
XX  ater  being  added  bx  precipitation  than  is  xxith- 
draxxn  bx'  exapotranspiration  bet\xeen  October 
and  March.  As  temperatures  xx-arm  in  March, 
exergreen  .species  nia\'  begin  draxxing  on  this 


19921 


Plant  Ai:)\rT\TK)\ 


205 


iiioistiiiT  resent",  ami  most  species  l)eii;iii  aetixc 
growth  ill  March  or  ApriL  Due  to  both  plant 
water  use  and  soil-surface  exaporation,  soil 
moisture  is  depleted  first  in  the  shallow  soil 
hncM's.  As  the  upper  layers  dr>',  plants  withdraw 
moisture  from  successively  deeper  soil  hners,  a 
proc(^ss  that  continues  in  e\ergreen  species 
throughout  the  summer  until  soil  moisture  is 
depleted  throughout  the  profile.  This  progres- 
sion of  seasonal  water  use  beginning  in  superfi- 
cial la\'ers  and  proceeding  to  deeper  soil  layers 
is  facilitated  In  the  pattern  of  fine  root  growtli, 
w  liicli  shows  a  similar  spatial  and  temporal  pat- 
tern (Fenuindez  and  (Caldwell  1975,  C'aldwell 
1976).  Root  growth  generalK  precedes  shoot 
growth  in  the  earl\-  spring  and  continues 
throughout  the  summer  in  e\ergreen  species, 
which  mav  appear  quiescent  abo\egroiind.  In 
annual  budgets  of  undisturbed  communities, 
.soil  moisture  withdrawal  almost  alwaxs 
approaches  measured  precipitation  o\  er  a  wide 
range  of  annual  fluctuations  in  total  precipita- 
tion, and  yew  little  moisture  is  lost  to  runoff  or 
deep  drainage  beneath  the  rooting  zone 
(Daubenmire  1975,  Caldwell  et  al.  1977, 
('ainbell  and  Harris  1977,  Sturges  1977).  Calcu- 
lati(  )ns  of  the  portion  of  exapotranspiration  actu- 
alK'  used  b\"  plants  in  transpiration  are  quite  high 
for  a  desert  enxironment  with  low  percent 
co\er;  they  range  from  50  to  75%  (Caldwell  et 
al.  1977,  Cambell  and  Harris  1977,  Sturges  1977). 
Competition  for  soil  moisture  is  not  neces- 
saril\-  limited  to  any  single  season.  Plant  water 
stress  is  highest  in  late  sunuuer,  and  siir\i\al  is 
most  likeK  to  be  influenced  at  this  time,  espe- 
cialK  if  one  plant  can  deplete  residual  soil  mois- 
ture below  the  tolerance  range  of  another.  This 
has  been  sliown  in  sexeral  cases  with  regard  to 
seedling  establishment  (Harris  1977,  DeLucia 
and  Schlesinger  1990,  Reichenberger  and  Pvke 
1990).  Growth  and  productivits"  are  more  likel\- 
to  be  affected  in  the  spring  and  earl\  summer 
growing  season.  This  is  because  most  of  the 
years  water  resource  is  alread\-  stored  in  the  soil 
in  earK  spring,  and  all  plants  are  drawing  on  a 
dwindling  resene  which  ultimateK  determines 
growing  season  length.  Competitixe  abilits'  is 
often  found  to  be  associated  with  an  abilit\  to 
begin  using  the  limiting  water  resource  earlier 
in  the  spring  (Eissenstat  and  Caldwell  19<SS, 
Miller  1988),  but  spatial  partitioning  is  also 
important.  Competition  ma\  be  most  intense  in 
shallower  soil  la\ers  because  grasses  and 
drought-deciduous    shrubs,   wiiicli    are    actixe 


oiiK  ill  the  spring,  are  shallower  rooted,  and 
lateral  root  spread  of  tlu^  e\ergreeii  species  is 
greatest  in  the  shallower  soil  la\ers.  The  more 
dominant  perennials  usualK  use  more  water 
o\er  the  whole  season  1)\-  tapping  deeper  soil 
la\ers  ((>line  et  al.  1977,  Sturges  1980)  and  are 
characterized  b\  higher  water-use  efficiencies 
(DeLucia  and  Sclilesinger  1990,  Smedlev  et  al. 
1991). 

Shnibs  appear  to  be  better  than  grasses  at 
withdrawing  water  from  deep  soil  laxers  for 
several  reasons.  In  shallow  soils  underlain  by 
fractured  or  porous  bedrock,  the  flexible,  mul- 
tiple taproot  structure  of  a  shrub  root  sxstem 
ma\"  be  better  suited  than  the  more  diffuse, 
fibrous  root  system  of  grasses  for  following 
cliinks  and  clea\age  planes  to  indeterminate 
depths.  This  should  allow  shnibs  to  better  cap- 
italize on  deep,  localized  pockets  of  moisture. 
Unfortunatelv  such  d\iiamics  are  rareK  studied 
quantitatixeK  because  of  the  difficult\"  of  mea- 
suring soil  moisture  budgets  or  root  distribu- 
tions under  such  conditions,  but  the\'  are 
implicated  b\'  plant  distribution  patterns  in 
man\  areas.  Indeed,  despite  the  \isiial  austeiit\' 
of  such  habitats,  rooting  into  major  joints  and 
cracks  in  bedrock  outcrops  can  create  sucli  a 
fa\"orable  microsite  b\'  concentration  of  ninoff 
in  localized  areas  that  ele\ational  limits  of  tree 
and  shrub  distributions  may  be  substantiallv 
lowered  as  would  be  expected  along  riparian 
corridors  or  other  unusnalK'  niesic  liabitats 
(Loope  1977).  Even  in  deep  soils,  shrubs  tend 
to  ha\e  deeper  root  svstems  than  grasses,  but,  in 
addition  to  this,  shiTibs  may  be  more  efficient 
than  grasses  at  extracting  deep  water.  Shiiibs  are 
sometimes  able  to  draw  on  deep  soil  moisture 
to  a  greater  extent  than  would  be  predicted  from 
root  biomass  distribution  alone  (Sturges  1980), 
and  this  is  due  in  part  to  an  intriguing  phenom- 
enon reported  b\-  Richards  and  Caldwell  ( 1 987), 
and  named  b\-  them  "Indraulic  lift."  During  the 
Iat(^  spring  and  earK  summer  most  ol  the 
remaining  soil  moisture  is  present  in  ckn-per  soil 
layers  wheic  rooting  (lensit\  ma\  be  relati\eK' 
low.  l^ue  to  low  (k'usities,  deep  roots  alone  ma\' 
be  unable  to  absorb  water  as  (juickl\-  as  it  is  lost 
l)\  the  tiaiisi)i  ling  shoot.  During  the  night,  water 
is  actnalK  ic^distributed  within  the  soil,  flowing 
from  deep  soil  lavers  through  the  roots  and  back 
out  into  shallower  soil  laxcrs.  This  is  the  phe- 
nomenon named  ■indraulic  lift."  and  the 
adxantage  to  the  plant  is  thought  to  be  a  reduc- 
tion in  the  rootiii'i  densitN    necessar\'  to  fully 


206 


Great  Basin  Naturalist 


[N'olume  52 


exploit  tlie  resources  of  the  deepest  soil  lavers. 
During  the  dav,  rates  of  water  uioxeiuent  from 
the  soil  into  the  roots  can  be  limiting  to  shoot 
activit)'.  Moistening  the  upper  soil  lavers  bv  noc- 
turnal h\draulic  lift  increases  the  root  surface 
area  for  al^soiption  during  the  periods  of  high 
transpiration.  Davtinie  water  use  can  he  sup- 
ported by  the  entire  root  system  and  not  just  b\' 
the  low-densitv  deep  roots  (Caldwell  and  Ricli- 
ards  f989). 

The  root  s\'stems  of  Great  Basin  shrubs  and 
Mojave  Desert  shrubs  differ  strongly  in  several 
ways.  (1)  Mojave  Desert  shiiilis  often  have  max- 
imal rooting  densities  at  soil  depths  of  0.1-0.3 
m,  and  maximmn  rf)ot  penetration  of  0.4-0.5  m 
(Wallace  et  al.  1980).  These  shallower  roots  are 
due  to  lower  rainfall  and  warmer  winter  temper- 
atures resulting  in  shallower  wetting  fronts  in 
the  soil,  and  the  de\  elopment  of  shallow  caliche 
layers.  (2)  Great  Basin  species  have  more  roots 
in  the  shallowest  0.1  m  soil  laver  (Wallace  et  al. 
1980,  Dobrowolski  et  al.  1990).  Differences  in 
soil  temperatures  ha\"e  been  used  to  explain 
these  patterns,  but  the  link  betvveen  cause  and 
effect  is  less  ob\ious  in  this  case,  and  conjec- 
tures should  be  \iewed  cautiouslv.  Much  hotter 
soil  temperatures  in  tlie  Moja\e  may  be  detri- 
mental to  roots  in  the  uppermost  few  centime- 
ters during  summer,  and  the  rapidly  di"ving  soil 
surface  may  be  too  ephemeral  a  moisture 
resoiu'ce  to  favor  large  investments  in  roots.  In 
contrast,  snowmelt  and  cooler  spring  tempera- 
tures may  result  in  less  rapid  evaporation  from 
the  soil  surface  in  the  Great  Basin  and  thus  fax  or 
more  rooting  l^v  perennials  in  that  zone.  (3) 
Because  of  the  greater  soil  volume  exploited,  as 
well  as  the  high  root  densitv  of  Great  Basin 
species,  their  ratios  of  rootishoot  biomass  are 
al)Out  twice  that  of  Moja\e  Desert  species 
( Bamberg  etal.  f  980,  Dobrowolski  et  al.  1990). 

Adaptation  to  salinity. — When  annual 
precipitation  levels  are  much  lower  than  poten- 
tial evaporation,  salts  are  not  leached  to  an\ 
great  depth,  and  they  can  accumulate  within  the 
root  zone.  This  is  especialK  important  in  associ- 
ation with  particular  bedrock  outcnps,  such  as 
the  Nhuicos  and  Ghinle  shales,  which  release 
high  concentrations  of  salts  during  weathering 
(Potter  et  al.  1985).  Precipitation  increases  with 
elexation,  and.  lollowing  major  storms  or  spring 
snowmelt,  there  may  be  surface  runoff  and 
recharge  of  groundwater  sy.stems  that  trans[)ort 
water  from  high  elexations  into  closed  basins. 
Streams  in  the  Great  Basin  usualK  terminate  in 


evaporati\e  pans  where  salinities  reach  extreme 
le\els  and  salts  precipitate  forming  a  surface 
crust.  The  water  table  near  these  evaporative 
pans  is  often  at  or  ven  near  the  sin-face  through- 
out the  \'ear,  l)ut,  if  there  is  no  groundwater  flow 
out  of  the  basin,  it  too  is  often  quite  saline 
(Dobrowolski  et  al.  1990).  Salinities  are  lowest 
on  slopes  and  at  higher  elevations  where  precip- 
itation exceeds  evaporative  loss,  and  they 
increase  on  more  level  terrain  and  in  lower-ele- 
\ation  basins  where  exaporation  exceeds  pre- 
cipitation. Sahnities  may  also  be  higher  in  areas 
where  wind-borne  materials  are  transported 
from  saline  playas  to  surrounding  slopes  (Young 
and  E\ans  f 986).  These  patterns  of  soil  salinitx' 
are  important  in  determining  plant  distribu- 
tions, with  more  specialized  salt-tolerant  spe- 
cies (halophvtes)  replacing  less-tolerant  species 
repeatedh  along  gradients  of  increasing  salinit)'. 
In  general,  species  diversity  is  low  on  saline 
soils.  The  vast  majorit)'  of  tolerant  shrub  species 
in  our  deserts,  and  all  the  shrubs  specifically 
mentioned  in  this  section,  lielong  to  a  single 
plant  family,  the  Chenopodiaceae  (goosefoot 
famiK).  Most  other  important  taxa  in  the  saline 
connmmities  are  grasses. 

In  the  most  extreme  case  of  h\persaline  salt 
flats  and  pans  there  may  be  standing  water  in 
the  wet  season  with  saturating  salt  concentra- 
tions. Under  such  conditions,  only  microflora 
consisting  of  a  few  species  of  photosMithetic 
flagellates,  cyanobacteria,  and  halobacteria  are 
commonly  found.  The  halobacteria  appear  to  be 
unique  in  having  adapted  in  an  obligate  manner 
to  the  high  salinities  of  these  environments. 
Thev  not  only  tolerate,  but  require,  high 
cvtoplasmic  salinities  for  membrane  stability 
and  proper  enzymatic  function  (Brown  1982). 
In  strong  contrast  to  this,  algae  and  all  higher 
plants  growing  in  hvper-saline  environments 
show  severe  inhibition  of  enzvnne  fvmction  at 
high  salinity,  and  thev  must  compartmentalize 
salt-sensitive  metabolic  processes  in  celhdar 
regions  of  low  ionic  strength  ( Muuus  et  al.  1982). 

The  best  definition  of  a  liahphvte  is  simply 
a  [)lant  tolerant  of  soil  salinities  that  would 
reduce  the  gi'owth  of  unspecialized  species.  This 
is  .somewhat  circular,  and  that  reflects  our  lim- 
ited understanding  of  how  halophv  tes  do  what 
thev  do.  Halophv  tes  are  more  likely  to  use  Na+ 
in  their  tissues  for  osmotic  adjustment,  while 
glvcophvtes  are  more  likely  to  have  high  K+ 
( Ilellebust  1976,  Flowers  et  al.  1977),  but  there 
are  munerous   exceptions.   Other  differences 


1992] 


Plant  Ai:)aptatk)n 


207 


max  he  nunc  (juaiititatixc  than  (|iialitati\ c  \  ar- 
ious  aspects  of  mineral  nutrition  in  halophx  tes 
are  less  sensitixe  to  high  soil  salinities,  hut 
unique  mechanisms  to  achiexe  this  tolerance 
ha\e  rareK'  heen  identified.  It  is  wideK  held  that 
the  ahilitv  to  compartmentalize  salts  and  restrict 
high  Na+  concentrations  to  the  \acuole  is  of 
crucial  importance  (Cakh\'ell  1974,  Flowers  et 
al.  1977,  linens  and  I.arhtM"  1982).  This  conclu- 
sion is  hased  primariK-  on  indirect  e\idence  of 
low  enz\nne  tolerance  of  salinitv;  howexer, 
rather  than  direct  measurements  of  actual  salt 
compartmentalization  (Munns  et  al.  19S2, 
Jefferies  and  Rudmik  I9S4). 

Haloplntes  differ  in  which  ions  reach  high 
tissue  concentrations  when  all  plants  are  grown 
on  the  same  medium  (Caldw^ell  1974).  Some 
will  concentrate  C1-,  for  instance,  while  others 
concentrate  S04~'.  These  differences  do  not 
necessarih'  determine  plant  distrilnitions,  such 
as  occurrence  in  soils  dominated  h)'  NaCl  \'ersus 
NaSOa,  but  rather  seem  to  reflect  different  reg- 
ulatoiA'  specializations  in  plant  metabolism 
(Moore  et  al.  1972).  A  strong  requirement  for  a 
uni([ue  composition  of  soil  salts  is  the  exception 
rather  than  the  mle,  and  the  most  important 
effect  of  soil  salinitv'  seems  to  be  a  disniption  of 
plant  water  relations  from  low  soil  osmotic 
potentials  rather  than  toxic  effects  of  specific 
ions.  Halophvtes  tolerate  these  conditions  bx' 
ha\ing  better  regulatoiA'  control  o\er  ion  mo\e- 
ment  within  the  plant,  ion  compartmentaliza- 
tion at  both  tissue  and  subcellular  lexels,  and 
better  homeostasis  of  other  a.spects  of  mineral 
nutrition  in  the  presence  of  ver\'  high  Na-K. 

Salinit\  poses  three  major  problems  for 
higher  plants.  First,  salts  in  the  .soil  solution 
contribute  an  osmotic  potential  depressing  the 
soil  water  potential,  and  this  ma\'  be  aggra\"ated 
as  salts  become  concentrated  with  soil  drving. 
E\en  when  sul)stantial  moisture  is  present, 
[)lant  tissues  must  endure  \  t-n  low  water  poten- 
tials to  take  it  up,  and  this  recjuires  a  specialized 
metabolism.  Second,  an\' salts  entering  the  plant 
with  the  transpiration  stream  will  be  left  behind 
in  the  leaf  intercellular  fluids  as  water  ('\a])()- 
rates  from  the  leaf.  This  can  result  in  salt 
buildup  in  the  intercellular  solution  causing 
water  moxement  out  of  the  cells  and  leading  to 
cellular  dehxdration.  Third,  salts  entering  the 
cxtoplasm  in  high  concentration  will  disrupt 
enz)ine  function.  Haloph\1:es  are  able  to  deal 

with  all  of  these  factors  over  a  wide  range  of  soil 
1.   .  . 

salinities.  Haloph\tes  show  a  greater  capacit\' 


for  osmotic  adjustment,  and  positixe  phot()s\n- 
thetic  rates  can  be  measured  in  the  leaxes  of 
man\  haloplntes  at  leaf  water  potentials  as  low 
as  -90  to  - 120  bars  (Caldwell  1974),  well  below 
the  range  that  would  result  in  death  of  e\en 
desert-adaj)ted  gl\coph\tes.  Tliis  is  accom- 
plisluHl  in  part  1)\'  transforming  the  available 
salts  in  the  enxironment  into  a  resource  and 
using  them  for  osmotica  in  j)lant  ti.ssues  (Moore 
et  ak  1972,  Bemiert  and  Schmidt  1984).  Many 
haloplntes  actualK  show  stimulation  of  growth 
rates  at  moderate^  en\  ironmental  salt  levels. 

Halophvtes  too  must  deal  with  the  problem 
of  salt  buildup  in  the  leaves,  and  the\'  do  so  by  a 
wide  \ariet\'  of  processes.  There  is  a  great  deal 
of  interspecific  \  ariation  in  which  method!  s )  are 
used.  All  the  methods  appear  to  incur  substan- 
tial energetic  costs  associated  with  maintaining 
high  ion  concentration  gradients  across  key 
membranes  (Kramer  1983).  Exclusion  of  salts  at 
the  root  is  possible;  this  is  the  method  most 
employed  by  winterfat  (Ceratoides  Janata).  Salt- 
bush  (Atriplex  spp.)  has  specialized  hair-blad- 
ders on  the  leaf  surface  into  which  e.xcess  salts 
are  actively  pumped.  The  hairs  e\'entualK'  nip- 
ture,  excreting  the  salts  to  the  outside.  Other 
plants  may  transport  salts  back  to  the  root  \ia 
the  phloem.  Man\-  plants  exhil)it  increased  leaf 
succulence  when  growii  under  high  salinit\;  and 
this  increase  in  cell  xolume  can  create  a  sink  for 
salts  within  the  leaf  without  raising  salt  concen- 
trations or  furtlier  lowering  leaf  osmotic  potential. 

hi  strong  contrast  to  the  exident  importance 
of  temperature  and  rainfall  pattern  in  favoring 
C:5  versus  C4  grasses,  Ci  shnibs  tend  to  belong 
to  edaphic  comnumities  as.sociated  with  saline 
soils.  The  same  species  ma\'  occur  in  both  warm 
and  cold  deserts,  and  in  areas  with  both  winter 
and  summer  rainfall  patterns.  This  is  an  intri- 
guing difference,  but  the  phwsiological  basis 
linking  C,  shrubs  with  high  salinitv'  is  less  well 
understood  than  the  tradeoffs  associated  with 
temperature  and  controlling  C5  and  C^  grass 
distributions.  Sjx'cies  number  and  percent 
cover  b\  shrubs  sucli  as  saltbush  {Ahiplcx  spp.) 
and  inkA\'eed  (Siicda  spp. ),  wliich  possess  the  C4 
pathwav,  usualK  inc-rc^ise  drainaticallv  with 
increasing  salinitv  on  w(41-drained  soils.  In 
marshx'  habitats  or  soils  with  a  shallow,  saline 
water  table,  howex  er.  haloplntic  (>-,  species  such 
as  pickleweeds  {Allen rolfia  spp.  and  Saliconiia 
spp.)  and  greasewood  [Sarcohatus  ver- 
micnloicles)  regain  dominance.  It  has  been  sug- 
gested that  hitrher  water-use  efficiencv  bv  C4 


208 


Great  Basin  Naturalist 


[\  nluiiie  52 


species  niav  be  acKantageous  on  saline  soils  to 
help  avoid  salt  bnildnp  in  the  leaf  tissues.  How- 
ever, it  has  not  been  showii  that  transpiration 
rate  is  an  important  factor  controlling  salt 
buildup  in  the  leaves  of  halopln  tes  when  com- 
pared wath  other  regulaton'  mechanisms 
(Osmond  et  al.  1982),  nor  does  this  Inpothesis 
explain  the  dominance  of  C3  species  in  wet 
saline  soils.  In  the  greasewood  and  pickleweed 
commimities,  soil  salinities  are  extreme,  but 
soils  remain  wet  throughout  the  growing  season, 
or  else  groundwater  is  available  within  the  root- 
ing zone  (Detling  1969,  Hesla  1984).  As  a  con- 
sequence, plant  water  potentials  do  not  reach 
the  extreme  low  values  of  the  saltbush  commu- 
nities. Over  a  wide  range  of  soil  salinities,  plants 
such  as  greasewood  appear  to  draw  on  readily 
available  deep  soil  moisture,  and  high  leaf  con- 
ductances are  maintained  throughout  the 
summer  (Hesla  1984,  Romo  and  Hafercamp 
1989).  The  highest  whole-plant  water-use  rates 
may  occur  in  the  presence  of  high  soil  salinitv"  in 
mid-summer  (Hesla  1984).  The  communities  in 
which  C4  shRil:)s  are  most  prevalent  have 
summer  stress  from  both  high  soil  salinitv  and 
mid-sunnner  soil  water  depletion  combined. 
These  species  reach  much  lower  plant  water 
potentials  during  summer  than  either  nonsaline 
communities  or  wet-saline  ccnnmunities.  The 
role  of  C4  pliotosviithesis  in  tolerating  these 
conditions  remains  to  be  determined,  but  it 
could  he  related  to  avoiding  excessively  low  leaf 
water  potentials  either  liy  (1)  retarding  soil 
moisture  depletion,  which  both  lowers  the  soil 
matrix  potential  and  concentrates  soil  salts,  or 
(2)  avoiding  exacerbation  of  low  soil  water 
potentials  due  to  high  flux  rates  and  large  water 
potential  gradients  between  the  leaf  and  root. 
Water  mo\ement  in  the  x-xlem  occurs  under 
tension,  and  anatomical  features  that  avoid  cav- 
itation in  the  xylem  at  extreme]\  low  water 
potentials  usually  reduce  the  hydraulic  conduc- 
tivity of  the  x"\'lem  per  unit  cross-sectional  area 
(Davis  et  al.  1990,  Speny  and  Tyree  1990).  Low 
specific  c()nducti\it\'  of  the  xTlem  will,  in  turn, 
predispose  the  plant  system  to  large  water 
potential  gradients  between  roots  and  shoots, 
causing  an  even  greater  depression  of  leaf  water 
potential.  This  problem  could  be  ameliorated 
either  by  increased  cross-sectional  area  of  the 
xylem  by  increased  allocation  to  wood  growth, 
or  by  features  such  as  C.|  photos\Tithesis  that 
reduce  the  flux  rate  of  water  associated  with 
photosN  nthetic  acti\it>  under  warm  conditions. 


Nutrient  Relations 

Acquisition  of  mineral  nutrients. — 
Apart  from  the  veiy  high  elevation  montane 
zones,  water  appears  to  be  the  most  limiting 
resource  in  the  Great  Basin  and  Colorado  Pla- 
teau communities.  Productixit)  is  usualK  well 
correlated  with  yearlv  fluctuations  in  precipita- 
tion and  spring  moisture  recharge  over  a  wide 
range  of  \alues  (Daubenmire  1975,  Kindschy 
1982),  and  competitive  success  has  more  often 
been  associated  with  soil  water  use  patterns 
than  nutrient  budgets.  Nonetheless,  addition  of 
mineral  fertilizer  sometimes  does  result  in 
modest  increases  in  producti\it\',  and  studies 
ha\e  shown  strong  effects  of  neighboring  plants 
on  nutrient  uptake  rates  (Caldwell  et  al.  1987). 
These  dynamics  have  been  less  studied  than 
have  plant  water  budgets,  and  broad  ecological 
relationships  are  just  now  being  worked  out  in 
detail.  Nutrient  acquisition  has  been  showni  to 
be  a  major  factor  determining  communits'  com- 
position only  in  veiy  special  habitats  such  as 
large  sand  dunes  (Bowers  1982)  or  unusual  bed- 
rock (DeLucia  and  Schlesinger  1990). 

MiCROPHYTIC  CRUSTS. — Throughout  the 
Great  Basin  and  Colorado  Plateau,  it  is  common 
for  the  exposed  soil  surface  to  l)e  covered  by  a 
complex  connniuiit\'  of  nonvascular  plants 
including  dozens  of  species  of  algae,  lichens, 
and  mosses  (West  199()).  These  organisms  often 
form  a  biotic  ciTist  in  the  upper  few  centimeters 
of  the  soil  and,  when  undisturbed,  may  result  in 
a  vei"y  conx'oluted  microtopograplu'  of  the  sur- 
face. While  a  detailed  discussion  of  the 
microplutic  crusts  is  bcNond  the  scope  of  this 
review,  it  is  important  to  realize  that  percent 
cover  by  such  crusts  often  exceeds  that  of  the 
vascular  plants,  and  their  contribution  to  total 
ecosvstem  prochicti\itA'  is  consitlerable.  Perhaps 
most  important  to  co-occurring  \ascular  plants 
are  the  nutrient  inputs  to  the  soil  b\'  nitrogen- 
fixing  cnist  organisms  (c\anobacteria  and 
lichens).  These  inputs  will  be  particularlv 
important  in  the  cold  deseit  where  fewxascular 
plants  form  sMiibiotic  relationships  with  nitro- 
gen-fixing bacteria. 

Nurse  plants  and  fertile  islands. — In 

man\  des(Mt  areas,  including  both  the  Mojave 
and  the  Great  Basin,  establishment  of  newindi- 
\iduals  may  occur  preferentialK'  under  the  exist- 
ing canopies  of  alreadv  established  indi\iduals. 
Tliese  pre\iousl\'  established  indixidnals  mav 
tlieu  be  referred  to  as  nurse  plants.  Preferential 


1992] 


Pl.WT  Ai:)MT\TION 


209 


estahlisliiiK'nt  inulcr  iiiirsc  plants  nia\  ocfiir  in 
spite  of  the  fact  that  759ic  or  more  of  tiie  gromid 
area  nia\'  he  liare  interspaces  b(^t\\'een  plant 
canopies.  The  phenomenon  can  he  important  in 
both  steadx-state  commnnitA  cl\  namics  and  also 
snccessional  patterns  following  distnrbance 
(Wallace  and  Ronme\-  1980,  Exerett  and  Ward 
1984).  Two  .somewhat  distinct  factors  contiibntc^ 
to  the  nnrse-plant  phenomenon.  The  first  has  to 
do  the  with  beneficial  effects  of  partial  shading 
and  rednced  wind  nnder  existing  canopies 
resulting  in  cooler  temperatnres  and  possibK' 
moister  soil  conditions  in  the  snrface  huers. 
These  interactions  depend  directk-  on  the  pres- 
ence of  the  nnrse  plant  in  creating  a  fa\orable 
microsite,  and  ha\e  been  studied  with  particular 
reference  to  pin\on  and  juniper  establishment 
in  the  Great  Ba.sin.  A  second  factor  inxoKes  the 
creation  of  fertile  islands  bv  the  prolonged  occu- 
pation of  the  same  microsite  b\'  man\'  genera- 
tions of  plants;  this  can  make  the  fertile  island  a 
preferred  site  even  if  the  previous  occupant  is 
alreacK  deceased.  This  microsite  impro\'ement 
occurs  due  to  preferential  litter  accumulation 
and  more  e.xtensixe  root  growth  directK  under 
a  plant  canopw  and  deposition  of  aeolian  mate- 
rials under  reduced  wind  speeds  in  plant  cano- 
pies. In  time,  soils  nnder  existing  plants  mav 
come  to  be  slightK'  raised  above  the  interspace 
level,  have  a  lighter,  loaniier  texture,  higher 
organic  matter  content  and  better  soil  structure, 
less  surface  compaction,  better  aeration  and 
more  rapid  water  infiltration,  and/or  higher 
l('\els  of  available  mineral  nutrients  than 
immediatelv  adjacent  interspace  soil  (A'est  1962, 
Wood  et  ai  1978,  Homnev  et  al.  1980,  Hesla 
1984,  West  1989,  Dobrowolski  et  al.  1990). 
Direct  effects  of  nurse  plants  and  indirect 
effects  of  fertile  islands  should  complement  and 
reinforce  each  other  in  maintaining  selective 
spacial  patterns  of  seedling  establishment.  Sur- 
face soil  nnder  haloplntes  mav  also  show- 
increased  salinitv  (Richard and  (]line  1965)  due 
to  excretion  ol  excess  salts  bv  the  canopv  or 
translocation  and  re-excretion  Ironi  the  roots. 

DiXER.SITY  OP^  Ghowtii  Foinis 

One  of  the  striking  features  of  the  cold  desert 
vegetation  is  the  uniformlv  low  stature  of  the 
vegetation.  This  is  undoubtedlv  due  to  several 
factors,  and  few  studies  have  specificallv 
addressed  the  role  of  plant  stature  in  these  com- 
munities.   Since   low  temperatures   mav    limit 


photosx  nllicsis  in  tlic  cool  spiiirj;.  and  earlier 
growth  on  limited  soil  moisture  resen(\s  mav  be 
c-()mp(titi\c'l\  advantageous,  occupving  warm 
microsites  mav  be  favored.  Substantial  increases 
in  air  temperature  and  reductions  in  wind  speed 
will  exist  in  the  lowest  meter  next  to  the  ground, 
and  especiallv  in  the  lowest  decimeter.  Low 
cushion  plants  oi-  low.  dense  shrub  canopies 
should  have  vvarmei"  spring  leaf  temperatures  by 
virtue  of  being  short  and  bv  virtue  of  leafing  out 
first  in  a  dense  clump  of  old  dead  leaves  and 
twigs  ( Smith  et  al.  1983,  Wilson  et  al.  1987).  This 
advantage  mav  be  partiallv  offset  by  overlv  high 
temperatures  in  summer  for  species  remaining 
active  all  sununer.  Stature  is  also  likelv  to  affect 
aeolian  deposit  of  materials  under  the  shrub 
canopies  (W^ood  et  al.  1978,  Young  and  Evans 
1986),  snow  accumulation  (Branson  et  al.  1981, 
West  and  Caldwell  1983),  and  the  likelihood  of 
winter  desiccation  under  cold,  windv  conditions 
(Nelson  and  Tienian  1983).  All  of  these  could 
be  important  factors,  but  few  detailed  studies 
have  been  done. 

Having  considered  tlie  relationships  of  the 
dominant  plant  habits  and  phenologies  to  cli- 
mate, it  is  perhaps  instructive  to  consider  whv 
.some  of  the  other  famous  desert  life  forms  are 
so  poorlv  represented  in  this  region.  Three  life 
forms  vvliich  are  prominent  features  of  the  warm 
desert  but  inconspicuous  elements  of  the  cold 
desert  are  (1 )  large  CAM  succulents  (e.g.,  cacti 
and  agave),  (2)  opportunistic  drought-decidu- 
ous shnibs  specialized  for  rapid  knif-flnshing, 
and  (3)  animals.  Definitive  work  explaining  the 
structural  nnilormitv  of  the  vegetation  is  not 
available,  but  the  environment  is  well  enough 
understood  to  identifv  at  least  some  of  tlu^  likelv 
causes. 

CAMSrcci'I.KXTS — Most  of  the  large  C.\.\I 
succulents  are  not  tolciant  ol  freezing  temper- 
atures, and  most  extant  species  would  be 
excluded  from  the  (jrcat  Basin  bv  this  factor 
alone.  Thei'e  ai"e,  however,  a  sulfitienl  mimbcr 
of  species  which  have  adapted  to  tolerate  cold 
temperatures  that  we  are  justified  in  asking  whv 
thev  have  not  undergone  more  adaptive  radia- 
tion, or  claimed  a  more  prominent  role  in  these 
communities.  The  most  important  factor  limit- 
ing this  life  form  is  probably  the  importance  of 
the  cool  spring  growing  season.  CAM  succu- 
lents generallv  ( 1 )  allocate  ven  little  biomass  to 
root  (root/shoot  ca.  0.1),  (2)  are  shallow  rooted, 
(3)  store  moderate-sized  (compared  to  soil 
v\ater-liolding  capacitv )  water  resenes  inside 


210 


Great  Basin  Naturalist 


[Volume  52 


their  tissues  wheu  water  is  available  in  the  sur- 
face soil  layers,  and  (4)  use  their  stored  water  in 
photosynthesis  with  unparalleled  water-use  effi- 
ciency by  opening  their  stomata  only  at  night 
when  temperatures  are  cool  (Nobel  1988).  They 
are  fa\ored  bv  (1)  very  warm  days  (30-40  C), 
which  allow  them  to  have  higher  photosyiithetic 
rates  and  cause  competing  species  to  ha\-e  very 
low  water-use  efficiencies;  (2)  large  diunial  tem- 
perature fluctuations  allowing  for  cool  nights 
(10-20  C)  which  allow  them  to  have  high  rates 
of  CO2  uptake  with  high  water-use  efficiency; 
and  (3)  intermittent  rainfall  \\'hich  onl\' wets  the 
upper  soil  layers  so  that  the  limitations  of  their 
shallow  roots  and  water-hoarding  strategy  are 
compensated  foib\  the  ephemeral  natiu-e  of  the 
soil  water  resoiu'ce.  These  conditions  are  some- 
what poorK"  met  in  the  cold  desert.  The  impor- 
tant water  resource  is  one  of  deep  soil  recharge 
that  favors  deep-rooted  species  and  confers 
much  less  advantage  on  internal  water  hoarding. 
Freezing  tolerance  in  CAM  succulents  appears 
to  be  associated  with  low  tissue  water  contents, 
and  this  mav  inhibit  uptake  of  water  when  it  is 
plentiful  in  the  siu-face  layers  in  the  thermalK' 
x'acillating  eark'  spring  (Littlejohn  and  Williams 
1983).  Furthermore,  water-use  efficiencies  of 
C3  and  C4  species  are  quite  high  in  the  cool 
spring. 

Nonetheless,  even  moderate  amounts  of 
summer  rain  in  the  southern  and  eastern  por- 
tions of  the  Great  Basin  result  in  numerous 
species  of  cacti.  Due  to  the  open  nature  of  the 
understoiy,  many  of  these  species  ha\e  a  large 
elevational  range,  and  they  are  often  more 
common  in  the  pinvon-juniper  or  even  the  mon- 
tane zone  than  on  the  desert  piedmont  slopes. 
Almost  all  of  these  cacti  are  small,  usually  5-20 
cm  liidi,  with  a  small,  globose  (e.g.,  Pediocactiis 
siinpsonU),  prostrate  (e.g.,  Opiintia  pohj- 
cantha),  or  low,  caespitose  habit  (e.g., 
Echinoccreus  tn<4ochidi(itus).  This  allows  them 
to  take  acKantage  of  the  warmer  da\time  tem- 
peratures near  the  ground  in  the  sj^ring  and 
facilitates  an  insulating  snowcover  during  the 
coldest  winter  periods.  The  number  of  and  total 
cover  by  cacti  increase  considerabK  with 
increased  summer  rainfall  on  the  Colorack)  Pla- 
teau, but  oulv  in  the  eastern  Mojave  with  both 
summer  rain  and  warm  spring  tempcratun^s  do 
we  find  the  larger  barrel-cactus  (e.g.,  Fcrocddiis 
acanfhoidcs)  and  tall,  shnibb)^  chollas  (e.g., 
Opinilid  (ic(i)ithiH'arpa). 


Opportunistic   drought-deciduous  / 

MULTIPLE  LEAF-FLUSHING  SPECIES. — This 
habit,  like  that  of  the  succulents,  is  favored  by 
( 1 )  intermittent  rainfall  wetting  only  shallower 
soil  layers,  and  (2)  warm  temperatures  allowing 
for  rapid  leaf  expansion  in  response  to  renew/ed 
soil  moisture.  Again,  these  requirements  are  not 
well  met  in  the  Great  Basin.  The  priman'  mois- 
ture resource  is  a  single,  deep  recharge  in  the 
winter.  Most  shiaib  species  are  deep  rooted,  and 
rather  than  experiencing  \acillating  water  avail- 
abiHtv',  they  have  actixe  root  grow1:h  shifting  to 
deeper  and  deeper  soil  kners  during  the  season, 
thus  producing  a  gradual  and  continuous 
change  in  plant  water  status.  This  allows  manv 
spring-active  shrubs  to  remain  partially  ever- 
green throughout  the  summer,  and,  in  regions 
where  it  occurs,  the\'  are  able  to  make  rapid  use 
of  anv  moisture  availalole  from  simimer  precip- 
itation without  the  need  for  renewed  leaf  pro- 
duction. The  only  shrub  reported  to  ha\'e 
)iiultiple  leaf  flushes  in  response  to  late  spring 
or  summer  rain  in  the  Great  Basin  is  the  dimin- 
utive and  shallow^- rooted  Artemisia  spinescens 
(Everett  et  ak,  1980).  Some  species  found  in  the 
Great  Basin  are  reported  to  have  multiple 
growth  c\'cles/year  where  they  occur  in  the 
Mojave  (Ackerman  et  ak  1980). 

ANNUALS  AND  LIFE-HISTORY  DIVERSITY. — 
The  spectacular  wildflower  show^s  displayed  in 
favorable  years  in  the  Mojave  Desert  do  not 
occm-  in  the  cold  desert  of  the  Great  Basin 
(Ludwig  et  ak  1988).  Annu;il  species  are  few  in 
nimiber,  and,  except  in  earK"  succession  after 
fire  in  woodlands  or  on  \en  disturbed  sites,  the)' 
rarely  constitute  a  major  fraction  of  total  com- 
munit>'  biomass.  This  is  undoubtedly  related  to 
sexeral  complex  factors,  but  various  aspects  of 
precipitation  patterns  are  likeK'  to  be  among  the 
most  important.  To  begin  with,  the  paucits'  of 
summer  rain  in  some  parts  of  the  Great  Basin 
ma\  largeh'  eliminate  an  entire  class  of  C4 
siuumer  annuals  important  in  the  floras  of  other 
regions  including  the  Cok)rado  Plateau.  Other 
aspects  than  seasonalih'  are  also  cnicial,  how- 
e\er.  Ver\  low  means  oi  annual  precipitation  are 
conunonh'  associated  with  large  annual  floras, 
but  correlated  with  low  mean  precipitation  is 
high  \ear-to-\ ear  \ariation  in  precipitation 
which  some  authors  have  argued  is  equally 
important.  The  coefficient  of  xariation  (CV)  in 
precij)itation  shows  a  r(4ationship  to  mean  pre- 
cipitation in  the  (wvat  Basin  and  Colorado  Pla- 
teau (Fig.  2)  veiv  similar  to  that  found  in  warm 


19921 


Plant  Ad  AinviioN 


211 


0.7 

o 

0.6 

t    ■ 

m 

';z 

O.S 

ro 

> 

B 

0.4 

,    ■ 

r 

CD 

0.3 

o 

■ , 

CI) 

0.2 

o 

O 

0.1 

, 1 -     T                             1 

3 

r2  =  .57 

- 

O 

- 

\oO                    cj^ 

s     ° 
o 

^ 

o   o°o         ^ — ~-- 
o 

o 

- 

" 

0.0 

100  200  300  400  500 

Mean  annual  precipitation,  mm 

Fig.  2.  The  relatioii.ship  hctween  mean  precipitation  and 
the  \ariabilit\'  of  rainfall  between  years  as  nie;isured  h\-  the 
coefficient  oi  \ariation  in  annual  precipitation.  The  data 
inchide  points  scattered  throughout  the  Great  Basin  in  Utali 
and  Xe\ada  and  the  Colorado  Plateau  in  Utah  and  Arizona. 
The  line  shown  is  the  least  squares  best  fit  for  the  data:  C\' 
=  1.27  -  0.403  °  log(niean  annual  precipitation,  mm)  {ii  = 
69 sites./)  <  .001). 

deserts  (Ehleringer  1985).  Although  mean  pre- 
cipitation has  tlie  greatest  single  effect,  there 
are.  aclclitionalK;  important  geographic  influ- 
ences on  the  CV  of  precipitation  which  are 
independent  of  mean  precipitation.  A  multiple 
regression  of  the  CV  of  precipitation  on 
logdnean  annual  precipitation),  latitude,  and 
elexation  in  the  Great  Basin  has  an  r  of  .81  and 
indicates  that  each  \arial)le  in  tlie  model  is 
highl\-  significant  (j)  <  .001  or  better).  For  a 
given  mean  precipitation,  the  C\^ increases  with 
decreasing  altitude  in  the  Great  Basin,  but  an 
ind(^p(Mident  effect  of  elevation  was  not  signifi- 
cant in  the  Colorado  Plateau.  The  CV  also 
increases  from  north  to  south  in  the  Great  Basin 
and  increases  from  south  to  north  in  the  CJolo- 
rado  Plateau,  which  results  in  a  latitudinal  band 
of  greatest  annual  \arial)ility  nuniing  through 
southern  Nevada  and  Utah.  This  l)and  is  related 
to  two  major  as]:)ects  of  regional  climate.  Mo\ing 
southward  in  the  Great  Basin,  temperatures 
gradually  increase,  favoring  moister  air  masses 
and  more  intense  storms,  but  sites  are  morc^ 
remoxed  from  the  most  common  winter  storm 
tracks,  and  the  number  of  rainv  daws  per  Near 
decreases  (Houghton  1969).  Moving  northward 
from  Arizona  and  New  Mexico,  the  southern 
Nevada  and  Utah  band  of  highest  precipitation 
variabilit\-  also  corresponds  to  the  northenunost 
extent  of  summer  storms  associated  with  tlie 


0  2  0  3  0  4 

CV  for  mean  annual 
precipitation 

F"ig.  3.  The  relationship  between  relial)ilit\-  of  annual 
precipitation  and  life-histon'  strategy  of  herbaceous  plants. 
The  site  with  greatest  representation  of  annuals  is  Deadi 
\'allev  in  the  Moja\e  De.sert,  the  second  highest  is  C^auNon- 
lands  in  the  Colorado  Plateau  of  southeastern  Utah,  and  the 
other  three  sites  aie  Great  Basin  Cold  Desert  or  shrub- 
steppe  (data  were  collected  b\'  Kim  Haiper  and  pnniously 
published  in  Schaffer  and  Gadgil  197.5). 


Arizona  monsoon,  and  the  region  where  the 
fraction  of  summer  rain  increases  substantially 
moving  southward.  This  zone  also  has  some  of 
the  most  arid  sites  of  the  entire  region  located 
along  the  transition  to  the  Mojave  Desert  in 
southern  Nevada  and  the  canyon  countiy  of 
southeastern  Utah,  and  these  sites  can  be 
expected  to  have  the  highest  variabilit\  due  to 
both  low  mean  rainfall  and  geographic  position 
correlated  with  rc^gional  weather  patterns. 
Becau.se  the  (ireat  Basin  and  Colorado  Plateau 
are  only  .semiarid,  the  CV  ol  annual  precipita- 
tion is  not  usually  as  high  as  in  manv  of  the  more 
arid  warm  deserts  (Beatley  1975,  Ehleringer 
1985),  but  particular  sites  may  be  both  arid  and 
highlv  unpn^dictable. 

Ilaiper  (cited  in  Schaffer  and  Ciadgil  1975) 
found  that  the  prevalence  of  annuals  was  posi- 
ti\eK  associated  with  the  C\'  in  annual  precipi- 
tation for  five  sites  located  in  the  Great  Basin, 
Colorado  Plateau,  and  Moja\e  Desert  (Fig.  3). 
The  largest  annual  populations  occurred  in 
Death  Valley  (Mojave),  followed  by  Canyon- 
lands  (Colorado  Plateau  in  southeastern  Utah). 
One  inteipretation  of  this  relationship  is  that 
high  \ariabilit\-  in  total  precipitation  between 
vears  mav  be  associated  v\  ith  high  rates  of  mor- 
tality and  therefore  favor  earlv  reproduction  and 
an  annual  habit  (Schaffer  and  Gadgil  1975). 
Manv  desert  annuals  are  facultativ  elv  perennial 
in  better-than-averaee  vears,  and  some  have 


212 


Great  Basin  Naturalist 


[Volume  52 


perennial  races  or  sister  species  (Ehleringer 
1985).  The  dynamics  and  distributions  of  these 
closely  related  annual  and  perennial  taxa  should 
receive  further  study  in  regard  to  their  expected 
life  span,  reproductive  output,  and  relationships 
to  climatic  predictability'.  Another  perspecti\'e  is 
to  ask  how  competition  between  very  distinct 
shnib  and  annual  species  is  affected  by  precip- 
itation variability.  While  in  many  respects  com- 
plementaiy  with  the  optimal  life  histoiy 
arguments,  this  approach  emphasizes  how  large 
differences  in  habit  affect  resource  capture  and 
competition  rather  than  focusing  on  subtler  dif- 
ferences in  mortalit)'  and  reproductive  sched- 
ules. The  lower  variability  of  precipitation  in 
much  of  the  Great  Basin  compared  to  the 
Mojave  and  Sonoran  deserts,  as  well  as  the  more 
reliable  accumulation  of  moisture  during  the 
winter-recharge  season,  may  favor  both  stable 
demographic  patterns  and  growth  of  perennials. 
Annuals  tend  to  be  shallow  rooted  (most  roots 
in  upper  0.1  m  depth),  and  they  are  poorly 
equipped  to  compete  with  shrubs  for  deep  soil 
moisture.  If  shrub  density  is  high,  and  years  of 
unusually  high  mortality'  are  rare,  then  shiaibs 
may  largely  preempt  the  critical  water  and  min- 
eral resources  and  suppress  growth  of  annuals. 
The  dominant  shrubs  of  the  warm  deserts  do  not 
have  high  root  densities  in  the  upper  10  cm  of 
the  soil  profile  (Wallace  et  al.  1980),  have  lower 
total  root  densities,  and  have  lower  total  cover 
when  compared  with  Great  Basin  perennials. 
Annuals  are  therefore  likely  to  experience  more 
intense  competition  from  shnibs  in  the  Great 
Basin.  This  conjecture  is  finther  supported  by 
considering  that  perennials  in  the  Great  Basin 
generally  transpire  50%  or  more  of  the  ammal 
moisture  input  over  a  wide  range  of  yearly  vari- 
ations. In  the  Mojave  this  fraction  may  average 
only  27%  and  vary  between  years  from  15  to 
50%  at  the  same  site  (Lane  et  al.  1984),  or  even 
be  as  low  as  7%  (Sanimis  and  Gay  1979).  The 
reduced  overlap  in  rooting  profiles  and  the 
greater  availability  of  unused  moisture 
resources  may  have  favored  the  development  of 
annual  floras  in  the  Mojave  Desert  more  than  in 
the  Great  Basin.  With  severe  distin-bance  from 
grazing  and  other  anthropogenic  activities, 
exotic  annual  species  have  invaded  many  Great 
Basin  communities.  Once  established  following 
distiubance,  these  annuals  are  not  always  easily 
displaced  by  short-tenu  shrub  succession.  While 
this  discussion  has  been  presented  in  the  con- 
text  of  annuals   versus   perennials,   tradeoffs 


lietween  short-  and  long-lived  perennials  may 
be  influenced  by  very  similar  climatic  parameters, 
sometimes  operating  over  different  time  scales. 

Other  factors  that  may  be  important  in  the 
ecolog)'  of  Great  Basin  annuals  include  the 
effects  of  the  very  well  developed  ciyptogam 
soil  cRists  or  vesicular  horizons  on  seed  preda- 
tion  (abilit)'  of  seeds  to  find  safe  sites),  seed 
germination,  and  seedling  establishment.  The 
restriction  of  winter  growth  by  cold  tempera- 
tures could  also  be  of  crucial  importance,  inhib- 
iting the  prolonged  establishment  period 
enjoyed  by  winter  annuals  in  warm  deserts.  Fall 
germination  followed  by  low  levels  of  photos)ii- 
thesis  throughout  the  mild  winter  is  essential  for 
\igorous  spring  growth  of  winter  annuals  in  the 
Mojave,  and,  while  heavy  spring  rains  may  cause 
germination,  such  late  cohorts  rarely  reach 
maturity  (Beatley  1974).  Annuals  are  common 
in  transition  zone  sites  of  the  ecotone  between 
Mojave  Desert  and  Great  Basin  plant  commu- 
nities in  southern  Nevada,  but  associated  with 
changes  in  perennial  species  composition  along 
decreasing  mean  temperatiu'e  gradients  in  that 
region  are  decreases  in  annual  abundance 
(Beatley  1975). 

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Sturges.  D.  L.  1977.  Soil  water  wididrawal  and  root  char- 
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under  gnibbed,  spraved,  and  undisturbed  big  sage- 
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W'ilev  ;ind  Sons,  Inc.,  New  York. 

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Wallace.  A.,  and  E.  M.  Ro.mney  1980.  The  role  of  pioneer 
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Wallace  A.,  E.  M.  Romney.  and  J.  \\'.  Cha  1980.  Depth 
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wildliuid  ectjsv'Stems  of  arid  to  semi-arid  regions.  Pages 
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West,  N.  E.,  and  J.  Gastro  1978.  Phenologv-  of  aerial 
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Wilson,  C.,  J.  Grace.  S.  Allen,  and  F.  Sl.\ck.  1987. 
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Wood,  M.  K.,  E.  H.  Blackburn,  R.  E.  Eckert,  Jr  ,  and 
F.  F.  Peterson  1978.  Interrelations  of  the  phvsical 
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Wyn  Jones.  R.  G.,  and  J.  C^jrham.  1986.  Osmoregulation. 
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Enviromaents  10:  10.'3-115. 

Received  17  August  1992 
Accepted  25  October  1992 


Creat  Basin  Naturdist  52(3),  pp.  216-225 

LIFE  HISTORY,  ABUNDANCE,  AND  DISTRIBUTION 
OF  MOAPA  DACE  (MOAPA  CORIACEA) 

G.  Gaiv  Scoppettone  ,  Howard  L.  Biirt^e    ",  and  Peter  L.  Tuttle  ' ' 

Abstract — Moapa  dace  {Moapa  roriaccii)  is  a  teder;iliv  listed  endangered  fish  endeniie  to  the  spring-fed  iieadwaters 
of  die  Muddv  River,  Clark  Connty,  Nevada.  Speeies  life  history;  abundance,  and  distribution  were  studied  from  March  1984 
to  JanuiUT  1989.  Reproduction,  which  was  obsei"ved  yetu-round,  peaked  in  spring  and  was  lowest  in  fall.  It  occurred  in 
headwater  tributaries  of  the  Muddy  Ri\er,  within  150  ni  of  warm  water  spring  discharge  in  water  temperatures  ranging 
from  30  to  32  C.  Feni;iles  matured  between  41  and  45  mm  in  fork  length  (FL).  Egg  abundance  increased  with  female  size 
(r"  =  .93);  counts  ranged  from  60  for  a  45-mm-FL  female  to  772  for  one  90-mm  FL.  The  oldest  of  eight  fish,  aged  by  the 
opercle  method,  was  a  90-mm-FL,  4+-year-old  female.  Adults  are  omnivorous  but  tended  toward  caniivory';  75%  of  matter 
by  N'olume  consumed  was  invertebrates  and  25%  pkints  and  detritus.  Fish  size  was  generally  commensurate  with  flow,  the 
largest  fish  occurring  in  the  greatest  flow.  Adults  were  near  bottom,  in  focal  velocities  ranging  from  0  to  55  cm/s.  Jn\'eniles 
occupied  a  narrower  range  of  depths  and  velocities  thim  adults,  and  lai^vae  occupied  slack  water.  From  December  1984  to 
September  1987,  the  total  adult  population  ranged  from  2600  to  2800.  Although  these  numbers  are  higher  than  prexiouslv 
believed  for  Moapa  dace,  they  are  still  sufficiently  low  to  warrant  its  end;uigered  status.  The  dependency  of  Moapa  dace  s 
different  life  histoiy  stages  to  \arious  areas  and  habitat  t\pes  of  the  Warm  Springs  area  suggests  that  all  remaining  habitat 
is  necessary  for  their  sumval. 

Ki'i/  icord.s:  Moapa  coriacea,  Moapa  dace,  life  liislonj.  rcpnulniiioti  l)iolo^y.jccmi(littj.  agc-i^n>ictli,Jo(Hl  habits,  habitat 
use,  bodij size,  Mitdch/  Riiei;  Nevada. 


Tlie  Moapa  dace  [Moapa  coriacea)  i.s  a  tlier- 
mophilic  niiniiow  endeniie  to  the  Mndd\'  Ri\  er 
system,  Clark  Counts,  Nexada.  First  collected 
in  1938,  it  has  lustorically  been  relegated  to  the 
headwater  area  where  the  Miiddv  River  origi- 
nates from  a  series  of  warm  springs  (Hubbs  and 
Miller  1948).  La  Rivers  (1962)  cafled  the  Moapa 
dace  and  its  coinhabitant,  Moapa  White  Ri\er 
springfish  [Crenichthijs  baileiji  nioapac),  ther- 
mal endemics  becanse  of  their  apparent  affinit\ 
for  warm  water.  Rarely  exceeding  12  cm  in  iork 
length  (FL),  Moapa  dace  ha\e  moiphological 
similarities  to  ronndtail  chnb  (Gila  roJ)iista)  and 
speckled  dace  (Rliinichtlujs  osctiln.s),  wliich  also 
inhabit  the  Muddy  River  (Hubbs  and  Miller 
1948).  They  are  more  similar,  however,  to  the 
genus  Agasir/,  which  occurs  in  other  lower  Col- 
orado River  drainages;  the  two  genera  are  spec- 
ulated to  have  a  conunon  ancestor  (Hubbs  and 
Miller  1948).  Moapa  dace  are  distinguished  In 
small  embedded  scales  and  a  bright  black  spot 
at  the  base  of  the  caudal  fin. 

Little  was  known  of  Moapa  dace  life  histor\ 


prior  to  this  studv  La  Rixers  (1962)  identified 
them  as  methodical  schoolers;  a  curson'  gut 
examination  bv  him  indicated  that  they  foraged 
primariK'  on  arthropods  and  some  vegetative 
matter.  In  a  systematic  sampling  effort,  Deacon 
and  Bradlev  (1972)  collected  Moapa  dace  in 
28-30  C  water;  one  specimen  was  collected  in 
19.5  C  water.  Within  the  confines  of  its  limited 
distribution,  Moapa  dace  ha\e  been  captured  in 
a  variety  of  habitats,  including  spring  pools  and 
slow-  to  fast-mo\ing  water,  and  in  association 
with  \arious  substrates  and  submergent  \egeta- 
tion  (Hubbs  and  Miller  1948). 

l^ast  ichtlnofaimal  siuvevs  suggested  a 
declining  Moapa  dace  population  (Deacon  and 
Bradle\'  1972,  Cross  1976).  These  suivevs  were 
([ualitatixe  and  produced  neither  an  estimate  of 
the  number  of  dace  remaining  nor  the  relati\e 
population  decrease  between  suneNS.  Ono  et  al. 
(1984)  tliought  that  ouK  sexeral  lumdred 
M()a[)a  dact^  persist(xl  and  that  their  distribution 
had  been  hirtlier  restricted  within  the  alread\ 
liiiiited  historic  habitat,  conlininsj;  them  to  the 


nj.S.  Fish  and  Wildlife  Senitc,  Nation.i 
^Present  address:  U.S.  Tisli  and  W  iMIil,- 
■^Present  addirss;  U.S.  Kisli  and  W  ildlil, 


(<-s.'aivhC:rii 
.(.rsli.ik  FisKr 
■  at  H.isin  (  ni 


.  Siilistatliai,  H.-iio.  Nrvada.  L'S.X  Sm02. 
.tancvOrinr.  Misalika.  Idalui.  L'S,\  S:«2(). 
•nn,  NiA.id.i,  i:s\  S9.5()2, 


216 


19921 


MOAPA  Dace 


217 


main  stem  of  the  upper  Muddy  River  and  a 
semi-isolated  headwater  spring  system  about 
130  m  long.  The  puq)ose  of  this  study  is  to 
expand  information  on  Moapa  dace  life  histon\ 
abundance,  and  distribution.  Life  histoiy  infor- 
mation includes  reproductiye  biologv',  habitat 
use,  food  habits,  and  age  and  growth. 

Study  Area 

The  Mudd\'  River  is  at  the  northern  edge  of 
the  Mohave  Desert,  where  average  annual  pre- 
cipitation is  15  cm  usualK^  in  the  form  of  rain. 
Caipenter  (1915)  described  historic  terrestrial 
vegetation  which  included  greasewood 
{Sarcohatus  vennicidatiis),  shadscale  (Atriplex 
confei'fifolia),  creosote  bush  {Larrea  triclen- 
tata),  and  mescjuite  (Prosopis  .sp.).  Stream  banks 
were  lined  with  willows  {Salix  sp.),  screw-bean 
(Prosopispubescens),  cottonwood  (Populus  sp.), 
and  mesquite  (Carpenter  1915,  Harrington 
1930).  Prior  to  the  completion  of  Hoover  Dam 
(aka  Boulder  Dam)  in  1935,  the  Muddy  (aka 
Moapa)  River  was  about  48  km  long  and  dis- 
charged into  the  Virgin  River,  which  joined  the 
Colorado  Rixer  (Hubbs  and  Miller  1948). 
Today,  it  is  about  40  km  long  and  discharges  into 
the  Overton  arm  of  Lake  Mead  (Fig.  1).  Source 
springs  of  the  Mudd\  River  probably  originate 
from  Paleozoic  carbonate  rocks  (Garside  and 
Schilling  1979)  and  occur  within  a  2-km  radius. 
As  is  t}pical  of  warm  springs,  the  water  is  rela- 
ti\ely  rich  in  minerals.  Garside  and  Schilling 
(1979)  list  sodium  and  calcium  as  predominant 
cations,  and  carbonate  and  sulfate  as  predomi- 
nant anions;  total  dissolved  solids  were  854  ppni 
and  pH  was  7.7.  Water  emerges  at  32  C  and 
cools  and  increases  in  turbidit)'  downstream 
(Cross  1976).  Although  spring  discharge  is  rela- 
tively constant  at  about  1.1  mVs,  the  Mudd\ 
Rixer  flow  fluctuates  because  of  rain,  agricul- 
tural diversions,  e\aporation,  and  transpiration 
(Eakin  1964).  The  headwater  region,  the  his- 
toric range  of  the  Moapa  dace,  is  known  as  the 
Warm  Springs  area  (Fig.  1).  During  our  stud\' 
the  area  was  used  primariK'  for  agriculture,  and 
up  to  0.25  m  Vs  of  river  discharge  was  being 
diverted  to  irrigate  alfalfa,  barley,  and  pasture. 
Spring  outflows  had  been  channelized,  and  se\- 
eral  were  converted  into  irrigation  ditches, 
some  lined  with  concrete.  Earthen  tributan- 
channels  had  scant  to  thick  riparian  corridors  of 
fan  palm  {Washingtonia  filifera),  tamarisk 
(Tamarisk  sp.),  ash  trees  {Frazinus  sp.),  and 


arrow  weed  (Pluchea  sericea).  Two  nonnative 
fishes  successfulK  established  in  the  Warm 
Springs  area:  mosquitofish  {Ganihiisia  affinis), 
present  when  Moapa  dace  were  discovered  in 
1938  (Hubbs  and  Miller  1948),  and  shortfin 
moUy  (Poecilia  mexicana),  introduced  in  the 
earlv  1960s  (Hubbs  and  Deacon  1964).  Besides 
Moapa  dace  and  springfish,  roundtail  chub  and 
speckled  dace  are  the  only  native  fishes  occur- 
ring within  the  Warm  Springs  area,  but  they  are 
rare  and  in  greater  abundance  downstream 
(Cross  1976,  Deacon  and  Bradley  1972). 

In  1979  the  Moapa  National  \Vildlife  Refuge 
(NW^R)  was  established  in  historic  habitat  at  the 
southern  edge  of  the  Warm  Springs  area  for  the 
preservation  and  peipetuation  of  the  Moapa 
dace  (Fig.  1 ).  The  refuge  stream  originates  from 
five  small  springs  occurring  in  a  radius  of  70  m 
and  having  a  cumulative  discharge  of  abut  0.09 
mVs  (Fig.  2).  Fan  palms  are  the  predominant 
riparian  vegetation.  In  1984  Moapa  dace  larx'ae 
and  adults  were  reintroduced  into  the  upper 
Refuge  Stream,  and  by  Januaiy  1986  there  was 
a  stable  reproductive  population  of  120  adults 
(authors,  unpublished  data).  Thev' were  isolated 
by  a  75-cm-high  waterfall.  Springfish  were  the 
only  other  fish  present,  and  they  were  abundant. 

Materials  and  Methods 

RepR0DUCTI\'E  BIOLOCY. — Among  our 
objectives  was  to  quantify'  duration  of  the  repro- 
ductive period  and  the  season  of  peak  laivae 
recruitment.  To  this  end,  a  segment  of  the  upper 
Refuge  Steam  system  was  snorkeled  at  30-  to 
90-da\  intenals  from  Febnian"  1986  to  |amiar\' 
1989  and  laivae  were  enumerated  (Fig.  2).  This 
is  the  area  in  which  virtually  all  reproduction  on 
the  Moapa  NWR  occurred.  Dace  7-15  mm  TL 
were  considered  larvae.  This  range  approximates 
the  proto-  to  metalanae  stages  of  the  similar- 
sized  speckled  dace  (Snyder  1981).  Snorkeling 
enabled  us  to  locate  reproduction  sites  in  the 
headwater  Muddv  River  .system  and  to  deter- 
mine the  abundance  and  distribution  of  adult 
Moapa  dace  as  well  as  to  (juantifv  hal^itat  u.se  for 
all  life  stages.  Areas  with  lanae  close  to  swim-up 
size  (about  7  mm  TL)  were  considered  repro- 
duction sites.  Fish  used  for  food  habit  analysis 
and  aging  were  also  used  to  detemiine  fecundit)^ 

H\BITAT  use. — We  defined  habitat  use  in 
terms  of  stream  depth  and  velocitv'  at  foraging 
sites  and  at  suspected  spawniing  areas.  Depth 
measurements  included  focal  and  total,  while 


218 


Great  Basin  Naturalist 


[\blume  52 


Colorado 
River 


115 


Fig.  1.  Map  showing  relatioiisliip  of  the  Miiddv  to  the  Xiigin  River  and  Lake  Mead,  Ne\'ada,  ;uid  relationship  of  the 
Warm  Springs  area  to  tlie  Mnd(l\-  liixer  (helowV  \\'ann  Springs  area  or  headwaters  of  the  Muddv  River  showing  tribntaiy 
streams  to  tlie  upper  Mnildv  Ri\er  and  relationship  ot  the-  Moapa  National  \\ildlile  Rehige  (above). 


1992] 


MoAPA  Dace 


219 


Upper 

Refuge 

Stream 


Fig.  2.  Map  of  Moapa  Nationd  VMldlife  Rcriigc:  shaded  site  indicates  the  reaeli  of  the  upper  Refuge  Streaiu  where 
liUAae  snorkel  counts  were  made  from  Februan  19S6  to  January  1989. 


\  elocih'  nieasurenient.s  included  focal  and  mean 
water  column,  as  prescribed  b)'  13()\ee  (1986). 
DissoKed  oxxgen  and  temperature  were  also 
measured.  Fish  were  located  using  mask  and 
snorkel.  A  Marsh  and  McBiniex  model  20 ID 
digital  flow  meter  mounted  on  a  calibrated  rod 
was  used  to  measure  depth  and  velocitN-,  and  a 


Yellow  Springs  Instrument  model  57  dissolved 
oxvgen  meter  for  temperature  and  dissolved 
owgen.  Sampling  occurred  from  1984  to  1986. 
Adult  habitat  was  also  defined  b)'  contrasting 
bod\-  size  with  (juantitv  of  stream  flow;  it  was  our 
subjective  evaluation  tliat  larger  fish  were 
inhabiting  lariier  water  \ olumes.  We  tested  this 


220 


Great  Basin  Naturalist 


[Volume  52 


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to  i~-  'o  9J 

00  CO  CO  CO 

o>  en  CT)  O) 

Month  /  Year 

Fig.  3.  Abundance  of  Moapa  dace  laivae  from  Februaiy  1986  to  janiiaiy  1989  in  tlie  Muddy  River  system  on  the  Moapa 
National  Wildlife  Refuge,  Nevada.  Bars  represent  a  single  dav's  count  for  the  month.  NS  indicates  not  sampled. 


h)^othesis  in  the  summer  of  1986  when  samples 
of  adults  \\'ere  minnow-trapped  from  the 
Muddy  River,  Muddy  Spring  Stream,  Refuge 
Stream,  and  Apcar  Stream  and  their  length  fre- 
quencies compared.  Discharge  for  each  stream 
was  measured  usino;  standard  U.S.  Geological 
Survey  methods  (Rantz  et  al.  1982)  near  each 
fish  sample.  A  one-way  factorial  ANOVA  was 
used  to  test  whether  there  was  a  significant 
difference  between  length  frequency  among 
fishes  and  different  water  volumes. 

Ace  and  GROVNTH. — The  opercle  bone  was 
used  for  estimating  age  as  described  by  Cassel- 
man  (1974).  Eight  specimens,  collected  in 
summer  1985  and  1986,  were  aged.  Flesh  was 
scraped  with  a  scalpel  and  the  bone  allowed  to 
dry'.  Glycerin  was  used  to  highlight  the  more 
transparent  region  of  the  bone,  which  was 
assumed  to  have  the  greatest  calcium  concen- 
tration and  to  have  been  formed  in  the  winter 
when  food  is  scarce.  The  more  opaque  region 
signifies  greater  concentration  of  protein  asso- 
ciated with  growth  (Casselman  1974). 

Food  habit— Food  habit  anaKses  were 
made  from  10  Moapa  dace  taken  9-1 1  Novem- 
ber 1984  from  each  of  three  uppc-r  Muddv  Riv(M- 
tributaries  (Apcar,  South  Fork,  and  Muddv 
Spring).  They  were  captured  by  seining  and 
with  unbailed  minnow  traps  fished  no  longer 
than  10  minutes.  Ranging  from  42  to  71  mm  FL, 


they  were  preserved  in  10%  formalin  solution. 
Contents  in  the  anterior  third  ol  the  gut  were 
examined  using  a  dissecting  microscope  and 
quantified  by  frequency  of  occiuTence  (Windell 
1971)  and  by  percent  composition  (H)iies  1950). 
Abundance  and  distribution. — The 
abundance  and  distribution  of  adult  Moapa 
dace  (>4()  mm  FL)  were  determined  by  snor- 
keling  the  upper  Muddy  River  svstem  begin- 
ning from  200  m  downstream  of  Warm  Springs 
Road  bridge  (Fig.  1).  Except  for  1984,  the  sur- 
veys included  5.3  km  of  the  upper  Muddy  River 
and  7.5  km  of  its  spring- fed  tributaries  (Refuge 
Stream  svstem,  Apcar  Stream,  Muddv  Spring, 
South  Fork,  and  North  Fork).  In  1984  the 
survev  area  was  the  same  except  that  only  the 
upper  130  m  of  the  Apcar  Stream  was  snorkeled 
rather  than  its  entire  stream  length.  Snorkeling 
was  conducted  over  periods  of  foiu"  to  six  days 
when  turbidit)'  was  low  (between  1.4  and  5.0 
NTU)  because  no  agricultural  return  flows  were 
entering  the  stream.  Coimts  were  made  6-10 
December  1984,  6-10  June  1986,  and  16-22 
September  1987.  Each  observer  enumerated 
Moapa  dace  twice  at  three  areas  of  relatively 
high  concentrations  (30-60  fish),  and  the  range 
of  results  was  then  calculated.  These  sites  were 
chosen  because  the  greatest  variation  among 
obsei-vers  was  expected  among  them.  For  the 
three  sites,  variation  was  less  than  15%  in  counts 


1992] 


MoAPA  Dacb: 


221 


1,000 


800   - 


c/5 
LU 


CD 

n 

E 

13 


600 


400   - 


200   - 


30 


- 

r  ^=  .93 

n    =23 

D 

D^     /^ 

■ 

/-"m 

y^ 

-^D 

D 

- 

f^ 

D 

D 

- 

□      02^ 
1                       1 

1 

1 

1              1 

40  50  60  70  80 

Fork  Length  (mm) 


90 


100 


Fig.  4.  Moapa  dace  fecundit)-  iis  a  tuiictioii  of  fork  length. 


between  indmduals;  thus,  we  consenathelv 
estimated  a  15%  \ariation  in  o\\\  population 
counts. 

Results  and  discussion 

Reproducti\e  Biolog)' 

Moapa  dace  lanae  were  found  vear-round, 
iudicating\ear- round  reproduction.  On  the  Moapa 
NW'R  peak  lanal  reciiiitment  was  in  spring,  tlie 
low  in  autumn  (Fig.  3).  Fish  at  other  reproduc- 
tive sites  in  the  Warm  Springs  area  exliibited  this 
same  general  trend.  Seasonal  fluctuation  in  lanal 
recruitment  was  probabh"  linked  to  a\  ailabilitv' 
of  food.  In  the  upper  Mudd\  Ri\  er  system  the 
abundance  of  benthic  and  drifting  invertebrates 
is  much  lower  in  winter  than  in  spring  (Scop- 
{)ettone,  un]:)ubli,shed  data).  Naiman  (1976) 
documented  substantial  seasonal  fluctuation  in 
primar)'  producti\it\'  in  anothei"  southwestern 
warm  springs  where  production  is  lowest  in 
winter;  presumably  most  invertebrate  popula- 
tion fluctuates  with  priman'  production. 

Recenth'  emerged  lanae  were  found  within 
150  m  of  spring  discharge  over  sandy  silt  bot- 
toms in  temperatures  of  30-32  C  and  dissoKed 
ox)gen  of  3.8-7.3  mg/L.  Whether  spawning 
occurs  only  at  these  head\\'ater  sites  or  is  suc- 
cessful onl\-  at  these  sites  is  unknown.  Visual 
cues  such  as  sexual  dichromatism,  pronounced 
male    spawiiing  tubercles,    or   o\ertly   gra\id 


females  were  not  readily  apparent,  and  spawiiing 
was  not  observed  during  our  stud\".  Howexer,  we 
indirectly  identified  and  quantified  spawning 
habitat.  The  presence  of  hundreds  of  proto- 
lanae  in  a  concrete  irrigation  channel 
immediately  downstream  of  the  Baldwin 
springhead  (Fig.  1)  indicated  that  reproduction 
had  taken  place.  Progenitors  apparentlv  came 
from  the  South  Fork,  entering  Baldwin  Spring 
outflow  through  a  diversion  channel  (Fig.  1). 
The  concrete  irrigation  channel  had  homoge- 
neous water  depth  and  \elocit\',  and  substrate 
was  sandv  silt.  Se\eral  depressions  in  the  sand 
were  similar  to  "redds"  described  for  longfin 
dace  (Agcw/V/  clm/so'^aster;  Minckle\'  and  Wil- 
lard  1971).  Depth  and  \  elocity  at  the  suspected 
redds  were  representative  of  the  outflow  chan- 
nel and  similar  to  other  suspected  spawning 
areas  in  the  Warm  Springs  area.  Depth  ranged 
from  15.0  to  19.0  cm,  near-lx^d  \elocities  from 
3.7  to  7.6  cm/sec,  and  mean  water  colunm  veloc- 
it\-  from  15.2  to  18.3  cm/sec. 

Similar  to  the  longfin  dace,  which  repro- 
duces during  much  of  the  year  (Kepner  1982), 
eggs  in  the  skein  of  Moapa  dace  were  in  differ- 
ent stages  of  development.  All  visible  eggs  were 
counted,  but  because  they  are  intermittently 
deposited  and  develop  throughout  a  gi\ en  year, 
our  counts  do  not  represent  absolute  annual 
fecundity.  How^ever,  egg  production  increased 
with  fish  size  (r  =  .93,  n  =  25;  Fig.  4).  Counts 


999 


Great  Basin  Naturalist 


[\^olnme  52 


80 

60 

40 

20 
0 

gso 

>»60 
O 

O  40 

3 

0 


Li. 


80 
60 
40 

20 

0 


80 
60 
40 

20 

0 

^80 

O  60 

c 
0  40 

S"20 
III     0 

80 
60 

40 

20 
0 


Dace 

Adults 

;                            n  =  564 

H,.. 

Dace 

Juvenile 

n  =  148 

I 

Dace 

Larvae 

n  =  201 

.■, ,, 

80  - 

60- 

40 

20 
0 


0    10  20  30  40  50  60  70  80  90  100110 

Total  Depth  (cm) 


Dace 
Juvenile 
n  =  148 


'        '       * I L. 


Dace 
Larvae 
n  =  201 


-1 1 *     * 


Dace 

Juvenile 
n  =  147 


Dace 
Larvae 
n  =  199 


0    10  20  30  40  50  60  70  80  90  100110 

Focal  Depth  (cm) 


0       10      20      30      40      50      60      70 

Mean  Water  Column  Velocity  (cm/sec) 


80 

60 

40 

20 
0 

80 

60 

40 

20 
0 

80 

60 

40 

20 
0 


Dace 
Juvenile 
n  =  147 


i 


Dace 
Larvae 
n  =  201 


0       10      20      30      40      50      60      70 

Focal  Velocity  (cm/sec) 


Fig.  5.  Mean  water  coliiinn  and  loeal  point  xelocitics,  total  depth,  and  local  point  depth  nsed  l)\  Moapa  daei'  adn 
juveniles,  and  larvae  in  the  upper  Mudd\  Rixcr  system  (Warm  Springs  area),  Ne\ada.  UiS-l  through  liJSB. 


1992] 


MoAPA  Dace 


223 


Tabi.K  1.  Fork  IcMigth,  sex,  ;iiul  cstiinatecl  age  of"  eis^ht  TaBI.K  2.  Fckk\  items  ingested  In  2]   Moapa  dace  b\' 

Moapa  dace  collected  from  the  upper  Miiddv  Hi\ cr  s\  stem.  percent  conn^xjsition  ( H\iies  1950)  and  percent  frequence 

Ne\ada.  in   1985  and   19S6.  Age  was  (Ictcnniind  1)\    the  ol  occnrrence  (W'indell  1971).  Nine  odier  guts  examined 

opercle  method.  w  tre  empty. 


FL 

(nnn) 


Sex 


Collection 
date 


Food  items 


Age 


45 
55 
61 
67 
69 

SO 
90 


Unknown 

4/86 

Unknown 

7/86 

Unknown 

7/86 

Female 

4/86 

Female 

04/22/86 

Unknown 

10/09/85 

Unknown 

10/11/85 

Female 

10/08/85 

0+ 

1+ 
1+ 

2+ 
2+ 
3+ 
3+ 
4+ 


ranged  from  60  in  a  45-nini-FL  indiviclnal  to  772 
ill  a  9()-nini-FL  dace.  Eggs  were  just  developing 
in  a  41-nnn-FL  female  and  were  matnre  in  a 
45-mm-FL  fish,  suggesting  that  females  mature 
at  lengths  in  this  range. 

Habitat  U.se 

Again,  Moapa  dace  larvae  were  found  exclu- 
sixely  in  the  upper  reaches  of  spring-fed  tribu- 
taries, while  juveniles  occurred  primarilv  in 
tributaries  but  were  more  far-ranging.  Adults 
were  present  in  tributaries  and  in  the  main  ri\er, 
with  larger  fish  generalK'  found  in  the  larger 
water  volumes.  There  were  significant  differ- 
ences in  length  frequencies  among  adults  from 
different  water  \olumes  {p  <  .006).  In  the 
MudcK  Rixer,  in  a  flow  of  about  0.50  m  Vs,  mean 
FI.  was  73  mm  (/]  =  78,  SD  =  16  mm);  Muddy 
Spring  had  a  flow  of  0.20  m  Vs,  and  the  mean  FL 
was  64  mm  (n  =  72,  SD  =  14  mm);  the  Refuge 
Stream  flowed  at  0.17  m'/s,  and  mean  FL  was 
56  mm  (/;  =  64,  SD  -  8  mm);  the  Apcar  Stream 
llowed  at  0.06  mVs,  and  mean  FL  was  51  mm  (n 
=  89,  SD  =  5mm). 

Lar\ae occurredand  fed  in  tlu^  mid-  to  uppc^" 
region  of  the  column.  They  were  found  most 
frequentk'  in  zero  water  velocit\"  ( Fig.  5).  As  size 
increased,  individuals  tended  to  occupy  faster 
water  and  occur  lower  in  the  water  column, 
juvenile  Moapa  dace  occupied  focal  and  mean 
water  column  velocities  ranging  from  0  to  46 
cm/s.  Adults  were  found  in  a  wide  range  of  water 
depths  and  velocities,  but  they  tended  to  orient 
at  the  bottom  in  low  to  moderate  current.  Water 
column  depth  ranged  from  15  to  113  cm  and 
focal  point  depth  from  9  to  107  cm.  Mean  water 
column  \elocit)-  ranged  from  2  to  77  cm/s  and 
focal  point  velocit)'  from  0  to  55  cm/s.  Water 
temperatures  within  adult  habitats  ranged  from 


%  composition        %  of  occurrence 


Gasthopoda 

Tt/ronui  clathnita 

1.1 

Olk;()(:iiaf.tk 

27.0 

AMI'IIII'ODA 

Hi/dlli'la  aztcra 

1.7 

IlKMIITKHA 

Pclocoiis  shoshoiw 

4.5 

HOMOITKIU 

Apiiiidac 

9.0 

Tkiciioitf.ha 

Dolophilodcs 

5.1 

Necfop.si/clie 

4.5 

LEPlDOn'F.HA 

Para'^i/ractis 

4.5 

COLEOPTKKA 

Steucluiis  ralicla 

1.1 

Dijtiscidae  (lan'ae) 

9.0 

DiPTElU 

Chlronomidae 

4.5 

Unidentified  insect  parts 

3.3 

Filamentous  algae 

18.5 

N'ascniar  plants 

3.4 

Detritus 

2.8 

4.8 
23.8 

9.5 

4.8 

4.8 

9.5 
9.5 

9.5 

4.8 
4.8 

4.8 
9.5 

42.3 
9.5 

14.3 


27  to  32  C  and  dissoKed  o.wgen  from  3.5  to  8.4 
mg/L. 

Age  Crowth 

Annulus  formation  is  t\picalh'  associated 
with  an  annual  period  of  slower  growtli  cau.sed 
bv  seasonal  changes  in  environmental  condi- 
tions such  as  temperature  or  food  resources 
(Tesch  1971).  Although  seasonal  water  temper- 
atures do  not  change  sul)stantiall\  in  the  Warm 
Springs  area,  there  is  an  apparent  reduction  of 
potential  food  during  the  winter  (Scoppettone, 
unpublished  data).  We  were  unsuccessful  in 
aging  Moapa  dace  bv  the  scale  method  because 
scales  were  small,  embedded,  and  extremely 
difficult  to  remove  from  live  specimens.  Also, 
einiroiinuMita!  conditions  in  waters  of  the  Warm 
Springs  area  were  sufficientlv  constant  that 
aiinuli  were  not  readilv  apparent.  Assmiied 
annuli  on  opercular  bones  were  presumed  to  be 
associated  with  slow(^r  growth  dining  the  winter. 
Ages  of  the  eight  fish  examined  ranged  from  O-l- 
for  a  43-mm-FL  individual  to  4+  for  a  90-mm- 
FL  female  (Table  1). 

Food  Habit 

Nine  of  30  guts  examined  were  emptv'  and 
the  remainder  generalK  contained  few  items, 


224 


Great  Basin  Naturalist 


[Volume  52 


Table  3.  Estimated  number  of  Moapa  dace  adults  in  six  tributan-  streams  in  the  Warm  Springs  area,  Muddv  Ri\er 
system,  Nevada,  6-14  December  1984,  1.3-18  June  1986,  iuid  16-22  September  1987. 


Stream 

December 

X'ariation 

June 

\'ariation 

September 

Variation 

name 

1984 

in  count 

1986 

in  count 

1987 

in  coimt 

Muddy  River 

475 

±71 

1230 

±185 

1165 

±175 

Refuge  System 

370 

±56 

406 

±61 

806 

±121 

Apcar 

200 

±30 

565 

±85 

475 

±72 

South  Fork 

300 

±45 

185 

±28 

100 

±15 

Nortli  Fork 

15 

±2 

30 

±5 

60 

±9 

Muddv  Spring 

1450 

±218 

160 

±24 

200 

±30 

Total 

2810 

±422 

2581 

±387 

2806 

±421 

"Onh  till-  iippi-r  loO  in  of  stiva 


but  what  had  been  consumed  indicated  Moapa 
dace  to  be  omnivorous  tending  toward  caniiv- 
ory;  75%  by  composition  was  invertebrates 
while  25%  was  plant  material  and  detritus 
(Table  2).  Among  21  dace  guts,  oligochaetes 
represented  the  largest  \'ohune  (27.0%)  of  food- 
stuffs consumed,  followed  by  filamentous  algae 
(18.5%).  In  terms  of  frequency  of  occurrence 
filamentous  algae  occurred  in  42.3%  of  the  guts 
while  oligochaetes  were  in  23.8%.  The  stmcture 
of  the  pharyngeal  teeth  also  suggests  an  omniv- 
orous diet;  they  are  strongly  hooked  but  ha\e  a 
well-developed  grinding  surface  (La  Rivers 
1962).  The  presence  of  detritus  and  gastropods 
indicates  at  least  some  foramne;  from  the  ben- 
thos,  and  we  obsened  fish  in  the  field  occasion- 
ally pecking  at  substrate.  However,  the  greatest 
time  in  foraging  is  expended  on  drift  feeding 
(authors,  unpublished  data),  although  our  data 
set  does  not  strongh'  support  this  obsenation. 

Abundance  and  13istribution 

Moapa  dace  were  more  widespread  and 
numerous  than  had  been  previously  rej)orted 
(Ono  et  al.  1984);  they  were  in  five  headwater 
tributaries  and  the  upper  Muddy  River  to  about 
100  m  downstream  from  the  Warm  Springs 
Road  bridge  (Fig.  2).  Numbers  ranged  from 
about  2600  in  1986  to  2800  in  1984  and  1987. 
The  numerical  distribution  for  the  three  years 
suggests  movement  by  the  adult  population 
(Ttible  3).  In  f984  the  Muddy  Spring  stream 
supported  about  50%  of  the  population  (1450 
adults),  with  only  ]6%>  (450  adults)  foimd  in  the 
river  In  June  1986  we  could  account  for  only  7% 
of  the  population  in  the  Muddy  Spring  stream, 
while  almost  50%  of  the  total  was  in  the  river.  In 
1987  the  mainstream  river  again  supported 
most  adult  Moapa  dace  (1200).  The  distribution 
of  adult  Moapa  dace  was  patchy  and  clumped. 
For  example,    during  the   snorkel   suive)'  in 


summer  1986,  79%  of  the  observed  dace  in  the 
main  stem  Muddy  River  were  in  groups  of  10  or 
more,  and  37%  were  in  groups  of  30  or  more.  In 
tributaries,  groups  were  generally  smaller,  with 
52%  of  the  adults  in  groups  of  10  or  more  and 
only  13%  in  groups  of  30  or  more. 

Conclusion 

Moapa  dace  are  dependent  upon  the  link 
between  the  upper  ri\'er  and  its  tributaries.  The 
main  stem  river  typically  harbors  the  largest, 
and  presumably  the  longest-lived,  and  most 
fecund  fish;  yet  tributaries  are  important  for 
reproduction  and  as  lanae  and  juvenile  nurser)^ 
habitat.  Age  and  growth  information  suggests 
that  three  years  is  the  mean  age  of  fish  in  the 
river  and  that  adults  in  smaller  tributaries  are 
one  to  two  vears  old. 

Although  the  Moapa  dace  population  is 
more  widespread  and  abundant  than  previously 
beliexed,  its  existence  remains  in  jeopardy. 
Widespread  movement  and  obligator'  spawTi- 
ing  near  warm  water  spring  discharge  suggest 
that  species  survival  depends  on  access  to  the 
entire  headwater  Mudd\'  River  svstem  (Warm 
Springs  area),  river  and  tributaries  alike.  Everv 
effort  should  be  made  to  presene  all  of  its 
remaining  habitat. 

Ac:  K  N  OW  L  E  D C;  M  E  N TS 

William  Burger  and  Dana  Winkleman 
assisted  in  snorkel  surveys,  and  Michael  Parker 
and  Nadine  Kanim  assisted  in  estimating  fish 
populations.  Peter  Rissler  helped  to  determine 
habitat  use.  Michael  Parker  conducted  gut  iuialv- 
sis.  Glen  Clemmer,  Randy  McNatt,  and  Tom 
Strekal  reviewed  the  manuscript.  Linda  Hallock 


1992] 


MOAPA  Dace 


225 


ln'Iped  with  editing  and  Steplianic  Byers  with 
graphics. 

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Received  1  August  1991 
Accepted  15  September  1992 


Creat  Basin  Natur;ilist  52(3),  pp.  226-231 

CONDITION  MODELS  FOR  WINTERING  NORTHERN  PINTAILS 
IN  THE  SOUTHERN  HIGH  PLAINS 

Lorcn  M.  Smith  ,  Douglas  G.  Sheelev",  and  Da\i(l  B.  Wester 

AbstiucT. — Three  condition  models  ior  wintering  Northern  Pintails  (Anas  acuta)  were  tested  for  their  abiiit)  to  predict 
fat  mass,  logarithm  of  fat  mass,  or  a  condition  index  (CI)  incoiporating  fat  mass.  Equations  generated  to  predict  fat  mass 
and  the  logarithm  of  fat  mass  accounted  for  more  than  69%  of  the  variation  in  these  dependent  variables.  Log  transforma- 
tions of  body  mass,  wing  length,  and  total  lengdi  explained  at  least  60%  of  the  variation  in  CI.  All  models  performed  better 
on  an  independent  data  set.  Mean  prediction  error  was  minimal  (<8%  of  measured  variables)  and  negative  for  all  models. 
Regression  models  apply  to  live  and  dead  pintails  and  thus  represent  tools  that  have  utilit)'  in  a  wide  variet)-  of  studies  on 
pint;ul  condition. 

Ki-if  words:  Niiillirni  Pintails.  Anas  acuta,  Ixxli/  n>ii(iitii>)i.  predictive  models.  Texas,  ivateifoiel. 


Biologists  have  used  variotis  indices  for 
assessing  waterfowl  nutritional  status.  Initially, 
only  body  mass  was  used  (Hanson  1962,  Folk  et 
al.  1966,  Street  1975,  Flickinger  and  Bolen 
1979),  but  later  stmctural  variables  were  incor- 
porated to  adjust  for  individual  size  differences 
(Oven  and  Cook  1977,  Bailey  1979,  Wishart 
1979).  Ringelnian  and  Szvmczak  (1985)  and 
Johnson  et  al.  (1985)  re\"iewed  a\ian  condition 
indices  and  noted  the  value  of  an  accurate  index 
of  lipids  in  migratoiy^  bird  management.  These 
studies  noted  that  scaling  moiphological  \ari- 
ables  with  body  mass  provided  tiseful  indices  to 
avian  body  condition. 

Northern  Pintails  {Anas  aciifa)M-e  one  of  the 
most  widespread  waterfowl  species  in  North 
America  (Bellrose  1980),  but  recently  their  pop- 
ulations have  declined,  making  them  a  species 
of  special  concern  (Smith  et  al.  1991).  Our 
objectives  were  to  pro\ide  an  ecjuation  to  pre- 
dict total  carcass  fat  (b()d>-  condition)  of  North- 
ern Pintails  and  to  test  that  index  on  an 
independent  data  set.  The  auatonn'cal  \ariables 
tested  are  suitable  for  field  studies. 

Study  Area 

The  stud\  was  conducted  in  the  Southern 
High  Plains  (SUP)  of  Texas,  an  82,88()-km-  area 
that  is  one  of  the  most  intensixel)-  cultivated 


regions  in  the  Western  Hemisphere  (Bolen  et 
al.  1989).  Twents'  thousand  pla\as  are  present  in 
the  SHP  providing  winter  habitat  for  waterfowl 
(Haukos  and  Smith  1992).  At  least  one-third 
(>300,000)  of  the  Northern  Pintails  wintering  in 
the  Central  FK'w^ay  wdnter  on  the  SHP  (Bellrose 
1980). 

Methods 

Northern  Pintails  were  collected  using 
deco\'s  and  b)'  jtmip-shooting  on  plavas  and 
associated  tailwater  pits  in  the  SHP  from  Octo- 
ber through  March  of  1984-85  and  1985-86. 
Tarsal  length  (measured  from  the  junction  of 
the  tibiotarsus  and  tarsometatarsus  to  the  point 
of  articulation  bet^veen  the  tarsometatarsus  and 
middle  toe,  0.01  mm),  flattened  wnng  chord 
(measured  from  the  insertion  of  the  ahila  to  the 
tip  of  the  tenth  priman',  0.1  cm),  and  total  body 
length  (measured  from  the  tip  of  the  bill  to  the 
end  of  the  p\'gost\le,  thus  avoiding  complica- 
tions due  to  tail  feather  growth,  0.1  cm)  were 
recorded  for  each  bird.  During  1985-86  an 
additional  wing  measurement  was  recorded 
from  the  insertion  of  the  alula  to  the  tip  of  the 
ninth  priman'  because  the  ninth  primary  maybe 
slightK-  longer  than  the  tenth.  Birds  were 
])lucked  and  frozen. 

Ingesta  and  intestinal  contents  were 
remoxed  in  the  laboratoiy.   Birds  then  were 


^  Department  of  Range  and  Wildlife  Management,  Texas  Tecli  Ui\iversi(\ .  I  .iilihoek.  Texas  T9K)9. 
-Box  464,  Eldora,  Iowa .50627. 


226 


1992] 


PlNTAII,C:()\'niTK)\  MODKLS 


00' 


TaHI.K  1.  X'arialile.s  u.st'd  in  prcdictixc  models  ol  IxxK condition  lor  Xortlicrn  TiiitaiLs  [Anas  aciitd}  on  tlie  Soiitlicrn  IIi<j;li 
Plain.s,  Texas. 


Adult 

Adult 

Ju\en 

ile 

Jmenile 

miJes  (n 

= 

140^ 

females  in 

=  69 ! 

males 

in 

=  58) 

lemales 

(11  =  49) 

Variable 

X 

SE 

X 

SE 

.V 

SE 

X 

SE 

Ma.ss  (g) 

963.93 

10.94 

8.35.07 

12.60 

911.97 

16.41 

786.68 

14.90 

Tarsal  length 

(mm) 

41.15 

0.17 

.38.68 

0.23 

41.13 

0.25 

38.90 

0.30 

Wing  len 

igtli  ( 

cm) 

26.5S 

O.Ofi 

24.69 

0.08 

25.92 

0.10 

24.23 

0.09 

Total  len 

gth  ( 

em) 

49.72 

0.12 

43.37 

0.14 

49.53 

0.22 

4.3.14 

0.19 

1  ,ipid  mass  (g 

:) 

171.57 

6.27 

173.20 

8.33 

147.93 

11.07 

148.21 

9.56 

reweighed  (neare.st  0.01  g)  to  determine  a  net 
carcass  mass  and  refrozen  (Table  1 ).  Frozen 
hi  i-ds  were  sectioned  with  a  meat  saw  and  passed 
twice  through  a  meat  grinder.  The  homogenate 
was  dried  to  a  constant  mass  in  either  a  forced- 
air  o\en  (60  C)  or  freeze  dner.  Dried  pintails 
were  regronnd  to  insure  a  uniform  mixture. 
Lipid  was  extracted  from  10-15  g  samples  using 
petrolevmi  ether  soKent  in  a  Soxhlet  apparatus 
(36-48  hrs).  Fat-free  diy  mass  (FFDM)  was 
calculated  by  subtracting  water  and  lipid  from 
total  carcass  mass  (body  mass  minus  feathers 
and  ingesta).  Total  carcass  mass  minus  water 
mass  \ielded  diy  mass  (DM). 

Three  models  were  e\aluated  to  predict  (1) 
fat  mass,  (2)  a  condition  index  (CI)  incorporat- 
ing fat  mass,  and  (3)  the  logarithm  of  fat  mass  of 
wintering  Northern  Pintails.  First,  pintails  were 
sorted  b\'  sex  (age  was  not  significant;  multiple 
regression,  P  >  .05).  A  predictive  model  for  fat 
was  generated  ff)r  each  sex  using  total  bodv 
length  (TOTAL),  wing  length  (WING),  tarsal 
length  (TARSAL),  and  bocK  mass  (MASS)  as 
cxplanaton- \ariables. 

In  model  1 ,  regression  coefhcients  of  cxpian- 
atoiy  variables  between  sexes  w(m-(>  iu)t  different 
(P  >  .05).  A  predictixe  e(juati()n  ap[)licable  to 
I)oth  .sexes  was  therefore  constnicted  which 
included  a  dunun\-  xariable  for  sex  (DSFX)  as 
well  as  stnictuial  \ariables. 

Th(^  second  model  was  constnictcnl  follow- 
ing John.son  et  al.  (1985);  a  Lipid  Index  was 
dehncd: 

Lipid  Index  =  Fat  /  FFDM. 

Fat-free  dn'  mass  is  included  to  correct  for  size 
tlifferences  between  indi\iduals.  Lipid  Index 
was  transformed  to: 

CI  =  log  (Lipid  Index  +  1) 

because  the  structural  measurements  are  allo- 
nietric  and  because  logarithms  can  be  used  to 
linearize  ratios  (Johnson  et  al.  1985).  The  con- 


stant 1  was  added  to  smooth  tlie  function.  CI  can 
be  simplified  to: 

CI  =  log(DM/FFDM) 

because 

DM  =  Fat  +  FFDM. 

Log  FFDM  was  modeled  as  a  function  of  the 
logarithms  of  structural  variables  (LTOTAL, 
LWING,  and  LTARSAL)  and  log  DM  as  a  hmc- 
tion  of  these  plus  the  logarithm  of  bodv  mass 
(LMASS)  (Johnson  etal.  1985).  Unlike  Mallards 
{Anasplatyrhynchos;  Ringelman  and  Szxniczak 
1985)  and  Canada  Geese  {Brantn  canadensis; 
Ra\'eling  1979),  water  content  of  wintering 
Northern  Pintails  fluctuated  widel\-  (Smith  and 
Sheeley  1993).  Therefore,  we  did  not  test  fat- 
free  mass  as  an  index  to  structural  size  (Ringel- 
man and  Sz)'mczak  1985). 

Johnson  et  al.  (1985)  used  k)garithms  of 
structural  \ariables  to  model  logarithms  of  car- 
cass fat  mass  (log  fat).  A  separate  equation  was 
estimated  for  each  age/sex  group  (model  3) 
using  dummv  \ariables  for  age  (DACE)  and  sex 
(DSEX)  because  regression  coefficients  for 
explanaton  \ariables  differed  (P  <  .05)  among 
these  four  groups. 

Predictixe  equations  were  \alidated  on  a 
data  set  of  40  randomlv  selected  pintails  not 
inchuknl  in  the  generation  of  models.  Percent- 
ages of  each  age/sex  class  of  pintails  in  the  inde- 
pendent sample  were  consistent  with  their 
occurrence  in  the  sample  collection. 

Pn^liction  (MTor  (PF)  was  calculated  as  an 
additional  test  of  model  jx'riormance.  PE  is 
defined  as: 

PE  =  .Measured  \'  -  Predicted  Y, 

where  Y  is  the  dependent  \ariable.  Mean  PE  is 
an  axerage  \alue  for  all  members  of  the  \alida- 
tiou  data  set.  Finallv,  predicted  fat,  CI,  and  log 
fat  wen^  correlated  with  Lipid  Index  in  the 
\  alidation  data. 


228 


Great  Basin  Naturalist 


[Volume  52 


T.\BLE  2.  Regression  equations  and  associated  statistics  tor  predicting  carcass  fat  (model  1)  content  (g)  in  Northern 
Pintiiils  (Anas  acuta)  collected  on  the  Soudiem  High  Plains  of'Texas,  October-Mtuch  198-1— S6. 


r' 

Exjilanatory  variables 

Equation 

Intercept 

MASS 

WING 

TOTAL 

DSEX 

1.1 

.779 

P;xrameter  estimate         191.854 

0.560 

-13.386 

-4.136 

_ 

(Male;  n  =  198) 

SE                                          — 

0.022 

3.894 

1.901 

— 

\'ariance  inflation  factor       — 

1.181 

1.231 

1.221 

— 

Piuti;il  R-                                — 

0.741 

0.013 

0.005 

— 

1.2 

.711 

Parameter  estimate         145.570 

0.570 

-9.516 

-4.953 

— 

(Female;  »i  =  118) 

SE                                          — 

0.035 

5.561 

2.994 

— 

Variance  inflation  factor       — 

1.125 

1.212 

1.174 

— 

PartiJ  R-                             — 

0.691 

0.007 

o.oor 

— 

1.3 

.757 

Parameter  estimate         190.494 

0.563 

-12.068 

-4.409 

-22.513 

(ComJDined;  tt  =  316) 

SE                                          — 

0.018 

3.178 

1.600 

10.536 

Variance  inflation  factor       — 

1.492 

3.164 

6.842 

5.987 

Partial  R'                              — 

0.726 

0.011 

0.006 

0.004 

•'Not  .significant  (P  >  .0.5). 

T.\BLE  3.  Regression  equations  and  associated  statistics  for  predicting  Condition  Index  (model  2)  in  Nortliern  Pintails 
(Anas  acuta)  collected  on  the  Southern  High  Plains  of  Te.xas,  October-March  1984—86. 


r2 

Expl 

anatory'  v 

txriables 

Equation 

Intercept 

LMASS 

LWING 

LTOTAL 

DSEX 

2.1 

.673 

Parameter  estimate          —0.816 

1.371 

-1.025 

-0.909 

_ 

(Male;;)  =  198) 

SE                                        — 

0.069 

0..343 

0.312 

— 

Vari;uice  inflation  factor       — 

1.190 

1.233 

1.229 

— 

Partid  R-                               — 

0.656 

0.015 

0.014 

— 

2.2 

.599 

Parameter  estimate          —0.725 

1.316 

-1.179 

-0.710 

— 

(Female;/)  =  118) 

SE                                          — 

0.101 

0.512 

0.486 

— 

Variance  inflation  factor       — 

1.123 

1.206 

1.176 

— 

Partiiil  R~                                — 

0.595 

0.019 

0.008 

— 

2.3 

.657 

P;u"ameter  estimate          —0.761 

1.350 

-1.080 

-0.834 

-0.041 

(Combined;  n  =  316) 

SE                                        — 

0.057 

0.286 

0.264 

0.016 

Varimice  inflation  factor       — 

1.496 

3.207 

7.035 

6.141 

P;utiiil  R-                                — 

0.610 

0.016 

0.011 

0.007 

Stepwise  multiple  regression  (iiiiLximum  R' 
improvement  technique)  was  used  to  generate 
and  test  all  models  (SAS  Institute,  Inc.  1985). 
Variables  were  eliminated  that  did  not  contrib- 
ute significantly  (F  <  .05)  to  a  model.  Partial  R- 
values  were  calculated  for  each  variable  in  a 
model.  A  sum  of  scjuares  (Ty|:)e  II)  for  each 
model  variable  was  divided  by  the  total  sum  of 
squares  in  the  model.  A  partial  R-  value  for  a 
given  variable  represents  the  uni(|ue  contribu- 
tion of  that  variable  when  all  other  \ariables  are 
already  present  in  the  model.  Partial  H"  values 
are  not  additive,  and,  therefore,  their  sum  will 
not  equal  the  total  model  K~.  Differences  in 
variation  accounted  for  by  ninth  \ersus  tenth 
primar\-  length  were  evaluated  using  the  K"  pro- 
cedure (SAS  Institute,  Inc.  1985). 


Results 

In  model  1  (Table  2)  bodv  mass  ex|:)lained  a 
major  portion  of  \ariation  in  carcass  fat  content 
in  males  (equation  1.1)  and  females  (equation 
1.2).  Total  length  did  not  account  for  a  signifi- 
cant (P  >  .05)  portion  of  variation  in  fat  content 
for  females  as  it  did  males.  Based  on  low  vari- 
ance inflation  factors  (\TF),  regression  coeffi- 
cient estimates  for  each  sex  were  stable.  When 
sexes  were  combined  through  use  of  a  dummy 
xariable  (equation  1.3),  the  \TF  for  TOTAL  and 
DSEX  were  relatixeK'  high;  this  is  largelv  attrib- 
utable to  the  hitrh  correlation  between  length 
and  sex  of  bird  (point  biserial  correlation  coeffi- 
cient ecjual  to  0.91). 

LTOTAL,  LWING,  and  LTARSAL  ex- 
plained variation  in  log  FFDM.  For  modeling. 


1992] 


Pintail  Condition  Models 


229 


Table  4.  Regression  equations  and  associated  statistics  for  predicting  log  carcass  fat  (model  3)  in  Nortlieni  Pintails  (Anas 
acuta)  collected  on  the  Southern  High  Plains  of  Texas,  October-March  19S4-86. 


Explanator)'  variables 


Equation 


Intercept 


LMASS 


LWING 


3.1  .727 
(Adult  male; /)  =  140) 

3.2  .693 
(Adult  female;  it  =  69) 

3.3  .722 
( |u\eiiile  male;  ii  =  .58) 


3.4  .745 

(Ju\enile  female;  n  =  49) 


Parameter  estimate  —3.410 

SE  — 

N'iiriance  inflation  factor  — 

Partial  R-  — 

Parameter  estimate  —  1 .61 1 

SE  — 

\'iU"iance  inflation  factor  — 

Partial  R-  — 

Parameter  estimate  —11.066 

SE  — 

Vtiriance  inflation  factor  — 

Partial  R-  — 

Parameter  estimate  —.5.444 

SE  — 

\'ariance  inflation  factor  — 

Partial  fi"  — 


3.412 

-3.209 

0.182 

0.993 

1.156 

1.156 

0.697 

0.021 

3.687 

-4.998 

0.303 

1.472 

1.034 

1.034 

0.687 

0.054 

5.028 

-1.223 

0.422 

2.009 

1.015 

1.015 

0.719 

().()()2 

3.968 

-2.834 

0.348 

1.844 

1.109 

1.109 

0.720 

0.013 

Table  5.  Coefflcients  of  determination  (R'}  and  predic- 
tive error  estimates  from  the  xiilidadon  (n  =  40)  of  predic- 
ti\e  equations  to  measured  \ariables  and  Lipid  Index  for 
wintering  Northern  Pintails  (Anas  acuta)  on  the  Southtni 
Iliiih  Plains  of  Texas,  October-March  1984-86. 


Mean  prediction'' 

Lipid  Index 

Equation 

R- 

error  (±  SE) 

R- 

1.1  Mid  1.2 

.785 

-9.921  ±  5.850'' 

.662 

(fat) 

6.16% 

1.3 

.765 

-9.043  ±  5.853 

.659 

(fat) 

6.24% 

2.1  and  2.2 

.697 

-0.0192  ±  0.0091 

,671 

(Condition  1 

Index' 

I 

7.87% 

2.3 

.7(X) 

-0.019  ±  0.0()92 

.675 

(Condition  ] 

[ndexl 

1 

7.79% 

3.1-3.4 

.7.33 

-0.050  ±  0.()()()9 

.634 

(log  fat) 

2.41% 

■"Prediction  error  expressed  ;is  a  percentage  of  the  mean  in  the  validation  data 
set. 
Negative  prediction  error  iiuhcates  o\erestimation  of  the  true  \alue. 

log  DM,  LMASS,  LWING,  and  DSEX  were 
significant  (F  <  .05).  TlnLs,  CI  was  modeled  with 
LTOTAL,  LWING,  LTARSAL,  and  LMASS  for 
sexes  separately  and  combined  (Table  3).  As  in 
model  1,  regression  coefficient  estimates  were 
stable  in  equations  2. 1  and  2.2;  when  se.xes  were 
combined,  mnlticollinearitv  betsveen  TOTAL 
and  DSEX  resulted  in  relati\cly  high  MFs  for 
these  variables. 

Age  and  sex  effects  were  significant  when  log 
fat  was  regressed  on  the  same  e.xplanaton' \ari- 
ables  used  in  model  2.  Furthermore,  the  struc- 
tural \ariables  LMASS  and  LWING  were  the 


onlv  variables  that  contributed  siguilicantK 
(P  <  .05),  but  they  were  not  homogeneous 
(F  <  .05)  between  age/sex  groups.  Therefore, 
four  equations  were  estimated  (Table  4).  DAGE 
explained  variation  in  log  fat  but  not  CI. 

Given  other  model  variables,  bodv  mass 
(MASS  and  LMASS)  consistentlv accounted  for 
the  largest  portion  of  variation  in  carcass  fat 
(Table  2),  CI  (Table  3),  and  log  fat  (Tiible  4)  of 
wintering  Northern  Pintails.  Wing  length 
(WING  and  LWING)  explained  1-5%  of  the 
variation  in  carcass  fat,  log  fat,  and  (>I  when 
other  variables  were  itlready  in  the  models. 
TARSAL  did  not  contribute  to  any  model.  Vari- 
ation accounted  for  b\'  ninth  and  tenth  pri man- 
lengths  always  differed  by  less  than  1%.  Conse- 
quently, ninth  priman'  length  was  not  tested  in 
any  model. 

In  the  \alidation  data  set  all  models 
accounted  for  69%  or  more  of  \ariation  in  car- 
ca.ss  fat  mass,  (]I,  and  log  fat  (Table  5).  All 
models  explained  less  than  70%  of  the  xariation 
in  Lipid  Index  for  \  alidation  data-set  birds.  Bias 
in  all  models  was  relatively  low  and  negative. 
Predictive  e(|uations  overestimated  fat  mass, 
CI,  and  log  fat  of  \  alidation  data-set  pintails. 

DISCUSSION 

A  useful  condition  index  will  sa\e  fimds  b\ 
eliminating  the  need  for  expensive  laboraton 
analyses  and  will  lessen  the  need  to  sacrifice 
birds  for  direct  nutrient  anaKses.  The  problems 


230 


Great  Basin  Naturalist 


[Volume  52 


associated  with  using  body  mass  alone  as  an 
index  to  condition  of  migratory  birds  have  been 
noted  (Bailev  1979,  Wishart  1979,  Iverson  and 
Volis  1982.  Johnson  et  al.  1985).  Because  indi- 
viduals \aiy  in  stnictural  size,  bod\'  mass  will 
reflect  that  \ariabilitA'  in  muscle  and  bone,  in 
addition  to  variation  in  lipids. 

Models  have  been  dexeloped  that  predict  fat 
content  in  waterfowl,  but  these  require  sacritice 
and  dissection  of  the  bird  (Woodall  1978,  Chap- 
pell  and  Titman  1983,  Thomas  et  al.  1983, 
Whvte  and  Bolen  1984).  These  equations  may 
incorporate  skin  (subcutaneous),  abdominal 
(omental),  and/or  intestinal  (visceral)  fat  mass, 
and  often  account  for  most  of  the  variation  in 
total  body-fat  content.  Our  study  was  designed 
to  develop  models  using  explanatoiy  variables 
that  could  be  applied  to  live  as  well  as  dead 
pintails. 

Miller  (1989)  developed  regression  models 
to  predict  carcass  fat  on  live  pintails  from  Sac- 
ramento Vallev,  California,  but  cautioned 
against  their  use  outside  that  region.  Our  regres- 
sion models  for  carcass  fat  provided  better  esti- 
mates of  fat  (K"  >  .71)  for  live  pintails  than  those 
developed  for  California  birds  IR'  <  .66).  How- 
ever, similar  to  Millers  (1989)  studv,  bodv  mass 
alone  accounted  for  most  of  the  variation  (R'  > 
.69)  in  pintail  carcass  fat. 

The  possibility  of  a  condition  bias  among 
water-fowl  captm-ed  in  baited  traps  versus  the 
general  population  has  been  addressed 
(Weatherhead  and  Ankney  1984,  1985, 
Buniham  and  Nichols  1985).  Hypothetically, 
birds  in  poor  condition  may  be  hungrier,  less 
wary,  and  more  likely  to  enter  a  trap  contiiining 
food.  Condition  models  could  be  used  to  test  for 
evidence  of  a  body-condition  bias,  given  that 
samples  of  pintails  captured  both  in  baited  traps 
and  bv  presumably  less-biased  methods  (e.g., 
net  gun)  are  available. 

Models  could  be  u.sed  to  test  for  annual 
variation  in  bodv  condition  and  for  chans;es  in 
condition  across  the  winter.  Ringelman  and 
Szymczak  (1985)  demonstrated  the  potential  of 
condition  indices  in  determining  spatial  differ- 
ences in  body  condition  and  the  preferabilitv  of 
condition  indices  to  use  of  body  mass  alone. 
Heppetal.  (1986)  also  used  condition  indic-esto 
docmnent  a  po.sitive  relationship  betAxcen  con- 
dition and  sun  ival  in  mallards. 

The.se  pintail  condition  models  should  be 
useful  to  waterfowl  biologists.  However,  models 
should  be  verified  when  used  outsick^  the  eeo- 


graphical  range  in  which  they  were  developed. 
For  comparisons  between  age  and  sex  classes 
we  encourage  use  of  model  3.  Research  also  may 
refjuire  knowledge  of  absolute  fat  content. 
Importance  of  accuracy  and  precision  will  affect 
model  selection.  Care  should  be  exercised  to 
restrict  model  use  to  winter  when  changes  in 
bod)'  mass  primarily  reflect  fluctuations  in  fat, 
not  fat-free  diy  mass  (i.e.,  protein  and  mineral 
fractions). 

Acknowledgments 

We  offer  our  thanks  to  A.  R  Leif,  CD. 
Olavv/sky,  D.  G.  Cook,  R  J.  Grissom,  and  R  N. 
Gray  for  field  assistance.  E.  G.  Bolen,  L.  D. 
Vangilder,  D.  H.  fohnson,  and  C.  B.  Ramsey 
provided  comments  on  the  manuscript.  The 
project  was  supported  by  the  Caesar  Kleberg 
Foundation  for  WildHfe  Consen'ation  and  the 
Texas  State  Line  Item  for  Noxious  Brush  and 
Weed  Control.  This  is  manu,scriptT-9-488  of  the 
College  of  Agricultin-al  Sciences,  Texas  Tech 
Universitv'. 

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Received  1  fuiic  /.9.9/ 
AreepU-d 22'}\nie  IW2 


Great  Basil)  Natunilist  52(3).  pp.  232-236 


EVALUATION  OF  ROAD  TRACK  SURVEYS 
FOR  COUGARS  (FELIS  CONCOLOR) 

\\  alter  D.  \  an  Sickle    aiul  Frederick  Ct.  Lindzev 


.'Kli.sTlucr — Road  track  sui"\'e\s  were  a  poor  index  of  eoutjar  ileiisitA  in  .sontlieni  LUiili.  The  weak  rekitionship  we  iound 
betsveen  track-finding  frequency  and  coug;u"  den.sit}'  imdouhtedK'  resulted  in  piut  from  the  fact  that  aviiilable  roads  do  not 
sample  properK'  from  the  nonuniformlv  distributed  cougar  population.  Howe\er,  the  significantly  positi\e  relationship  ir" 
=  .73)  we  found  between  track-finding  fre(jueuc\  and  number  of  cougar  home  langes  crossing  the  sur\('\  load  suggested 
the  technique  may  be  of  use  in  monitoring  cougar  populations  where  road  abundance  and  location  allow  tlie  population  to 
bv  sampled  properK.  The  amount  of  variance  in  track-finding  frequency  unexplained  1)\  number  of  iiome  r;uiges  o\erlapping 
suiACN  roads  indicates  the  index  ma\  be  useful  in  demonstrating  onl\-  relati\'el\'  large  changes  in  cougar  population  size. 


Kci/  uiirds:  cDiti^cir.  Felis  coneolor.  truck  .s»rrr//.  l^tali. 

Sign  left  by  animals  ha.s  been  connnonly 
u.sed  b\-  wildlife  managers  to  make  inferences 
about  population  characteristics  (Neff  1968, 
Lintlzevet  al.  1977,  Novak  1977).  This  approach 
is  appealing  becan.se  it  .seldom  recjuires  special- 
ized equipment  and  is  usually  nnich  less  costlv 
than  other,  more  intensive  techni(jues.  The 
approach  requires,  however,  that  the  relation- 
ship between  sign  and  the  population  character- 
istic of  interest  (e.g.,  size,  composition)  be 
understood. 

Track  counts  have  been  used  to  indicate 
cougar  (Fclis  co)}color)  abundance  or  change  in 
abundance,  but  population  estimates  were 
seldom  available  to  evaluate  the  validitv  of  these 
indices  (Koford  1978,  Shaw  1979,  Fitzhugh  and 
Smallwood  1988).  \'an  Dyke  et  al.  (1986)',  how- 
e\er,  conducted  road  track  sunews  in  an  area  of 
known  cougar  densit\  and  found  a  weak  rela- 
tionship (/■"  =  .18)  between  track-finding  fre- 
(jueucv  and  densitv.  Because  of  the  potential 
value  ol  this  techni(}ue  to  agencies  charged  with 
management  of  cougars,  our  objectixe  was  to 
test  again  the  relationship  between  track-find- 
ing f re(|U(Micy  and  cougar  densit\  follow  iug  [)ro- 
cedures  of  Van  l>ke  et  al.  (1986).  AdditionalK. 
we  examined  the  influence  cougar  distribution 
patterns,  as  measured  b\  cougar  home  ranges, 
had  on  track-finding  lre<|U(mc\. 


Study  Aiuv\ 

The  Bonlder-Escalante  stud\'  area  comprises 
4500  knr  of  Garfield  and  Kane  counties  in  south 
central  Utah.  Boulder,  Escalante,  and  Canaan 
moimtains  dominate  the  area  topographicallw 
and  elevation  ranges  from  1350  m  to  3355  m. 
Hot,  dry'  weather  is  characteristic  of  )nne  and 
July,  with  rains  beginning  in  August  and  contin- 
uing through  September.  Annual  precipitatit)n 
ranges  from  18  cm  at  low elexations  to  60  cm  at 
high  elevations;  axerage  temperatures  for 
Escalante  in  januan  and  juK'  are  -2.8  C  and 
24.5  C,  respectixcK"  (U.S.  Department  of  Com- 
merce 1979). 

Desert  grass  and  shrub  communities  domi- 
nate the  \egetation  with  a  sparse  o\"erstoi^'  of 
piu\()n  pine  {Finns  cdiilis)  and  juniper 
ijunipcrus  (isti'Dspcniui)  between  1350  m  and 
1800  m.  Dense  pinxon-juniper  stands  with  a 
sagebmsh  {Aiicinisid  tridciitatti)  underston' 
dominate  the  xegetation  between  1800  m  and 
2400  m.  Ponderosa  pine  {Finns  poiidcro.sa)  and 
oakbrush  {Qucrciis  <j^(ii)ihclii)  are  pn)miuent 
abo\  ('  2400  m  w  here  rock"\,  \ertical-w  ailed  can- 
Nons  with  large  areas  of  bare  sandstone  charac- 
terize the  topographv  Subalpiue  meadows  with 
suuill  stands  of  Eugelmann  spruce  (Picea 
cn<i('hn(nni),  ([uaking  aspen  (Popnhis  li\-ninl(>i(lcs). 


WyoiningCoi.iu-ratiw  Fisli  unci  Wildlife-  Hi-sr.ucli  I'liil.  I5i)\  .-^Kifi,  Uiiivcrsitv  St.itiiiii.  1„ 


.  \\voiiinii;.S207r 


O-T,--) 


19921 


CoucarThack  Slknkv 


233 


a\id  w  liitc  lir  (Al)i('s  concolor^  occur  ahoxc  2700 
111.  Hi\ er  camons  transxcrse  the  area  with  asso- 
ciated \e<z;etatioii  consistinti;  primariK'  of  Fre- 
inout  Cottonwood  {Fopulus  jrcniojitii)  and 
willow  (.SV///.V  sppJ  (Ackcrnian  19S2.  Heinker 
19S2). 

The  human  population  of  about  800  is  con- 
centrated in  the  towns  ot  Escalante  and  Boul- 
der. I.i\estock  grazing,  timber  harvesting,  and 
energx"  exploration  are  tlie  priman^  land  uses  in 
the  area.  Road  densit\'  is  about  25  km  oi  road 
per  100  km-  (\an  Dyke  et  al.  1986).  Hunting  of 
cougars  is  prohibited  on  the  stud\'  area. 

Methods 

Capture  and  \h)nitoring  Procedures 

Cougars  were  tracked  on  horseback,  treed 
with  the  aid  of  trained  hounds,  and  immobilized 
with  an  intramuscular  injection  of  ketamine 
Indrochloride  and  wlazine  h\ch"ochloride 
(Hemkeretal.  1984).  Each  immobilized  cougar 
was  fitted  with  a  collar  containing  a  motion-sen- 
sitixe  radio  transmitter  (Telonics,  Inc.,  Mesa, 
.\rizona).  Radio-collared  cougars  were  moni- 
tored with  portable  radio  telemetrv  equipment 
on  the  groimd  and  from  the  air.  All  radioloca- 
tions were  assigned  UTM  coordinates  and 
recorded  to  the  nearest  100  m.  An  attempt  was 
mad(^  to  locate  all  radio-collared  cougars  a  min- 
iiiiuin  of  once  each  week. 

Tlie  Bould(M--Escalante  stucK  area,  including 
areas  occupied  b\  collared  cougars,  was 
searched  periodicalK'  for  sign  of  new  cougars 
(e.g.,  tracks,  scats,  scratches).  When  detectetl, 
uncollared  cougars  taking  up  residence  and 
tnuisi(^nts  were  captured  and  radio-collared 

Hoad  IVack  Surxcws 

C-ougar  densit\  was  measured  as  both  the 
number  of  known  cougars  per  km"  in  the  siua  ex 
area  and  the  number  of  home  ranges  ol  inde- 
pendent cougars  oxcrlapping  the  suiacx  road. 
We  conducted  both  s\stematic  (Fitzliugh  and 
Smallwood  1988)  and  random-svstematic  (\'an 
Dyke  et  al.  1986)  road  tiack  sune\s.  Onl\'  dii1 
roads  were  sunexed. 

For  the  sxstematic  sunev  the  study  area  was 
dixided  into  three  sune\'  areas  spatialK'  and 
behaxiorally  (home  range  boundaries)  isolated 
from  the  others.  One  11.3-kni  secticMi  of  road 
xx'as  chosen  in  each  area;  roads  xx'ere  similai"  in 
elexation  change,  habitat  t\pe.  and  condition 


(substrate,  surface  condition).  Suncx  aieas  dil- 
lered  in  cU^isitx  (indejH'Uck'ut  adult  cougars  per 
km")  and  the  number  ol  home  ranges  that  inter- 
sected road  sc^'tions:  2-3  in  the  first,  4—5  in  the 
second,  and  6-7  in  the  third. 

Roads  xx'ere  sunexcnl  from  a  pickup  truck  at 
8-12  kph.  Each  road  including  both  shoulders 
x\  as  dragged  xvitli  a  conifer  tree  pulled  from  the 
rear  of  the  tnick.  The  folloxving  dax"  both  sides 
of  the  road  xvere  searched  for  cougar  track  sets 
bx  drixing  on  one  side  and  returning  on  the 
other.  A  track  set  xxas  defined  as  a  continuous 
set  of  tracks  created  bx  one  cougar  on  a  single 
occasion.  Three  to  10  days  later  each  road  xxas 
again  sin"xexed  and  dragged.  We  felt  that  after  3 
daxs  the  effect  of  dragging  xxould  be  minimal, 
antl  moxements  of  cougars  in  the  area  (Heniker 
et  al.  1984)  suggested  this  intenal  xxould  be 
sufficient  to  proxide  independent  sampling 
periods.  Dust  ratings,  determined  from  imprint 
characteristics  of  the  obseiver's  shoe  (\'an  D\ke 
et  al.  1986),  xxere  conducted  ex^ery  km  before 
and  after  dragging  to  (juantif\"  road  surface  con- 
dition. At  each  stop  the  obsener  took  10  steps, 
5  on  each  shoulder;  then  each  impressic^i  xxas 
given  a  point  xalue  from  1  to  4.  Simple  regres- 
sion anaK'ses  xvere  used  to  examine  the  relation- 
ship betxx'een  track  sets  per  km  surveyed  and 
both  measures  of  densitx.  Track  .sets  per  km 
surxexed  xx'ere  considered  the  independcMit 
xarial)le  because  onlx'these  (kita  xxould  be  ax  ail- 
able  to  the  manager. 

The  random-systematic  road  track  suiaox- 
inxolxed  dixiding  the  studx  aica  into  four  sun  ex' 
areas.  Again,  the  four  ai"(^as  were  spatially  and 
behaxiorallx  isolated  from  eacli  other.  Txx'o 
sune}"  areas  had  2 — f  cougar  liome  ranges  oxer- 
lapping  roads  and  t\x'o  had  5-7.  Each  area  had  a 
different  dcnsitx'  of  cougars  (0.0 1 7, 0.032, 0.042, 
0.057  cougars/km").  A  16-km  stretch  ot  roadxx-as 
landonilx  selected  in  each  area,  and  the  first 
ai"ea  to  be  suiACxcd  xxas  randomix  chosen.  Sur- 
x"exs  xx'ere  run  as  described  lor  sxstematic  sur- 
xexs  except  that  an  all-terrain  xeliicle  xxas  used 
and  onlx'  on(>  shoulder  ol  the  road  xxas  dragged 
Once  ;ill  tour  Avvds  had  be(^n  surxexed,  xx'e 
returned  to  the  first  aica,  randomly  selected 
different  16-km  suncx  routes  for  each  area  and 
l)egan  the  se(juence  again.  Sunex ed  roads  xxere 
not  eligible  for  resampling  until  all  dirt  roads 
xxithin  an  area  had  bcx'u  sampk'd  once.  For 
analxses.  each  16-km  section  ot  road  xxas 
di\  ided  into  segments  xai"xing  in  length  from  1 
to  10  km  depending  on  the  numlx^r  of  home 


234 


Cheat  Basin  Natlhalist 


[\i)l 


r2=0.73 
df=3 
P  =  0.066 


0  0.005  007  0075  0.02 

TRACK  SETS  PER  KILOMETER  SURVEYED 

Fig.  1.  Relationship  behveen  cougiu'  track  sets  per  kilo- 
meter and  cougars  with  home  rmiges  overlapping  tlie  snr\i\ 
road  on  the  Boulder-Escakuite  stuch'  area,  lltah,  19SS. 

ranges  o\erlapping  the  segment.  Each  segment 
tlien  had  a  home  range  oveHap  \'ahie  (2-7)  and 
was  assigned  one  of  the  ionr  densitv  xahies. 

We  examined  the  relationship  between  traek 
sets  found  per  km  sunexed  and  the  t\vo  niea- 
sin"es  of  densit\"\\dth  simple  regression  anaKsis. 
Road  segments  with  the  same  home  range  o\  er- 
lap  values  were  eombined  to  obtain  km  sur- 
veyed, as  were  road  segments  representing  the 
same  densities.  Data  points  entered  into  the 
regression  etjuations  were  the  siun  of  traeks 
found  in  eaeh  of  the  six  home  range  overlap  or 
four  densitv  categories  divided  bv  the  sum  of  km 
surveyed  in  the  respective  categories. 

We  evaluated  whether  dragginti  would 
improve  suivev  roads  with  a  simple  regression 
of  pre-drag  dust  ratings  against  post-drag  rat- 
ings. Data  from  both  road  track  sune\\s  were 
combined  to  increase  sample  size,  and  regres- 
sion slopes  were  tested  against  1.  The  number 
of  track  sets  found  on  dragged  and  undragged 
roads  was  also  compared  b\'  dividing  the  total 
luuiiber  of  track  sets  in  each  by  the  total  km 
searched  in  each. 

Multiple  regressiou  analysis  was  used  to 
examine  the  effect  of  rainfall  and  traffic  on 
one-day,  post-drag  dust  ratings.  I're-drag  dust 
ratings,  rainfall,  and  traffic  were  the  in(k^pen- 
dent  variables  considered.  We  used  two  indica- 
tor variables  to  code  the  three  levels  of  rainfall 
and  two  to  code  the  three  levels  of  traffic.  The 
three  road  surface  categories  related  to  increas- 
ing rainfall  intensit)' were:  unchanged,  dimpled 
(individual  raindrop  impressions  distinct),  and 


deformed.  Traffic  categories  were:  no  traffic, 
traffic  on  one-h;i]f  the  length  of  the  road,  and 
traffic  on  more  than  one-half  the  length. 

Results 

The  systematic  njad  track  siuAevs  were  con- 
(hicted  Mav-june  1988.  During  this  period  407 
km  of  road  v\'as  surveyed  and  two  track  sets  were 
found.  One-hundred  thiitv-five  km  (12  surveys) 
of  road  was  sun'ev'ed  in  an  area  where  2-3 
ranges  overlapped  the  suney  road,  146  km  (13 
sui"vev's)  where  4-5  ranges  overlapped,  and  126 
km  (11  sunevs)  where  6-7  ranges  overlapped 
the  survey  road.  Unequal  survev  numbers 
resulted  from  weather  or  ecjuipment  problems 
precluding  surveys  being  run.  Each  road  (11.3 
km)  was  sruveyed  in  three  hoiu's,  v\ith  tv\'o  areas 
being  surveyed  the  first  day  and  the  third  the 
next  da\\  The  two  track  sets  were  found  on  a 
road  overlapped  bv  4-5  cougar  home  ranges. 
Because  of  the  small  number  of  track  sets  found, 
these  results  were  not  regressed  against  either 
measure  of  densits'. 

Random-sv'stematic  road  track  siu-veys  were 
nni  in  Inly  and  August  1988.  During  this  period 
684  km  v\as  siu'veved  and  seven  cougar  track 
sets  v\'ere  found.  Three  hundred  fiftv  km  (37 
road  segments)  was  located  in  an  area  of  lovv- 
home-range/road  overlap  and  334  km  (42  road 
segments)  in  high.  The  number  of  km  searched 
per  day  was  16. 

We  identified  no  relationship  between  den- 
sitv, as  measured  in  cougars  per  km",  and  track 
finding  frequency  (r  =  .00,  P  -  .886,  n  =  4). 
However,  the  relationship  (Y  =  2.23  +  197X,  r 
=  .73,  P  =  .066,  ROOT  MSE  =  1,  /i  =  5)  betvyeen 
number  of  cougars  knov\ni  to  have  home  ranges 
overlapping  die  road  and  track-finding  frequency 
was  positive  (Fig.  1 ).  Tlu^  data  point  associated 
v\ ith  the  home  range  ov  erlap  value  of  7  was  drop- 
ped because  <20  km  of  road  v\'as  suneved. 
Results  from  both  one-dav  periods  and  three  or 
more  days  were  combined  for  these  analvses. 

Because  of  the  small  number  of  track  sets 
lound,  we  did  not  statisticalK  evaluate  the  rela- 
tionship beh\'een  track-tinding  frequency  and 
dust  rating  categories  or  dragged  and 
undragged  roads.  We  found  a  positive  relation- 
ship between  post-drag  dust  ratings  (Y)  and 
pre-drag  ratings  after  one  (AT)  and  three  or 
more  (X2)  days  (r  =  .54,  Y  =  6.05  +  0.875X1.  P 
<  .001,  ROOT  MSE  =  10.4,  n  =  43)  (r  =  .34,  Y 
=  3.14  +  0.707X2,  P  <  .01,  ROOT  MSE  =  4.6, 
n  =  20).  However,  we  ftiiled  to  reject  the  null 


1992] 


CorcAH Track  SrH\i:Y 


235 


li\ potlicsis  islopi'  =  1  Hii  both  cases,  iiidicatiiisj; 
that  our  iiictluKl  ol  road  drasfs^iiiu;  did  little  to 
iiiiproM'  ttaekiiiij;  inediuiii  or  that  dust  iatiu'j;s 
were  uot  sensitixe  enough  to  detect  changes  in 
the  tracking  medium.  Data  associated  with 
heaxA  rainfall  \\  t're  omitted  Irom  these  anaKses. 
Multiple  regression  anaKsis  (onc^  da\ )  relating 
[)()st-drag  dust  ratings  to  pre-drag  dust  ratings, 
lain  tall,  and  traffic  Nielded  a  three-variable 
model  that  contained  onl\- pre-drag  dust  ratings 
(A'l  1  and  rainfall  (X2,  X3)  as  the  independent 
\  ariables  (r  =  .67,  Y  =  7.65  +  0.838X1  +  0.76X2 
-  5.65X3,  P  <  .000[X1],  P  <  .583[X2],  P  < 
.001  [X3].  ROOT  MSE  =  9,  /i  =  43).  Moderate 
rainfall  had  little  effect  on  post-drag  dust  rat- 
ings. Howexer,  heaxA'  niintall  resulting  in  road 
surlace  deformit\"  had  a  deleterious  effect  on 
post-drag  dust  ratings.  The  effect  of  traffic  on 
post-drag  dust  ratings  was  not  signiHcant(F>  .05). 


location  in  determiiu'ni^  umuberof  tracks  found, 
use  of  index  \alues  to  compare  cougar  density 
betx\eeu  areas  in  tenuous.  The  probabilitx  of 
existing  road  net\\'orks  in  t\\T)  area.s  sampling 
similarh'  from  tiu^  tA\'o  po])ulations  seems  small. 
U.se  of  track  suiacns  to  document  cougar  pres- 
ence is  feasible,  but  again,  the  approach  ulti- 
mateK  relies  on  loads  intersecting  a  cougar 
home  range. 

IdealK;  roads  with  suitable  trackin*!  surface 

o 

should  be  abundant,  as  in  paits  of  the  Northwest 
where  logging  is  connnon,  and  located  .so  that 
the  home  range  of  each  cougar  would  be  inter- 
cepted. Even  in  an  ideal  situation,  howe\(M\  the 
index  maxpnne  sensitixe  onlv  to  relativeK'  large 
clianges  in  cougai"  [lopulation  size.  Twentx- 
sexen  percent  of  the  xariauce  in  number  of 
tracks  found  xx-as  unexplained  bx'  number  of 
cougar  hoiiu^  ranges  ()xerlap[)ing  sunex*  roads. 


Discussion 

The  ntilitx  of  road  track  sunex's  for  monitor- 
ing cougar  abundance  is  limited  bx' the  generallx' 
])()()r  relationship  betxveen  cougar  density  and 
track-finding  frequencx'.  Both  our  results  {>"  - 
.00).  although  based  on  a  small  sample,  and 
tho.se  of\'an  Dxke et al.  (1986)  {r  =  .18)  inilicate 
a  weak  relationship  bet\xeen  cougar  densit)  and 
track-finding  frequencx'.  The  strongest  signifi- 
cant relationship  found  bx  \'an  Dyke  et  al.  {r  - 
.61 )  resulted  from  a  nuiltiple  regression  model 
with  track-finding  frecjuencx'  the  dependent 
\  ariable  and  female  densitx;  good  tracking  con- 
ditions, aud  proxiuiitx  of  cougars  to  sunex  road 
the  iud(q)endent  \ariables.  As  the  authors 
noted,  hoxxexer.  a  biologist  xvould  sekUjin  haxe 
kuoxxledge  of  cougar  distribution  in  regard  to 
sunex'  roads. 

The  poor  relationship  documented  betxx'een 
track-finding  frequencx  and  cougar  densitx 
appears  tlie  n^snlt  of  sampling  problems,  largelx 
bexond  the  coutiol  of  the  biologist.  (Cougars  an^ 
rarelx  uuiloruiK  distribut(nl  (Hemkc^r  et  al. 
I9S4!.  and  axailable  roads,  the  sampling  sti'ata. 
are  sekkim  abundant  enough  or  optimalK 
located  to  sample  from  a  nonnniforui  distribu- 
tion. .\xailable  roads,  for  example,  could  fail  to 
intercept  anx'  cougar  home  ranges  or  could  be 
found  ()ul\  in  the  areas  occupied  bx  cougars,  in 
both  scenarios,  the  index  (tracks  found)  could 
easilx  proxe  to  be  a  poor  measure  of  change  in 
cougar  numbers  o\-er  time  in  an  area.  Likexxise, 
because  of  the  potential  importance  of  road 


AcKNow  i.i:i:)(;.\iENTS 

This  research  xvas  fimded  bv  the  Utali  Dixi- 
sion  of  Wildlife  Resources  and  administered  bx' 
the  \\\"oming  and  Utah  Cooperatixe  Fishen 
and  Wildlife  Research  Units.  We  thank  W.  j. 
Bates  for  coordinating  oiu"  project  thnigh  the 
UDWR.  H.  J.  Harioxx;  R.  A."  Poxvell,  L.  L. 
McDonald,  aiul  D.  G.  Bonett  rexiexved  initial 
drafts  of  the  manuscript.  \\^e  offer  special  thanks 
to  C.  S.  .Mecham  and  M.  (].  Mecham  for  field 
assistance  and  fimctioniug  as  houndsmen. 

LlTl'lHATlM^I-:  ClTK.n 

.AcKKHMAN,  H.  H.  19S2.  (^ouiiar  pivdation  and  ecological 
energetics  in  sontliem  Utiili.  Unpublished  masters 
tliesis,  Utah  State  University,  Logiui.  95  pp. 

FiTZHUcar  E.  L.,  and  K.  S.  Smai.i.uood  198S.  Teclnii(jue.s 
for  monitoring  monntmn  lion  population  le\eLs.  Pages 
69-71  /■;/  H  II  Smith,  ed..  Proceedings  of  the  Third 
Moimtaiii  I, ion  Workshop,  .\rizona  Came  and  Fish 
Department. 

Ill  Mkl-K  T.  v.  19S2.  Pi  )[)nlat  ion  characteristics  and  mo\e- 
ment  patterns  (jf  cougars  in  southern  Utah.  Unpub- 
lished masters  thesis.  Utah  State  Unixersitv  Logan.  66 
pp. 

ill  Aikii;  T.  P..  1'  (;.  I,lMr/.l•,^  and  B.  B.  Ac  kkhnian  19S4. 
Population  characteristics  and  moxement  patterns  of 
cougars  in  .southern  Utdi.  journal  of  Wildlife  Manage- 
ment 48:  1275-12S4. 

KoiOlU).  (J.  B.  1978.  The  welfiueol  the  puma  in  (!aliloniia. 
Camixore  I:  92-96. 

I.ixnzKY.  F.  C.  S.  K.  Thompson  and  J.  1.  nou(a:s  1977. 
Scent  station  index  of  black  bear  abundmice.  Journal  of 
Wildlife  .Management  41:  151-1.53. 

.\kfk  D.  ).  1968.  The  pellet-group  count  technique  for  big 
game  trend,  census,  and  distribution:  a  re\iew.  Joumal 
of  Wildlife  Manauement  .32:  .597-614. 


236  C;reat  Basin  Naturalist  [\oIiime52 

NOXAK.M.  1977.  Determining  the  a\erage  size  and  coinpo-  presence.  |ounKiI  (.rWildlifr  .Management  50:   102- 

.sition  of  beaver  familie.s.  |()urnal  of  Wildlife  Manage-  109. 

ment41:  751-754.  U.S.  Dhpahtmiat  OF  Conlmkiu:!'.   1979.  CJimatokweal 

Sll.uv.   II.  C.  19,9    A  monntam  lion  Held  gnid<>.  .Arizona  data    annual    snmnian.    Climatologieal    Data    Utdi 

Game  and  Fi.sli  Department  Special  Report  No.  9.  27  Sl(  1.3) 

pp. 
\\\  DvKK.  F.  G.,  R.  II.  Bkockk   and  II.  G.  Shaw    19S6. 

U.se  ol  road  track  connts  a.s  indices  of  monntain  lion  Received  U)  \oiemh  ■  ■  IMl 

Accepted  16  April  1992 


Great  13asiii  .\atiir;Ji.st  52(3),  pp.  237-244 

LEAF  AREA  RATIOS  FOR  SELECTED  RANGELAND  PLANT  SPECIES 

Mark  A.  Welt/,',  Wilhcrt  H.  Blackhuni".  and  J.  Hosier  Sin  laiitoii' 

AHSTKACr — Leaf  area  estimates  are  re([iiiretl  In  Indrolojiie,  erosion,  and  'j;ro\\  tli A  ii'kl  siniukition  models  and  are 
important  to  the  nnderstanding  ol  trtuispiration,  interception,  COo  fixation,  and  tlie  energ\  balance  for  native  pkmt 
connnnnities.  Leaf  l)iomas.s  (g)  to  leaf  area  (nim")  linear  regression  relationships  were  e\alnated  for  15  perennial  grasses, 
12  shruhs,  .md  1  tree.  The  slope  coefficient  ((So)  of  the  linear  regression  eqnation  is  a  ratio  of  leaf  area  to  leaf  hiomass  and 
is  definetl  as  the  leaf  area  ratio  [LAR  =  one-sided  leaf  area  (nim~)/()\en-dr\-  leafweight  (g)].  LAR  represents  (3(1  in  each 
regression  eqnation,  where  Y  =  P{|(X).  Linear  regression  relationships  lor  leaf  area  were  compnted  (r~  =  .84-.9S)  for  all 
28  natixe  nuige  species  after  fnll  leaf  extension.  Within-pkint  estimates  of  leaf  lU'ea  for  niesquitc  iProsojns  ^Idiidulosa  Torn 
\Ar.<^hni(liiIosa  [Torr.]  Cockll.)  or  liinepricklviish  (Zanthoxt/hnn  fag^ara  [L.\  Sarg.)  were  not  significantK' different  (P<  .05). 
LARs  for  three  of  the  shnibs  and  the  tree  were  established  at  fonr  different  phenological  stages.  There  were  no  significant 
differences  {P  <  .05)  in  LARs  for  lime  prickh-  ash,  niesqnite,  and  Texas  persimmon  {Diospijras  texana  Scheele)  after  fnll 
leaf  extension  dnring  the  growing  season.  The  LAR  relationship  forTe.xas  persimmon  changed  significantly  after  fnll  leaf 
extension.  LAR  relationships  for  Texas  colnbrina  (Cohtbrina  texemis  [T  &  G.]  Gray)  changed  in  response  to  water  stress. 


Kct/  tcanls:  h'tij  diva  index,  drought  response.  Icafhioiiiaw 

Eighh"  percent  of  the  world's  rangeland  is 
classified  as  arid  or  seniiarid  (Branson  et  al. 
UJSl  I.  i.e..  precipitation  is  less  than  e\"apotrans- 
piratioii.  Under  these  conditions  water  axail- 
al)ilit\'  is  tile  most  important  en\ironniental 
factor  controlling  plant  production  and  snni\ al 
t Brown  1977).  E\apotranspiration  (ET)  is  the 
major  component  of  the  water  balance  and  is 
estimated  to  accomit  for  96%  of  annnal  precip- 
itation for  rangeland  ecos\stems  (Branson  et  al. 
1981,  C^arlson  et  al.  1990),  with  surface  rinioff 
accounting  for  most  of  the  remaining  4% 
(Gifford  1975,  Lauenroth  and  Sims  1976.' Carl- 
son et  al.  1990). 

Ex'apotranspiration  has  Ixn^n  irieasiired  for 
selected  rangeland  plant  coimnunities  with 
Ksimcters  and  tlu^  Bow  en  ratio  method  (\\'ight 
1971,  Hanson  1976,  C;av  and  Frit.schen  1979. 
Carlson  etal.  1990).  Estimates  of  ET  for  mnnea- 
sured  rangeland  plant  connnmuties  are  usualK' 
simulated  from  hydrologic  models  (Lane  et  al. 
1984,  \\'ight  1986).  For  luclrologic  simulation 
models  to  be  biologicalK'  meaningful,  inipnned 
metliods  of  sinnilating  exapotranspiration  from 
rangeland  plant  connnnnities  are  needed.  Two 
different  approaclies  are  currently  being  used. 
One  approach  is  to  use  a  crop  coefficic^nt  (Kc) 


(W'ight  1986).  Kc  is  defined  as  the  ratio  of  actual 
exapotranspiration  to  e\apotranspiration  when 
water  is  nonlimiting.  This  empirical  method  is 
extremeh'  difficult  to  parameterize  for  range- 
lands  because  water  is  often  limiting  and  esti- 
mates of  transpiration  are  confounded  h\  soil 
water  exaporation  (Wight  and  Hansen  1990). 
Thus,  \Vight  and  Hansen  (1990)  reporied  that 
Kc  \alues  were  not  transferable  across  range 
sites.  The  second  method  is  based  on  leaf  area 
inde.x  (LAI)  (Ritchie  1972).  LAI  is  defined  as  the 
foliage  area  per  unit  land  area  (Watson  1947). 
The  LAI  method  is  uiore  process-ba.sed  than  the 
Kc  approach  and  has  Ikhmi  siiccesshdK  used  in 
se\eral  rangeland  Indrologic,  erosion,  and 
growtli/\ield  sinnilation  models  (Wight  and 
Skiles  1987,  Lane  and  Nearing  1989,  Arnold  et 
al.  1990). 

A  limitation  in  using  natural  Resource 
models,  like  the  \\'ater  Erosion  Prediction  Proj- 
ect (WTPP)  (Lane  and  Nearing  1989),  is  in 
dexeloping  L.\I  c-oefficients  for  rangeland 
[)lants.  LAI  is  difficult  to  measiu-e  because  of  the 
drought-deciduous  nature  of  certain  shrubs,  in 
wliicli  sexcral  c\cles  of  leal  initiation  and  defo- 
liation occur  within  a  single  growing  season 
(C;anskoi)p    and    Miller    1986)    and    seasonal 


.,USD,\.  .Xgriciiltural  Rp.search  Senice.  Southwest  V\atersliecl  Researcli  Center.  2()()()  F,;Lst  Allen  Road.  Tucson.  .Arizona  8.57194.596. 
"Northern" Plains  Area  Adniinistratne  OITice.  2625  Redwing  Road.  Suite  ^50.  Fort  Collins,  Colorado  80.526. 


231 


23(S  (;hkat  Basin  Naturalist  [Volume  52 

TaBI.K  1.   DfSfriptioii  of  studv  sites,  raii^e  sites,  and  soil  series  oC  species  exaliiated  (or  leaf  area  to  leaf  hioiiuiss 
relationsliips. 


Frost- 

.Mean 

i>p'r 

In 'c 
[)eriod 

Location 

Range  site 

(mm) 

(days) 

Soil  series 

Soil  famiK 

T<)iiihstoii(\  AZ 

I,iine\  upland 

.■35(i 

239 

Stronghold 

('oarse-loaiiiN,  mixed 
thermic,  Ustollic  Calciorthid 

Meeker.  CO 

(;Ia\('\  slopes 

200 

ISO 

Degater 

Clav,  montmorillonitic, 
mesic,  Tvpic  Caniborthid 

Sidnev.  MT 

Siltv 

.300 

130 

\ida 

Fine-loamv,  rui.xed,  T\pic 
Argboroll 

Chickaslia.  OK 

Loani\  praiiie 

927 

200 

(;rant 

Fine-silty.  nii.xed,  Udic 
.\rgiustoll 

Cliiekaslia.  OK 

iM'oded  prairie 

927 

200 

Eroded 
C;rant 

Fine-siltv,  niLxed,  Udic 
Argiiistoll 

Ft.  SiippK.  OK 

Dnne 

.597 

200 

Pratt 

Sandy,  mixed,  thermic, 
Psammentic  Haplustalf 

Wooilward.  OK 

Shallow  praii'ie 

5S4 

200 

Oiiinlan 

Loam\-,  mixed,  thermic, 
shallow  T\pic  Ustochrept 

Alice.  TX 

Fine  sand\  loam 

710 

2S() 

Miguel 

Fine,  niLxed,  h\perthermic, 
Udic  Palenstalf 

Soiiora,  TX 

Shallow 

009 

240 

Punes 

Fine-loam\-,  mixed,  thermic. 
T\pic  Calciustoll 

cliaiiti;c.s  ill  leaf  .size,  shape,  antl/or  tliickues.s  i.s  with  the  leaf  area  ratio  (LAR)  method  (Rad- 

re.siilt    IVoni    water,    nutrient,    and    chemical  ford  1967).  LAR  is  defined  as  the  ratio  of  leaf 

.stresses  ((>utler  et  al.  1977,  (>urtis  and  Luchli  areaper  unit  weight  ofplant  material.  The  slope 

19S7).  Foliar  surface  area  of  irregular-shaped  coefficient  On)  of  the  linear  regression  e(juatit)n 

tree  leaxes  has  l)een  estimated  b\-  coating  the  is  a  ratio  of  leaf  area  to  leaf  biomass  and  is 

Iea\es  witli  a  monolayer  of  glass  heads  and  mea-  defined  as  the  leaf  area  ratio  [LAR  =  one-sided 

suring  displacement   (Thompson   and   Le\ton  leaf  area  (nnn-)/oven-diy  leaf  weight  (g)].  LAR 

1971)    and   1)\    estimating   from    photographs  represents    Po    in    eacli    regres.sion    equation, 

(Miller  and  Scliultz  1987).  Miller  et  al.  0987)  where  Y  =  P(,(X).  LAI  can  be  calculated  as  the 

estimated  total  surface  area  of  juniper  foliage  product  of  LAR  and  live  biomass  per  unit  area, 

from  projected  leafar(\i determined  from  a  leaf  Tlie  objectixe  of  this  study  was  to  determine 

area  meter.  Miller  et  al.  suggested  this  method  LARs  for  selected  rangeland  species, 
underestimated  leaf  area  by  10%  diic  to  leaf 

owrlaj).  Cregg  (1992)  reported  that  knif  area  MATERIALS  AND  METHODS 
could    be    satisfactoriK'    (\stimate(l    from    leaf 

weight  or  xolume  ior  Juiiipcnts  vir^iiiiaiia  and  The  study  area  included  nine  range  sites  in 
J.  .sco})til<>niiit.  llowexcr.  leafar(>a  r(>lationships  fUe  states  and  was  part  of  the  USDA  Water 
differed  In  crown  position  and  seed  source.  Erosion  Prediction  Project  (WEPP)  (Table  1). 
Sapwood  area,  stem  diameter,  trec^  height.  The  dominant  plants  on  each  range  site  were 
canopy  area,  and  canopy  \()lume  ha\e  been  exaluated.  LARs  for  15  grasses,  12^ shrubs,  and 
correlated  to  total  .shrub  biomass  and  leaf  bio-  1  tree  were  deyeloped  (Table  2).  Selected 
mass  (Ludwiget  al.  1975,  Brown  1976.  Ritten-  rangeland  .species  were  sampled  once  during 
house  and  Snexa  1977.  Whi.senant  and  Burzlaff  the  sununcM-  of  1987  near  Tombstone,  Arizona; 
1978.  Cianskopp  and  Miller  1986,  Hughes  etal.  and  in  1987  near  Meeker,  Cok)rado;  Sidney, 
1987).  In  contrast,  onl\  a  few  studies  ha\('esti-  Montana;  Chickaslia,  Ft.  Supply  and  Wood- 
mated  leaf  area  and  LAI  for  rangeland  plant  ward,  Oklahoma;  and  Sonora,  Texas,  sites.  Sea- 
communities  ((;olf  1985,  (;au.skopp  and  Miller  sonal  fluctuations  in  LAR  for  du'ee  shrubs  and 
1986,  and  Ansley  et  al.  1992).  oiu^  tree  were  exaluated  near  Alice,  Texas,  in 

An  eftecti\(>  method  is  needed  to  iinpro\e  1 985  and  1986. 

LAI  estimates  lor  natural  resource  models.  One  Vov  k^af  area  (k'termination  grass  leaf  bioma.ss 

potential  a[)pr()acli  lor  impnning  LAI  (>stimates  from  10  raii(k)mK  located  ().25-nr  (jnadrats  was 


19921 


Ranc;ela\u  Leaf  Area  K.vnu.s 


239 


TMii.!-;  2.  Ixjcation  of  stucK'  sites,  sample  dates,  Iieitjlit  class,  iiniiiher  of  samples,  and  species  exaliiated  for  leaf  area  to 
at  liiomass  relatiousliips. 


Height  class  (iii) 


Species 


Location  Sample         0-11-2  2-3  3—4  >4  (ionimon  name  Scientific  name 

date 


ihston 


Meeker,  CO 


AZ  Aucr.  1983 
Au>i.  1983 
An<i.  1983 

Aug.  1983 
Ano;.  1983 
Aug.  1983 
fmie  1987 
June  1987 


6  6 

7  8 


Si(lnc\.  M  r  |uK  1987 

|iil\  1987 

Chiekaslia.  OK  |une  1987 
I  line  1987 
"liiiie  1987 

Chifkaslia.  OK  "|mie  1987 
June  1987 

I  line  1987 


10 
15 
15 
10 
10 

10 
10 
10 
10 
10 
10 
10 


10 


Ft.  SuppK.OK     |nne  1987 

10 

|une  1987 

10 

|une  1987 

10 

Woodward.  OK    |inie  19S7 

10 

|une  1987 

10 

Mice.  TX               .\Ia\  1985 

4 

4 

4 

4 

Aug.  1985 

2 

2 

2 

0 

No\-.  1985 

2 

-) 

2 

■-> 

Jan. 1986 

NA' 

Apr.  1986 

2 

2 

2 

2 

Ma\- 1985 

5 

5 

5 

5 

Aug.  1985 

3 

3 

3 

3 

Nov.  1985 

3 

3 

3 

3 

Jan. 1986 

3 

3 

3 

3 

Apr.  1986 
\la\- 1985 

3 
5 

3 
5 

3 

3 

Aug.  1985 

5 

5 

Nov.  1985 

5 

5 

Jan.  1986 

5 

5 

Apr.  1986 
.\Ia\-  1985 

5 
5 

5 
5 

.\ug.  1985 

5 

5 

Nov-.  1985 

5 

5 

Jan. 1986 

NA 

A]K-.  1986 

5 

5 

Soiiora,  TX           |niie  1987 

10 

June  1987 

10 

June  1987 

10 

Little  l("af  sumac 
Tarbusli 

Hrooiii  snakeweed 

Creosotel)usli 
Desert  /.iimia 
Mariola 

Shatiscale  saltl)usli 
\\'\()ming  big  sagehnisli 

Needle-and-tl  u'ead 

Western  wheatgrass 

Indiangrass 

Big  hluestem 

Little  bluesteni 

Buffalograss 

Seribners  dicliai 1 1 1  lel i 1 1 m 

Sand  paspalnm 

Sand  sagebrusli 
Tall  dropseed 
Sand  lo\egriiss 
Haii^v  grama 
Sideoats  grama 
I  lonex  mesiiuite 


Hliiis  inicr()j)liijll(i  Kngelm. 
Floiireiisia  ccmiia  DC. 
(Uiticrrczia  sarotlirae  (Pursh) 

Hritt.  6c  Rusb\. 
Ijirrcd  tridoitata  (DC.)  Coxille 
'Aiimhi  puiuila  Cra\ 
hnihciiiiDu  iiicanitm  H.B.K. 
.\lri])lcx  cotifeiiifolki  (Torr.  &  Frem. )  Wats. 
Aiicinisia  trklenlala  sulwp. 

ut/(>inin<iensis  Beetle  &  Young 
Stipa  coDKita  Trin.  &c  Hupr. 
.\<^r()j)i/r()ii  fiinithii  R\db. 
S(>r<i^lui.slniin  iiiitmis  (L.)  Nash 
.\iiclr<>i)i><^()n  gcrardii  Vitnuui 
Scliizaclii/hiiin  scoparium  (Mich.x.)  Nasli 
Biichlof  (Idcti/loklcs  (Nutt.)  Fngelm. 
niclunilhcliitm  olif^osaiUlies  (Scluilt.) 

(aiikl  \ar.  scrihiicrianuin  (Niish)  Could 
I'dspaliim  sctdcenm  Miclrc.  \ar. 

strdiniiu'iiiii  (Nash)  D.  Banks 
.A livmisid  jihfolia  Torr. 
Sporoholus  aspcr  ( Michx. )  Kunth 
Erogro.s/Z.s-  tiicliodcs  (Nutt.)  Wood 
Boiitcloiid  liirsuta  Lag. 
Boittchnui  aiiiipc'iulula  (Michx.)  Torr. 
Prosopis  gidiululosd  Torr.  \iu". 

"laiuhilosd  (Torr.)  C-tx-kll. 


5        Lime  priekK  ash  Zdiillioxi/liiin  faodra  (L.)  Sarg. 


Texas  colubrina  Colnbrind  tcxciisi.s  (T.  6c  C.)  Gray 


Texas  persimmoTi  Diospyms  texaiui  Scheele 


White  tridens  Tridtiis  dlhe.sccns  (N'asev)  \Vo<it.  &  Standi. 

(>"urK  mescjuite  llilarid  l)clan<ieria  (Steud.)  Nash 

Texas  wintergrass  Slijxi  Iciicotiiclui  Trin.  &  Riipr. 


'No. sample 


cted  (or  dtxidiions  shrubs  and  tp 


240 


Great  Basin  Naturalist 


[\  oluine  52 


Tablf,  .3.  Mean  and  standard  error  oflcat  hioniass  and  leaf  ;irea.  and  linear  regression''  model  slope  eoettieients  (LAR  ' 
relating  leafiu-ea  to  leafbionuLss  for  selected  rangeland  grasses  and  shnibs  sampled  after  f'nil  le;if' extension. 


Species 


Grass  Ks 
Needle-and-tliread 
Western  wiu-atgrass 
Indiangrass 
Little  l)ln(^stem 
Big  hlnesteni 
Buff;ilo  grass 
Scrihners  dicliantheliiuu 
Sand  paspdnm 
Tiill  dropseed 
S;uid  lovegrass 
Hain  grama 
Sideoats  grama 
White  tridens 
Texas  wintergrass 
(JurK'  mesqnite 

SllKlBS 
Desert  zinnia 
Mariola 

Broom  snakeweed 
Little  leaf  snmac 
Tarhnsh 
Oeosott-husli 
Sand  sagebrnsh 
Shadscale  saltbnsli 
Wyoming  big  sagebnish 


Leaf  biomass 


3.6 
2.0 
S..5 
2  7 

1.3 
1.5 
1.3 
1.5 
0.9 
0..S 

0.  r 
o.r-) 

0.7 
1.2 

o.s 


1.6 

3.5 
•3.7 
3.9 
3.7 
.3.0 
3.2 
3.9 
5.3 


SE 


O.SO 
0.33 
1.56 
0.3S 
0.45 
0.22 
0.21 
0.23 
0.15 
0.12 
0.13 
0.22 
0.16 
0.24 
0.15 


0.10 
0.40 
0.51 
0.71 
1 .00 
0.19 
0.58 
O.Sl 
0.S.3 


Leaf  area 
(nmi") 


SE 


3,580 

5,760 

82,670 

28,030 

11.290 

6,820 

15,300 

7,580 

8,500 

8,650 

4,360 

5,240 

.3,980 

8,.32() 

5,270 


9,440 
19.410 
11.160 
22,0.50 
2.3,.360 
16,790 

5,9.50 
10,5,30 
18,220 


900 

902 
1,3.50 
4,710 
2,213 
1,091 
2,601 
1,1.36 
1,3.34 
1,3.S3 

769 
2,8.36 
1,007 
1,.361 

925 


580 

1,2.S(I 

920 

331 

20.3 

910 

1 .257 

2,047 

2,715 


LAR 

{nim"g    ) 


1.040 

.98 

2,910 

.98 

9,440 

.96 

10,780 

.98 

12,970 

.86 

5,680 

.97 

16,110 

.96 

6,890 

.95 

9,.390 

.99 

11.380 

.98 

5,890 

.99 

10,210 

.98 

5,8.30 

.98 

6,720 

.95 

6,620 

.99 

5,700 

.89 

5,690 

.84 

2,700 

.96 

4.700 

.91 

6,100 

.97 

3,660 

.86 

2,010 

.98 

2.640 

.98 

3,340 

.97 

''.All  areuiweiglit  regressions  were  sigiiilkaiil  at  /'  « 
'Ix'af  area  ratio  ( 1.AR)  represents  ^n  in  eaih  ici^re 


■  V  =  p„iXi, 


used.  Cirass  hioinass  in  each  (juadrat  was  clipped 
to  a  2()-mni  stubble  height  and  separated  by 
species  into  lix'e  or  dead  leaves.  Li\'e  lea\"es  were 
placed  in  plastic  bags  on  ice  for  later  determina- 
tion ot  leal  area.  The  lea\es  were  flattened  and 
[)laced  between  clcnir  plastic  sheets  and  then 
processed  tlnough  a  leaf  area  meter.  Leaf  area 
was  determined  with  a  Li-Cor  .3()()()'  leaf  area 
meter  to  the  nearest  1  mm"^.  The  samples  were 
then  oven-dried  at  fSO  C  for  threc^  da\s  and  dn 
mass  determined. 

To  ensure  that  .samples  ol' shrubs  and  trees 
represent(Hl  the  full  range  of  size  of  plants  pres- 
ent, a  .stratific^d  random  sampling  procedure  w  as 
used.  Height  classes  of  1  m  were  adiitrariK 
chosen,  and  plants  were  selected  randoniK  from 
each  class.  As  a  result,  total  number  of  plants 
.sampled  \  aried  among  .speci(>s  depending  upon 
the  range  of  plant  heights  (Table  2). 

An  open-ended cul)e  (250  mm  on  a  side)  was 
used  to  sample  shrub  and  tree  leaf  biomass.  The 


The  ii.se  ot  a  trade  or  linn  name  in  this  papi-r  is  tor  reader  inlornialion  .ind 
does  not  in)pl\  endorsement  In  the  U.S.  Department  of  .-Vgrienhnre  ol  .in\ 
prixlnet  or  service. 


sample  cube  was  placed  in  an  area  considered 
representatixe  of  the  entire  canop\',  and  the 
lea\es  within  the  area  were  remoxed  In*  hand. 
LARs  were  determined  in  the  same  manner  as 
for  grasses. 

Within-plant  \ariabilit\'  of  LARs  was  e\alu- 
ated  for  four  mes({iiite  trees  and  foiu"  lime 
prickh'  ash  shnibs  in  Mav  1985  near  Alice, 
Texas.  Fifteen  sample  cubes  were  randomlv 
located  and  sampkxl  from  each  of  the  four  raes- 
(jiiite  trees.  For  the  lime  prickK  ash  shrubs  12 
sample  cubes  were  hanested  from  each  of  the 
four  shrubs.  LAR  was  determined  in  the  same 
inanntM-  as  pre\ionsl\  described.  A  one-wax' 
anal\ sis  of  \ariance  was  used  to  test  for  differ- 
ences (F  <  .05)  among  the  slopes  of  the  regres- 
sion e(juations  within  plant  canop\'  b\  species 
(Ste(>l  and  Torrie  1980).  Within-plant  LARs 
were  not  significanth'  different  for  lime  prickly 
ash  and  mescjuite  hi  May  1985.  Based  on  these 
relationships,  one  sample  per  plant  was  utilized 
during  the  reinaind(M-  of  th(>  stiuK. 

Three  shrubs,  lime  prickK  ash,  Texas  per- 
simmon, and  Texas  colubrina,  and  one  tree. 


19921 


Raxcklam)  Li-:af  Area  Kviios 


241 


Table  4.  Mean  and  standard  error  of"  leaf  biomass  and  leaf  area,  and  linear  regression''  model  slope  coefficients  (LAU 
lating  leaf  iUX'a  to  leaf  bionuiss  for  selected  rangeland  shrubs  and  trei'  on  a  line  sandv  loam  range  site  near  /Vlice.  Texius. 


Species 

Date 

Leaf  liiomass 

SE 

Leaf  area 

(mm") 

SE 

LAR 

(nnn"g'  ) 

r' 

Unie  priekK  ash 

Max-  19S5 

4.7 

0.73 

45,180 

1,450 

8,760  a'' 

.99 

Ang.  19S5 

4.2 

0.63 

40,330 

1,530 

8,730  a 

.98 

N(n-.  1985 

5.6 

0.89 

43,360 

1,460 

8,670  a 

.98 

Jan. 19S5 

4.9 

0.76 

44,310 

1,450 

8,870  a 

.98 

Apr.  19S6 

5.3 

0.65 

52,730 

1,580 

8,690  a 

.98 

\h'S<jnite 

Mav  1985 

6.5 

0.87 

57,830 

1,610 

8,990  a 

.98 

Aw^.  1985 

5.7 

0.64 

56,040 

1,470 

8,780  a 

.98 

Nov.  1985 

5.5 

0.70 

48,460 

1,410 

8.630  a 

.98 

Jan.  1985 

\A'' 

Apr.  1986 

6.4 

0.81 

59.100 

1,470 

9,290  a 

.98 

Texas  persimmon 

Max  I9S5 

4.6 

0.64 

49.960 

1,940 

10,590  b 

.96 

An<j;.  1985 

4.1 

0.65 

41.670 

1,780 

10,.360b 

.98 

No\-.  1985 

4.8 

0.59 

51.060 

1.790 

10,1.30  b 

.98 

Jan.  1986 

4.6 

0.68 

44.720 

1,900 

10,020  b 

.98 

Apr.  1986 

4.7 

0.69 

64, 150 

2,070 

12,660  a 

.97 

Texas  colnbrina 

Max- 1985 

4.9 

0.78 

55,070 

2,020 

10,310  b 

.98 

.-\ng.  1985 

5.2 

0.89 

57,010 

1,720 

10.110  b 

.98 

Nox-.  1985 

3.8 

0.65 

55.380 

2.090 

13.360  a 

.98 

Jan.  1986 

NA 

Apr.  1986 

4.1 

0.71 

41,760 

1,880 

10.230  1) 

.98 

''.\11  area:  ueiglit  regres.sions  were  signilitaiit  at  P  <  .05. 
Leaf  area  ratio  (LAR)  represents  |3ii  in  each  regression,  w'liere  V  =  3ii(X). 

'Parameters  in  the  columns  by  species  sliaringa  common  letter  are  not  signilicaiitK  different  if  <  .0.51  lia.sctl  < 
'  No  sample  was  collected  for  deciduous  slirnbs. 


;)f  slope  test. 


honex"  nie.s({uite,  were  .selected  for  e\  iiluation  ot 
.seasonal  fluctuation  in  LAR.  Hone\  mesquite, 
Texas  j3ersininion,  and  Te.xas  colnbrina  are 
drought-deciduous  while  lime  prickK  ash  is  an 
exergreen.  Sauij:)le  dates  were  selected  to  cor- 
resjiond  to  the  jihenological  stages  of  ( 1 )  niaxi- 
niuni  leaf  area,  (2)  peak  drought  defoliation,  (3) 
autunui,  just  prior  to  winter  leaf  fall  and  dor- 
mancy, and  (4)  after  winter  leaf  fall  for  the 
ck'ciduous  shnili. 

The  Statistical  Analxsis  Sxsteni  (SAS  1982) 
was  utilized  to  tnaluatc  linear  regression  rela- 
tionships,  Y  =  Pi,  +  Pi(,X),  between  leaf  biomass 
and  leaf  area.  Where  Y  is  estimated  leaf  area 
(nmr),  p,,  is  the  intercept,  pi  is  the  slojoe  (LAR 
coefficient  as  defined  bv  Radford  1967  in  mnr 
g  ),  and  X  is  leaf  biomass  (g).  The  intercept  was 
tested  to  determine  if  it  w^as  significantK  differ- 
ent (P  <  .05)  from  zero.  The  intercej^t  was  not 
significantK  ditfert^nt  from  zero  for  all  sjoecies. 
Therefore,  tht^  data  were  reanal\y,ed  and  pre- 
.sented  using  a  linear  regression  model,  Y  = 
P(i(X).  similar  to  that  reported  by  Coombs  et  al. 
(1987)  and  Ansley  et  al.  (1992)  for  estimating 
LAR.  All  statistical  tests  were  judged  significant 
at  P  <  .05  unless  otherwise  stated.  A  homogene- 
ih  of  sloj^e  test  was  used  to  test  for  differences 


among  the  slojx^s  of  the  regression  equations 
(LAR)  between  sample  periods  within  spcH'it^s 
(Steel  and  Torrie  1980). 

Results  a.\d  Discussion 

Leaf  area  of  graminoids  was  highl\-  corre- 
lated with  leaf  biomass  for  all  species  within 
samj3le  dates  (Table  3).  The  LAR  for  perennial 
grass  leaf  area  ranged  from  2910  to  16,110  mnr 
g~'.  The  LAR  for  shnibs  and  trees  ranged  from 
2010  to  13,360  nun-  g  '.  Goff  (1985)  also 
reported  significant  lin(^ar  regression  relation- 
sliips  (r  =  .83-. 97)  for  LAR  for  1  1  natixe  grass 
species  in  southern  .\rizona.  Golf  rejiorted  that 
the  lintnir  regression  coefficients  for  stem  area 
to  stem  biomass  (SAR)  ranged  from  32  to  739f 
of  the  LAR  and  the  mean  SAR  was  44%  of  the 
mean  L,'\R. 

There  was  no  significant  seasonal  \ariation 
in  L\R  for  lime  prickK-  ash  and  mesquite  (Table  4 ). 
Although  there  was  no  significant  sea.sona]  dif- 
ference between  mescjuite  L.\R  relationships,  a 
gradual  decrease  in  the  LAR  from  May  through 
Noxember xx'as apparent  in  1985.  Furthermore, 
the  LAR  xxas  larger  in  April  1986,  though  it  xxas 
not  significantK-  different  from  1985  sampling 
dates.  Moonex-  et  al.  ( 1977)  found  that  the  sj)ecific 


242 


Giiivvi"  Basin  Naturalist 


[\  olume  52 


leaf  densitv'  (nig  innT")  of  niesquite  leuM's 
increased  over  the  growing  season.  The  densit}' 
ranged  from  0.{)()()4  nig  nnn "  in  the  spring  to 
0.01 7  nig  mm'  in  die  fall.  This  corresponds  with 
a  leaf  area  change  of  5880  to  25,000  nnii'  g  '. 

Ansle\-  et  al.  ( 1992),  working  in  north  central 
Texas,  reported  that  LAR  of  niescjnite  ranged 
from  9916  to  5944  ninr  g'.  Mesquite  LAR 
declintnl  from  Ma\  throngli  Angust  1987,  hnt 
stabilized  from  Angnst  through  September  fol- 
lowing substantial  precipitation.  In  1988 precip- 
itation was  substantialK'  less  than  in  1987,  and 
the  mean  I.AR  was  significantly  lower  than  in 
1987.  LAR  followed  the  same  pattern  in  1988, 
declining  from  a  high  of  6877  in  the  spring  to  a 
low  of  4996  mm"  g'  in  October.  Anslev  et  al. 
(1992)  speculated  that  the  decline  in  LAR  was 
caused  b\-  cell-wall  thickening  in  response  to 
dning  conditions,  based  on  the  work  of  Kramer 
and  Ko/.k)wski  (1979). 

The  siinilarit\' in  LAR  across  sampling  dates 
f  ron  1  th  is  stud\'  may  be  partially  explained  in  th  at 
sam[)ling  was  not  initiated  until  all  leaves  were 
lulK  expanded  (or  approximatelv  lour  weeks.  In 
addition,  Ajiril,  Ma\;  June,  and  September  pre- 
cipitation was  significantlv  above  the  k)ng-terni 
average  ])recipitation  and  no  noticeable  water 
stress  was  apparent  in  the  trees  sampled.  Nilsen 
et  al.  (1986)  indicated  that  relati\e  leaf  area  of 
phreatoplntic  mesfjuite  {P.  olanchilosa  \ar.  tor- 
rcijana)  in  tlu^  Sonoran  desert  of  southern  (iai- 
ifoniia  remained  nearK  constant  from  Ma\ 
through  \o\ember.  Maximum  leaf  area  was 
maintaiiK^d  throughout  the  liottest  and  driest 
months  ol  the  wav  \ia  access  of  deep  stored  soil 
water  by  taproots.  When  water  availabilit}'  to  the 
normally  phreatophytic  mesquite  was  reduced, 
total  leaf  area  was  reduced  (Nilsen,  Virginia,  and 
Jarrell  1986).  We  hvpothesi/xMl  that  nies(juite 
lea\-es  reach  a  stable  weight  at  niaturit\  and  the 
lack  of  water  stress  during  the  growing  season 
prevents  the  changes  in  leaf  weight  (o  leaf  area 
reported  by  Ansley  et  al.  (1992).  Changes  in  leaf 
weight  as  a  result  of  translocation  ol'  sugars, 
.starches,  other  compounds,  and  insect  damage 
could  not  be  detected  or  .separated  from  cell- 
wall  thickening  from  water  stress  witliin  the 
precision  of  sampling  in  om-  stud\. 

Texas  persimmon  LAR  in  April  1986  was 
significantl)-  greater  than  for  sampling  dates  in 
1985.  Meyer  (1974)  reported  that  Texas  persim- 
mon produces  two  tvpes  of  leaxcs:  a  large  leaf  in 
the  center  of  the  canopy  and  a  smaller  leaf 
around  the  i^erimeter  of  the  plant.  The  leaxes 


arc*  initialK  light  green  in  color  and  become 
glabrous  after  elongation  ceases.  As  the  leaf 
matures,  the  x)'leni  and  bundle  fibers  become 
increasingly  lignified  and  the  leaf  tunis  dark 
green,  with  the  underside  becoming  densely 
covered  with  trichomes.  Leaf  modification  is 
complete  by  early  July.  The  lower  LAR  of  Texas 
persimmon  leaxes  in  1986  was  attributed  to  the 
leaxes  not  being  fullv  elongated,  with 
incomplete  development  of  trichomes  and  lig- 
nification. 

LAR  relationships  forTexascolubrinaxaried 
seasonally.  LAR  was  similar  during  the  early 
growang  seasons  in  May  1985  and  April  1986, 
and  in  August  1985.  In  November  the  LAR  was 
33%  greater  than  during  other  sample  dates 
(Table  4).  Rasal  leaves  of  Texas  colubrina  are 
approxiniateK  10  times  larger  than  the  outer 
canop\  leaves.  In  response  to  an  extended  diy 
period  in  fuK  and  August,  Texas  colubrina 
dropped  95%  of  its  leaves.  The  onl\-  leaves 
I'etained  during  this  diy  period  were  the  large 
basal  leaves  in  the  center  of  the  shnib.  The 
significant  difference  in  LAR  between  the 
sample  dates  was  attributed  to  the  different 
proportion  of  leaf  tspes  and  not  the  change  in 
specific  weight  of  the  leaves. 

(Tunskopp  and  Miller  (1986)  reported  sim- 
ilar significant  seasonal  changes  in  LAR  for 
Wxoming  big  sagebrush.  Tlie\'  speculated  that 
the  greatest  proportion  of  seasonal  \  ariation  was 
due  not  to  the  development  or  alterations  in 
starch  and  sugar  accuniulations  but  rather  to 
changes  in  the  proportion  of  larger  persistent 
leaves  to  smaller  ephemeral  leaves. 

Shiiib  leaf  biomass  to  leaf  ai'ea  was  liighlv 
correlatetl  for  the  nine  other  shrubs  sampled 
(Table  3).  The  LAR  for  slinib  leaf  area  ranged 
from  2010  to  6100  nuir  g"'.  Other  researchers 
have  also  reported  satisfacton  results  in  relating 
l(\il  biomass  to  leaf  area  (Schilesinger  and 
(^habot  1977,  Kaufmann  et  al.  1982,  Ganskopp 
and  Miller  1986)  within  sample  date.  Based  on 
the  seasonal  xaiiabilitvin  LAR  for  Texas  persim- 
mon and  Texas  colubrina  in  this  stiuK  and  the 
findings  ol  (Tunskopp  and  Miller  (1986)  in  ea,stern 
Oregon  tor  Wvoming  big  .sagebni.sh,  we  c;ui  state 
that  s(\i.s()nal  \ariabilit\  in  tlie.se  and  other 
(h'ought-deciduous  shmbs  is  an  important  source 
of  xaiiation  tliat  needs  to  be  accounted  for  when 
simulatinti  LAI  owv  the  entire*  tirowiii<i  sea.son. 


19921 


Raxcelam)  Lkaf  Ahka  Ratios 


243 


Conclusion 

For  tlic  spt'cics  saiiipli'd.  leal  hioiiiass  is  a 
reliable^  estimator  of  leaf  area,  llowexer,  for 
some  slinil)  species,  seasonal  differences  in 
cle\('loi)m(Mit  and  shedding  of  different  t\pes  of 
l(^a\es  and  leal  nioiphological  de\elopnient  c-an 
prodnce  significant  temporal  flnctnations  in 
LAR.  Caldwell  et  al.  (198f)  reported  that  for 
semiarid  hnnciigrasses,  leaf  blades  of  regrowing 
tillers  had  grc\it(^r  photos\nthetic  capacit\'  than 
blades  on  nnclipped  plants.  This  resulted  in 
greater  carbon  gain  for  clipped  plants  and  an 
increased  photosMithesis/transpiration  ratio. 
Nowak  and  Caldwell  (1984)  reported  that  the 
photosvnthetic  rate  for  both  clipped  and  nn- 
clipped plants  decreased  with  age  of  the  lea\es. 
Cnrrent  rangeland  Indrologic  simnlation 
models  do  not  account  for  changes  in  LAR  or 
exapotranspiration  rates  as  a  function  of  age  of 
the  leaf.  [)r()poition  of  leaf  t\pe,  or  compensa- 
ton  photosNiithesis  rate  increases  following 
defoliation  due  to  grazing.  Models  currently 
utilize  a  fixed  coefficient  for  calculating  LAI.  If 
significant  adxances  in  modeling  e\'apotranspi- 
ration  on  langelands  are  to  b(^  made, 
improxements  in  the  relationships  used  to  sim- 
ulate exapotranspiration  that  incoiporate  these 
processes  will  he  needed.  The  LAR  method  of 
calculating  LAI  exaluated  in  this  studx"  proxides 
a  fast,  reliable  method  of  estimating  LAI  neces- 
san  to  [)arameterize  these  hydrologic  simula- 
tion modc^ls.  To  account  for  the  seasonal 
diffen^nces  in  L.\R  for  Texas  persimmon  and 
Texas  colubrina,  a  xx'eighted  average  based  on 
season  of  xear  is  recommended  for  parameter- 
izing tlu^WT.PP  model.  For  plants  like  m(^s(|uite 
and  lime  piicklx  ash,  one  LAR  xalue  can  be  u.sed 
in  non-drought  vears.  For  xears  xxith  significant 
dn  periods,  a  decrease  in  LAR  of  10^0%  max 
need  to  be  accounted  for  xxith  non-phreato- 
piixtic  nies(|nite.  as  indicated  bx  this  xxork  and 
thatof  Ansleyetal.  U992j. 

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244 


Great  Basin  Naturalist 


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nitrogen  trratiiicnts.  Water  Rcsijurce  Rescardi  12: 
437-442. 

li  i)\\  k;.  J.  A.,  J.  F,  Rf.y\oi.])s  and  !'  1).  Wiiitson  1975. 
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Mkykh.  R.  E.  1974.  Moipliolo<);\-  and  anatomy  of  Texas 
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WiiisENANT,  S.  G.,  and  D.  F  Burzlaff.  1978.  Predicting 
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WiciiT  J.  R.,  G.  L.  Hanson,  and  K.  R.  Gooley  1986. 
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Idaho. 


Received  r,  June  19^1 
Accepted  U)  Septeniher  1992 


Cwrat  Basin  NatiinJist  52(3),  pp.  245-252 

ECOLOGY  AND  MANACiEMENT  OE  \IEi:)US AHEAD 
(TAENIATHERUM  CAPUT-MEDUSAE  SSP.  ASPERUM  [SIMK.J  MELDEKIS) 

|ames  A.  Youiuj; 

AhsTKaci'.  —  Mediisahcad  is  aiiotlicr  in  the  cxlcnsixc  list  ol  annual  herbaceons  spciirs  to  invade  the  temperate  desert 
raniielands  ol  tlie  Great  Basin.  Medusaliead  is  not  preferred  1)\  lar^e  herl)i\i)i(s  and  apparentK'  is  not  preferred  In' 
'j;rcnn\ores.  I  lerl)age  of  tills  annual  grass  enlianees  ignition  and  spread  ofwildl'ii'es.  Mcdus.ilie.id  is  liigliK  eonipetitixc  witli 
the  seedlings  of  native  species  and  is  prohabK'  tlie  greatest  threat  to  thi'  l)iodi\crsit\  of  ihc  natnral  \fgetation  that  has  \et 
been  aeeidentalK  introduced  into  the  Great  Basin.  Hespite  the  ob\ions  biologii-al  disni|)tions  that  are  associated  with 
niedusahead  imasion.  the  species  offers  awealtli  ol  oppoit unities  for  stndents  to  exaniiiie  the  nieehanisni  b\- which  this 
species  is  so  successful.  Stutlents  of  cNoIntion.  |ilant  pli\  sioli>g\.  and  ecologx  max  find  this  species  to  be  an  excellent  model 
lor  colonization. 

Kci/  uonl.s:  uwdusdhfdd.  Tat'niatherum  ca]")ut-mednsa(^  aiiiiiKil  tr^rass.  coloiiiziit^siwrii's.  uihlfircs.  grr/z///g. 


Ill  the  nianagemeiit  of  natural  resources 
tlieii'  are  certain  problems  that  1)\  their  persis- 
tence, inagnitnde  of  ecological  disniption.  and 
eeononiic  impact  refuse  to  dissipate  as  a  result 
ol  being  ignored  and  neglected.  UnFortunateK 
tor  range  management,  niedusahead 
iTaoiidtlicrunt  c(ipnt-})icdus(ic  [L.]  Nevski)  is 
that  t\pe  ot  problem.  During  the  1950s 
niedusahead  was  considered  among  the  most 
pressing  problems  on  the  rangelands  of  C^alifor- 
nia,  Idaho,  and  Oregon.  A  great  deal  of  research 
effort  was  dexoted  to  solving  the  niedusahead 
problem,  \aliiable  information  was  learned 
about  the  ecoplnsiologs  and  s\niecolog\"  of 
iiiechisahead.  (Control  methods  were  dexeloped 
using  herbicides.  The  fatal  link  in  integrated 
])rograms  for  the  suppression  of  niedusahead 
populations  pro\ed  to  be  artificial  rexegetation 
technologies  after  niechisahead  was  controlled. 
The  nature  of  tlu^  sites  infested  had  more  to  do 
with  this  lailiire  tliaii  the  weed  itself,  especialK 
in  the  bitermountain  area.  The  recent  discox'en 
ol  niedusahead  in  northern  Utah  has  renewcxl 
interest  in  suppressing  tilis  rangeland  weed. 

M\  purpose  in  this  review  is  to  relresh  our 
c()llecti\-e  memories  about  medusaliead  ecologx 
and  management. 

Ta.xoxomv 

As  is  olten  the  case  with  an  introduced  s])e- 


cies,  there  has  bec^i  coulusion  about  the  c'orrect 
scientific  taxon  lor  medusaliead.  Tlie  first 
description  ol  niedusahead  in  a  Nortli  American 
flora  used  the  tiixon  Eh/iims  caput-nwdiisac  L. 
(Howell  1903).  There  is  apparent  agreement 
that  niedusahead  is  a  member  of  the  trilx^ 
Triticeae  of  the  grass  tamil\-.  There  is  also  appar- 
ent agreement  among  moiphologists  and  c\  to- 
geneticists  that  niedusahead  does  not  fit  in  the 
genus  Elipmis.  N'arious  autliors  haxc  placed 
niedusahead  in  Hordciiin  or  Hordcli/iitits. 
Newski  (1934)  proposed  tliat  medusaliead  was 
tniK"  a  different  genus  and  published  the  name 
Tdcnidfhcrum.  jack  Major  ol  the  I  iii\ cisilx  ol 
(California  suggestcHl  in  19f-)()tliat  material  intro- 
duced to  the  United  States  was  Taeiiiathcnini 
(ispcriiiii  (Major  et  al.  1960).  Based  on  the 
European  and  Hussian  literature.  Major 
reported  tliat  I'dcuidlUcnitn  contained  three 
geograpliic  and  moiphologicalK  tlistinct  ta\a,  T. 
cdjntt-incdnsdc.  T.  dsfxTiiiii.  and  T.  crinituiii. 
Tlie.se  three  sjx'cies  are  loiiiid  in  the  Mediterra- 
nean region  and  extend  eastward  into  central 
.\sia.  Alter  examiiiiiiij;  the  European  material, 
growing  in  place.  .Xhijor  decided  the  I  iiited 
States  introduction  was  T.  d.spcruin. 

The  Danish  scientist  Signe  Frederikseii 
rexi.sed  the  genus  in  19Sfl  He  kept  the  same 
three  taxa,  but  reduceil  them  to  subspecies  of 
Tdciiiddici'iini  cdpul-nicdusdc.  Positixe  identifi- 


.\griciiltmal  Hesearcli  Senice.  U.S.  Dep.irttm-iit  c>IAi;ntiiltun-.  920  \all<-\  Hoad.  Hi-iio.  \\-\a(la  Sy.5I2. 


245 


246 


c;riv\t  Basix  Naturalist 


[Volume  52 


cation  to  the  lowest  le\el  possible  is  ahsolntely 
essential  for  am  proposcxl  biological  control 
program  for  medusaheacl.  According  to 
Frederiksen's  revision,  subspecies  crinituiii  has 
a  \'er\'  strict  spike.  Subspecies  captif-nicdiisae 
lias  a  large  open  spike  with  straight  awais.  The 
spike  of  subspecies  a.spcniDi  is  intermediate 
with  angled  awns.  Subspecies  (ispcnim  is  die 
only  one  of  the  three  witli  pronounced  barbs 
coated  with  silica  on  the  awns.  Apparently,  the 
correct  taxon  for  the  medusahead  of  western 
North  America  is  Tacniatlicnuu  capiit-nicdiisac 
ssp.  aspeniin  (Simk.)  Melderis  (Frederiksen 
1986). 

Taeniathenun  caput-niedusae  ssp.  capiit- 
nicdiisae  is  mostK  restricted  to  Portugal,  Spain, 
southern  France,  Morocco,  and  Algeria.  It  has 
been  collected  outside  this  area  in  Europe  and 
Asia,  but  Frederiksen  considers  it  adxentitions 
in  the.se  areas.  Subspecies  chnitnt)i  is  found 
from  (ireece  and  Yugosla\ia  eastward  into  Asia. 
Subspecies  aspcniiii  completely  overlaps  the 
distribution  of  the  other  two  subspecies.  All 
three  subspecies  integrate  with  each  other. 
ApparentK'  onlv  the  one  subspecies  occurs  in 
Nortli  America.  Does  this  indicate  one  or  vev\ 
limited  introductions? 

.Mechi.saliead  is  predominanth'  self-polli- 
nated. Genetically  the  genus  appears  to  stand 
alone  in  genomic  relations  within  the  Triticeae 
(Schooler  1966,  Sakamoto  1973).  ApparentK 
Tacniafhcrinii  has  a  genome  that  is  distinct,  but 
faintK'  related  to  those  of  Fsadii/rostachi/s, 
Dasijpi/nitii,  Erciiiopiptim,  or  Hordcmii 
(Frederiksen  and  Hot  hue 'r  1989). 

IIlSTOm-  IN  NOHTII  Amkhica 

.Medusahead  was  first  collected  in  the 
United  States  near  Roseburg,  Oregon,  on  24 
June  1 887  by  Thomas  Jefferson  Howell  ( 1903). 
It  was  next  collected  ncnu-  Steptoe  Butte  in  east- 
ern Washington  in  1901  b\- George  Xixsex  (Piper 
and  Beattie  1914),  followed  by  a  collection  n(>ar 
Los  Gatos,  California,  in  1 908  In  Charles  I  litch- 
cock  (Jepson  1923).  Medusahead  certaiuK 
attracted  the  noted  agrologi.st.  McKell.  Hobin- 
.son,  and  Major  (1962)  commented  on  diis 
.strange  initial  distribution  reaching  390  miles 
north  and  450  miles  south  from  the  point  of 
initial  collection.  EaH\  lied)arium  .specimens 
show  a  rapid  spread  to  the  .south  into  California. 

J.  F.  PechantH-  made  die  first  collection  in 
Idaho  in  1944  near  Payette  or  about  ISO  miles 


.south  ol  Steptoe  Butte  (Sharp  and  Tisdale 
1952).  Fred  Rennertold  jack  Major  he  had  seen 
medusahead  near  Mountain  Home,  Idaho,  as 
early  as  1930,  and  Lee  Sliaq)  had  reports  from 
ranchers  that  the  species  occurred  in  Idaho  as 
early  as  1942.  The  medusahead  infestation  in 
Idaho  increased  to  30,000  acres  b\'  1952.  Min 
Hironaka  estimated  that  150,000  acres  were 
infested  by  1955,  and  the  Bureau  of  Land  Man- 
agement estimated  700,000  acres  were  infested 
by  1959.  At  that  rate  of  spread  it  appeared  that 
all  of  Idaho  would  be  infested  by  the  end  of  the 
next  decade.  The  spread  of  medusahead  slowed 
and  nearly  continuous  infestations  remained 
confined  to  Gem,  Payette,  and  Washington 
counties  in  southwestern  Idaho.  There  were 
several  spot  infestations  in  surrounding  counties 
(Hironaka  and  Tisdale  1958). 

Medusahead  spread  soutli  in  California  to 
Santa  Barbara  on  the  southern  coast  and  Fresno 
Count\'  in  the  interior  vallexs.  The  rapid  spread 
from  southwestern  Oregon  through  northern 
and  central  California  occiuTed  in  annual-dom- 
inated grassland,  oak  {Qtierciis)  woodland,  and 
chaparral  commimities.  These  areas  lia\e  a 
Mediterranean  t\pe  climate  with  hot,  di")'  sum- 
mers and  cool,  moist  falls,  winters,  and  springs. 
Germination  occurs  in  the  fall  and  flowering 
and  seed  set  in  the  spring. 

In  northea.steni  California,  east  of  the  Sierra 
Ne\ ada-Cascade  rim,  medusahead  inxasion 
occiuTed  at  a  much  slower  rate.  In  the  Pitt  Ri\'er 
drainage,  vegetation  is  an  intergrade  of  Oregon 
white  oak  (Qucrciis  ^(irnjaiui)  woodlands, 
cismontane  California  species,  western  juniper 
ifiiiupcnis  occidental  is),  ponderosa  pine  {Pi)uis 
pondcrosa)  woodlands,  and  sagebrush  {Aifeini- 
.s7V/)/buncli grass  communities  more  tspical  of 
the  Intennountain  area. 

Medusahead  was  discoxt'red  in  the  Great 
Basin  at  \erdi,  Nevada,  in  the  earK  1960s.  Iso- 
lated inf(\stations  were  subsequentK  found 
along  the  eastern  front  of  the  Sierra  Ne\ada  in 
ar(>as  wliere  range  sheep  bauds  used  to  concen- 
trate^ wliile  waiting  for  mountain  summer  pas- 
tures to  be  Iree  of  snow. 

In  northeastern  C'alifoinia  in  the  CTreat  Basin 
duiing  (h("  earl\'  1960s,  tluM'e  were  two  small 
inlestations  in  citv  lots  in  Snsanxille  and  a  small 
infestation  at  the  old  slu^ep-shearing  site  of 
\iew  land  along  the  niilroad  above  Wendel,  Cal- 
ilornia.  .Another  isolated  infestation  occurred  at 
die  mouth  of  Fandango  Pass  in  Suiprise  Valley. 
B\    the  earK    1970s,   medusahead  was   uearK" 


19921 


ECOlXKiY  AND  MANA(;KMENT()F  MKDI  SAIlKAl) 


247 


continuous  ox'er  al)out  60. ()()()  at'ics  of  tlic 
Willow  (]reek-Tal)l('Ian(ls  northeast  ol  Susan- 
\illc.  ('uncntK.  alter  lour  \ears  ol  extreme 
(lrouu;iit.  uiedusahead  s[)()t  iutestatious  occur 
o\tM-  [)erliap.s  an  additional  uiillion  acres  on  the 
westcM'u  maitjiu  ol  the  (weat  Basin. 

HlOl.OCV  OF  MEI^USAIIIvM; 

Medusaliead.  in  some  wavs,  is  a  rerun  of 
clieatgrass  {Bronms  tectoniin)  imasion. 
(dieatgrass  dominates  secondan'  succession  in 
a  majorit)'  of  sagehnisli/bunchgrass  communi- 
ties in  the  Great  Basin  and  proxides  a  significant 
portion  of  the  forage  base  for  lixestock  grazing. 
Howe\er,  there  are  hiiihK'  si(j[nificant  differ- 
ences  in  the  ecolog\-  of  the  t^vo  grass  species 
(Harris  and  Wilson  1970,  Al-Dakheel  1986). 

Germination. — The  canopsis  of  medusa- 
head  is  less  than  a  millimeter  wide  with  a  \en 
shaip  callus  and  an  elongated,  non-geniculated 
awii.  The  medusaliead  caiyopsis  is  covered  with 
small  barbs  of  silica.  \^cious  is  the  best  descrip- 
tion for  this  grass  canopsis.  Bo\e\"  et  al.  (1961) 
determined  that  medusaliead  had  a  much 
higher  ash  content  (o\er  10%)  than  other  grass 
species  and  the  ash  was  about  7o7c  silica.  Hea\A' 
deposition  of  .silica  occurs  on  the  barbs  of  awns 
and  the  epidermis  of  leaxes. 

For  the  \ast  majorit\  of  collections  of 
cheatgrass  from  the  Intermountain  area,  seeds 
are  ready  to  germinate  when  tlun  are  mature. 
No  pregermination  treatments  are  necessar\ 
(Young  and  Exans  1982).  For  collections  from 
the  Great  Plains  and  perhaps  the  Columbia 
Basin,  seeds  may  have  a  brief  afterripening  dor- 
mancy. In  contrast,  seeds  of  medusaliead  have  a 
temperature-related  afterripening,  and  germi- 
nation will  not  occur  except  at  cold  incubation 
temperatures  for  about  90-120  daws  after  matn- 
ritx  (Young  et  al.  1968).  Nelson  and  Wilson 
( 1969)  found  this  (loi-manc\  was  eontiolled  In 
niat(M-ials  located  in  the  awn. 

The  high  silica  content  on  the  herbage  of 
medusahead  makes  the  litter  xen  slow  to 
decompose.  Harris  (1965)  described  Hie  chok- 
ing accumulations  of  medusahead  litter  that 
built  up  for  sexeral  \ears.  We  exalnated  the 
germination  of  seeds  of  \arious  annual  grass 
species  in  medusaliead  litter  (Young  et  al.  1971a). 
Allelopathy  was  not  suspected,  but  rather  the 
ph\  sical  holding  of  seeds  out  of  contact  with  the 
surface  of  the  seedbed.  Medusahead  seeds  ger- 
minate \-er\-  well  without  the  callus  end  of  the 


seeds  touching  a  moisture-supplving  substrate, 
bi  this  situation,  germination  of  medusahead 
seeds  is  controlled  In'  the  relatixe  humiditx 
within  the  litter  and  tlie  incubation  tempera- 
ture, which  of  course  influences  the  relatixe 
humidity.  The  needlelik(\  xitreous  carxopses  of 
medusahead  appear  hxdrophobic  rather  than 
hygroscopic.  Not  ouK'  can  medusahead  seeds 
germinate  under  diese  conditions,  but  thex  can 
be  dried  until  the  priman-  root  is  dead;  then, 
lolloxxing  remoistening.  a  nex\- adxcntitious  root 
xvill  dexelop. 

Raxuiond  Exans  and  I  demonstrated  x\  hat  a 
great  modifxing  influence  litter  coxer  can  be  to 
the  surface  of  seedbeds  on  temperate  desert 
rano;elands  in  terms  of  n^dncing  extremes  in 
temperature  and  consening  moistm-e  (Exans 
and  Young  1970,  1972).  (^anopses  of 
s(juirreltail  {Eh/nuis  In/strix)  are  xcn-  similar  in 
moqihological  appearance  to  those  of 
medusahead.  As  I  xxill  discuss  later,  s(juirreltail 
seedlings  are  one  of  the  fexx-  natixc  species  that 
can  become  established  in  undisturbed 
medusahead  stands.  Both  Tacniaflicniin  and 
Ely  nuts  are  members  of  the  tribe  Triticeae,  but 
thex"  do  not  share  the  same  genome. 

Medusahead  populations  easiK-  exceed  1000 
plants  per  square  foot,  and  thex-  are  phenotxpi- 
callx'  plastic  enough  that  a  population  of  1  plant 
per  square  foot  can  exceed  the  seed  production 
of  1000  plants  per  square  foot  (unpublished 
research,  ARS,  Reno,  Nexada).  Huge  seed 
banks  dexelop  in  medusahead  conunuuities  in 
the  litter  and  .soil.  Medusahead  seetl  accjuires  a 
dormancx  in  the  field  similar  to  that  of 
cheatgrass  (see  Young  et  al.  1969).  The.se  dor- 
mant seeds  respond  to  eiuichment  of  the  seed- 
bed xxith  nitrate  and  gibberellin  (Exans  and 
^bnug  1975). 

Life  cycle. — Medusahead  seeds  can  ger- 
minate in  the  fall,  xxinter,  or  spring;  and  seed- 
lings liom  all  seasons  can  j^roduce  fioxx'ers  and 
seeds  earix  in  the  sunnner.  The  striking  thing 
about  the  medusahead  life  cxcle  is  that  it 
matures  from  2  to  4  xveeks  later  than  other 
annual  grasses.  All  those  famous  botanists  and 
range  scientists  xx'ho  xxere  out  on  the  range  di.s- 
coxering  nexx-  infestations  of  medusahead  xx'ere 
led  to  the  populations  In  the  bright  green  color 
xx'lien  all  other  aimuals  in  either  cisniontane 
Galifoniia  or  the  Great  Basin  xvere  broxxn. 

R.  L.  Piemeisel  recognized  the  dominance 
of  alien  plant  species  in  the  secondan  succes- 
sion of  disturbed  satiebrush  communities  in  the 


248 


(;rk,\t  Basin  Naturalist 


[\  oluiiie  52 


InterniounUiin  area  (PicMiieisel  1951).  Wbrkiiiii; 
on  the  Snake  Rixer  plains  of  Idaho  (hirin<j;  the 
1930s.  Piemeisel  enumerated  (k)niinance  honi 
Russian  thistle  (Salsola  austral  is)  to  tumble 
mustard  {Sisipnhriuni  altissiinuin)  to  eheat- 
grass.  Continued  disturhanee  tended  to  per- 
petuate cheatgrass  donn'nance.  According  to 
Piemeisel,  the  animal  species  that  germinates 
first,  reaches  nuL\imum  growth  and  maturit\ 
first,  lias  the  capacit\'  to  withstand  crowding, 
and  has  high  seed  production  is  the  one  that  will 
occup\'  and  persist  in  serai  sagehmsh  plant  com- 
munities. Piemeisel  always  noted  that  no  one 
species  had  a  clear  tlominance  on  all  these  char- 
acteristics, hut  on  balance  cheatgrass  was  the 
clear  winiKM'. 

Medusahead  contradicts  sexeral  of 
Piemeisels  criteria.  Medusahead  seeds  are  ini- 
tiall\-  ck)rmant  with  temperature-related 
afterripening  requirements,  while  cheatgrass 
seeds  ha\e  no  such  restraints.  This  works  only 
for  initial  establishment  because  once  seed 
banks  are  established  with  seeds  with  ac(|uired 
dormancy  our  research  indicates  that 
cheatgrass  and  medusahead  seeds  ha\e  e(jual 
chances  of  germination  with  the  initial  moisture 
exent  in  the  tall.  Medusahead  does  take  iruich 
longer  to  mature  than  cheatgrass  and  perhaps 
tumble  mustard.  Min  Hironakaand  his  students 
hax'c  conducted  a  series  of  excellent  experi- 
ments comparing  the  cumulatiye  growth  cunes 
for  roots  and  aerial  structures  of  medusahead 
and otlier grasses  (Hi ronaka  1961.  Hironakaand 
Sindelar  1973.  1975).  Dr.  Ilironaka  concluded 
from  these  studies  that  the  comparati\e  growth 
phenokjgx  restricts  medusahead  to  areas  with 
suiplus  .soil  moisture  alter  cheatgrass  normally 
matures. 


Soils 

Ha\ni()nd  Eyans  noted  in  the  195()s  when 
medusahead  first  inxaded  Glenn  and  Colusa 
counties  in  the  northern  Sacramento  \alle\-  of 
C^alifoniia  that  medusalunid  appeared  to  be 
restricted  to  clay-textured  soils  (personal  com- 
munication). Malloiy  (1960)  reported  on  this 
relationship  at  the  1960  meeting  of  the  California 
.section  of  the  Societx'  for  Range  Manag(Mn(Mit. 
Burgess  Kay  made  the  cliilling  obsenation  that 
after  a  cotiple  of  decades  this  relationship  disap- 
peared and  medusahead  occupied  many  sites 
with  coarser-textured  soils  (personal  communi- 
cations). 


In  the  Intermomitain  area.  Ma\narcl 
Fosbergof  the  Unix  ersit\'of  Idaho  reported  that 
the  medusahead  infestations  along  the  Colum- 
bia l^ixer  in  Washington,  Idaho,  and  Oregon 
were  restricted  to  clay-textured  soils  (Fosberg 
1965).  He  suggested  that  the  greater  soil  mois- 
ture-holding capacity  of  these  soils  allowed 
medusahead  to  complete  its  life  c\cle. 

Building  on  the  work  of  Fosberg  and 
Ilironaka,  I  sampled  the  plant  communities  in 
the  medusahead  in\asion  area  along  the  western 
edge  of  the  Great  Basin  (Young  and  E\ans 
1970).  Medusahead  was  foimd  on  the  margins 
of  man\'  degraded  meatlows  where  moisture 
relationships  probabK  fa\ored  it  oyer 
cheatgrass.  A  much  larger  area  of  infestation 
was  sagebnish/grass  communities.  The  sage- 
brush communities  consi.sted  of  mounttiin  big 
sagebrush  (Aiiciuisia  tridcntata  ssp.  vaset/ana) 
on  .soils  with  sand\  loam  to  loam-textured  sur- 
face horizons  and  often  well-dexeloped  argillic 
horizons.  A  second  series  of  sagebrush  commu- 
nities consisted  of  low  sagebiTish  (A.  arbuscida) 
growing  on  soils  wdth  clay-textured  surface  hori- 
zons. Harn"  Simimerfield  (retired  soil  scienti.st, 
Soil  Consenation  Senice  and  Forest  Senice, 
USDA)  suggests  the  low  sagebrush  soils  share 
the  same  development  as  the  big  sagebrush 
soils,  but  the  surface  horizons  have  been 
removed  by  erosion  (personal  commimication). 
On  the  Modoc  Plateau  of  northeastern  Califor- 
nia these  two  series  of  plant  conununities  divide 
the  landscape  about  ecjuallv  (Young  et  al.  1977). 
In  the  northern  Crreat  Basin  low  sagebnish  con- 
stitutes onK  about  lO^f  of  the  total  sagebrush 
vegetation. 

On  the  western  edge  of  the  Great  Basin, 
medusahead.  in  nonmeadow  situations,  is 
largely  restricted  to  low  sagebrush  potential 
plant  coimnunities.  Would  this  restriction  to 
cla\  soils  change  over  time  as  appears  to  have 
happened  in  cismontane  California?  Remem- 
ber the  studies  of  Raymond  Evans  tliat  showed 
competition  in  the  cismontane  portion  of  the 
Califoiuia  annual  grasslands  is  initiallv  for  light, 
while  in  chcMtgrass  communities  of  the  Inter- 
mountain  area,  competition  is  oyenvhelmingly 
foi-soil  moisture  (  Pa  aiis  ct  al.  1970.  1975). 


WiLDFlHKS 

Accumulations  of  litter,  on  areas  where 
medusahead  is  t\stablished,  will  bum.  McKell, 
Wilson,  and  Kav  (1962)  had  initial  results  tliat 


19921 


Ecoi.ocv  A\i)  M  \\  \(:i:\ii:ntof  Mkdusaiikad 


249 


seeiiu^d  to  iiulicatc  that  hiirniiiii;  \\;is  tlic  answer 
to  the  control  of  nu'diisalicad.  Ilic  idea  was  to 
hui'ii  stands  wliilo  coinpctinij;  annnal  (2;rassrs 
were  tulK  mature  and  niedusiiliead  seeds  were 
still  in  the  inflorescences.  This  stucK'  showed 
hm^ned  seeds  would  not  (germinate.  Ilowexer, 
the  hurned  seeds  were  apparentK  incubated  at 
20  (-",  and  unburned  fresh  seed  would  not  ha\e 
germinated  at  that  temperatiu-e.  We  tried  a 
series  of  burning  experiments  on  the  Pitt  Ri\er 
bidian  reservation  and  found  burning  taxored 
medusahead  (Young  et  al.  1972 1.  We  helped 
Forest  Sen  ice  range  consenationists  evaluate 
burning  treatment  on  low  sagebiiish  communi- 
ties on  the  Silver  Lake  district  of  Fremont 
National  Forest  in  Oregon;  the  off-season  burns 
appeared  to  favor  remnant  perennial  grasses 
over  medusahead. 

Low  sagebnish  comnumities,  because  of 
lack  of  herbaceous  cov  er,  are  relativelv  resistant 
to  the  spread  of  wildfires.  Big  sagebrush  com- 
munities, especiallv  those  with  cheatgrass 
undenstories,  are  ven  subject  to  the  spread  of 
wildfires.  Invasion  of  medusahead  into  low 
sagebnish  communities  introduces  wildfires  to 
these  communities,  perhaps  for  the  first  time 
since  they  were  in  pristine  condition.  Perennial 
grass,  forb,  and  shrub  cover  are  all  negativelv 
correlated  with  medusahead  cover  in  the  west- 
em  Great  Basin  (Young  and  Evans  1970). 

Grazixc  Preference 

It  is  obvious  from  the  above  discussion  that 
preference  bv  grazing  animals  plays  an  impor- 
tant part  in  the  successional  dynamics  of 
medusahead  coiiiinunities.  One  of  the  few  stud- 
ies of  medusahead  palatabilitv  was  conducted 
on  the  northern  coast  of  California  using  sheep 
in  small  hurdle  plots  (Lusk  et  al.  1961).' Under 
the  conlinetl  conditions  of  thc^  studv.  sheep  uti- 
lized medusahead  when  it  was  green.  When 
faced  with  no  choice,  thev  used  some  herbage 
after  the  medusaliead  matured.  How  nuich  uti- 
lization of  medusahead  would  occur  in  temper- 
ate desert  situations  is  unknown. 

C'heatgrass  .stands  [)ut  a  tremendous  produc- 
tion of  grass  canopses  into  a  local  eco.svstem. 
\ertebrate  granivores  have  adapted  to  this  food 
source.  Savage  et  al.  (1969)  showcxl  in  feeding 
trials  that  Chukar  Partridges  {Alcrtoris  ^raeca) 
could  not  utilize  the  caiyopses  of  medusahead 
as  a  food  source.  These  birds  are  dependent  on 
cheatgrass  seeds  in  the  fall  and  winter.  We  do 


not  know  what  the  iulluence  of  medusahead 
inv  asion  would  be  on  other  granivores.  Seeds  of 
other  recently  introduced  weeds  in  temperate 
ck\sert  coimnunities,  such  as  those  of  barbvvire 
Russian  thistle  {Salsola  paulsvnii),  are  heavily 
prey(xl  upon  by  granivores.  I f  cheatgrass  popu- 
lations crash  because^  of  replacement  bv 
medusahead,  what  ha]-)jx'us  to  cheatgrass  seed 
predators':^ 

A  studv  c'onductetlat  Washington  State  Uni- 
versitv  illustrates  that  granivore  preference 
works  both  ways  in  plant  succession.  Bird  pop- 
ulations prefer  the  seeds  of  native  perennial 
grass  species  over  tho.se  of  clu^itgrass  and 
medusahead  (Goebel  and  Bern  1976). 

Utilization  of  medusiiliead  bv  large  herbi- 
vores of  infested  ranges  results  in  increased 
incidence  of  injun  from  the  seeds.  Data  on  the 
level  of  injun'  are  not  available  for  domestic 
livestock  and  certainlv  not  available  for  wildlife. 

Control  of  Medu.saheai:) 

Kavdev  eloped  highlv  technical  and  vorv  suc- 
cessful control  and  revegetation  techni(jues  for 
the  annual-dominated  rangelands  of  cismon- 
tane  California  using  the  herbicide  [paraquat 
( I,l'-dimethvl-4,4'  bipvridinium  ion)  and  spe- 
cialized seeding  equipment  (Kav  1963,  1966, 
Kay  and  McKell  1963). 

This  technique  was  not  successful  in  the 
Intermountaiu  an^a  because  medusahead 
[)lants  were  not  susceptible  to  paracjuat  in  the 
temperate  desert  environment  antl  the  annuiil 
legumes  that  proved  so  adapted  to  (ismontane 
California  were  not  adapted  to  the  sagebmsh 
environment  (Young  et  al.  1971b'.  Ilerbicidal 
fallow  techni(jues  using  atrazine  (6-chIoro-N- 
ethv  1-N '-[  1 -methv  letlivi  1- 1 .3,5.-tria/,ine-2,4-di 
amine)  or  dalapon  i2.2-dichl()ropropanoic 
acid),  and  mechanical  fallow  techni(jues  were 
developed  lor  use  in  the  (ireat  Basin.  Milken 
and  .Miller  ( I9S())  provide  a  summarv  of  lierbi- 
cidal  control  measures  applied  experimentallv 
for  tlie  control  of  medusahead.  A  large  part  of 
the  area  infested  with  medusahead  in  the  west- 
ern Great  Basin  was  never  adaptcnl  to  these 
treatments  because  of  surface  rock  cover  that 
prohibited  tillage  or  seed-drilling  techniques. 
The  current  mass  cancellation  of  federal  regis- 
tration for  uses  of  herbicides  on  rangelands  and 
the  failure  of  federal  land  management  agencies 
to  a(k)pt  the  use  of  herbicidal  revegetation  tech- 
ni(jues  have  made  the  use  of  these  techniques 


250 


Great  Basin  Naturalist 


[\'olunie  52 


impossible.  Landfornis  and  soils  ol  the  sites 
where  niedusahead  is  spreading  into  temperate 
desert  rangelands  are  eritieal  laetors  in  the  eeo- 
logical  suppression  ot  this  speeies. 

Nature  of  Medusaiikad-infested 
Landscapes 

The  landseape  ol"  the  western  Great  Basin 
where  medusahead  has  in\aded  is  eomposed  of 
a  series  of  fairly  reeent  basalt  flows  that  eom- 
prise  the  Modoc  Plateau  and  the  extreme  south- 
ern extension  of  the  Columbia  Rixer  Basalts. 
Superimposed  on  the  flows  are  clays  from  a 
Tertiary-age  lake.  This  lake  was  much  older  than 
pluvial  Lake  Lahontan,  which  lapped  at  the 
lower  margins  of  the  flows.  The  old  lake  left 
thick  beds  of  cla\'-textured  sediments  occasion- 
all)'  interbedded  with  diatomaceous  earth.  The 
clay  minerals  are  predominantly  double  lattice 
forms  that  expand  and  contract  with  moisture 
content.  This  exj^iansion  and  shrinkage  has 
sorted  basalt  rock  from  the  buried  fk)ws  into 
giant  polygons  and  pressure  ridges  until  por- 
tions of  the  landscape  resemble  arctic  ice  packs 
that  are  black  instead  of  white. 

There  are  a  host  of  topoedaphic  situations 
within  this  wilderness  that  support  specific 
assemblages  of  plants;  however,  the  landscape  is 
characterized  by  upland  areas  of  residual  soils 
with  loam-textured  surface  soils  that  support  big 
sagebrush  and  clay-textured  surface  soils  that 
support  low  sagebrush.  \'ast,  nearK  lexel 
benches  of  lake  sediments  support  swirling 
mosaics  of  basin  big  sagebrush  (Arfeinisid 
triclentata  ssp.  trident ni(t)  and  a  recentl\- discov- 
ered t}pe  of  sagebrush,  a  subspecies  of  low 
sagebnish  known  as  Lah(jntan  sagebrush.  The 
basin  big  sagebrush  occurs  in  depressions  whei'e 
erosional  products  accumulate  on  .soils  with 
cla\-textured  surface  horizons,  a  ven  unusual 
occurrence  for  the  Great  Basin.  The  Lahontan 
sagebrush  communities  occur  on  the  lake  bed 
clay  sediments  that  are  veneered  with  thin 
layers  of  subaerially  deposited,  coarser-textured 
soil. 

Wind  erosion  products  accunuilate  under 
the  shnib  canopies  and,  coupled  with  organic 
matter  from  leaffall,  build  mounds  under  the 
shrubs  while  miniplayas  develop  in  the  inter- 
spaces. Eckert  et  al.  (1989)  have  described  and 
experimented  with  the  seedbeds  of  these 
mound  interspace  situations,  particularly  the 
vesicular  crust  that  forms  in  tlie  interspaces  and 


limits  establishment  of  perennial  grass  seed- 
lings. 

The  area  of  medusahead  inxasion  in  the 
western  Great  Basin  is  a  microcosm  where 
events  in  soil  and  plant  ecolog\'  that  influence 
millions  of  acres  in  the  Intermountain  area  are 
brought,  b\  fortuitous  combinations  of  ph\sical 
and  biological  parameters,  into  shaip  focus.  In 
the  medusaliead  in\  asion  area,  lake-deposited 
red  clay  is  in  obxious  disc()ntinuit^•\\^th  the  thin, 
gravish  surface  soil.  \n  imdisturbed  profiles  of 
this  situation  the  influence  of  alle\iation  of  sub- 
aerial  deposited  material  is  apparent  on  the 
structure  of  the  clav  subsoil,  indicating  the 
antiquity  of  this  process  (personal  communica- 
tion, Robert  Blank,  soil  scientist,  ARS,  USDA). 

Accumulations  of  medusahead  litter  change 
wildfire  characteristics,  and  the  shiiib  compo- 
nent of  the  plant  communitv'  is  eliminated.  Con- 
tinued grazing  of  medusahead-dominated 
grasslands  is  extremely  deleterious  on  remnant 
perennial  grasses  because  of  differential  grazing 
preference.  In  contrast  to  medusahead, 
cheatgrass  is  seasonalK'  preferred  forage  spe- 
cies, and  even  the  dn'  herbage  of  cheatgrass  is 
utilized  bv  li\estock.  This  dilutes  the  effect  of 
grazing  as  far  as  the  native  perennials  are  con- 
cerned. Lack  of  preference  for  medusahead 
concentrates  the  effects  of  herbi\'oi-y.  Subaeri- 
ally deposited  surface  soil  is  extremeh"  erodible 
once  protection  of  the  shnib  canopy  and  its 
dependent  microph\tic  cnist  is  lost.  Loss  of  the 
surface  leads  to  exposiu'e  of  the  cla\'  sediments 
that  then  function  as  Vertisols,  shrinking,  crack- 
ing, and  swallowing  the  surface  and  reexpand- 
ingwith  moisture.  Medusahead  is  one  of  the  few 
plant  species  adapted  to  these  Vertisols.  Perhaps 
some  of  the  soils  of  these  landscapes  were 
always  Vertisols  where,  in  wet  \'ears,  annual 
sunflowers  {Helianthus  annuus)  and  turkey 
mullein  {Erenwcorjms  seti^^erus)  formed  the 
onl)'  nati\e  vegetation.  Perhaps  excessive  graz- 
ing conxerted  some  of  these  soils  to  \ ertisols 
before  medusahead  arri\ed.  The  important 
point  is  that  medusahead  is  actixek  attacking 
assemblages  of  natixe  vegetation  and  changing 
the  physical  and  biok)gical  potential  of  the  sites. 

Management  of  Medusahead 
Infestations 

It  is  difficult  to  rexegetate  \^ertisols  in  desert 
enx ironments  xxith  both  seedlings  of  xx'oodv  and 
herbaceous  species,  natixe  and  exotic.  Not  only 


19921 


Ecology  and  Ma.\ac;i;me\t  of  Mkdl  saiiead 


251 


establishment  but  also  subsefjuent  growth  are 
problems  on  these  soils  despite  both  tremen- 
dous eation  exchange  capaeit\  and  moistnre- 
holding  capaeitA".  The  tremendous  matrie 
potential  of  these  Hue  cla\' soils  is  al\\a\  s  suipris- 
ing.  Moisture  is  not  axailable  loi-  normal  plant 
growth  when  soils  still  stick  to  \our  boots. 

NaTIH  \l,  SUCCESSION' 

Dr.  Mill  Hironaka  suggests  that  o\er  pro- 
longed periods  perennial  seedlings  might  estab- 
lish in  medusahead-intested  sites,  especialK  the 
short-li\ed  perennial  grass  squirreltail  (Hiro- 
naka 1963).  Dr.  Hironaka  and  his  students  fol- 
lowed this  aspect  of  medusahead  succession  in 
several  studies.  He  demonstrated  that  squirrel- 
tail  can  establish  in  medusahead  communities, 
but  he  found  the  perennial  grass  populations  to 
be  CN'clic.  When  the  squirreltail  plants  die,  the\' 
are  replaced  b\'  medusiiliead,  not  longer-li\ed 
perennial  grasses  (personal  communication). 

hi  the  western  Great  Basin,  Dr.  Hironaka's 
work  is  borne  out  b\'  gradual  increases  in 
squirreltiiil  plant  densits'  as  grazing  manage- 
ment systems  ha\e  been  implemented.  This  has 
been  especially  noticeable  during  the  past  four 
years  of  extreme  drought.  Densities  of  one 
squirreltiiil  plant  per  10  square  feet  began  to 
change  the  aspect  of  medusahead-dominated 
sites,  but  the  fragile  nature  of  this  impro\ement 
is  apparent  when  bioassay  of  seed  banks  shows 
250-500  viable  medusahead  seeds  per  square 
foot  (down  frcjm  1 000  per  square  foot  before  the 
drought)  and  fails  to  detect  am  viable  squirrel- 
tail seeds  (unpublished  research  AHS,  USDA, 
Reno,  Nevada). 

As  you  look  at  medusahead-infested  areas  on 
the  X'ertisols  of  the  western  Great  Basin,  vou 
have  a  nagging  thomj-ht  that  something  is  miss- 
ing.  The  Lahontan  and  big  sagebmsh  comnui- 
nities  of  the  ancient  lake  sediments  have  as  their 
most  frequent  perennial  grass  Sandberg  blue- 
grass.  This  species  is  completelv  absent  from  the 
medusahead  stands  and  is  missing  from  the 
stands  where  scjuirreltail  has  begun  to  return. 
What  factors  of  seedbed  qualitv  exclude^  the 
native  invader  Sandberg  bhungrass  and  are  the 
same  factors  related  to  the  failure  of  higher-level 
perennial  grasses  to  become  established  in 
squirreltail/medusahead  communities? 

The  striking  difference  between  nativ  e  and 
medusahead  communities,  other  than  loss  of 
shnib  canopies,  is  loss  of  subcanopv  mounds 


and  microphvtic  cmst  that  covers  the  mounds 
to  extend  down  to  mingle  with  vesicular  crust  in 
the  interspaces.  The  thalloplntic  crust  of 
mosses,  lichens,  and  livenvorts  is  obviouslv 
gone,  and  we  can  onlv  speculate  on  the  fate  of 
tlie  microscopic  crust  of  algae,  fungi,  and  bacte- 
ria. Prolonged  medusahead  dominance  mav 
decrease  populations  of  nncorrhizae  spores  in 
the  soil  and  thus  inflnenc-e  growth  of  artilicialK 
established  perennial  seedlings  (personal  com- 
munication, Jim  Trent,  soil  microbiologist.  .\liS, 
USDA,  Reno,  Nevada). 

Specific  plant  pathogens,  developt^d  and 
marketed  bv  biotechnological  companies,  mav 
have  a  role  in  range  weed  control.  Perhaps  a 
Fusarittni  species  exists  that  would  be  highly 
specific  for  medusahead  (personal  communica- 
tion, Joe  Antognini,  National  Program  scientist. 
Weed  Science,  ARS,  USDA). 

Taxonomists  and  greneticists  who  have 
worked  with  medusahead  have  commented  on 
how  variable  individual  collections  mav  be. 
Common  garden  studies  have  shown  this  to  be 
tnie  for  collections  from  the  American  \\'est 
(McKell,  Robinson,  and  Major  1962,  \bung  et 
al.  1971b).  We  found,  in  common  garden  stud- 
ies, a  collection  from  northern  California  that 
matured  4  weeks  earlier  than  the  average  for 
other  collections  or  on  or  before  the  maturitv  for 
cheatgrass.  As  medusahead  evolves,  we  have  vet 
to  see  the  limits  of  its  potential  on  the  vxesteni 
range.  The  recent  discoven'  of  medusahead  in 
Utah  illustrates  that  portions  of  the  eastern 
Great  Basin  have  the  potential  to  be  iii\  adcd  bv 
this  weed  (Horton  1991). 

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Rrccivnl  2:Ulai/  Um 
Arrrpird  22  Jinir  I9h)2 


(ireat  Basin  Naturalist  .")2i.'Vi.  pp.  25o-2Hl 

R(X)ST  SITES  USED  BY  SANDHILL  CRANE  STA(;iNG 
ALONG  THE  PLATTE  HI\  EK.  NEBRASKA 

Bra(lle\'  S.  NOrliiiii;  ,  Staiilcx  11.  .VikIitsoii  ,  and  Wiune  A.  Iliihcrf 

.•\bstiu(X — We  iLssessed  the  influence  of  water  depth,  extent  of  unobstructed  \ie\\\  and  huiuan  disturbance  features  on 
use  of  roost  sites  h\  Sandliill  Crimes  along  the  Platte  Ri\er,  Nebraska,  during  .spring  niigraton  stopox  er.  .Xt'riai  photos  tiikcn 
near  dawn  were  used  to  determine  areas  of  flock  use  and  habitat  a\ailabilitv  in  four  sample  reaches,  and  measurements 
were  made  on  the  ground  at  flock  roost  areas.  In  general,  depths  of  1-13  cm  were  used  bv  s;uidhill  cranes  in  greater 
proportion  than  those  a\ailable.  Exposed  sandbars  ;uid  depths  >20  cm  were  a\()id(^d,  wliile  depths  of  14-19  cm  were  u.sed 
in  proportion  to  their  a\ailal)ilit\-.  Sites  11-50  m  from  the  nearest  \isual  t)bstruction  were  used  significantly  greater  than 
their  availabilit);  while  sites  ()-4  and  >50  m  from  \isual  obstnictions  were  avoided.  Sandliill  Cranes  avoided  sites  near  pa\  etl 
roads,  gra\el  roads,  single  dwellings,  and  bridges  wjien  .selecting  roost  sites:  howewr.  thex'  did  not  appear  to  be  disturiied 
b\  private  roads,  groups  of  residential  buildings,  graxel  pits,  railroads,  or  electrical  trausniissioii  lines. 

Kc'i/  words:  Sdiulliil!  Crane.  Cirus  canadi-nsis,  river  rocistn.  habitat  .sclcclioii.  water  di])th.  (li.slurhanrc.  saiulhars.  Platte 
Hirer 


Tlie  impact  of  water  re.sourcc^  clex'elopnieiit 
on  the  Platte  Rixer  i.s  well  described 
(Kroonemever  197S,  Williams  1978,  Eschneret 
al.  19S].  Kii-cherand  Kariinger  198L  U.S.  Fish 
andWildlite  Senice  1981.  Krapn  1987,  Sidle  et 
al.  1989).  The  major  impact  lias  come  from 
irrigation  projects  along  the  North  Platte  Ri\er 
(Krapn  et  al.  1982),  which  remo\e  approxi- 
mately 70%  of  the  annual  How  of  the  Platte 
Ri\er  before  reaching  sonth  central  Nebraska 
(Krooneme\er  1978).  Concomitant  with  chan- 
nel shrinkage,  woocK' vegetation  has  encroached 
on  thonsands  of  hectares  of  former  channel 
area,  contributing  to  further  changes  in  channel 
features  and  altering  habitat  for  numerous  .spe- 
cies of  migraton- birds  in  tlie  Big  f^end  Reach  of 
the  Platte  River  in  Nebraska  (U.S.  ImsIi  and 
Wildlife  Sei-vice  1981).  The  Big  Bend  l^eacli  of 
the  Platte  Ri\er  in  Nebraska  is  an  area  of 
importance  to  numerous  sjiecies  of  migraton 
birds  ol  the  Central  I^1\\\a\  ( If.S.  Fish  and  Wild- 
life Senice  1981 ). 

This  area  is  an  important  stojioxcr  area  lor 
most  of  the  midcontin(mt  population  of  Sandhill 
Cranes  (Cms  i-ditddciisis^  i  4(H ).()()( )--6( )().()()() 
birds),  which  roost  in  the  riwr  and  feed  in 
neadn  com  fields  (Krapn  et  al.  1981,  Krapn 
1987).  The  endangered  Wliooping  Cj-ane  (C^, 


(iincricdiui)  also  uses  the  area  during  migration, 
and  the  tlu'eatened  Bald  Eagle  {Haliacctus 
lei(coccpJialiis)  is  a  common  winter  resident 
(U.S.  Fish  and  Wildlife  Senice  1981).  The  area 
is  also  important  habitat  for  the  endangered 
interior  population  of  Least  Tern  {Sfcnia  aiitil- 
lantm)  and  the  threatened  Piping  Ploxer 
(Charadriiis  niehxhis),  both  of  which  nest  along 
the  Platte  Ri\er  (U.S.  Fisli  and  Wildlife  Senice 
1981,  Sidle  etal.  1989). 

Considerable  attention  lias  been  gi\en  to  the 
impact  of  changing  channel  conditions  on  the 
midcontinent  population  of  Sandhill  Cranes 
{Gnis  canadensis)  that  congregate  along  the 
riverfront  earl\'  March  to  mid-,\pril  during  their 
animal  spring  migration  (Lewis  1977,  Krapn 
1978,  U.S.  Fish  and  Wildlife  Senice  1981). 
During  this  time  approximatcK  4()(). ()()()  Sand- 
hill Cranes  use  tins  an^a  while  euroiile  to  their 
breeding  grounds  in  (Canada,  Alaska,  and  eastern 
Siberia  (U.S.  F^ish  imd  Wildlife  Senice  1981). 

In  Nebraska  various  facets  of  Sandhill  Crane 
roosting  habitat  re<juirements  ha\e  been  stud- 
ied (Frith  1974,  Lewis  1974,  U.S.  Fi.sh  and 
W  ildlife  Senice  1981,  Krapn  etal.  1982.  1984). 
I  i()\\(n cr.  these  studies  ha\(^  not  considered  the 
infhience  of  habitat  axailabilitA  in  ndation  to 
habitat  use.  The  i)un)ose  ot  this  stud\  was  to 


W'yoinini;  (;o()ptTati\e  Fish  unci  W'ikllirc  Hescarch  Liiit.  Bo.\  :!166.  L'niversih'  Station.  Laramie,  Wyoming  S207r 


2o.i 


254 


Ghi:at  Basin  Naturalist 


[X'olunie  52 


Reach  1 


Reach  3 


■  Reach  4 


Intensive  Study  Area    ^1^* 
0       1       2      3      4       5  km 


Fiii.  I.  StiuK  sites  in  the  Platte  Hi\'er,  Nehraski] 


(Ic'tcniiiiic  tlic  influence  oF  habitat  axailahilitx, 
as  well  as  habitat  use,  on  the  selection  of  roost 
sites  b\'  Sandhill  Cranes. 

This  stiicK'  was  designed  to  assess  the  infhi- 
ence  of  three  tvpes  of  habitat  features  on  roost 
sites  used  by  Sandhill  Cranes:  (1 )  water  depth, 
(2)  magnitude  of  unobstructed  \iew.  and  (3) 
disturbance  features. 

STrnv  Ahka 

The  study  area  is  locatcxl  in  south  central 
Nebraska  in  Hall  and  l^ufTalo  counties  in  die 
eastern  halfofthe  Big  Bend  Hcnich  of  the  Platte 
Kiver.  It  encompasses  a  36-km  stretch  of  the 
Platte  Kiver  beginning  4  km  west  of  Shelton  to 
(^rand  Island  (Fig.  1).  All  held  measurements 
were  in  four  1 .6-km  reaches  along  the  main 
channel  of  the  Platte  Ri\er. 

Spiing  precipitation  in  Nebraska  contributes 
to  the  Platte  Kiver  Basin  flow,  but  most  of  the 
flow  is  derixed  from  spring  runoff  that  originates 
as  snownu'lt  in  tlie  Kocky  .Mountains  (E.schner 
et  al.  I9(S1).  Spring  runoff  flows  into  both  the 
North  and  South  Platte  ri\ers,  which  flow  nortli- 
east    and    southeast,    resi)(>ctivel\.    across    the 


Cireat  Plains  to  their  confluence  near  North 
Platte,  Nebraska. 

The  stnd\'  area  is  characterized  b\  numerous 
braided  channels  interspersed  with  imxege- 
tated  sandbars  that  fre(juentl\"  shift.  Most  of  the 
land  within  and  adjacent  to  the  stuch'  area  is  in 
private  ownership.  Land  use  in  the  area  is  pre- 
dominantly agriculture  and  includes  approxi- 
mately 60%  cropland  (mostly  com),  5%  tame 
pasture,  20%  nati\e  grassland,  and  15%  riparian 
woodland  (Keinecke  and  Krapn  1979). 

The  riparian  woodland  comprises  eastern 
Cottonwood  (Poptthis  deJioidcs)  forests  with 
(k)minant  understoiA  species  of  red  cedar 
(Jiinij)rrii.s  lir^inidiia)  and  rough-leaf  dogwood 
(doniu-s  (Innnmotidii).  On  low  islands  and  \eg- 
etat(^(l  sandbars,  peach-leaf  willow  {Salix 
aini/i^ddloidcs).  ccnote  willow  {S:  cxig^nal),  and 
indigo  bush  (Aniorplia  fnitirosa)  are  the  domi- 
nant species  (U.S.  Fish  and  Wildlife  Senice 
1981,  Currier  1982). 

MKTHODS 

.Aerial  photographx  was  used  to  determine 
flock  locations  and  delineate  flock  boundaries  of 


19921 


CiiWE  Roost  Sites 


roosting  Saiulliill  Cranes  along  a  36-kni  stretcli 
ot  the  Platte  Hi\{M-.  Photograpln"  was  restricted 
to  mornings  with  less  than  10%  cloud  coxer  and 
ceilings  abo\e  975  m.  Flights  were  begun  30 
minutes  ht^fore  sunrise  ])ecanse  ol  the  need  to 
pli()t()gra[)h  Sandhill  C'ranes  before  the\'  lea\e 
the  roost  in  earl\'  morning.  Light  was  adequate 
to  piMinit  photograph\'  10-15  minutes  before 
sunrise. 

A  Hasselblad  500  P.L,  70-mm  camera  was 
used  to  photograph  the  stud\'  area.  The  camera 
was  mounted  in  a  standard  camera  hatch  in  a 
Cessna  172  fixed-wing  aircraft  and  was 
equipped  with  an  SO-mm  focal  length  Zeiss  lens. 
Exposures  were  made  at  1/60  and  1/125  second 
at  f2.8  using  Kodak  Tri-X  640  AFS  Aerographic 
film.  The  camera  was  equipped  with  a  70  expo- 
sure back  loaded  with  5.5  m  of  film  allowing  <S0 
ex[)<)sures. 

The  aircraft  was  flown  at  approxiiiiatcK  140 
km/hr  at  an  initial  altitude  of  790  m  aboxe 
ground  lexel  for  the  first  two  flights.  During  the 
last  two  flights  the  altitude  was  increased  to  910 
HI  al)o\e  ground  le\el.  These  altitudes  provided 
a  0.48-km"  and  0.64-km"  coxerage  on  each 
frame,  respectively.  Frame  rate  was  controlled 
1)\  an  intenalometer,  calibrated  for  309^  oxer- 
lap,  to  pnnide  continuous  photographic  co\er- 
age  of  the  study  area. 

Shortly  after  each  flight  the  film  was  custom 
pr()ces.sed  by  hand  agitation  in  a  single  solution 
tank,  xaning  time  and  (kn'eloper  temperature 
to  obtain  optimum  dexelopment.  Approxi- 
matcK  150  frames  were  e.xposed  from  each 
flight.  Frames  were  examined  under  SX  magni- 
fication to  identifx  crane  flocks  and  were 
enlarged  to  41  X  51  cm  ( 16  X  20  in)  and  printed 
on  Kodak  PoK'  contract  RC  paper.  Processed 
photogiaplis  wen^  stored  for  later  anaKsis  of 
\isual  obstructions  and  disturbance  features. 

Each  of  the  tour  1 .6-km  reaches  was  marked 
on  both  sides  of  the  rixer  bank  with  16.  1-nr 
markers  mack'  of  white  cloth.  The  markers, 
placed  100  m  apaii  at  the  edge  of  the  rixerbank, 
were  positioned  in  such  a  wa\'  that  markers  on 
tlie  opposite  sides  of  the  channel  were  parallel 
to  the  channel.  The  markers  enabled  accurate 
scale  measurements  to  be  taken  from  photos 
and  proxided  position  reference  for  tiansects 
across  the  channel  xx'hen  sampling  water  depths. 
Aerial  photographs  coxering  each  reach  xxere 
used  to  determine  the  position  of  transects 
through  flocks.  Transects  were  positioned  so 
that  each  flock  studied  on  a  photo  x\as  dix ided 


into  general  areas  of  ecjual  size  with  txvo  to  fix-e 
transects  depending  upon  flock  si/.e.  A  flock  x\'as 
(k'fined  as  a  continuous  distribution  of  birds  or 
an  aggregation  of  birds  sjxitiallx'  independent  of 
other  birds  separated  bx  a  distance  >2()  m. 
Flocks  usnallx'  occurred  in  configurations  that 
a[)pear{>d  distinct  from  other  flocks  in  the  xic  initx. 

After  transects  xx'cre  located  on  [)hotograplis, 
thex" xxere  measured  and  laid  out  on  the  ground 
in  relation  to  marker  locations  using  \inx  1  flag- 
ging placed  on  each  side  of  the  channel.  Water 
depths  xxere  measured  to  the  nearest  3  cm  at 
3-m  inten  als  and  plotted  on  acetate  oxerlaid  on 
aerial  photogra]:)hsx\ith  delineated  flock  bound- 
aries. Width  and  depth  data  xx^ere  combined  to 
gix  e  mean  estimates  for  each  of  the  four  reaches. 

Each  1.6-km  reach  xvas  sampled  as  soon  as 
possible  after  each  flight,  alxvavs  xvithin  three 
dax's.  Staff  gauges  xxere  placed  in  each  area  to 
measure  anx* changes  in xxater  lexel  between  the 
time  each  reach  xx'as  photographed  and  the  time 
it  xvas  sampled.  Detectable  changes  in  xxater 
lex'el  xx^ere  recorded  and  used  to  con-ect  dt^pth 
distributions. 

Discharge  xvas  measured  on  each  flight  dax' 
in  close  proximity'  to  the  study  areas  folloxxiug 
the  techni(jue  of  Buchanan  and  Somers  (1969). 

Contact  prints  xxere  made  from  each  roll  of 
film.  Indixidual  frames  xx'ere  cut  out  and  glued 
onto  posterboard  to  form  a  mosaic,  proxiding  a 
continuous  coxerage  of  the  rixer  channel.  Scale 
was  determined  bx'  comparing  bridge  segments 
and  transect  locations  on  the  contact  prints  xxith 
measurements  of  these  locations  niadc^  on  the 
ground.  Scale  e,stimates  were  made  along  2-  to 
3-km  segments  of  rixer  Photograph  scales 
ranged  froiu  1 :8,681  to  1:1 0,334  for  the  first  txxo 
flights,  and  1 :  10,595  to  1:11, 857  for  the  last  txx'o 
flights. 

A  binocular  zoom  iuicro,scope  (1-4X)  xx^as 
used  to  ick^ntifs  flocks  and  delineate  flock 
boundaries  on  the  contact  prints  covered  xxith 
ac(Tatc.  Flocks  wcic  delineated  and  subse- 
(juentK  nmubered  on  the  acetate  oxerlax'S  on 
contact  photos.  The  distance  from  the  edge  of 
each  flock  to  the  nearest  xisnal  obstniction  x\as 
measured  to  the  nearest  0.5  nun  on  the  photos 
(ground  distance  =  ^6  m)  using  a  drafting  cal- 
iper \  isual  obstructions  inclnck'd  xegetation,  a 
rixer  bank,  or  anx  otlier  'xisualK  solid"  object 
>1  m  in  height. 

Kandom  points  were  plotted  on  contact 
photos  to  (\stimate  the  featm-es  of  ax  ailable  hab- 
itat. Ranck)m  points  xxere  determined  bx  a  .series 


256 


G  H  EAT  B  AS  I N  N  ATU  R  A  LI  ST 


[\ bluiiie  52 


of  random  numbers  identifying  point  coordi- 
nates on  gridded  overlay  coxering  contact 
prints.  Points  outside  the  rixer  channel  were 
discarded.  Onl\-  random  [X)ints  located  in  water 
were  u.sed  because  points  on  sandbars,  islands, 
or  the  ri\er  bank  were  not  considered  poten- 
tiall\-  usable  roosting  habitat.  A  total  of  339 
random  points  within  the  ri\er  channel  were 
identified  on  the  contact  prints.  Grid  squares 
were  1.25  mm"  to  ensure  a  representative 
sample  of  locations  on  the  ri\  er.  As  with  flock 
locations,  the  distance  from  each  random  point 
to  the  nearest  \isual  f)bstruction  was  measured 
on  the  photos  t(;  the  nearest  0.5  nun  using  a 
drafting  calipei-. 

For  analvsis  oi  human  disturbance  features, 
flock  locations  and  random  points  along  the 
entire  36-km  stud\'  area  were  transferi-ed  from 
70  nun  contact  prints  to  acetate  overlays  of  color 
infrared  aerial  photographs  (scale  1:25,595) 
using  a  zoom  transfer  scope.  The  photographs 
taken  in  April  1989  were  obtained  from  the 
Bureau  of  Reclamation  in  Cirand  Island, 
Nebraska.  Distances  were  measured  from  the 
edge  of  each  flock  and  individual  random  points 
selected  b\'  placing  a  card  over  the  photograph 
to  the  nearest  human  disturbance  features. 
These  features  included  pa\ed  roads,  gravel 
roads,  prixate  roads,  urban  dwellings,  single 
dwellings,  railroads,  connnercial  development, 
highwa\s,  and  bridges.  Distances  were  mea- 
sured to  the  nearest  0.5  mm  on  photos  (ground 
distance  =  13  m)  with  a  drafting  caliper. 

Data  AnaK  sis 

FrequencN'  histograms  were  plotted  for  mea- 
sured distances  from  the  edge  of  a  flock  and  for 
random  distances  to  the  nearest  visual  obstruc- 
tion and  disturbance  features.  Frequencv  distri- 
butions were  plottc>d  for  axailable  and  used 
selected  water  depths.  Fre(jU(Mic\-  distributions 
of  available  and  used  selected  water  depths  for 
each  1 .6-km  reach  were  determined  bv  combi  u- 
ing  flock  data  for  each  reach  for  a  given  flight. 
Available  depths  were  defined  as  all  depth  mea- 
surements taken  along  a  transect,  and  used 
depths  were  those  depths  where  birds  were 
present  along  a  tran.sect.  Habitat  selection  v\as 
computed  by  dividing  the  proportion  of  habitat 
u.sed  within  a  depth  intenal  bv  the  proportion 
of  depths  available  in  that  same  intenal  (Bovee 
1986).  Depths  used  less  than  their  availabilitv' 
were  defined  as  being  av  oided,  while  those  used 
more  than  their  availabilitv  were  defined  as 


being  selected.  Habitat  avail abilitv,  use,  and 
selection  were  summarized  within  reaches, 
across  flight  dates,  and  from  data  pooled  across 
reaches  and  flight  dates.  Data  were  pooled  to 
generalize  the  selection  of  depths  over  the 
course  of  the  sampling  period. 

The  chi-sqiuire  of  homogeneity  (Marcum 
and  Loftsgaarden  1980)  was  used  to  test 
whether  differences  existed  between  the  distri- 
l)utit)n  of  random  points  and  those  locations 
used  bv  Sandhill  Cranes  relative  to  visual 
obstructions  and  distiu'bauce  features.  It  was 
also  used  to  determine  if  there  were  differences 
between  the  proportion  of  used  and  available 
water  depths  among  and  within  reaches.  Confi- 
dence intervals  were  calculated  using  the 
Bonferroni  Z-statistic  to  test  which  intenals 
within  the  distributions  were  used  more  or  less 
than  exjDected  (Byers  et  al.  1984).  Differences 
between  selection  functions  were  tested  wdth  a 
Z-test.  Analysis  of  variance  (ANOVA)  was  used 
to  determine  if  visual  obstructions  had  an  effect 
on  the  disturbance  potential  created  by  various 
tvpes  of  disturbance  features.  Significance  for 
all  statistical  inferences  v\'as  P  <  .05. 


Results 

A  total  of  four  sampling  flights  were  made: 
one  each  on  21  and  31  NIarch  and  4  and  10  April 
1989.  A  total  of  285  flocks  were  identified 
during  the  four  flights.  Folkming  the  flights,  20 
flock  sites  vwre  selected  and  sampled  and  a  total 
of  5109  depth  measurements  were  recorded  in 
the  field. 

Sampling  areas. — Reaches  I  and  II  were 
the  narrowest,  with  mean  channel  widths  of  254 
m  (range  =  225-319  m)  and  249  m  (range  = 
241-263  m),  respectivelv,  while  reaches  III  and 
I\',  located  upstream,  were  wider.  Reach  III  had 
a  mean  channel  width  of  413  m  (range  =  387- 
440  m),  while  reach  W  had  a  mean  channel 
v\idth  of  357  m  (range  =  296-445  m). 

Reaches  I  and  II  had  similar  discharge  (17 
mVs),  while  reaches  III  and  l\  had  greater 
values  (27  and  44  m  Vs)  on  21  March  (Table  1 ). 
Discliarge  in  rcnich  III  was  tvpicalK  tv\ice  as 
high  as  reaches  I  and  II.  Reach  1\'  had  the 
highest  discharge  of  the  four  reaches,  often 
three  times  greater  than  in  reaches  I  and  II 
(Table  1).  Reaches  I,  II,  and  III  were  located  in 
a  braided  portion  of  the  surface  along  the  south 
chaimel  and  coutaineil  onlv  partial  river  flow. 


19921 


CiuxE  Kous'i' Sites 


25' 


TaBI.F,  1.  Disc-lianji;e  in  cubic  meters  per  second  (m   )  for  saiii|ile  reaclu's  on  tlifTerent  i'\\'j}]\  dates  alon<4  tlie  Flatte  Ui\er 
Nebraska,  during  spring  19S9. 


Flight  date 


Reacli  I 


Reach  II 


Reach  III 


Reacli  I\' 


21  March-' 
31  March 
4  April 
10  April 


17.4 

11.1 

10.6 

7.9 


17.4 


10.6 
7.9 


27.5 
18.6 

13.7 


44.6 

32.1 
2S..S 
21.7 


■'Discliarge.s  for  all  reaches  on  21  March  were  nieasiircd  cm  24  Maich.  TIids.  a  lhii'i'-ila\  la;;  |)rrio<!  fxisted  betxM'cii  the  lime  each  reach  u'a.s  flouTi  and  (he  tin 
each  reach  w;is  measured  for  discharge. 


Reach  1\  was  located  aloiiL:;  tlic  iiiaiii  cliaiiuci 
and  contained  total  ri\er  flow. 

IlARIT\T  WAILABILITY. — The  distribution 
of  a\ailal)le  water  depths  differed  among 
ivaches.  On  21  March  1989,  82%  of  tlie  avail- 
able habitat  in  reaches  I  and  II  consisted  of 
depths  0-25  cm.  In  contrast,  53%  and  66%  of 
the  axailable  habitat  in  reaches  III  and  I\', 
respectixelw  consisted  of  depths  0-25  cm. 

An  increased  freqnenc\  of  shalhnv  depths 
(0-19  cm)  and  a  decreased  frequenc\-  of  deeper 
depths  (>20  cm)  occurred  o\er  the  stud\ 
period.  This  di\ision  is  made  because  cranes 
seldom  used  depths  greater  than  20  cm.  The 
increase  in  exposed  sandbars  (depth  =  0  cm )  was 
most  pronounced  in  reaches  I  and  II,  which 
showed  increases  of  13%  and  11%,  respecti\el\ . 
Reaches  II  and  III  showed  increases  of  12%  and 
19%,  respectiveK',  in  axailable  depths  of  1-4  cm 
betAveen  the  first  and  last  flight.  Reaches  III  and 
W  showed  decreases  of  10%  and  7%,  respec- 
tixcK;  in  depths  >38  cm  for  the  same  period. 
During  the  stiuK  period  a  progressixe  decrease 
in  discharge  occurred  (Table  1),  causing  more 
shallow  areas  (0-19  cm). 

HabiT.AT  use. — Frequency  di.stributions  of 
roosting  habitat  use  hv  cranes  indicated  the 
liighest  proportions  of  used  water  deptlis  were 
from  the  1-4  and 5-7  cm  increments.  This  range 
ot  water  depth  accounted  for  65%  of  the  mea- 
sured depths.  There  was  no  discernible  \'aria- 
tion  in  the  frequency  of  water  dejiths  us(^d 
among  the  four  reaches. 

There  was  a  small,  but  significant,  difference 
in  tli(^  distribution  of  depths  used  bet^\•een  the 
beginning  and  end  of  the  study  period  (F  <  .05). 
Depths  of  0  cm  showed  a  significant  decrease  in 
use,  while  depths  20-22  cm  sliowed  a  signihcant 
increase  in  use  {P  <  .05).  The  data  showed  a 
significant  difference  between  the  distribution 
of  used  and  available  water  depths  for  all  foui- 
sampling  periods  (P  <  .001).  Sandhill  Cranes 
used  progressiyely  deeper  water  depths  as  the 


stud\  season  progressed.  Depths  >20  cm  were 
used  significanth-  less  than  expected  during  the 
first  flight;  but,  b\-  the  last  sune)',  only  depths 
>29  cm  were  used  less  than  expected  {P  <  .05). 
Depths  of  0  cm  were  generally  avoided  bx 
Sandhill  (>ranes  during  the  last  two  sur\e\  s  and 
were  used  less  than  woidd  be  expected  bx 
chance  (P  <  .05). 

Habitat  selection  was  assessed  using  both 
habitat  use  and  axailabiHtv'  data  for  specific 
water  depths.  The  most  frecjuentK  occurring 
depth  intenals  for  which  selection  occurred 
were  5-7  cm,  fcjllowed  by  1-4,  8-10,  1 1-13,  and 
14-16  cm  in  decreasing  order  of  preferenc-(\ 

Visual  (obstructions. — There  was  a  sig- 
nificant difference  between  the  distribution  of 
flock  locations  and  random  points  reiatixe  to  the 
distance  from  the  nearest  \isual  obstruction 
(F  <  .001).  Proportional  u.se  of  sites  0-50  m 
from  the  nearest  \isual  obstruction  was  signifi- 
canth' greater  than  a\ailabilit\  (F  <  .05),  while 
sites  >50  m  from  a  \isual  obstruction  were 
avoided  (F<  .05). 

The  0-25  m  interval  was  di\ided  into  six 
increments:  0,  1— f.  5-10,  11-15.  16-20.  and 
21-25  m.  There  v\as  a  significant  difference 
lK^t^\'e(^u  the  distribution  of  flocks  and  random 
point  distances  (F  <  .001 ).  Sites  as  close  as  5-10 
m  from  the  nearest  visual  obstruction  were  us(^d 
by  Sandhill  Cranes.  Onlv  sites  0 — f  ni  from  a 
\  isual  obstruction  v\ere  avoided  (F  <  .05),  while 
sites  1  1-25  m  from  a  visual  obstruction  were 
used  more  than  expected  (F  <  .05). 

Msual  obstructions  v\ere  divided  into  three 
categories:  (1 )  unvegetated  bank.  (2)  \egetated 
bank,  and  (3)  xegetaled  island.  There  v\'ere  no 
significant  differences  in  the  distribution  of  dis- 
tances b(^h\een  an  unvegetated  and  vegetated 
bank,  but  there  were  significant  differences  for 
tlie  distribution  of  distances  between  vegetated 
banks  and  vegetated  islands  and  between 
unvegetated  banks  and  vegetated  islands  (F  < 
.005).  Sandhill  Cranes  roosted  a  mean  distance 


258 


Cheat  Basin  Naturalist 


[\'()]unie  52 


of  45  Ml  from  umegetated  banks.  50  in  lioni 
wgetated  hanks,  and  27  ni  from  v{>o;etat('d 
ishmds. 

Channel  width. — ^There  was  a  ndation- 
ship  between  the  niinimnm  unob.stnicted  chan- 
nel width  and  distance  to  the  nearest  \isiial 
obstrnction.  The  distance  to  the  nearest  \'isnal 
obstrnctions  was  ahuiction  of  less  than  one-half 
die  niininnnn  unobstrncted  channel  width. 

There  was  a  significant  difference  between 
the  distribntion  of  flock  locations  and  random 
points  relati\e  to  minimnm  nnobstructed  chan- 
nelwidth  (P  <  .005).  Sandhill  Cranes  used  chan- 
nels 100-200  ni  wide  in  greater  proportion  than 
tho.se  generalK  a\ailable.  Channels  narrower 
than  100  m  were  axoided,  while  those  >200  m 
wide  were  used  in  proportion  to  their  axailabil- 
ih'.  The  mean  minimum  unobstioicted  channel 
wudth  used  by  roosting  flocks  was  196  m  (range 
=  34-445  m).'  Nearly  100%  of  the  flocks  we  re  hi 
channels  with  a  minimum  unobstructed  chan- 
nel width  of  >50  m,  and  over  97%  and  80%  of 
the  flocks  were  in  channels  with  a  minimum 
unobstructed  width  of  >]()()  and  >150  m. 
respectixeK'.  The  mean  relative  flock  size  (sui- 
face  area)  was  3883  m~  (range  =  19-55,354  nr). 
There  was  no  relationship  between  flock  size 
and  minimum  nnobstructed  channel  width. 
Both  large  and  small  flocks  were  located  in  wide, 
as  well  as  narrow,  channels. 

liiiinaii  Disturbance  Features 

PWKi:)  KdADS. — Sandhill  Crane  flocks  were 
not  distributed  randoniK  with  respect  to  dis- 
tance from  pa\ed  roads  (P  <  .001 ).  Sandhill 
Cranes  showed  avoidance  of  sites  closer  than 
500  m  from  the  nearest  paved  road  (P  <  .05), 
but  used  sites  as  clo.se  as  301-400  m.  Sites 
located  701-900  m  from  the  nearest  paved  road 
were  used  mon^  than  expected  (/'  <  .05). 
Sandhill  Cranes  roosted  a  mean  distance  of 
12fiO  m  from  the  nearest  paxed  road  when  a 
\isual  obstrnction  was  present,  but  a  uK^an  dis- 
tance of  1575  m  from  the  nearest  paved  load  in 
the  absence  of  \isual  obstnictions. 

C;ra\'EL  roads.— There  was  a  significant 
difference  behveen  the  distribution  of  used  sites 
and  random  locations  relative  to  tlistance  from 
gra\el  roads  (F  <  .01).  Sandhill  Cranes  showed 
a\()idance  of  sites  that  wert>  closer  than  400  m 
from  tiie  nearest  graxel  road  (F  <  .05),  but  flocks 
were  located  as  close  as  301-400  m.  Sites  that 
were  601-800  m  from  the  nearest  gravel  road 
were  used  more  than  expected  (F  <  .05).  The 


presence  of  \isual  ob.struction  between  a  roost- 
ing flock  and  the  nearest  gravel  road  did  not 
appear  to  reduce  the  disturbance  potential  cre- 
ated bv  gravel  roads. 

Sinc;LE  D\\ELLINC;S. — There  was  a  signifi- 
cant difference  between  the  distribution  of  used 
and  random  locations  relative  to  the  distance  to 
the  nearest  single  dwelling  (F  <  .01).  In  general. 
Sandhill  Cnuies  showed  an  axoidance  for  sites 
closer  than  400  m  from  a  single  dwelling  (F  < 
.05).  Sites  501-600  m  from  the  nearest  single 
dwelling  were  used  more  than  expected  (F  < 
.05).  The  presence  of  a  visual  ob.struction 
between  a  flock  and  the  nearest  single  dwelling 
did  not  affect  the  disturbance  potential  created 
bv  single  dwellings. 

Bridces. — Sandhill  Crane  flocks  were  not 
distributed  randoniK  with  respect  to  distance 
from  bridges  (F  <  .001).  They  showed  avoid- 
ance of  sites  closer  than  400  m  from  the  nearest 
bridge  (F  <  .05).  Similarly,  they  used  sites  >400 
m  from  the  nearest  bridge. 

Other  disturbances. — No  significant 
differences  were  found  between  urban  dwell- 
ings, gravel  pits,  commercial  development, 
transmission  lines,  and  the  distribution  of 
Sandhill  ('nine  flocks. 

Discussion 

Depth  Distribution. — Tliis  study  indicated 
that  Sandhill  Cranes  prefer  water  depths  of 
1-13  cm  for  roosting  but  roost  in  greater  depths. 
Lataka  and  Yahnke  (1986)  developed  a  predic- 
tive model  for  Sandhill  Crane  roosting  habitat 
and  stated  that  the  majorih'  roosted  in  water 
depths  between  0  and  12  cm,  which  is  presum- 
ably the  optimal  depth  for  roosting.  Similarly, 
Frith  ( 1986)  suggested  a  water  depth  of  2-15  cm 
as  optimum  for  roo.sting  sites.  Currier  (1982) 
reported  a  slightlv  deeper  range  of  depths  from 
10-1 5  cm  as  optimum  for  roosting.  Lewis  (1974) 
suggested  that  loost  sites  be  characterized  bv 
deiiths  10-20  cm,  and  Folk  (1989)  reported  an 
even  greater  range  of  depths  used  for  roosting: 
0.1-21.0  cm  for  Santlhill  (Cranes  along  the 
North  Platte  Ri\  t-r  in  Nebraska. 

D(>spit('  a  change  in  the  availabilitv  of  water 
(k'pths  with  over  a  50%  reduction  in  discharge 
oxer  the  period  of  study  (Table  1),  onlx'  slight 
differences  xvere  detected  in  the  oxerall  use  of 
specific  water  depths.  The  fact  that  habitat  use 
remained  tlu^  same  despite  a  change  in  habitat 
selection  sut£tiests  that  selection  indices  more 


19921 


Cham-:  Hoost  Sites 


259 


str()n>j;K  rclclcct  cliaiiiics  in  liabitat  a\ailal)ilit\ 
than  habitat  preference.  If  habitat  sc^lcn'tioii  had 
reflected  hal)itat  preference,  then  habitat  selec- 
tion indices  wonld  ha\e  been  more  similar 
between  tlie  beginning  and  end  of  the  stnd\ 
period. 

X'l.Sl  AL  ()B.STRUCTIO\S. — This  .stnd\-  indi- 
cated that  Sandhill  Cranes  will  not  roo.st  closer 
than  5  m  from  a  \isual  obstniction  and  that 
distances  from  II  to  25  m  are  the  most  fre- 
(jnc^ntK  used,  batkaand  Yahnke  (1986)  reported 
that  Sandhill  (>ranes  did  not  roost  <b5  m  from 
tlie  bank.  Folk  (19S9)  suggested  that  Sandliill 
Cranes  preferred  to  roost  >25  m  from  a  \isual 
obstruction,  but  he  obsened  roosting  as  close  as 
4  m  from  a  \isual  obstniction.  Our  results  indi- 
cate that  \arious  forms  of  visual  obstiiictions 
liaxe  different  impacts  on  roost  site  selection. 
0\erall,  \egetated  islands  ha\e  little  influence 
on  the  selection  of  roo.st  sites,  whereas  \ege- 
tated  banks  ha\e  greater  influence. 

It  is  generall\-  beliexed  that  Sandhill  (Cranes 
maintain  an  optimum  distance  from  a  \isual 
obstruction  to  increase  their  securits'  from  ter- 
restrial predators,  primariK' candids.  This  is  e\i- 
denced  by  the  fact  that  the  majorits'  of  flocks  are 
located  in  closer  pro.\imit\'  to  \egetated  islands 
than  to  unxegetated  or  \egetated  banks. 

Channel  moipholog\'  ma\'  also  be  a  factor 
influencing  the  distribution  of  roosting  areas 
relati\e  to  banks  or  islands.  This  assertion  is 
supported bv obsenations  from  depth  measure- 
ments which  suggest  that  water  (k^ptlis  and 
\elocities  near  l)anks  are  deeper  and  faster  than 
(k'pths  near  islands  due  to  bank  inidercutting. 
Thus,  sites  near  islands  mav  contain  a  greater 
proportion  of  suitable  roosting  depths  than  sites 
adjacent  to  banks. 

ClIAXXEL  WIDTH. — Sandliill  Cranes  .selec- 
tixely  used  channels  10()-2()()  m  wi{k\  while 
channels  narrower  than  lOO  in  were  axoided. 
Nearl\-  ]{){)7r  of  the  roosting  Sandhill  ( irane 
flocks  were  located  in  channels  with  an  unob- 
structed channel  width  >5()  m,  and  oxer  8()9f 
were  located  in  channels  >15()  m  wide.  Wuk^ 
channels  potentialK  proxide  mor(^  space  for 
roosting  Sandhill  Cranes,  more  securits  from 
predators,  and  more  axailable  water  tlepths  to 
choose  from.  However,  since  channel  width  w  as 
evaluated  independentK-  of  channel  depth,  it  is 
possible  that  use  of  narrow  channels  (<1()()  m 
wide)  is  limited  not  so  much  bv  a  requirement 
for  wider  channels,  but  b)-  deeper  water  that 


flows  through  these chaimc^ls  (  Latkaaiid  Valinke 
I9Sfi). 

Oui"  findings  corroborate  the  results  of 
Krapu  et  al.  (  1 9S4 ).  w  ho  reported  that  o\er  997f 
of  all  roosting  Sandhill  (Cranes  were  in  unob- 
structed channels  o\er  50  m  wide  and  almost 
7()9f  were  in  channels  >  150  m  wide.  In  contrast, 
data  from  nighttime  aerial  thermographx  In- 
Pucherelli  (19S8)  suggested  that  almost  half  of 
all  roosts  were  in  cliannels  <15()  m  wick'  and 
that  the  greatest  proportion  of  roosts  were  in 
channels  51-150  ni  wide. 

Folk  and  Tacha  (1990)  studied  roosting 
along  the  North  Platte  River  in  Nebraska  and 
reported  a  channel  width  criterion  that  was  dif- 
ferent from  this  study,  Thev  reported  that  82% 
of  the  roosts  were  in  channels  >48  m  wide  and 
18%  were  in  channels  from  16-47  m  wide. 

HUxMAN  DISTURBANCE. — Our  stnd\  (lemon- 
.strated  that  human  disturbance  features  influ- 
ence selection  of  roost  sites  b\-  Sandhill  Cranes. 
In  general,  the  greatest  disturbance  potentials 
were  attributed  to  roads  (pa\ed  and  graxel), 
bridges,  and  single  dwellings  where  irregular 
but  considerable  hiuiian  acti\it\-  might  occur. 
Cra\el  pits,  private  roads,  railroads,  and  power 
lines  had  infrequent  disturbances  and  did  not 
seem  to  affect  roost  site  selection.  In  iill  likeli- 
hood some  form  of  acclimatic^n  occurs  between 
the  constant  disturbance  on  comnuM-cial  and 
ui'ban  development. 

There  is  little  literature  that  objectively 
describes  the  zones  of  influence  exerted  bv  var- 
ious human  disturbance  features  on  the  selec- 
tion of  roost  sites  bv  Sandhill  ('ranes  along  the 
Platte  River.  Folk  (1989)  suggested  that  riparian 
forest  along  tlu^  river  provides  a  visual  barrier 
against  most  tv  pes  of  potential  disturbances  and 
that  Sandhill  ( Cranes  roost  in  sections  of  the  river 
as  close  as  SO  m  from  a  bridge.  In  contrast,  our 
studv  intlicales  that  Sandhill  Cranes  roost  in 
sc^ctions  of  the  river  that  are  >400  m  from  the 
nearest  bridge.  We  feel  that  our  results  provide 
an  ol)jectivi'  description  of  potential  zones  of 
influence  e.xerted  bv  various  disturbance  fea- 
tures and  tlu^  (4Tect  tliese  leatm-es  have  on  roost 
site  selection  bv  Sandhill  Cranes  along  the 
Platte  River. 

In  sunnnan.  our  studv  shows  the  importance 
of  sandbars  with  water  less  than  20  cm  in  depth 
surrounded  by  deeper  water.  These  sandbars 
must  be  at  least  5  m  from  some  form  of  visual 
obstruction  such  as  dense  vegetation.  This 
apparentlv   allows  the  Sandliill  Cranes  to  see 


260 


Gi^KAT  Basin  Natui^alist 


[\'()luiiie  52 


approaching  predators.  As  a  result.  Saiulliill 
Cranes  nonnalK'  roost  in  channc^ls  l()()-2()()  ni 
wide.  These  sites  are  general I\  a\\'a\-  from 
human  disturbances  such  as  roads,  bridges,  and 
prixate  dweHings.  Sandhill  (>ran(\s  could  toler- 
ate irregular  disturbances  such  as  private  roads 
and  railroads. 

Tlie  fact  that  80%  of  the  midcoutiueut  pop- 
ulation of  Sandliill  CJranes  uses  this  area  for 
staging  in  the  spring  indicates  its  importance.  It 
is  during  this  period  that  the  birds  apparently 
build  up  energ\'  resenes  allowing  them  to  con- 
tinue their  northward  migration.  If  the  area 
were  to  become  imlit  for  Sandhill  Cranes,  the 
population  would  likeK  sulfer  decline. 

ACKNOWLEDC;  M  E  NTS 

We  appreciate  the  help  ol  Delmar  Holz, 
Laura  Smith,  and  Craig  Schwieger  of  the  Grand 
Island,  Nebraska.  Office  of  the  U.S.  Bureau  of 
Reclamation,  who  supplied  needed  equipment 
and  assisted  in  the  field.  The  U.S.  Fish  and 
Wildlife  S(M-\1ce  personnel  in  Grand  Island, 
especialK"  Jerrv'  Brabander,  and  fohii  Sidle,  who 
piloted  the  plane  on  most  flights,  were  of  great 
assistance.  Tienu'  Parrish,  Gene  Maddox,  and 
Kexin  (Tlaubius  helped  gather  field  data.  We  are 
grateful  to  the  landowners  in  the  Platte  River 
Valle\w  ho  allowxnl  us  access  to  tlieir  properties; 
and  we  appreciate  the  help  of  Kenneth  and 
Marie  Strom.  We  also  wish  to  thank  Ronald 
Marrs  for  his  help  in  inteq)retation  of  aerial 
])hotographs.  Funding  was  proxided  b\  the  U.S. 
Bureau  ol  Reclamation  in  Cirand  Island, 
Nebraska,  and  the  U.S.  Fish  and  Wildlife  Ser- 
vice National  Ecology-  Research  Center  in  Fort 
Collins,  Colorado. 

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Khoonkmkvf.k.  K.  E.  1978.  The  U.S.  Fish  and  Wildlife 
Senice's  Platte  Ri\'er  national  wildlife  stud\.  Pages 
29-.32  //(  |.  C.  Lewis,  eil..  Proceedings  of  the  1978 
Crane  W(jrksliop,  Fort  Collins.  Colorado. 

LviKA  D.  C.  and  j.  W.  Yaunke  1986.  Simulating  the 
roosting  habitat  of  Sandliill  Cranes  and  validation  of 
suitabilih-of-use  indices.  Pages  19-22  in  J.  Winer, 
M.  L.  .Morrison,  and  C:.  J.  R;ilph.  eds..  Wildlife  2000: 
modeling  habitat  relationships  of  terrestrial  \erte- 
brates.  lhii\ersit\ of  Wisconsin  Press.  .Madison. 

Li  \\  Is  |,  C.  1974.  Ecol()g\'  of  the  Sandhill  Crane  in  soutli- 
lasteni  (.'I'litral  Fl\Avay.  Unpublished  doctoral  disser- 
tation. Oklahoma  State  University  Stillwater.  213  pp. 

.   1977.  Siuidhill  Cranes  (Gms  canadensis).  Pages 

5-43 //i  C.  C.  .Anderson,  ed.,  Management  of  migrator)' 
shore  and  upland  game  birds  in  North  .America.  Inter- 
national Association  of  Fish  and  Wildlife  .\gencies, 
Washington.  D.C. 

\I\i;(  IM  C.  1,.,  ,111(1  I).  ().  hiiis(.\  MiniA  1980.  .\  non- 
mapping  techiiKjuc  lor  stiuKiiig  habitat  preferences, 
journal  of  Wildlife  Mauageinent  44:  96.3-968. 

I'll  Ml  lii  I  I  I.  M.  |.  1988.  Measuring  channel  width  \ariables 
ol  S;milliill  Crane  roosting  habitat  sites  along  the  Platte 
Hi\ei  using  nighttime  aerial  thermograpln'.  Applied 
Scieiuv  R.'lcivncc  Memo.  No.  AD  88-4-5.  U.S. 
Departiiuiil  of  Interior.  Hiirt-an  of  Reclamation, 
I  )cn\cr,  ( Colorado. 


19921 


CRAM::  HoosT  Sites 


261 


Hki\K(  KK  K.  l.aiuIC  L.  KhaPL'  1979.  Spring  food  Iiahits 
of  Sandliiil  (.'laucs  in  Nebraska.  Pages  13-19  /)(  |.  ( .'. 
Lewis,  ed..  Proceedings  oi  tlie  197S  (iranc  Worksliop. 
F^oit  Collins.  C'oloratlo. 

Siiii.K.  J.  G..  E.  D.  .Mii.LKK.  and  P.  J.  Clhhii.h  19S9. 
Changing  habitats  in  tlie  Platte  Ri\er  \'alle\  oi 
Nebraska.  Prairie  Naturalist  21:  91-104. 


U.S.  Fish  .wn  Wh.di.ikk  Skhvice  1981.  The  Platte  Ri\er 
(X()log\  stndy.  Special  research  report.  Jamestown, 
North  Dakota.  I  ST  pp. 

Will, I  WIS  (;.  P  1978.  The  case  of  the  shrinking  channels — 
the  North  Platte  and  Platte  Rixers  in  Nebraska.  Circn- 
lar  781.  U.S.  Ceological  Sur\e\,  Reston,  \'irginia.  48 pp. 

Received  6  Drccinher  I^J^Jl 
Accepted  :V) }nhi  ]UH2 


Great  Basin  Naturidist  52{3),  pp.  262-26S 

POST-PLEISTOCENE  DISPERSAL  IN  THE  MEXICAN  VOLE 

{MICROTUS  MEXICANUS):  AN  EXAMPLE  OF  AN  APPARENT  TREND 

IN  THE  DISTRIBUTION  OF  SOUTHWESTERN  MAMMALS 


Russell  Diuis'  cUicl  J.  R.  Calkhan" 

ABSTR-Kcrr. — The  present  distribution  of  the  Mexican  \ole  {Micwtiis  mcxicanits)  is  not  entirely  the  product  of  post- 
Pleistocene  forest  fragmentation  and  extinction;  recent  dispersal  also  is  indicated.  Literature  records  further  suggest  that 
this  phenonient)n  nia\-  reflect  a  general  pattern  of  northward  lange  expansion  in  many  southwestern  mauuual  species. 

Kx-i/  iionl.s:  Microtns.  vole,  dispersal,  hi()<^e()^i-(ij)liy.  vieaiitiitce.  Pleistocene. 


Traditional  hiogeographic  tlieon  attributes 
tlie  niodeni  distribution  of  small,  nonll)ing 
niontant^  niamnials  in  the  Southwest  to  post- 
Pleistocene  climatic  change  (Brown  1971, 1978, 
Patterson  1984,  Patterson  and  Atmar  1986). 
Restriction  of  woodland  and  forest  habitat  to 
higher  elex^ations  is  assumed  to  have  stranded 
sucli  species  on  isolated  patches  of  montane 
habitat.  Although  it  is  recognized  that  local 
extinction  has  caused  further  range  reductions, 
post-Pleistocene  range  expansion  generalK'  has 
been  discoimted(  Brown  1971,  1978).  This  relict 
model  satisfactorih'  explains  the  distribution  of 
many  Great  Basin  species,  but  evidence  from 
else\\'here  in  the  Southwest  strongK'  supports 
recent  dispersal  (Da\is  and  Dunford  1987, 
Daxis  and  Ward  1988,  Da\is  et  al.  1988,  Daxis 
and  Bissell  1989,  Daxis  and  Brown  1989, 
Lomolino  et  al.  1989). 

In  this  paper  we  will  review  exidence  indi- 
cating that  manv  southwesteni  nuunmals — 
including  the  Mexican  xole  and  other  montane 
mammals,  as  well  as  nonmontane  species — 
have  shown  a  striking  northward  range  shift 
during  the  past  sexeral  decades.  For  some  spe- 
cies this  pattern  appears  to  reflect  mildei-  win- 
ters or  human  influences;  for  others  the  trend  is 
harder  to  explain.  If  xerified,  how  e\(M-,  this  trend 
presupposes  (among  other  things)  a  greater  dis- 
persal capability-  than  is  txpicalK  attributed  to 
small  mammals. 


DISPERSAL:  A  BRIKF  RK\1E\\ 

Post-Pleistocene  dispersal  has  been  \erified 
primarily  in  (1)  conspicuous,  diurnal  mammals 
such  as  sciurids  and  (2)  mammals  colonizing 
regions  that  were  previously  well  sampled  by 
collectors.  For  species  and  groups  that  do  not 
fall  into  either  categorv,  the  biogeographer  is 
left  to  interpret  broader  distribution  patterns 
ancPor  small  bits  of  indirect  evidence. 

As  an  example  of  the  first  situation,  Davis 
and  Browii  (1989)  and  Davis  and  Bissell  (1989) 
showed  that  recent  dispersal  has  significantly 
altered  the  distribution  of  Aberts  squirrel 
iSciunis  abciii).  Another  example  involves  the 
duskv  chipmunk  {Tamias  ohscunis),  which  was 
absent  from  Thomas  Moimtain  in  southern  Cal- 
ifornia at  least  between  1974  and  1976  (Calla- 
han 1977).  Bv  1979  the  species  had  recolonized 
this  peak,  which  is  isolated  from  the  San  facinto 
range  bv  a  lO-mile  stretch  of  semiarid  grass- 
lancPsagebrush  habitat  (Callahan,  in  prepara- 
tion). The  second  scenario  is  illustrated  bv  Davis 
and  Dunford  (1987)  and  Davis  and  Ward 
(1988),  who  found  evidence  of  recent  montane 
colonization  by  Signiodon  ochrog^nathus  in  a 
well-studied  area  of  southeast  Arizona. 

Since  manv  small  mammals  are  not  readily 
trapjx'd  and  manv  localities  have  not  l)een  sam- 
pled extensivc'lv,  it  is  easv  for  critics  to  "shoot 
down"  new  distribution  records  on  the  grounds 
of  inadequate  prior  sampling.  In  such  cases  it  is 


^ni-piirtiiifiit  <il  Kitiliiif\  ami  F.voliitidnun  Bioloi^.  L'niversilNof  Arizona,  Tiicsdii,  Arizcma  S572I 

"Miisi'iiiii  ()(  Soultmvslfni  Biolojij.  Univt-rsilyof  Nt-w  Mexico.  All)ii(|mr(|ije,  New  M<\k()  ST131    M.11I111.4  ailiir 


i-t,Caliloniia92o46. 


262 


\'()Lt:  DlSI'KHSAL 


263 


Fig.  1,  D\stii\n\t\i)u  of  Microtits  mcxicaiius.  At  tliis  sc;Je, 
()nl\  the  two  most  isolated  populations  in  the  United  States 
are  distin2;uished  (modified  from  Findle\-  et  ;il.  1975,  Hall 
19S1,  Finlevet  A.  1986,  Hoffmeister  19S6). 


necessaiy  to  look  at  broader  cli.stribution  pat- 
terns and  draw  some  reasonable  inferences. 
Da\is  et  al.  ( 19S8)  anaKzed  .soutliwesteni  mon- 
tane mammal  distributions  and  found  that  dis- 
tance from  the  source  was  a  significant  predictor 
of  species  richness — a  relationship  suggesting 
dispersal.  Lomolino  et  al.  (1989),  in  a  stud\' 
encompassing  nuich  of  the  Southwest,  con- 
firmed the  relationship  between  species  rich- 
ness and  isolation,  and  proposed  recent 
di.spersal  b\'  se\eral  montane  species  including 
Microtiis  mcxicaiuis. 

Mfxicax"  \'()ij-:  DisTHiHrriox 

The  rang(^  of  the  Mexican  \ole  (Fig.  I)  pres- 
■ntl\  extends  from  Mexico  into  Arizona,  New 
Mexico,  southern  Colorado,  and  Utah  (Durrant 
1952,  Armstrong  1972,  Findlev  et  al.  1975.  Phill 
1981,  floffmeister  1986).  The  species  t>picall\ 
nhabits  meadows  in  ponderosa  pine  and  mixed 
•ouiter  forests,  but  can  occup\-  pimon-jimiper 
woodland  if  suitable  understoiv  is  present 
Harris  1985,  Hoffmeister  1986). 'in  Arizona  it 
'ccurs  less  often  in  interior  chaparral  and  Cir(>at 
iasin  desertscrub  (Hoffmeister  1986). 

The  late  Pleistocene  distribution  of  this  spe- 
ies  probably  was  continuous  from  the  Mexican 
lateau   to  the  southwest   U.S.   (Findle\-  and 


Fig.  2.  Details  ol  tlie  distribution  oi  Microtti.s  incxicanus 
in  Arizona  showing  isolated  populations  and  three  subspe- 
cies A,  B,  and  C  (modified  from  Hoffmeister  19S6).  Open 
circles  indicate  records  added  b\-  Spicer  et  al.  (1985)  and 
Spicer  (1987);  subspecific  relationships  of  these  populations 
are  unknowni.  Papago  Springs  is  a  late  Pleistocene  fossil  site 
which  includes  a  tentative  record  (or  this  species  (Harris 
1985). 


Jones  1962).  Harris  (1985)  (jucstions  a  fossil 
record  from  southeast  Arizona  that  would  con- 
firm this  past  di.stribution,  but  the  present  dis- 
junct range  of  the  species  (Fig.  1)  implies  its 
former  presence  in  southeast  Arizona  regardless 
of  the  fossil  record.  Post-Pleistocene  climatic 
changes  fragmented  this  distribution,  and  local 
extinctions  in  southeast  Arizona  apparentK'  sep- 
arated the  Mexican  and  northern  populations. 
This  scenario  is  consistent  with  the  historical 
legacy  h\pothesis,  but  there  is  also  evidence  that 
the  pattern  has  been  modified  b\  recent  dis- 
persal as  disc-ussed  below. 

E\il)i:\(:K  FROM  Arizona. — The  Mexican 
\-ole  now  occurs  in  the  continuous  forests  of 
central  Arizona  and  on  isolated  mountains  to  the 
south,  southwest,  and  north  (Figs.  1.  2).  Four 
populations  occm-  on  mountains  connected  to 
the  central  high  countn  In  pimon-juniper 
woodland  and  interior  chaparral  (Brown  and 
Lowe  1983),  through  which  the  species  could 
(lis[)er,se:  the  Nantanes  Plateau,  the  Sierra 
Ancha,  the  Bradshaw  Moimtains,  and  the  South 
Kaibab  (Fig.  2).  Three  other  populations  occur 
at   sites    that    are   isolated   b\-   grasslands   but 


264 


Great  Basin  Naturalist 


[\V)Iuiiie  52 


lOOkm 


F"ig.  3.  Details  ol  the  clistrihiitioii  ot  Microtiis  iiu'xicanit\ 
in  N't'w  Mexico  and  .southern  Colorado  showing  some  iso- 
lated populations  (modified  IVom  Findlev  et  al.  1975;  some 
data  from  Hall  I9S1).  Open  eireles  indicate  records  listed 
bv  Finlev  et  al.  (1986). 


interconnected  hy  pinNon-jnniper  woodland 
and  interior  cliaparral:  Pro.spect  \alle\,  the 
Music  Monntains,  and  the  llnala[)ai  Monntain.s 
(Fig.  2). 

Since  -lif  iiualapai  Monntain.s  and  Prospect 
X'alley  still  contain  small  ])atches  ol'  forest,  the 
vole  populations  at  these  sites  might  be 
Plei.stocene  relicts  in  Forest  refugia.  Bnt  the 
population  in  the  Music  Mountains,  a  site 
midway  bet\veen  the  other  two,  consists  of OnK- 
pinyon-juniper  woodland  (Sj^icer  ot  al.  19S5). 
Tliis  habitat  interconnects  all  three  l()caliti(>s 
and  is  more  likely  to  sene  as  a  dispersal  corridor 
than  as  a  post-Plei.st(X-ene  refnginm.  The  spe- 
cies was  recorded  in  the  Uualapai  Mountains  in 
1923  and  in  Prospect  \'alley  in  1913,  but  it  was 
not  found  in  the  Mu.sic  Mountains  until  1981 
(Spiceretal.  1985). 

When  the  rate  of  dispersal  exceeds  that  of 
extinction,  a  species  should  be  present  on  those 
luontane  islands  closest  to  the  .source,  assuming 
the  species  can  cross  the  intenening  habitat 
(Mac.Vrthur  andWilson  1967).  The  distiibution 
of  the  Mexican  xole  in  the  Southwest  generalK 
fits  this  model  (Fig.  2;  Lomolino  et  al.  1989).  In 
Arizona  tlie  most  closely  related  i.solate  popula- 
tions occur  in  geographic  jiroximitx  (Hofhuc>ister 


1986).  Recent  dispersal  is  not  the  onK  possible 
explanation  for  this  pattern,  but  it  is  the  most 
parsimonious  one;  ancient  relicts  in  dissimilar 
habitats  would  be  expected  to  show  more  e\i- 
dence  of  di\'ergence  after  sexeral  thousand 
vears. 

There  is  exidence  of  a  recent  range  expan- 
sion in  northeast  Arizona.  The  Mexican  vole  was 
first  recorded  in  the  Navajo  Mountains  in  south- 
ern Utah  and  northern  Arizona  in  1933  (Benson 
1935).  Although  this  locality  seems  isolated, 
since  1986  the  .species  has  turned  up  at  se\eral 
other  sites  on  Black  Mesa  in  northeast  Arizona 
( Spicer  1987).  These  sites  fall  on  a  line  southeast 
from  Na\ajo  Mountain  to  the  southwest  foot- 
liills  of  the  Chuska  Mountains. 

At  Black  Mesa  (Fig.  2)  the  habitat  is  pin\'on- 
juniper,  with  ponderosa  pines  and  a  few  Doug- 
las-firs on  north-facing  slopes,  draws,  and  other 
protected  areas  (Spicer  1987).  Again,  this  is 
relati\el\'  poor  habitat  for  this  species,  and  it 
seems  unlikeK'  that  the  population  could  have 
siuvived  in  isolation  for  sexeral  thousand  \ears. 
Between  these  sites  and  Naxajo  Mountain  is 
mostlv  pinyon-juniper,  with  narrow  strips  of 
northern  grassland  and  Great  Basin  desertscrub 
(Browii  and  Lowe  1983).  The  Mexican  \ole 
occupies  these  habitats  elsewhere  and  presum- 
abl\  can  disperse  through  them.  This  scenario 
implies  that  the  Chuska  Mountains,  now  unoc- 
cupied bv  the  species  (Hoffmeister  1986),  will 
eventualK'  be  colonized  (or  recolonized)  from 
the  northwest. 

E\'idenc:e  from  New  Me.mco  and  Colo- 
rado.— Findle\et  al.  ( 1975)  suggested  that  the 
range  of  Microtus  iiwxicaniis  in  New  Mexico 
could  lia\  e  expanded  as  a  result  of  recent  dis- 
})ersal.  In  the  Sandia  Mountains,  trapping  from 
1950  to  1970  re\ealed  onl\-  M.  hn^iicaudus. 
Mexican  xoles  were  first  taken  there  in  1970  and 
soon  became  the  dominant  species.  While  the 
species  could  ha\e  been  overlooked  earlier,  dis- 
persal from  the  Manzano  Mountains  (Fig.  3)  is 
an  ecuuilK'  likel\-  scenario.  Until  1 975  these  were 
the  northernmost  records  east  of  the  Rio 
( wande  Ri\er  in  New  Mexico.  The  Mexican  vole 
has  since  been  recorded  from  five  sites  in 
extreme  northeast  New  Mexico  (Dakjuest  1975, 
Finlex-  et  al.  1986). 

In  C>olora(k)  the  first  .specimens  were  taken 
in  1956  at  Mesa  Verde  (Rodeck  and  Anderson 
1956).  Later  the  species  was  found  at  seven 
more  (Colorado  sites  (Fig.  3;  Mellott  and  Choate 
1984,  Finle\  et  al.  1986).  \  trapping  studv  in 


19921 


\'()i.i".  DisrKP.SM. 


265 


'l^vm.l-:  1.  Sontlirni  inaiiniial  sprcics  lorwhicli  there  ise\i<lcncc  ol  a  recent  nortliwanl  raiim' e\])aiisioii.  Unless  iiulicated 
otherwise,  e\icleiice  is  based  on  directionality  and  clironolo<;;\  ol  records:  1,  Arizona  distribntion  in  Cockruin  U960)  vs. 
Ilolfnieistcr  (19S6);  2,  distribution  in  Hall  luul  Kelson  (1959)  \s.  Hall  (19S1);  3,  Texas  distribntion  in  Tavlor  and  Davis 
(1947),  Davis  (1960).  ;uk1  Davis  (1974).  Nomenclature  follows  Jones  et  al.  (1986). 


Species 


Region  and  direction 
of  expansion 


l'".\idence  ami  Helerences 


DicMphis  virainiana 

Mi>nit(>(ij)\  iit('iial(>iilii/ll<i 

Cluunmi/ctcris  iiifxiraiui 

Ij'jiloiiiictcris  sanhonii 

I.tisiiinis  ciid 
hlidiniciiris  phi/llotis 

Tddiirkid  jcinorosaccci 
Tadfihdd  ituicrotis 
Diisifpiis  lunciutinrtiis 

Lcptis  allciii 
S(iuni\  (ihciii 

Bdiounjs  tdiiliiii 
SigtHixloii  lii.spitlu.s 

Siginodoii  fuliiicnlcr 
Sigmodan  ocJirognathus 

\licr(>tits  iiivxicdiiiis 

^dsiid  UdsUd 


'onrpatiis  mcsolcunis 
njds.su  Idjdcii 


N  throusili  E  U.S.;  N  into 
S  .Arizona  tniiii  \  Nh-xico 

\  in  Texas 

Now  a  winter  resident  in  S 
.Arizona 

Now  a  winter  resident  in  S 
Arizona 

N  in  Texas 

N  in  .SW  U.S.  to  UtcJi 

N  in  .Arizona 

N  in  .Arizona;  also  Texas? 

N  froin  S  Texas  into  Okla- 
homa, ("olorado,  Kansas, 
and  Nebraska 

Limiti'dK  NE  in  Arizona 

NW  in  Colorado.  N  into 
\\\()niinti.  W  into  Utah 

N  from  SE  Texas  into 
Oklahoma,  and  NE  in 
New  .Mexico 

N  in  the  U.S.;  through 
Kansas  to  Nebritska.  and 
N  in  Rio  Grande  \'alle\  in 
New  Mexico 

N  in  New  .\h'xico 

.\\\  in  .Arizona  and  N  in 
Texas 

\arions  in  .Arizona;  N  in 
.New  .Mexico  into  S  Colo- 
rado 

N\\  in  Arizona  ;uh1  per- 
haps in  .New  Mexico 


N\\'  in  Arizona 

N    in    Arizona   and    New 
.Mexico 


I 


Udvardy  (1969).  McM;uius  (1974);  Y.  Petryszvii 
(personal  communii'ation ! 

3;  Tavlor  and  Da\is  0947);  Da\is  (1960);  Da\is 
(1974);  Mollhagen  (197.3) 

H.  Sidner  (personal  communication  '.  probabK  due 
to  humiuiugbird  feeders 

K.  Sidner  (personal  comiiiiinication  '.  [)rob;il)l\  due 
to  lumnningbird  feeders 

.Spencer  etal.  (198S) 

First  U.S.  record  was  in  19.55  in  SE  Arizona  (Cock- 
nmi  1956);  2 

1  and  2 

1;  Mollhagen  (197.3) 

i^uchamiau  and  Tahnage  (1954);  Ud\ard\-  ( 1969); 
Humphre\  (19741;  Meaneyetal.  (1987) 

1;  lack  of  records  in  N  (Cochise  ('o.  until  1976 
(Allen  1895,  Roth  and  Cockrum  1976) 

I3a\is  and  Bissell  (1989';  known  dispersal  ability' 
and  histon'  of  ponderosa  pine  distribution  i  l^avis 
and  Brown  1989) 

Diersing  (1979);  Stangl  and  Dakjucst  (1986); 
Tavlor  and  Daxis  (1947)  \s.  Da\is  (1974);  recent 
record  in  Ijuia  ('o..  New  Mexico  (\\.  CTannon, 
personal  comuHmicatiouh  (^hoate  et  al.  (1990) 

Cockrum  (1952);  Mohlenrich  ( 1961 );  Jones  (1960); 
Cameron  and  Spenci'r  ( 1981) 


.Mohlenrich  (1961) 

Davis  and  Dunford  ( 1987);  Davis  and  Ward  ( 1988); 
Da\is  et  al.  (1988);  Hollander  et  d.  (1990);  Stangl 
and  Dalcjuest  (1991) 

This  studv 


Not  reported  bv  eiirlv  explorers  (Davis  1982);  not 
recorded  in  .Arizona  until  1892,  in  extreme  S 
( HoffincMster  1986);  no  late  Pleistocene  record 
(Harris  19.85.  Tabor  1940);  Wallmo  and  Gallizioli 
(1954):  but  see  Kaufmann  et  al.  (1976) 

1;  recent  records  (Hoflmeister  1986) 

Indian  name  for  peccarv  is  of  Spanish  origin  (Sowis 
1984);  rarelv  encountered  bv  early  explorers 
(Davis  1982);  no  use  bv  earlv  prehistoric  cultures 
(Crosswhite  1984,  Sovvls  1984) 


266 


Ghkat  Basin  Naturalist 


[\  olunie  52 


1938,  and  others  prior  to  1975,  found  no  Mexi- 
can \-oles  near  Cimarron,  New  Mexico,  although 
otlier  \ole  species  were  taken  (Armstrong  1 972, 
Findlev  et  al.  1975).  The  Mexican  vole  is  now 
common  in  tlie  area  (Finley  et  al.  1986);  dius, 
the  northward  range  expansion  by  this  species 
ma\-  he  continuing  into  nottfieast  New  Mexico 
and  southeast  Colorado. 

Discussion  and  Conclusions 

The  historical  legacy  hypothesis  requires 
widespread  late  Pleistocene  distribution.  The 
fossil  record  documents  the  late  Pleistocene 
presence  of  Microtus  nwxicainis  in  jouthem 
New  Mexico,  adjacent  portions  of  Texas,  and 
(perhaps)  southeast  Arizona.  Despite  the  admit- 
tedlv  weak  fossil  record,  however,  there  is  no 
evidence  that  the  species'  range  formerly 
included  the  entire  area  where  populations  now 
exist  (Harris  1985).  Sex'eral  lines  of  e\idence 
support  post-Pleistocene  dispersal  lor  this 
species: 

1 .  Distance  as  a  predictor  of  pres- 
ence/absence (Lomolino  et  al.  1989). 

2.  Tlu^  clo.se  relationship  of  adjacent  Ari- 
zona populations,  isolated  bv  theoreti- 
calK'  crossable  habitat. 

3.  Its  presence  in  isolated  habitats  unlikeK 
to  have  served  as  post-Pleistocene 
refugia. 

4.  Recent  records  suggesting  dispersal  in 
n(nlh\\'est  and  northeast  Arizona,  cen- 
tral and  northeast  New  Mexico,  and 
southern  (Colorado. 

Although  the  distribution  of  the  Mexican  xole 
undoubtedly  has  been  influenced  by  historical 
events  and  by  local  extinctions,  it  is  difficult  to 
ignore  the  evidence  of  past  and  continuing  post- 
Pleistocene  dispersal. 

A  reviewer  of  this  paper  asked  win'  the  Mex- 
ican vole  and  other  small  mammals  took  4()()() 
years  to  reach  certain  localities  that  w(^  claim 
were  colonized  within  the  past  few  decades. 
This  point  re(juires  clarification.  First,  there 
have  been  local  changes  in  \egetation  and  cli- 
mate in  the  Southwest  during  the  past  50  to  1  ()() 
years,  and  these  conditions  max-  have  favored 
recent  dispersal  even  though  the  broader  pic- 
ture has  remained  constant  for  some  4000  vears. 
Second,  we  do  not  claim  that  these  recent 
records  represent  the//r.sf  colonizations  by  the 
Mexican  vole  or  other  species.  Thev  are  simply 


the  first  such  events  tliat  have  been  recorded  in 
the  literature.  If  these  animals  were  able  to  cross 
unsuitable  habitat  once,  then  thev  could  have 
done  so  repeatedl)'  in  the  course  of  centuries. 

Our  suggestion  of  recent  dispersal  bv  the 
Mexican  vole  should  be  evaluated  in  the  context 
of  a  more  general  pattern  involving  manv 
mammal  species.  Post-Pleistocene  dispersal  has 
influenced  montane  species  assemblages 
throughout  much  of  the  Southwest  (Lomolino 
et  al.  1989).  In  addition,  we  propose  a  second 
pattern  of  recent  northward  range  expansion 
involving  at  least  19  North  American  mammal 
species,  all  primarily  austral  in  distributicMi  but 
occupying  a  wide  range  of  habitats  (Table  1). 

This  pattern  of  northward  dispersal  is  not 
easily  exjolained,  and  there  is  unlikelv  to  be  a 
single  causative  factor.  For  some  species,  the 
shift  appears  to  result  from  climatic  change 
and/or  habitat  modification  b)'  humans.  Alterna- 
tively, the  pattern  can  be  viewed  as  one  smiill, 
reconiizable  northward  surtje  in  a  continuincr 
Holocene  c>cle  of  nortli/south  distribution 
shifts. 


ACKNOWLEDCMENTS 

Preliminaiv  drafts  of  the  manuscript  were 
read  bv  R.  Sidner,  M.  V.  Lomolino,  and  E.  L. 
Cocknun.  T  \'an  Devender  and  an  anonvnnous 
reviewer  also  provided  helpful  comments. 
Some  of  the  fieldwork  cited  was  conducted  by 
R.  B.  Spicer  and  C.  LaRue  with  support  from 
the  Arizona  Department  of  Game  and  Fish. 

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Received  12  October  1990 
Accepted  17  June  1992 


Creat  Basin  \atiiralist  52io),  pp.  269-277 

CAN  TOWNSEND'S  GROUND  SQUIRRELS  SURMVE 
ON  A  DIET  OF  EXOTIC  ANNUALS? 

Eric  Yensen    ami  Dana  I,.  OiiiniicN" 

Abstrvct. — Southwesteni  Idaho  dcsrrt  sliruh-bunchgras.s  raiiiicland  is  liciii",;;  iiuaded  In  liir-proiic  exotic  aniiiuds  that 
pcnnanentlv  dominate  the  landscape  iollowing  wildfires.  This  stiuK  \\as  luulertiiken  to  describe  diet.s  oi  Tow  nsiMid's  liround 
s(juirrels  (Spennophilus  totcii.sciulii  idtiJiocnsi.s)  at  four  stiuK  sites  with  xaning  degrees  ol'  exotic  annual  in\asion  to 
determine  if  tlie  S([uirrels  could  utilize  high  proportions  of  exotic  annuals  in  their  diets.  Townsend's  ground  s(juirrels  were 
collected  in  .March  and  Ma\  of  19(S7  and  1988,  and  stomach  contents  were  anal\7.ed  using  a  microhistological  techniciue. 
Grasses  comprised  37-87%  of  Townsend's  ground  squirrel  diets  at  the  four  sites.  N'ati\e  species.  especialK'  Sandberg's 
bluegrass  [Poa  secunch),  winterfat  (Ceratoklcs  lanata),  big  sagebrush  (Artemisia  tridentatu^  and  six-weeks  fescue  (Viilpia 
octojlora)  constituted  7-96%  (x  =  47.2%)  of  the  diet,  whereas  exotic  species.  especi;ill\'  cheatgrass  (Broinns  tedontm), 
tunibleweed  iSalsola  ihcrica).  iuid  tans\miLStards  [Descuminia  spp.)  made  up  4-68%  (\  =  48.0%)  of  the  diet.  At  each  site 
2-A  species  comprised  >90%  of  the  diet.  There  was  no  apparent  correlation  betw  een  the  importance  \  alues  of  exotic  species 
at  a  site  and  their  importance  in  Townsend's  ground  squirrel  diets. 

Kvij  uonl.s:  .Spermophilus  t(n\nsendii,  fixxl  habits,  dictani  anali/sis.  Idaho,  urotuid  stjiiirrcls. 


The  Snake  Ri\er  Birds  of  Pre)'  Area  i.s  a 
243,()()()-ha  tract  of  multiple-u.se  shrub-steppe 
rangeland  achuinistered  bv  the  U.S.  Biu-eau  of 
Land  Management.  Towaisends  ground  squir- 
rels (Speniu)pliihis  townsendii  idaltoensis)  are 
important  prey  of  raptors,  and  continued  exis- 
tence of  the  area's  dense  breeding  populations 
of  raptors  depends  upon  dense  Townisends 
ground  scjuirrel  populations  (U.S.  Department 
of  Interior  1979). 

hnasion  of  southwestern  Idaho  rangeland 
by  e.xotic  annuals  such  as  cheatgrass  (Broinus 
tectonun).  tumblemustard  {SisijDihhuiii  aliis- 
siiiiuni\  pinnate  tan.symustard  {Descuraiiiia 
piniuila).  and  tunil)I(n\-e(Hl  (Salsola  iheiica)  has 
resulted  in  trecjueut  and  destructive  wildfires 
that  kill  natix  e  shrubs  and  weaken  natixe  bunch- 
grasses.  Oxer  time,  fires  ha\e  n\sulted  in  the 
permanent  replacenu'iit  ot  luitiw  slirul)-  and 
bunchgrass-dominated  comnumities  b\  e.xotic 
annual-dominated  comnumities  (YenscMi  1980. 
Kochert  and  Pellant  1986). 

Townsend's  gromid  S(juirrel  populations  are 
niucli  less  stable  in  exotic  annual-dominated 
connnnumities  than  in  natixe  shrub  communi- 
ties (Yensen  et  al.  1992).  Native  perennial  f()d)s, 


bunchgrasses,  and  shrubs  apparentK'  proxide  a 
more  constant,  stable  food  soinx-e  than  exotic 
annual  species  that  ma\-  xan-  in  producti\1t>' 
between  xxet  and  dn"  xears  b\-  sexeral  orders  of 
magnitude  (Young  et  al.  1987). 

Like  other  ground  squirrels  of  subgenus 
Spenuopluhis.  Townsend's  groimd  s(juirrels  eat 
green  xegetation  earK'  in  their  fom-  to  iixc- 
month  actixe  .season,  then  eat  seeds  of  gras.ses 
and  forbs  to  fatten  up  for  hibernation  ( Hoxxell 
1938,  Rickart  1982).  In  southwestern  Idaho, 
Toxx'usend's  groimd  squirrels  are  in  (^stixa- 
tion/hibeniation  from  |une  or  (ulx'  until  the  fol- 
lowing Januan  or  Februarx  with  loxv  sunixal 
rates  (ca.  289f ;  Smith  and  lohnson  1985).  Food 
quantitx  and  cjualitx'  could  influence  oxerwin- 
teiing  surxixal  as  xx'ell  as  reproductixe  success 
the  folloxxing  spring. 

Toxxnsend's  ground  s(|uirrels  are  known  to 
eat  natixe  forbs  {Sphdcnilcca:  Daxis  1939 ),  bunch- 
grasses  {Pod  sp.;  fune  grass,  Kiwlciia  sp.;  Daxis 
1939),  and  desert  shnibs  (big  sagebrush,  Arte- 
uiisia  Iridcntata:  budsage,  Artemisia  spinescens; 
shadscale,  Atriplex  coufeiiifolia;  Daxis  1939, 
lohnson  1961),  as  xvell  as  in.sects  such  as  grass- 
hoppers and  cicadas,  and  occasional!)'  carrion 


,  Museum  of  Natural  II  istorv-.  .Mbert.sou  College,  Caldwell,  Idaho  8,360.5. 

'Bureau  of  Land  Mauanenient.  .3948  Uevelopmeut  Ave..  Boise.  Idaho  8370.5.  Present  aildress:  Idaho  .Xnii\  National  Cinard.  Department  of  En\iroinnent. 
Box  45,  Gowen  Field.  Boise,  Idaho  8370.5. 


269 


270 


Great  Basin  Natuhaljst 


[W^lume  52 


Tabi.K  1.  \e2;etation  importance  values  (%  relative  cover  plus  %  fVeciiiencv)  in  May  1987  and  1988  at  four  study  sites 
near  Co\ote  Butte  in  the  Snake  River  Birds  ofPrev  Area,  southwestern  Idaho. 


Study  Site 

Bis 

Nativ 

e 

Exotic 

Rehabilitation 

Species 

sagebmsh 

grasses 

annuals 

seeding 

1987 

1988 

1987 

1988 

198' 

r           1988 

1987 

1988 

Ghassks 

'Broiniis  tectoniin 

25 

2 

11 

33 

86 

35 

0 

31 

Poo  seen  11(1(1 

67 

60 

90 

58 

45 

45 

85 

60 

Viilpid  octojlora 

16 

"■ 

24 

0 

2 

8 

12 

3 

Sitanioii  liijstrix 

16 

11 

14 

14 

8 

21 

0 

28 

°.\<ir<>i)ijron  dcsciioni ni 

0 

0 

0 

0 

0 

0 

29 

0 

SlIIUBS 

Cc rat (7 ides  Unuita 

29 

47 

3 

5 

(1 

0 

( 

0 

Artemisia  trideiitafd 

33 

39 

0 

0 

0 

4 

0 

0 

Atni)lex  iiitttiillii 

0 

0 

0 

0 

0 

0 

0 

20 

FOKBS 

"Sdlsola  iberiea 

0 

0 

33 

26 

0 

0 

40 

18 

"Descumiuia  sopliia 

0 

0 

3, 

0 

0 

0 

0 

0 

'Sistjiiihriiiin  (illissimiiin 

0 

11 

8 

31 

34 

14 

0 

5 

"iMctucd  serriold 

0 

0 

0 

0 

0 

0 

4 

0 

Other  forbs 

0 

5 

0 

9 

0 

■-) 

0 

2 

Total  Co\F.K(%) 

35 

24 

26 

15 

21 

10 

18 

14 

*ex<)tic  species 

(Howell  1938.  Alfoni  1940).  However,  they  do 
eat  introcluced  clieatgrass,  tunihleiiiustard, 
peppergrass  {Lcpkliuiii  pcifolidfiiin;  Da\is 
1939)  as  well  as  crop  species  like  alfalfa,  wheat, 
bade\;  potatoes,  beets,  carrots,  and  lettuce 
(Howell  1938). 

Johnson  (1980)  and  Rogers  and  Gano  (1980) 
studied  diets  of"  Sj)('niioj)liilu.s  lowiisciulii 
townsendii  in  Washington  and  found  natixe 
bluegrass  {Foa  sp.,  26-29%)  and  lupine 
{Lupinus  hixifloais,  11-25%)  to  be  dietaril\- 
important,  whereas  Dcscurainin  was  the  onl\- 
e.\(;tic  eaten  in  quantit\'  (15-33%);  cheatgrass, 
tnnibleweed,  tuuiblouustard,  and  peppergrass 
constituted  0-4%  of  the  diet.  Johnson  et  al. 
(1977)  estimated  the  percent  \-olunie  of  food 
categories  in  174  Towiisend's  ground  scjuirrel 
stomachs  in  the  Snake  Ri\-er  Birds  of  Prey  Area. 
They  found  grasses,  including  cheatgrass,  were 
most  important,  followed  In  forbs  and  winterfat 
{Ccnitohlcs  laiuila). 

Because  cheatgrass.  tuiiiblewced.  tinnble- 
nuistard,  and  peppergrass  are  becoming 
increasingly  dominant  in  the  Snake  River  Bircls 
of  Prey  Area,  tliis  study  was  designed  to  leaiii  if 
Townsends  ground  s(juirrels  were  substituting 
these  exotics  for  natixe  species  in  their  diets.  We 
also  wished  to  learn  if  cousnmplion-introdnced 
plant  species  increased  with  increases  in  the 


proportion  of  exotic  aiuuial  species  in  the  habi- 
tat. Hf)wever,  the  studx'  was  not  designed  to 
stud\'  dietaiv  preference  as  such. 

Study  Sites 

Four  stud\  sites  were  located  near  Go\ote 
Butte,  approxiuiateK'  19  km  south  of  Kuna,  Ada 
Gountx,  Idalio,  in  the  Snake  Ri\er  Birds  of  Prev 
Area.  The  sites  described  l:)elow  were  selected 
for  progressively  greater  dexiation  from  undis- 
turbed native  vegetation. 

Unburned  big  SA(;EBRL)S1I. — This  site 
(TIS.  RIW  Sec.  24;  elev.  850  m)  is  a  big  sage- 
brush-winterfat  mosaic  and  represents  the 
uubunied  condition  of  thc^  other  three  sites.  Big 
sagebrush,  winterfat,  and  native  grasses 
(Sandbergs  bluegrass  [Poa  sccunda],  squirrel- 
tail  \Sit(nii()ii  lii/sfrix],  and  sLx-weeks  fescue 
[Viilpia  ()cl()fh)ni\)  dominate  the  site;  cheatgrass 
is  the  main  exotic  annual  (Table  1). 

Nvri\  !■:  GRASS.— This  site  (TIS,  RIW,  Sec. 
13;  ele\'.  850  m)  is  <1  km  northwest  of  the 
unburned  big  sagebrush  site  in  a  former  big 
sagel)rush-wint(M"fat  conuiiunitv  burned  by  a 
human-caused  wildfire  on  26  August  1983.  The 
fin^  killed  thc^  shnibs,  and  the  site  was  domi- 
nated subse(juentl\'  b\  uatixx-  Sandbergs  blue- 
grass,  six-weeks  fescue,  and  scjuirreltail,  with 


19921 


TowxsKXDs  (iHorxn  Soiikhki,  Dikts 


271 


some  introduced  tuiiil)I('\\('('d.  tlicatgrass,  and 
other  (Aotic-  animals  present  lTal)le  1  V 

fclXOTIc  AXXIALS— This  site  (TIS.  HIW, 
Sec.  13;  ele\'.  850  ni)  is  adjacent  to  the  natixe 
grass  site  and  was  similar  to  it  prior  to  the  1983 
burn  (13.  L.  Quinnex;  unpnbHshed  data).  Both 
sites  were  bunied  b\"  the  same  fire.  However, 
since  the  fire,  the  exotic  annuals  cheatgrass  and 
tumble  mustard,  with  some  remnant  natixe 
glasses,  especiallx  Sandbergs  bluegrass  (Table 
1),  haxe  dominated  the  site. 

RKHABILITATION  seeding. — This  site  is 
located  fi  km  east,  2.5  km  south  (TIS,  RIE,  Sec. 
27:  elex'.  885  m)  of  the  unburned  big  sagebnisli 
site.  The  area  burned  in  1981,  xxas  reseeded 
with  deseit x\iu\itgrass  (Ag^ropi/roii  desertoniin) 
in  1982,  but  burned  again  in  1983.  In  1987  and 
1988,  tlie  area  xx'as  dominated  bx'  Sandbergs 
bluegrass,  desert  xxheatsrass,  tumblexxeed,  and 
other  natixe  and  exotic  forbs  (Table  1). 

Methods 

To  determine  the  degree  of  exotic  annual 
inxasiou  at  each  site,  vegetation  analxsis  xxas 
conducted  in  earl)' June  1987  and  late  Max'  1988 
x\  hile  Towiisend's  ground  squirrels  xvere  being 
collected.  At  each  site  xve  used  a  transect  xxith 
foitx"  1-m'  quadrats  spaced  at  lO-m  intenals 
(Daubenmire  1959).  Percent  coxer  of  each  spe- 
cies xx'as  estimated  using  a  1-nr  (jiiadrat  frame 
dixided  into  tiMiths  to  facilitate  estimation.  To 
gixe  a  better  approximation  of  the  axailabilitx  of 
each  plant  species,  percent  relatixe  coxer  and 
jiercent  relative  frequencv  xx^ere  conxcMted  to 
importance  values  (Cox  1990). 

Squirrels  xvere  collected  bx  tia])ping  and 
shooting  at  all  four  sites  in  Max  and  June  1987 
in  =  75)  and  in  March  and  Max  1988  (//  =  42) 
except  from  the  rehabilitation  seeding  site  in 
May  1988.  Squirrels  xx'ere  aged  in  the  field  using 
pelage  and  bodx  xxeight  ciiteria  (Bureau  oi 
Land  Management,  unpublislied  data).  Hepre- 
sentatix-e  specimens  xxere  prepanxl  as  ( 1 )  stan- 
dard stiidx  skins  xxith  skulls  (/;  =  12),  (2) 
skeletons  [ii  =  3),  or  (3)  skulls  only  (/;  =  25)  and 
deposited  in  the  Albert.son  College  Museum  of 
Natural  Histon.  Tooth-xx-ear  patterns  (Yenseii 
1991 )  xx'ere  consistent  xxith  the  ase  assi";imi(Mits 
for  all  specimens.  Based  on  these  criteria,  all 
1987  specimens  xx'ere  juxeniles  since  thex  xxere 
collected  late  in  the  active  season  xx'hile  the 
adults  xx'ere  entering  seasonal  torpor:  all  1988 
specimens  xx^ere  either  yearlings  or  adults. 


Stomachs  xx'ere  remoxed  from  the  animals 
iimn(xliat(^lx- postmortem  andpresened  in  70% 
cthaiiol.  In  the  lab,  stomach  contents  xvere 
remoxed  from  ethanol,  diluted  50%  xxith  xxater, 
and  homogenized  1  min  in  a  Waring  blentler  to 
produce  fragments  of  uniform  size.  The  homog- 
enate  xvas  xvashed  through  a  l-mm  siex-e 
(Hansen  1978)  and  collected  in  aO.l-mm  screen 
to  remoxe  tiny,  unidentifiable  fragments.  The 
material  xvas  then  mounted  on  microsco])e 
slides  using  Hertxvigs  and  Hoxers  media 
( Sparks  and  Malechek  1 968 ). 

Plant  .species  in  the  diet  x\er(^  identified  bx' 
compari.sons  to  a  reference  collection  of  micro- 
scope slides  using  microhistological  characters. 
All  reference  slides  xx'ere  made  from  catalogued 
specimens  in  the  Albertson  College  Harold  M. 
Tucker  Herbarium  andxx'ere  prepared  using  the 
technicjue  described  aboxe. 

For  food  habits  analysis,  one  slide  xvas  exam- 
ined per  stomach.  Occurrence  of  food  catego- 
ries (frequencx')  xx^as  recorded  from  each  of  20 
microscope  fields  per  slide  using  a  phase- 
contrast  microscope  at  lOOX.  Frequencv/20 
fields  xx'as  tlien  converted  to  percent  relatixe 
density'  (Sparks  and  Malechek  1968)  using  a 
table  dex'ek)ped  for  fre(juencx-to-densitx  con- 
xersion  (Fracker  and  Brischle  1944). 

The  importance  of  each  dietarxcategorxxxas 
calculated  in  three  xvays:  (1)  percent  relatixe 
densitv;  a  standard  dn-xveight  conxersion  from 
frecpiency  data  (Spark's  and'Malechek  1968);  (2) 
percent  frecjnencx  in  stomaclis.  the  percentage 
of  stomachs  from  a  site  xxith  the  item;  and  (3) 
percent  fre(jnencx'  in  micr()sco[)ic  fields,  the 
percentage  of  all  microscopic  fields  from  a  site 
xxith  the  item. 

Txx'entx'  microscopic  fields  xxere  examined 
from  each  slide  using  a  predetermined  pattern, 
and  frequencx'  of  occurrence  of  each  spt^cies 
xxas  recorded.  The  frecjuency  of  each  dietarx 
categon/2()  fields  on  one  slide  xxas  compared 
xxith  other  slides  (or  replicate  counts  of  the  same 
slide)  using  the  Kulcvznski  Index  (Oo.sting 
1956)  (also  xxell  knoxxii  as  the  Brax-Curtis  simi- 
laritx  index  [Brax-  and  C>urtis  1957]) 

2w/  (a  +b) 

The  index  xxas  calculated  as  a  dissimilaritx  index, 

1  -  [2w/{a  +  h)] 

using  a  BASK'  microcomputer  program  pro- 
xided  bv  Ludxxig  and  Rexiiolds  ( 1988). 

\\'eather  data  xxere  from  the  National  Oce- 
anic and  .Atmospheric  Administration  monthly 


272 


CiHEAT  Basin  Nathhalist 


[N'olunie  52 


Tablk  2.  Late  season  (25  Mav-19  June)  1987  Townsend's  ground  stjuinel  diets.  Data  are  from  stomachs  of  juvenile  TGS 
at  four  sites  in  tlie  Snake  Ri\er  Birds  of  Prev  Area.  Adults  were  entering  toipor  and  none  were  collected  during  this  period. 
Dietar\-  composition  is  given  as  percent  relative  densitv'  (RD).  percent  frequency  in  microscope  fields  (MF),  and  percent 
frequency  in  stomachs  (PS)  for  each  dietary  categor\-.  Other  s\nil)()ls:  +  =  <17f,-  =  aiiscut,  and//  =  number  of  stomachs. 


Dietar)'  categor>- 


Unbumed 
big  sagebrush 

RD    MF    PS 


Native 
grasses 


Exotic 
annuals 


RD    MF     PS 


RD    MF     PS 


21 


GlUSSES 

Bromu.s  tcctonim  22  41 

Poa  sc'ciinda  24  35 

Sitanion  liystrix  +  2 

Oryzopsis  hijmcnoides  - 

Grass  seed  +  5 

Grass  root?  2  9 

Total  grasses  49 

Siihi;bs 

Ccratokh's  lanala  3  9 

Artemisia  thdciitata  +  2 

Atiiplex  niitttillii  -  - 

Clinfsotlunniitis  lisciiliflmus         -  - 

Total  shrubs  3 

FOHHS 

Sdlsola  ihciiai  39  69 

Sisi/inhiiiiui  (iltissiimiin  +  + 

Di'scitniiuia — 2  spp.  +  2 

Lcpiditiiii  pcrfoliatum  -  - 

C.njptdttthd  intcmtpta  +  + 

Rdnitunihis  tcsticuldln.s  +  1 
Ldctucd  scrriold 

Gheno[5odiaceae  -  - 

Uuidentilied  lorb  -  - 

Total  fod)s  40 

MlSCKLI-ANKOrS 

Insects  S  17 

Fungi  -  - 

Unknown  +  + 

Unidentified  seed  +  I 

Totiil  mi.scelliuieous  S 


71 

Sfi 
10 

19 
10 


52 
14 


91 
5 

24 

5 
5 


62 


20 


62      93 


19 
1 


20 


44 
4 


100 


+ 

+ 

5 

+ 

1 

5 

+ 
)4 

2 

5 

4 

5 

10 

1 

3 

10 

+ 

+ 

5 

40 

60 

5 

20 


5 


90 
10 


15 


31 

45 

87 

"■ 

S 

40 

+ 

+ 

13 

+ 

1 

13 

+ 

3 

13 

39 

43 

59 

67 

6 

19 

67 

2 

5 

13 

51 

1 

3 

20 

13 


7  21  ST 

+  1  ■; 

+  4  7 

8 


Rehabilitation 
seeding 

RD     MF     PS 


19 


95 
11 


57 

11 

17 

32 

+ 

■T 

5 

4 

12 

37 

+ 

2 

5 

16 

63 
21 


21 


+ 

0 

21 

6 

" 

11 

L8 

3 

11 

53 

+ 

2 

5 

4 

5 

5 

Idaho  (.'limatolopcal  Data  rcpoit.s  lortlic^  Kuna 
2  NNE  weather  station  ea.  20  km  N  oFthe  .stucl\- 
sites. 

Hksuits 

Vegetation  Analwsis 

The  \egetation  at  c-ach  site  (Tal)le  I)  \ane(l 
signifieantly  from  the  other  sites  (all  p  <  .01; 
H  X  C  G-tests  of  independenee;  Sokal  and 
Hohlf  1981).  Using  the  Knlc\zn.ski  lnckv\,  the 
similarity-  among  the  four  sites  averaged  48.7% 
(range  27-73%)  in  1987.  The  unburned  sage- 
brush site  was  more  similar  (60% )  to  the  native 
grass  site  and  less  similar  to  the  exotie  annual 
and  seeding  sites  (44  and  47%,  respecti\ely). 


The  \egetation  at  eaeh  ot  the  lour  sites 
\aiied  significantly  (all  p  <  .01;  R  x  C  G-tests 
of  independence)  between  \ears  (Table  1). 
Importance  \aliies  axeraged  65%  similar  (range 
48-77%)  at  a  site  between  years.  Total  percent 
coxer  decreased  on  all  sites  in  1988.  In  1988, 
when  thert^  was  less  herbaceous  co\'er,  the  sites 
were  slightK'  more  similar  (.\  =  61.3%,  range 
47-74%).  Thus,  each  site  differed  almost  as 
much  beh\'een  \ears  as  the  sites  differed  among 
each  other  in  a  gi\en  \ t'ar. 

8tomach  Anahses 

Although  the  three  measm-es  of  dietaiy 
iin[)()rtance  (percent  relative  densit\-  [  =  percent 
dry  weight],  percent  frequency  in  microscope 
fields,  percent  frequency  in  stoiuachs)  gave 


19921 


TOWNSKXDS   (;iU)lM)  S()L  lUKliL  DlKTS 


273 


(lilfci"iMit  iiiiiiu'i'ica]  results,  the  rank  orders 
among  ealegories  were  generalK  consistent 
(Tables  2 — 1).  Ho\\e\er,  percent  frecjuencx  in 
stomachs  was  \er\-  sensitixe  to  sample  sizes. 

There  were  1-9  food  cate<2;ories  per  stom- 
ach. Site  means  varied  from  3.S  to  4.4  categories 
per  stomach.  The  total  numher  ol  food  catego- 
ries usetl  1)\  all  Townsentl's  ground  stjuirreis 
sampled  at  a  site  \arietl  from  4  to  IT  on  the  three 
sampling  occasions  (Ma\-|nne  1987,  March 
1988,  May  1988).  However,  if  species  used  in 
trace  amounts  (<5%  relative  densits)  are  elim- 
inated, onlv  3-6  (x  =  4.0)  categories  were  used 
per  site  and  onK-  2-4  species  comprised  >10% 
ot  the  diet.  Species  comprising  >l()'yf  of  tlie  diet 
at  one  or  more  stud\'  sites  included  Sandberg's 
bluegrass,  cheatgrass,  six-weeks  fescue,  winter- 
fat,  bio;  sagebrush,  tumbleweed,  Descunibiui 
spp.,  seeds  of  bur-buttercup  [Rdiitiiiciiltts  tcs- 
ficiilatus),  and  insects. 

Grasses  were  important  constituents  of  the 
diet  in  both  1987  and  1988  and  often  comprised 
o\er  oiWc  of  tlie  diet  (37-889f  relative  density. 
Tables  2—4).  Sandberg's  bluegrass  and 
cheatgrass  were  both  heavih'  utilized,  especialK 
in  March  1988  (55-87%  of  diet).  Late  in  the 
Townsends  ground  scjuirrel  active  season  (Ma\ 
and  June)  use  of  grasses  declined  (except  at  the 
exotic  annual  site  in  1988).  Most  of  the  grass 
eaten  in  .\hi\-|une  consisted  of  seeds,  especiallv 
of  cheatgrass.  Sandberg's  bluegrass  leaves  were 
utilized  slightlv  more  than  cheatgrass  leaves 
(Tables  2-4),  and  the  tvvo  together  were  far 
more  important  than  all  other  grasses  com- 
bined. S(juirreltail  was  little  used,  altliough  it 
was  the  third  most  abundant  grass. 

Winteifat  (0-43%  relative  density  )  and  big 
sagebrush  (0-21%)  were  both  eaten,  and 
winterfat  was  especiallv  important  at  tiie  exotic 
site  where  it  was  least  abundant.  Winterfat  was 
utilized  at  all  sites  in  1987,  even  though  it  was 
not  abundant  enough  to  be  sampled  1)\  the 
vegetation  analvsis  at  the  exotic  annual  site.  In 
1988  it  was  eaten  onlv  at  the  unburned  big 
sagebnish-winterfat  site,  and  its  use  declined 
between  March  and  Ma\'  1988  (Table  2).  Big 
sagebnish  was  u.sed  in  March  at  all  sites  in  both 
vears  but  was  less  important  in  Max. 

Tumbleweed  and  tumblemustard  were  the 
most  important  forb  species  cc^nsumed.  Tans\- 
mu.stards  [Descurainia  sophia  and  D.  pimiaia). 
peppergrass,  seeds  of  bur-buttercup,  and  leaves 
of  pricklv  lettuce  {Lactuca  seniola)  were  of 
secondan  importance.  All  of  these  are  intro- 


duced annuals.  BristK  cnptantha  (Cn/pfantha 
inl('rnij)fa)  was  the  onK  nati\(^  forb  found  in 
Townsends  gn)und  s(|uirrel  stomachs.  Altliough 
1988  sample  sizes  were  small,  the  importance  of 
forbs  in  the  diet  increased  in  the  samples 
between  March  and  Mav  1988,  while  the  per- 
centage of  grasses  and  shnibs  decreased  (Tables 
3-4),  thus  .suggesting  large  seasonal  differences 
between  March  and  Nhiv  diets. 

A  sm"j)rising  number  of  insects  were  eaten, 
especially  in  Ma\-june  1987  (3-19%;  Table  2). 
However,  insects  wen^  not  important  in  1988 
(trace  amounts  at  the  big  sagebrush  site  onlv). 
Insect  remains  were  so  f  ragmentarv  that  identi- 
fication was  not  usuallv  possible.  However, 
abundant  Lepidoptera  lanae  could  be  recog- 
nized bv  the  soft  e.xoskeleton  and  prolegs,  and 
fragments  recognizable  as  beetle  antennae  and 
t4\tra  were  found. 

The  importance  values  of  exotic  species 
were  lowest  at  the  unburned  big  sagebrush  site 
in  both  vears  and  highest  in  the  exotic  annual 
site  in  1987  and  at  the  native  grass  site  in  1988. 
However,  there  was  no  correlation  between  the 
importance  values  of  all  exotic  annuals  at  a  site 
and  their  importance  in  the  diet  at  that  site  (r  = 
-.454;  Tables  1-4). 

Di.sc.'us.siox 

Th(^  data  show  tliat  lor  sites  with  varviug 
degrees  of  exotic  annual  invasion  sampled  over 
a  tAvo-v(\u'  period,  Tovvnsend's  ground  scjuirrels 
can  and  do  utilize  introduced  species  in  their 
diets,  and  that  cheatgrass.  tumbleweed.  and 
tumblenuistardare  the  most  impoitantof  the.se. 

Both  the  vegetation  at  a  site  and  Tmnsend's 
ground  squirrel  diets  varied  considerably 
between  years  and  among  sites.  Differences  in 
amount  of  precipitation  most  likelv  account  lor 
the  differences  in  vegetation  importance  values 
betA\-e(Mi  vears.  There  was  less  September-Muv 
pi-ecipitation  ( 192  nun  in  1986-87  and  170  nun 
in  1987-88  at  Kuna  ca.  20  km  \).  The 
Daubenmire  (juadrats  were  taken  on  the  same 
transect  in  botli  vx^ars  bv  the  same  technicians. 

The  substantial  annual  differences  in 
Townsends  ground  scjuirrel  diets  may  be  the 
result  of  (1)  vegetation  differences  between 
vears.  (2)  the  fact  that  juveniles  were  sampled 
in  1987  and  adults  and  vearlings  were  collected 
in  1988,  (3)  differences  in  collecting  dates  (25 
Mav-19  June  1987  versus  16-19  Mav  1988).  or 
(4)  small  .sample  sizes. 


274 


Cheat  Basin  Naturalist 


[Volume  52 


TA151.F.  3.  EarK'  season  (March)  1988  Towjiscnd's  ground  sciiiirrel  diets.  Data  are  from  stomachs  of  achilt  and  yearling 
TCS  at  four  sites  in  tlie  Snake  River  Birds  of  Prey  Area.  (Ju\eniles  were  not  a\ailahle  in  March.)  Dietaiy  composition  is 
given  as  percent  relatixe  den.sitv'  (RD),  percent  fre(iuency  in  microscope  fields  (MF),  and  percent  frequencx  of  stomachs 
(PS)  containing  each  dietaiT  categoiy.  Otlier. symbols:  +  =  <!%,-  =  ab.sent,  n  =  number  of  stomachs. 


U 

nbun 

lied 

Native 

Exotic 

Rehabilital 

:ion 

big 

sagebrush 

grasses 

annuals 

seeding 

Dietary  categor\' 

RD 

MF 

PS 

RD 

MF 

PS 

RD 

MF 

PS 

RD 

MF 

PS 

11 

4 

5 

7 

16 

Gn.XSSRS 

Broiniis  tcctonini 

.39 

3.5 

75 

fi7 

4fi 

100 

51 

Ifi 

71 

39 

70 

100 

Poa  scat  11(1(1 

48 

73 

100 

IfS 

92 

100 

4 

80 

100 

39 

71 

100 

Viilpia  octoflora 

+ 

1 

25 

- 

- 

- 

- 

- 

- 

+ 

+ 

6 

Sitaiiioii  lii/strix 

- 

- 

- 

- 

- 

- 

+ 

+ 

14 

- 

- 

- 

A}ir()j)t/roii  dcscifoni  m 

- 

- 

- 

- 

- 

- 

- 

- 

- 

1 

1 

13 

Total  grasses 

87 

83 

55 

79 

SlIHlBS 

Cenitokles  Idiiata 

- 

- 

- 

- 

- 

- 

24 

55 

86 

- 

- 

- 

Artemisia  tridetitata 

11 

41 

50 

15 

35 

60 

21 

40 

100 

3 

6 

13 

Atriplcx  iiuttallii 

- 

- 

- 

- 

- 

- 

- 

- 

- 

3 

8 

19 

TottJ  shrubs 

11 

15 

45 

6 

FOHBS 

Salsola  ihcricii 

- 

- 

- 

- 

- 

- 

- 

- 

- 

- 

- 

- 

Sistjinhhtiui  alti.ssiiiiiiiii 

- 

- 

- 

1 

5 

20 

+ 

1 

14 

2 

4 

56 

Dcscuraiiiia — 2  spp. 

2 

6 

25 

- 

- 

- 

- 

- 

- 

10 

25 

75 

Raiiiiuailus  testiculatus 

- 

- 

- 

- 

- 

- 

- 

- 

- 

- 

- 

- 

Cn/ptaiitha  iiitcrnipta 

- 

- 

- 

- 

- 

- 

- 

- 

- 

2 

4 

13 

Hdlofictoii  'jKvncratiis 

- 

- 

- 

- 

- 

- 

- 

- 

- 

1 

7 

13 

Lepidiuin  pojoliatiimy 

- 

- 

- 

- 

- 

- 

- 

- 

- 

+ 

+ 

6 

Crepis  dciuniiuitdr' 

- 

- 

- 

- 

- 

- 

- 

- 

- 

+ 

+ 

6 

Lactiicd  seiTJola? 

- 

- 

- 

- 

- 

- 

- 

- 

- 

+ 

+ 

6 

Chenopodiaceae 

- 

- 

- 

- 

_ 

- 

- 

- 

- 

+ 

+ 

13 

Tot;il  forbs 

2 

1 

+ 

15 

Tlie  (liffercMice.s  in  age  elasses  were  probabK 
not  important.  Fitch  (1948)  found  no  differ- 
ence.s  in  adult  and  juxcnile  diet.s  in  California 
ground  s(|uirreLs  (.S.  hccc-licifi),  nor  did  Hansen 
and  Jolin.son  (1976)  find  difference.s  in  W\o- 
ming  groimd  squirrel  (S.  elegons)  diets  bv  sex  or 
age  class.  D\iii  and  Yen.sen  (in  pn^paration) 
found  no  dietan  diflerences  between 
adultAearling  and  ju\-enile  age  clas.ses  in  Idaho 
(S.  hnimieiis)  or  (>oIumbian  (S.  cohinihidims) 
groiuid  squirrels.  On  the  other  hand.  Hie  19SS 
data  do  show  a  strong  seasonal  component. 
Thus,  the  obsened  annual  dietan  differences 
may  be  a  result  of  later  collecting  dat(>s  in  I9<S7. 
combined  with  annual  \egetation  differences. 

Although  1  17  stomachs  were  examined,  the 
sample  sizes  were  too  small  to  draw  many  con- 
clusions about  intersite  and  between-sea.son 
diets  in  19S8.  This  amount  of  collecting  Iiad  a 
delctcrii)iis  cKect  on  local  TowTiseud's  ground 
s(|uirrel  densities,  and  we  recommend  u.se  of 
olhcr  methods  if  sea.soual  or  annual  dietan 
shihs  are  ol  iuter(\st. 


At  each  site,  several  plant  species  were  found 
in  Townisend's  groimd  scjuirrel  stomachs  that  did 
not  appear  in  the  vegetation  analysis  for  that 
site.  The  Danbenmire  (1959)  method  of  vege- 
tation anaKsis  ga\'e  an  intuiti\el\'  acceptable 
estimate  of  dominant  \egetation,  but  for  estab- 
lishing a  close  link  between  plant  abundance 
and  herbivore  diets,  a  finer-scale  method  of 
resource  analysis  is  necessan'.  Since  indi\'idual 
Towusends  groimd  squirrels  ha\e  large  home 
ranges  (mean  =  1357  m";  Smith  and  john.son 
1985)  with  a  wide  foraging  radius,  it  is  not  sur- 
prising tliat  Townsends  ground  scjuirrels  were 
eating  species  not  recorded  b\'  the  \egetation 
anaKsis.  e\en  though  the  sites  were  relatixelv 
homogeneous. 

There  was  no  correlation  between  the  total 
abundance  of  exotic  annuals  at  a  site  and  their 
imjKutance  in  the  diet.  The  number  of  plant 
species  in  the  diet  not  recorded  In  the  vegeta- 
tion anaKsis  precluded  determining  dietar)' 
preference  indices  for  Townsends  ground 
squirrels.  However,  examination  of  vegetation 


1992] 


Tow  \si:\ns  C.Horxn  Soiihukl  Diets 


275 


Txni  I  4  I ,ate  season  (Ma\)  19cSS  Towiisciurs  lironiul  scjiiirrel  diets.  Data  are  troin  stoinaclis  of  adult  and  xcarlin^ TCIS 
at  lour  sites  in  the  Snake  Ri\er  Birds  of  Pre\-  Area.  Dietar\'  composition  is  gi\en  as  percent  relative  densit\  (HD).  percent 
fre(juenc\'  in  microscope  fields  (MF),  and  percent  frecjuency  of  stomaciis  (PS)  containinsj  each  dietan  cateiiorx.  Other 
sNUibols:  +  =  <1%,  -  =  absent,  ii  =  nimiber  of  stomachs.  Site  4  Wits  not  sampled  in  19SS. 


U 

nburned 

Native 

Exotic 

big 

sagebrush 

grasses 

annuals 

Dietar\  categon 

RD 

MF 

PS 

RD 

MF 

PS 

RD 

MF 

PS 

/) 

1 

5 

4 

Chvssf.s 

P(Ui  scciindd 

.-) 

10 

100 

19 

40 

so 

- 

- 

- 

Broimts  tcctoniiii 

22 

65 

100 

35 

57 

so 

24 

.34 

75 

Vtdpia  octoflora 

29 

75 

100 

21 

3fS 

100 

6 

IS 

100 

Sifanion  lii/sth\ 

1 

5 

100 

4 

6 

40 

5 

IS 

50 

A^ropi/nui  ilcsciioniiii 

1 

5 

100 

- 

- 

- 

- 

- 

- 

(Jrass  seed 

2 

10 

100 

4 

10 

20 

2 

1 

25 

(Ji'ass  root 

- 

- 

- 

5 

13 

«) 

- 

- 

- 

TotiiJ  grasses 

57 

88 

37 

SlIIUBS 

Cvratoiclcs  hiiiafa 

- 

- 

- 

- 

- 

- 

13 

25 

50 

Alii' mis  in  fiidcntata 

- 

- 

- 

- 

- 

- 

3 

S 

50 

Aliiplcx  iiiitlallii 

3 

15 

100 

6 

14 

20 

2 

4 

25 

Clin/sotliaiumis  viscidiflnnis 

- 

- 

- 

- 

- 

- 

+ 

1 

25 

Total  shrubs 

3 

6 

18 

FOHBS 

Stdsola  ihcriai 

22 

fiO 

100 

- 

- 

- 

- 

- 

- 

Sisi/uthiiuDi  (dtissiiiuiiii 

- 

- 

- 

6 

IS 

60 

I 

4 

75 

Dcscurainia — 2  spp. 

- 

- 

- 

- 

- 

- 

15 

33 

50 

Riitiuiiadus  fcsticuhitits 

17 

55 

100 

- 

- 

- 

28 

43 

50 

Forb  root 

- 

- 

- 

+ 

1 

20 

- 

- 

- 

Total  torbs 

39 

6 

44 

Mis(:i;ll.\.\kols 

ins(^ct 

- 

- 

- 

- 

- 

- 

+ 

1 

25 

L'nknowm 

- 

- 

- 

- 

- 

- 

+ 

( 

25 

Total  miscellaneous 

0 

0 

+ 

ahuiidance  (Table  1 )  iiicoiiipari.soii  toconsiiiup- 
tioii  (Tables  2-4)  indicates  that  most  of  the 
abundant  plant  species  were  also  important  in 
the  diet,  and  that  rare  plants  were  being  used 
onK  in  trace  amounts.  There  were  some  inter- 
esting exceptions  to  this,  however.  Cheatgrass 
was  di(4anl\  im])()itant  (39%  relative  densifs) 
but  not  rccordt^l  in  the  \egetation  analwsis  at  the 
rehabilitation  site. 

l^iets  became  more  di\erse  in  Max;  probabK 
as  a  result  of  gras.ses  curing  and  setxls  becoming 
a\  ailable.  Ground  squirrels  eat  large  amounts  of 
seeds  prior  to  entering  torpor  (Rickart  1982,  E. 
Yensen,  personal  obser\ation).  Perhaps  if  insuf- 
ficient seeds  are  a\  ailable  during  a  drought  \ear, 
Townsend's  ground  s(  juirrels  turn  to  insects  as  a 
fat  source.  However,  at  the  exotic  annual  site 
w  here  insect  use  was  highest  in  19S7,  cheatgrass 
(mostK'  seeds)  was  the  major  con.stituent  of  the 
diet.  This  relationship  should  be  explored  further. 

Although  halogeton  {HaJofH'ton  fjjotncratns) 


was  not  recorded  1)\  the  xcgetalioii  anaKsis, 
small  amounts  of  it  were  found  in  iwo  stomachs 
at  the  rehabilitation  site  in  March  1988  (Table 
3).  Halogeton  is  poisonous  to  livestock,  but 
sheep  can  eat  it  with  impunitx  in  winter,  proba- 
bly because  rains  hav  e  leached  the  oxalates  out 
of  the  dricnl  leavt-s  (C]ook  1977).  Presumably. 
Townsend's  ground  scjuirrels  were  eating  dried, 
rather  than  frc^sh,  leaves  in  March. 

Idaho  and  (-olumbian  groimd  squirrels  have 
higliK  varic-d  diets  of  1 1-25  plant  species  per 
fecal  pellet  group  (D\ni  and  Yensen.  in  prepa- 
ration). However,  in  that  stud\  onl\  2— i  plant 
species  (usualK'  grasses)  contributed  >10%  to 
the  diet.  Rogers  and  Gano  (1980)  found  duit 
onl\  three  plant  species  [Poa  spp.,  Dcscurainia 
piiiiKifa.  and  Liipiniis  laxiflonts)  contributed 
>]()'/(  of  the  di(^t  of  Townsend's  ground  squir- 
rels in  southeastern  Washington.  Hansen  and 
Ueckert  (1970)  found  1-.5  species  contributed 
>10%  in  the  di\erse  (47  plant  species)  diets  of 


276 


Great  Basin  Naturalist 


[\'olume  52 


WVoming  ground  squirrels  in  Colorado.  Hansen 
and  Johnson  (1976:750)  concluded  that 

Richardson  [=\\Vomiii<i]  ground  squirrels  graze  on 
a  varied  of  pUuits  as  they  fill  tlieir  stomachs  rather 
than  selecting  onlv  preferred  foods  when  their  stom- 
achs are  nearl^•  enipt).  Tliis  may  be  an  e\olutionar\- 
strateg)-  de\eloped  to  allow  them  to  consume 
vetches.  The  dilution  of  toxic  foods  b\-  non-toxic 
foods  decreiises  the  probability-  of  plant  poisoning. 

Frecland  and  Janzen  (1974)  re\ie\\ed  strat- 
egies of  herbi\  ()r\  h\  mammals  in  response  to 
secondan-  plant  compounds.  The\-  suggested 
that  a  generalist  herbi\ore  should  feed  predom- 
inanth'  on  one  or  hvo  foods,  but  continue  to 
sample  other  foods  present.  When  an  herbixore 
experiences  a  nutritional  deficiency.  :t  should 
sample  all  a\ailable  foods  until  it  finds  some- 
thing which  supplies  that  nutrient. 

The  feeding  strategies  proposed  by  Freeland 
and  Janzen  (1974)  and  Hansen  and  Johnson 
(1976)  appear  to  occur  in  sexeral  members  of 
the  subgenus  Spennophilus.  The  data  indicate 
that  ground  squirrels  specialize  on  2-4  highly 
nutritional  species,  but  supplement  them  with  a 
wide \ariet\' of  (jther species,  apparentK  as  "poi- 
soning insurance."  In  this  study,  Townsends 
ground  squirrels  similarly  depended  on  onl\  a 
few  species  for  the  bulk  of  the  diet,  but  a  wide 
\ariet\'  of  trace  species  was  not  axailable.  If  any 
of  these  species  should  proxide  insufficient 
(juantities  of  a  key  nutrient  (e.g.,  linoleic  acid 
necessaiy  for  hibernation),  then  the  limited 
selection  of  food  species  could  ha\e  negati\e 
population  consequences. 

The  question  of  whether  Tow -nsends  ground 
squirrels  can  utilize  exotic  annuals  as  dietai-y 
staples  is  an.swered  in  the  affirmatixe  b\  this 
study.  Native  forb  species  were  of  minor  impor- 
tance in  the  diet,  but  this  does  not  necessarily 
reflect  preference.  Nati\  e  forbs  are  now  so  rare 
at  the  foin-  sites  that  none  were  recorded  b\  the 
\egetation  anal\sis,  and  thus  they  may  not  haye 
been  a\ailable  for  consumption.  Only  one  native 
forb  iCnjptantlia)  was  found  in  the  stomaciis. 
The  consequences  of  limited  dietai-y  yariet}'  on 
tlie  long-tenn  nutrition  of  Townsend's  ground 
squirrel  are  unknowni. 

.VlKXOWLEDCMKXTS 

We  tliaiik  j.  \\'ea\cr  and  other  Bureau  of 
Land  .Management  pc^rsonnel  who  collected  the 
scjuirrels  and  the  \egetation  data,  B.  DMii  and 
T  Foppe  for  discussion  of  the  microhistological 
analysis  technique,  P.  L.  Packard  for  ac'cess  to 


herbarium  specimens,  and  we  especialK"  appre- 
ciate M.  P.  Luscher's  assistance  in  preparing  the 
slides.  R.  G.  Anthony,  D.  R.  Johnson,  S.  Knick, 
E.  .\.  Rickait.  K.  Steenhof,  and  B.  Van  Home 
made  helpful  comments  on  an  earlier  draft  of 
the  manuscript. 

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Cook.  C.  W.  1977.  Effects  of  season  and  intensit\-  of  use  on 
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centage dr\'  weight  in  diets  using  a  microscopic  tech- 
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Bureau  of  Land  Management,  Boise,  hlaho.  S2  pp. 
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population  fluctuations  in  southwestern  Idaho,  .\mer- 

ican  Midland  Naturalist.  In  press. 
YouNo.  J.  A.,  R.  A.  E\ANs  R.  E.EckEin   |i;    antl  B.  L.  k\v 

1987.  Cheatgrass.  Rangelands  9:  266-270. 

Rcceiied  5  Maij  1991 
Acccjilcd  1  September  1992 


Creat  Basin  Naturalist  52(3),  pp.  278-283 


AVIFAUNA  OF  CENTRAL  TULE  X'ALLEY, 
W  ESTERN  BONNEVILLE  BASIN 


Peter  Hoxiiiiih 


Ki'ij  ivnnls:  hirds.  inifdiaia.  dcscii.  (UjUdtir  liahitiit.  Great  Ba.siii.  uctlaiid-s 


Fautiii  (1946)  (k>scril)ed  the  flora  and  fauna 
of  sewral  northern  desert  biotic  coniniunities  in 
Tule  \alle\-,  located  80  km  west  of  Delia,  Utah, 
in  Millard  Countv  of  western  Bonnexille  Basin. 
His  study  durintj;  1939  (June  to  September)  and 
1940  (.'Vpril  to  September)  included  a  descrip- 
tion of  greasewood  {Sarcohatiis  vcniiicidatiis) 
and  pickleweed  {AUenwlfea  occiclentaUs)  com- 
nnuiities.  From  1980  through  1991  while  in ven- 
torxing  the  acpiatic  habitats  of  Tule  Valley,  I 
noted  the  axifamia  utilizing  wetlands,  springs, 
adjact^nt  greasewood  and  pickleweed  commu- 
nities, and  saline  flats.  This  note  reports  on  the 
a\ifauna  oc-curring  within  the  two  communities 
and  compares  the  1980-91  faunal  lisitng  with 
that  reported  prexiously  by  Fautin  (1946).  Com- 
parisons are  also  made  with  Fish  Springs 
National  Wildlife  Refuge,  located  50  km  north 
of  the  Tule  Valle\  springs.  This  study  identifies 
changes  in  raptors  and  songbirds  that  ha\e 
occurredoxer  40 years  and  notes  the  differences 
bet\yeen  natural  springs  and  wetlands  and  those 
dedicated  to  waterfowl  management. 

Description  of  tiik  Tiii.k  \'alley 

AgiATIC  EN\  IHONMKNTS 

Within  the  greasewood  and  ])icklc>weed 
connnunities  of  central  Tule  Valle\  are  some  25 
tissure-fault  springs  and  associated  wetlands. 
Saline  flats  coxered  in  part  by  water  from  saline 
seepage  springs  occiu"  to  the  east  and  west  of 
these  fissiu-e-fault  springs.  The  springs-wetlands 
yar\-  in  si/.e  from  100  nr  to  o\er  97.000  nr 
(Coyote  Springs)  with  a  total  of  195,000  nr. 
Couductixity  oltlie  aquatic  sxstems  \aries  from 
1200  (spring  .sources)  to  greater  than  93.000 
umbos    per   cm    (some    wetlands    and    saline- 


ponds).  Tln"ee-comered  bulrush  (Sciiyiis  aincr- 
ic(inus)  and  salt  grass  [Distichlis spicata)  are  the 
dominant  emergent  species,  with  Phra<j^inifcs 
aiistralis,  Tijplui  doDiiti^cnsis,  and  Scirpiis 
aciittis  occurring  in  highl\-  localized  stands. 
Tamarisk  (Tamarix  ramosissiina)  is  the  only 
shrub  gnnxing  within  some  springs-wetlands 
but  was  not  noted  by  Fautin  (1946). 

Methods 

A  total  ot  36  \isits  were  made  to  Tule  Villex' 
bet\yeen  1980  and  1991,  with  10  \isits  of  t\yo- 
day  dvu'ations  occvniing  in  1981.  Imentories 
were  conducted  during  each  month  (except  Jan- 
uan)  with  emphasis  during  March,  Maw  and 
June.  Birds  were  inxentoried  b\  random 
encounters,  and  unidentified  species  were  not 
pursued.  Nomenclature  follows  that  of  Peterson 
(1990). 

Results  and  Discussion 

Table  1  lists  the  80  species  of  birds  identified 
during  1980-91,  the  months  they  were  encoun- 
tered, and  those  species  also  reported  b\'  Fautin 
( 1946).  Mallard  (scientific  names  noted  in  Table 
1),  Northern  Harrier,  Horned  Lark,  Connnon 
Raxen,  and  Marsh  Wren  xxere  encountered 
year-roimd  and  are  considered  permanent  resi- 
dents. Almost  half  (31)  of  the  species  inxento- 
lied  during  this  studx'  xx'ere  obsened  t\yo  or 
fex\  t'r  times  (dates  included  in  Table  1 )  and  are 
considered  casual  or  transient  xisitors.  The 
single  Palm  Warbler,  a  casual  bird  in  Utah 
( Belile  et  al.  1985),  xx'as  identified  by  its  charac- 
teristic tail  moxement  as  prexiouslxobserxedby 
me    on    numerous    occasions    durinti    iuuiual 


721  Scttmcl  Avenue.  Sail  l,iil;e  (  \{\    Tl.ili  S  ll(K5. 


278 


19921 


Notes 


279 


migrations  in  the  Midwest.  Tlic  salint-  ])()ncls 
west  of  tlie  lissnre-tanlt  springs  Iiostod  gnlls, 
nunierons  waterfowl,  and  shorehirds  dnring 
migration. 

Fifteen  species  (Cw-eat  Bine  Heron.  Turke\ 
\  ultnrc.  Sharp-shinned  Hawk,  Coopers  Hawk, 
Swainsons  Hawk,  Red-tailed  Hawk,  l-Jnrrowing 
Owl,  Conmion  Xighthawk,  Western  kingbird. 
Northern  Mockingbird,  Yellow  Warbler, 
Yellow-breasted  Chat,  Cireen-tailed  Towhee, 
Brewers  Sparrow  and  Lark  Bnnting)  obsened 
b\  Fantin  (1946)  were  not  enconntered  in  this 
stndw  Bnrrowing  Owls,  while  nesting  in  the 
adjacent  shadscale  connnnnitw  were  not 
obsened  in  the  greasewood  commnnits.  The 
absence  of  raptors  (in  particidar  the  Swainsons 
Hawk)  and  the  Turkey  \^ulture  ("onl\'  occasion- 
alK  seen,  but  obser\ed  throughout  the  summer 
ill  one  conimunit\'  or  another,  "  Fautin  1946: 
2S5)  could  reflect  the  rangeland  predator  con- 
trol programs  occurring  in  Tule  \'alle\-  since 
Fautin  did  his  studies.  Absence  of  other  species 
mentioned  abo\e  could  reflect  the  loss  of  wil- 
lows (Sdli.x exi^^iia)  which  Fautin  ( 1946:257)  had 
noticed  as  being  prexalent.  Most  of  the  birds 
Fautin  reported  for  the  greasewood  communit)' 
that  were  not  obsened  during  the  present  study 
were  considered  transients  b\^  Fautin  (1946). 

Oxer  157  species  with  41  permanent  resi- 
dents (those  species  that  can  be  found  in  all 
seasons)  and  54  nesting  species  ha\e  l)een 
reported  for  Fish  Springs  National  Wildlife 
R(4uge  (U.S.  Department  of  the  Interior  19SS). 
This  contrasts  sharpl)- with  the  axifaunaof  Tule 


\all('\.  whicli  consists  of  5  [)ermaiient  residents 
and  a  total  of  17  summer  residents.  The  larger 
number  of  species  at  Fish  Springs  National 
Wildlife  Refuge  j)r()babl\  reflects  the  availabil- 
ity of  surface  water,  tlu>  presence  of  trec^s  and 
buildings,  and  tlie  proximitA'  of  the  springs-wet- 
lands to  the  momitaiiious  Fish  Springs  Range. 
Tule  N'allev  springs-wetlands  are  nndexeloped 
and  lack  the  man-made  features.  An  additional 
factor  that  mav  contribute  to  the  (hfference  in 
a\ifauna  constitnencx  of  Tule  \alle\-  and  Fisli 
Springs  is  the  contribution  over  man\'  \ears  of 
field  ornithologists  at  Fish  Springs  National 
Wildlife  Rehige. 

Two  birds.  Western  Sandpiper  and  Lincoln's 
Sparrow,  ha\e  not  been  reported  in  this  region 
in  the  Latilong  study  (\\iilters  and  Soren.son 
1983);  and  the  Lincoln's  Sparrow  was  not 
reported  at  Fish  Springs  (U.S.  Department  of 
the  Interior  1988).  Fish  Springs  and  Tule  X'allev 
are  in  the  same  Latilong  region,  and  Fish 
Springs  olisenations  o\en\'helm  the  Tule  \'alley 
obsenations  within  the  Latilone  studw 


CONCMA'SIONS 

A  listing  of  the  axilanna  lor  central  Tule 
\"alle\  is  reported.  Comparisons  are  made  to  the 
axifauna  List  reported  b>'  Fautin  (1946)  and  to 
the  species  list  prepared  b\-  the  Fish  Springs 
National  Wildlife  Refuge.  Dillerences  in  spe- 
cies are  noted  and  explanations  are  offered. 


Tablk  1.  Distribution  of  birds  in  the  grea.sevvood-wetland  coinmunitv  of  Tide  X'alle' 


Month  of  YeiU" 


J 


M 


M 


J        J 


O       N       D 


Specific  date.s°° 


PoniC:il'KDlDAK 

Pied-billed  Crebe 

Podircps  tii'^ricoUis 
Eared  (irebe 

PodiUjinhus  an riliis 

'  \kdkidak 
American  Bittern 

Bdtaunis  lcriti<iiii(isus 

<  •rcat  Blue  Heron 

\i(lca  iwrixlias 
Sii()\\-\  E(j;ret 

l'.-n-clt(i  tliiila 
Hlack-crowned  Night  Heron 

Sijc^ticorax  tn/ciicorax 

.'hreskiohmtiiidae 

Wliite4'aced  Ibis 

Plc'atlis  chilli 


I 


8/8/81:  6/li:y82 
6/20/81 

9/29/84 

6/1.3/82 
8/18/81:  10/20/90 

8/21/87;  8/2.3/91 


280 


(tHEat  Basin  Naturalist 


[\V)lume  52 


Tablf  1.  Continued. 


Vlonth  of  Year 


M       J        J 


O       N       D 


Specific  dates" 


Anatidak 
Canada  Cioose 

Bratitd  canadensi.H 
Creen-winged  Teal 

Aucis  crecca 

°M;JIard 

Anas  j)hitt/rl)  i/tichos 
Xortliem  Pintail 

Alias  acuta 
Cinnamon  Teal 

Anas  cijanoptcra 
American  Wigeon 

Anas  aincricana 
(>an\asback 

Ai/tlu/a  valisiuciia 
Redlif-ad 

Aijtlii/a  aiiwricana 
Merganser 

Mcrt^its  sj). 
Ruddy  Dnck 

Oxyti  ra  jainaicensis 

Caihahtidaf. 
"Turkey  X'ultnre 
Catliaiics  aura 

Acc;iPirHinAK 
"Northern  Harrier 

Circus  ci/anciis 
°Sliaq>-shimied  Hawk 

Accipitcr  striatus 
"('ooper's  Hawk 

Accipitcr  coopcrii 
"Swaiiison's  Hawk 

Butco  swainsani 
°Red-t;iiled  Hawk 

Butco  jainaicensis 
Rongli-legged  Hawk 

Butco  la<i(rpus 
"CJolden  Eagle 

Acju  ila  ch  n/sactos 
"Americiui  Kestrel 

Falco  sparvcrius 
"Prmiie  Falcon 

Falco  mcxicanus 

Rai.lidak 
Virginia  Rail 

Rallus  liinicola 
Sora 

Porz/ina  Carolina 
American  C'oot 

Fiilica  aincricana 

ClIAlUDHIIDAK 

"Killdeer 
CJiaradrius  vocifcnis 

Ri-x;i;n\in()STHii)Ai-; 
Black-necked  Stilt 
Himautopus  mexicann s 


3/7/87 
4/27/81 


3/22/82 


3/7/81 
3/20/90 

9/25/82:9/29/84 
4/4/82:5/11/88 


X  \  X  \  \  \  X 


X  X  X  X 


8/2 1/S" 


1992] 

T.^BLE  l.ConHnued. 


Notes 


SCOLOPACIDAE 

.S[X5tted  Siuidpiper 
Atiitis  manddria 
Westeni  SanclpijxT 
Cdlidris  inaiiri 
niiiiliii 

Calidri-s  alpiuii 
Common  Snipe 
Gdllina^o  acdlina^o 
Lahidae 
Gulls 
Liinis  sp. 

COLUMBIDAE 

'Mourning  Do\e 
Zcnaida  inacro\ira 
Strkmdae 
"Burrowing  Owl 
Athene  ciinicuUiria 
Caprimulgidae 
"Common  Niglithawk 
Chordciles  ininor 
Apodidae 
\\'hite-throated  Swift 
Aewnautes  saxatalls 

PlCIDAE 

Xorthem  Flicker 
Colalptcs  aiiriitiis 

TVIUXNIDAE 

"Western  Kingbird 
Tijmnnus  vciiicalis 
Aluimdak 
"Homed  L;uk 
Erenwphila  alpcsths 
HllU\DIMD.\E 
\  iolet-green  Swallow 

Tarhijciiuia  thdassinti 
"Barn  Swallow 
Hinindo  ru.stica 

COHMUAK 

°{>'onimon  Ra\cn 
Corvius  corax 

Trocu.ody'iidae 
Miirsh  Wren 

Cistothnriis  palustris 

I  MUSCICAPIDAI': 

Mountain  Bluebird 
Sialia  cumicoidcs 

MiMIDAK 
"Northern  Mockingbird 

Mimiis  pahjolottos 
*Sage  Thrasher 

Orcoscoptes  inoiitanus 

ViOTACILLlDAE 

American  Pipit 
Anthus  ndwsccns 

-AM  I  DAE 

.  "Loggerhead  Shrike 
Lanius  bidoviciamis 

1 

1 


281 


Month  of  Year 


]        ^       ^       ^       ^^       ]        JA        S       O       N       D  Specific  dates" 


8/21/87 
4/20/86 
4/20/86 


XXX 


X  X  X  X 


^  X  X  X  X 


X  X  X 


X  10/25/81:12/6/81 


'^  X  X  X  X  X 


X  X  X  X  X 


8/8/81;  6/13/82 
9/19/81 


X  X  X  X  X 


X  X  X  X  X 


X  X  X  X  X  X 


X  X  X  X  X  X 


8/24/81 


X  X  X  X 


X  X  X  X 


282 

Table  1.  Continued. 


Great  Basin  Naturalist 


[Volume  52 


Montli  of'Yeiir 


J        FMAMJ        J        ASOND 


Specific  dates" ° 


Stuhmdaf. 

Starling 

Stunuts  vulffiris 
Kmbkki/idak 

"Yellow  Warbler 

Dcndroica  petechia 
"Yellow-nimped  Warbler 

Dendroiai  roroiiata 
Palm  Warbler 

Dendroica  pal  mam  in 
"Common  Yellowthroat 

GeothUjpis  trick  as 
"Yellow-breasted  Chat 

Icteria  vireiis 
"Green-tailed  Towhee 

Pipilo  chloninis 
American  Tree  Sparrow 

Spiz-(dla  arhorea 
"Brewer's  Sparrow 

Spizella  hreweri 
°\'esper  Sparrow 

Fooecefes  ^raininrii s 
Lark  spiirrow 

Cfiondcstcs  iJ^ratnmacHs 
"Black-throated  Sparrow 

Amphispiza  bilineata 
"Sage  Sparrow  j 

Amphispiza  belli 
"Lark  Bunting 

Calamospizd  melaiioconis 
Sa\annah  Sparrow 

Passerculus  sandtcichensis 
Fox  Sparrow 

Passerella  iliaca 
"Song  Sparrow 

MeU)spiza  melodia 
Lincoln's  Sparrow 

Melospiza  lincolnii 
"White-crowned  Sparrow 

Zonotrichia  leucophnjs 
Junco 

Jiinco  sp. 
"Red-winged  Blackbird 

Afifilaius  phoeniceiis 
Western  Meadowlark 

Stumella  m'^lecta 
"Yellow-headed  Blackbird 

Xanthocq)haltts  xanthocephahis 
"Brewer's  Blackbird 

Enpha'^us  cijanoeephalus 
"Brown-headed  Cowbird 

Molothnis  ater 

FKrNCll.l.lDAF, 

American  Cokhincli 

Carduelis  tristis 
Passkkidaf. 
House  Sparrow 

Passer  doinesticus 


2/21/8L3/7/81 


9/19/81 


9/16/80;  12/6/81 


9/20/81 
5/2/S7 


10/20/90 

12/5/81 

4/4/81 


12/6/81 


10/25/81 


•Identified  \n  FaiKin  (1946), 

•"Dates  in  right  c-oliiinn  are  for  t%vo  or  fewer  obser\ations. 


19921 


Notes 


283 


ACKNOW  LEDGMENTS 

1  wish  t(j  thank  Da\  id  E.  )()\  irm"  and  C."Ia\t()n 
M.  White  for  reviewing  the  niannscript  and  tor 
snhsecjiient  discussions. 

References 

Bkiii.k  W.  II..  E.  D.  .SonKNSKN  aiulC.  M.  Wmitk  1985. 
Utdi  birtls:  a  revised  checklist.  Occasional  Fuhlication 
#4.  Utah  Museum  of  Natural  Histon-,  Salt  Lake  Cit\. 
108  pp. 


F.\L  TIN.  R.  W.  1946.  Biotic  communities  of  northern  desert 

shnib  biome  in  western  Utah.  Flcolos^ical  Monographs 

16:251-,31(). 
Pi;i  KK.SON,   R.  T.    1990.  ,'\  field  guide  to  western  birds. 

Houghton  Mifflin  Co.,  Boston.  432  pp. 
U.S.  Dkpahtmknt  of  i'iik  Intkhioh    1988.  Birds  ol  the 

Fish  Springs  National  Wildlilc  Heluge.  Dug\va\',  Utah. 

RFfi-6553i-2. 
W.M.TK.HS.  R.  E..  ;uid  E.  Sokknsf.n,  f.ds   1983.  Utah  bird 

distril)ution:   Latilong  study   1983.   Utdi   Division  of 

W  ildlife  Resources  Publication  83-10.  97  pp. 

Received  10  yovcniher  imi 
Accepted  22  jiuw  1992 


Great  Basin  XatimJist  52(3),  pp.  284-287 

WILDFIRE  AND  SOIL  ORGANIC  CARBON  IN 
SAGEBRUSH-BUNCHGRASS  X'ECETATION 

Ste\'en  A.  Acker 

Kc'ij  words:  soil  organic  matter,  soil  organic  carbon,  wildfire,  big  sagebrush.  Artemisia  tridentata  wyomingensis, 
Artemisia  tridentata  tridentata,  Imnchg^rass,  long-term  site  degradation,  Oregon. 


Soil  organic  matter  is  an  important  compo- 
nent of  the  enxironment  for  plants,  one  that 
enhances  a\ailabilit\'  of  water  and  nntrients 
(Nelson  and  Sonnners  1982),  contributes  to  a 
siiitai)le  seedbed  ( Monsen  and  McArthur  1985), 
and  enhances  seedling  emergence  (Wood  et  al. 
1978K  In  the  sagebnish  region  of  the  Inter- 
movmtain  West,  loss  of  organic  matter  due  to 
recurring  wildfire  may  be  a  mechanism  of  long- 
term  site  degradation,  ultimateK'  caused  b\ 
excessixe  li\estock  grazing  and  the  introduction 
of  aggressixe  annual  plants  (West  1988).  Loss  of 
organic  matter  or  plant  co\er  due  to  fire  ma\' 
increase  erosion  and  decrease  infiltration, 
therein'  decreasing  seedbed  quality'  (Monsen 
and  McArthur  1985).  Loss  of  organic  matter 
ma\-  also  render  soils  less  friable  and  more  likel\- 
to  form  crusts  upon  drving,  and  so  increase  the 
resistance  emerging  seedlintrs  nuist  o\ercome 
(Wood  et  al.  1978).  On  the  other  hand,  it  is 
concei\able  tliat  the  increase  of  the  introduced 
annual  cheatgrass  {Bromiis  tectonim  L.)  that 
mav  follow  wildfire  (West  1988)  iua\-  increase 
soil  organic  matter  over  the  long  nm,  due  to 
litter  accumulation.  Documentation  of  the 
response  of  soil  organic  matter  to  wildfire  in  the 
sagebnish  region  is  limited.  On  relatixeK  mesic 
big  sagebrush  {Artemisia  tridentata  Nutt.)  sites, 
tlie  occurrence  of  a  single  fire  apparently  does 
n(jt  decrease  organic  matter  in  the  siuface  soil 
layers  (Nimir  and  Payne  1978,  Humph re\ 
1984).  This  study  concerns  the  effect  of  wildhre 
on  soil  organic  matter  in  relatively  xeric  big 
sagebrush  sites  (Acker  1988). 


Methods 

I  studied  soil  organic  matter  at  two  ptiirs  of 
burned  and  adjacent  unbumed  big  sagebrush- 
bunchgrass  stands  in  northern  Hamev  Countv', 
Oregon,  USA.  The  stands  were  selected  along 
with  se\en  other  pairs  for  a  stucK  of  post-wild- 
fire big  sagebnish-bunchgrass  vegetation 
dynamics  (Acker  1988).  I  selected  as  study 
stands  bunied  and  adjacent  unburned  areas  in 
which  at  least  one  of  four  climax  bunchgrass 
species  was  present  (bluebunch  wheatgrass, 
Agropyron  spicatum  [Pursh]  Scribn.  &  Smith; 
Indian  ricegrass,  Onjzopsis  Jn/menoicles  [R.  & 
S.]  Kicker;  needle-and-thread,  Stipa  comata 
Trin.  &  Rupr.;  and  Thiu'ber's  needlegrass,  Stipa 
thurheriana  Piper)  (Hironaka  et  al.  1983).  The 
climate  is  semiarid  (28.9  cm  annual  precipita- 
tion on  axerage  for  Bums,  Oregon,  about  40  km 
north  of  the  study  area),  with  hot,  dn'  summers 
and  cold  winters  (Franklin  and  D\niess  1973). 
Soils  are  stony  and  shallow  o\er  lava  or  welded 
ash  deposits,  and  are  classified  as  Lithic  Xerollic 
Haplargids  mixed  with  Lithic  Torriorthents 
(Lindsax  et  al.  1969).  Within  pairs,  the  sites  are 
similar  in  elexation,  slope,  aspect,  and  surface 
soil  texture  (Table  1 ).  Other  than  incidental  use, 
none  of  the  four  stands  was  grazed  b\  domestic 
livestock  during  this  stud\  oi"  o\er  sexeral 
decades  (M.  Armstrong,  personal  communica- 
tion). Shrub  skeletons  were  present  on  all  the 
bunied  stands.  Thus,  prior  to  the  recent  fires, 
])aired  stands  probabi)  had  similar  fire  histories. 
The  initial  wildfire  occurred  in  August  1981. 
The  stands  were  sampled  in  the  earlx'  summer 


D<-partmeiit  of  Bolanv.  Uiiivt'rsit\  ofWisconsni-Madison,  M.ulison.  Wisconsin  53706  l>ivsi-iit  .uldivss   Drpartinent  c.f  Forest  Scit-nce  Collese  of  Forestry, 
Oregon  State  Universit)-.Corvallis,  Oregon  97331. 


284 


19921 


Notes 


285 


Tablk  1.  Enxironmental.  historical,  and  \egetation  data  for  burned  (odd  numbers)  and  adjacent  unbumed  (e\en 
numbers)  bis^  sas;ebnish-bunchgrass  stands,  IIarne\-  Coinih',  Oregon,  USA.  Soil  texture  determined  b\'  method  of"  Liegel 
etal.  (1980)^ 


Stand 
number 


Kiev. 

(m) 


Aspect 
categorv"' 


Slope 


Soil  texture, 
top  10  cm 


Dominant  plant 
species  (1985)'' 


1 

1325 

9 

17 

siuidv  loam 

2 

1325 

8 

12 

sandv  loam 

3 

1360 

3 

19 

loamx-  sand 

4 

1360 

2 

22 

loam\-  sand 

BRTE,  ERFI.  POSE,  PHHO,  ORHY 
ARTRW,  PHIIO.  ASFI 
BRTE,  STC02,  CH\I 
ARTRT,  BRTE,CHNA 


"1  =  SSW;  2  =  S.SW;  3  =  SSE,VVS\V;  4  =  SE,\V;  5  =  ESE.VVNW;  fi  =  K.NW:  7  =  ENE.NNW;  S  =  NE.N;  9  =  \NE  (baseil  o.i  .Vli.lr  and  Lotaii  1985).  Categories  1^ 
are  wanii  a.spects;  categories  5-9  are  ccxil  iispects. 

Plants  vsitli  at  le;ist  3%  co\'er,  in  descending  order  .\RTRT  =  Aiiiiidxia  tridcnInUi  ssp.  triikniata.  ARTRW  =  Artcinisiu  Irideiitata  ssp.  wijomiiigcnsls .  ASFI  = 
Astragalus  tilipes;  BRTE  =  Bwmm  tectoninu  CHNA  =  Cltnjsutltamnus  naiiseosiis  ssp.  aUiicaulK;  CUVT  =  Chn/sotlwmmis  tiscUliJlortis  ssp.  visddiflimts.  ERFI  = 
Eriaenm  fihfolius:  ORIIY  =  Orijzopsis  htjmeiwidcs .  PHHO  =  Phhx  Iwodii;  POSE  =  Pua  sccuiida;  STC02  =  Stipa  coinata.  \oiiclier  specimens  on  Pile  at  University 
of  Wisconsin — .Madison  I  lerbariuin. 


Table  2.  Comparison  of  organic  carbon  in  top  10  cm  of  soil  in  burned  and  adjacent  unbunied  big  sagebrush-bunchgrass 
stands,  northern  Hamey  Count\',  Oregon,  USA.  X'alues  are  mean  percentages  of  mass  of  oven-dried  .soil  (standard  errors 
in  parentheses).  Sttmdard  errors  were  computed  using  each  stands  variiuice  for  1987  and  the  number  of  subsamples  for 
the  vear  listed  (Petersen  and  CaKin  1986).  The  number  of  degrees  of  freedom  for  all  tests  is  30  (E.  Nordheim,  personal 
communication). 


Result  of  two-tailed  t  test. 

\'ear 

Organic  carbon 

N 

burned  \s.  unbimied 

Stands  1  ; 

ind  2 

1985 

burnetl: 

1.19(0.24) 

3 

.4  >  P  >  .2,  NS^' 

unbumed: 

0.83(0.23) 

3 

1986 

burned: 

1.17(0.21) 

4 

P  >  .5,  NS 

unbumed: 

1.34(0.20) 

4 

19S7 

bumed: 

1.31  (0.10) 

16 

.4  >  F  >  .2.  NS 

unbumed: 

1.15(0.10) 

16 
Stands  3 ; 

uid  4 

19S5 

bumed: 

0.63(0.17) 

3 

P  >  .5,  NS 

unbumed: 

0.68(0.16) 

3 

19S6 

bumed: 

0.60(0.15) 

4 

P  >  .5,  NS 

unbumed: 

0.65(0.14) 

4 

1987'' 

bumed: 

0.83  (0.07) 

16 

P  >  .5,  NS 

imbumed: 

0..S4  (0.07) 

16 

■'Not  significant 
''Both  stands  3  ai 


d  4  bumed  between  the  1986  and  19S7  samplings. 


of"  1985,  1986,  and  1987.  Stands  3  and  4  hnnunl 
again  in  a  wildfire  September  1986. 

I  collected  samples  from  the  top  10  cm  of 
soil,  3  samples  per  stand  in  1985,  4  in  1986,  and 
16  in  1987.  In  the  first  two  years  sampling  l(K'a- 
tions  were  laid  out  in  a  systematic  manner.  In 
1987  samples  were  collected  in  a  stratified 
random  manner.  The  randomization  for  tlu^ 
only  remaining  unburned  stand,  stand  2,  was 
further  restricted  so  that  the  area  under  slirub 
canopies  was  sampled  roughK  in  proportion  to 
the  co\er  of  shrubs  in  the  stand.  Shnibs  can 
nfluence  spatial  patterns  of  soil  chemistry  in  big 
sagebrush  yegetation  (Doescher  et  al.  1984). 

Organic  matter  of  the  soil  samples  was 
issessed  using  the  \Valkle\- Black  rapid  dichro- 


matc  oxidation  mctliod  of  organic  carbon  deter- 
mination (Nelson  and  Sommens  1982).  I  used 
the  standard  correction  factor  of  1 .3  to  adjust  for 
organic  carbon  not  oxidized  in  the  procedure. 
Giyen  the  uncertain  (juantitatixe  relationship 
betxyeen  soil  organic  carbon  and  soil  organic 
matter,  I  report  soil  organic  carbon,  as  Nelson 
and  Sommers  recommend  (1982). 

I  used  two-tailed  /  tests  to  compare  organic 
carbon  betxxeen  [)aired  stands  ( Sokal  and  liohlf 
1981).  For  1985  and  1986  I  used  the  .sample 
\ariance  from  the  1987  obserxations  and  the 
sample  size  from  the  year  in  f|uestion  to  deter- 
mine the  denominator  ot  (lie  test  statistic 
(Petersen  and  Calyin  1986).  This  wius  done  due 
to  the  larger  sample  size  and  the  (stratified) 


286 


Cheat  Basix  Naturalist 


[Volume  52 


random  arrangement  of  the  1987  samples 
(Greig-Smith  1983).  In  the  strictest  sense,  these 
observations  can  onI\  establish  clifferenees 
between  adjacent  stands.  .Applying these  results 
to  burned  and  unburned  big  sagebrush- 
bunchgrass  stands  more  generally  is  tenuous, 
due  to  die  lack  of  replication  (Ilurlbert  1984). 

RF..SU1TS  AND  DiSCUS.SION 

For  both  pairs  of  stands  there  was  no  signif- 
icant difference  in  organic  carbon  in  the  top  10 
cm  of  soil  in  any  of  the  three  years  (Table  2). 
None  of  the  individual  comparisons  is  sugges- 
tive of  such  a  difference  (P  >  .20  in  ail  cases). 
Although  I  did  not  test  statistically  for  a  tempo- 
ral trend,  soil  oi-ganic  carbon  does  not  appear  to 
ha\e  changed  oxer  the  course  of  the  studv  in  any 
of  the  stands.  Thus,  the  recurrence  of  fire  at 
stands  3  and  4  does  not  appear  to  have  altered 
soil  organic  carbon. 

Changes  in  organic  matter  are  b\^  no  means 
the  onK'  ecologically  important  soil  changes 
wildfire  may  cause  in  big  sagebrush  vegetation 
(e.g.,  increa.se  of  organic  acids  in  burned  soil; 
Blank  and  Young  1990).  Furthermore,  the  short 
duration  and  small  sample  size  limit  the  gener- 
alitv  of  conclusions.  However,  these  stands  are 
not  unlike  others  in  the  general  \icinit)'  where 
climax  buuchgrasses  persist  (Acker  1988).  In 
addition,  tlie.se  stands  offer  a  rare  opportunit\- 
to  obsene  l)i<i  sao;ebrush-bunch'j;rass  vegeta- 
tion  processes  in  the  absence  of  lixestock  grazing. 

Wildfire  apparently  has  not  decreased  or 
increased  soil  organic  matter  on  these  stands. 
From  other  studies,  1  ha\e  concluded  that  post- 
wildfire  vegetation  dynamics  in  these  stands  and 
similar  ones  nearb\-  is  dominated  by  cheatgrass 
and  does  not  feature  increasing  abundance  of 
climax  buuchgrasses  (Acker  1988).  To  explain 
tliese  trends  may  require  invoking  sonu^liing 
other  tlian  irreversible  site  degradation,  as  indi- 
cated by  k)ss  of  soil  organic  mattcM". 

A(:k\()\\i,ki)(;.\ik\ts 

liesearch  was  si ip[)orted  by  graduate  fellow- 
ships and/or  research  grants  from  the  National 
Science  Fountlation,  the  University  of  Wiscon- 
sin Craduate  School,  the  Da\is  Fund  of  the 
UniversitAofWi.sconsiu  Departments  of  Botanv 
and  Zoologx;  and  Sigma  Xi.  I  tliank  Nhilheur 
Field  Station  and  the  Bureau  of  Land  Manage- 
ment, Bums,  Oregon,  for  logistical  support  of 


fieldw'ork,  and  Professor  Jim  Bockheim  and 
Kurt  Schulz  for  acKice  on  soil  analysis  and  access 
to  laboratoA'  facilities. 


Literature  Cited 


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and  ob.se nations.  Unpublislied dissertation,  Universitv- 
of  Wisconsin-Madison.  204  pp. 

Blank  R.  R.,  and  J.  A.  Young.  1990.  Chemical  changes  in 
the  soil  induced  bv  fire  in  a  conniinnit\'  dominated  by 
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DoEscilER.  P.  S.,  R.  F.  MiLLEK  and  A.  H.  Winwakd 
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1983.  Sagebnish-grass  habitat  tvpes  of  southern  Idaho. 
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succession  after  fire  on  Arteinisia-gvdss  sites  in  south- 
eastern Idaho.  Vegetatio  .57:  91-101. 

HlRl.BERT,  S.  11.  1984.  Pseudoreplication  and  the  tlesign  of 
ecological  field  experiments.  Ecological  Monographs 
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EiKcEi,  E.  .A...  C.  R.  SiNH)N  and  E.  E.  Scmiulte.  1980. 
Wisconsin  procedures  for  soil  testing,  phuit  anaKsis 
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Extension-Madison.  51  pp. 

LiND.sAY,  M.  G.,  B.  B.  Lo\ELi„  |.  \.  N()Rc:re\.  G.  H. 
SiMONsoN.  B.  R.  Tiio.MAS,  and  D.  W  .\ndehson. 
1969.  Malheur  Lake  Drainage  B;isin  general  soil  map 
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long-range  reijuirements  for  water  State  of  Oregon 
Water  Resources  Board,  Salem.  79  pp. 

MoNSKX  S.  B.,  and  E.  D.  McAhtulr  1985.  Factors  influ- 
encing establishment  of  seeded  broadleaf  herbs  and 
shrubs  following  fire.  Pages  112-124  //!  K.  Siuiders.  J. 
Durham,  et  al..  eds..  Rangeland  fire  effects:  a  s\nipo- 
siuni.  U.S.  Department  of  Interior.  Bukmu  of  Land 
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Mill!  P.  S.,  and  [.  E.  I^orw  1985,  Distmbancehiston  and 
scrotiuN'  of  Films  roiiloiin  in  western  Montana.  Ecol- 
ogx  66:'  1 6.58-1 66S. 

Nei.sox,  D.  W.,  and  L.  E.  .So\imi:hs  1982.  Total  carlion. 
organic  carbon,  and  organic  matter.  Pages  .539-579  in 
A.L.  Page,  R.  1 1.  Miller,  and  D.  R.  KeenW  eds..  Meth- 
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1992]                                                                   NOTKS  287 

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son,  Wisconsin.  .ind  woodliuids.  Pages  209-230  /;;  M.  C.  Biuhonr  and 

\i\ilK    \l    H..  and  Ci.  F.  Paynk    197S.  Eft'ects  of  spring  W.  D.  Billings,  eds.,  \ortli  American  terre.stri;il  vege- 

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Accepted  22  Jinic  1^2 


Great  Basin  Naturalist  52(3),  pp.  2.S8-2S9 

STRUCTURE  OF  A  WHITE-TAILED  PRAIRIE  DOG  BURROW 

Lvnii  A.  Cooke    and  Steven  R.  Svviecki" 
Kcii  words:  (  Aiioiins  Iciiciinis,  hiirnm-  stniiiiin'-  liilirnuitiilmn,  iic.st. 


Mttlc  piil)lislie(l  intonnation  is  ux'ailable  on 
the  striietuie  of  white-tailed  prairie  dog  (Cijn- 
onujs  Ictiainis)  burrows.  Clark  (1971,  1977) 
described  the  stnicture  of  two  partially  exca- 
vated burrows  in  WX-oniing,  and  Bums  et  al. 
(1989)  described  structure  and  function  of 
another  burrow  in  Montana.  Neither  of  these 
studies  reports  finding  either  hibernating  ani- 
mals or  remains  of  known  hibernators  who  died 
o\er  wint(M-.  This  note  describes  the  structure  of 
a  burrow  sxstem  in  Colorado  that  had  a  known 
histon  of  prairie  dog  use  for  two  years  prior  to 
excavation.  Burrow  excavation  was  undeitaken 
to  establish  fates  of  t\vo  juveniles  who  hiber- 
nat(nl  in  tlie  burrow  in  19(S8  but  were  not 
resightedin  19S9. 

The  excavated  burrow  is  located  on  the 
Arapaho  National  Wildlife  Refuge,  Walden, 
Colorado  (Jackson  Count),  T8N' R79W  S5). 
Dominant  shrub  species  include  greasewood 
{Sarcohatiis  vennindafus),  rabbitbrnsh  (Chn/so- 
iluniuuis  natiseosiis).  and  sagebrush  {Aiieniisia 
Ihdculala  K  Dominant  grasses  are  wheatgrasses 
(A<iropijroit  spp, ).  The  burrow  sv.stem  was  exca- 
vated by  hand  in  |mie  1989.  During  excavation 
measurements  were  taken  periodicallvof  tlepth 
and  dimejisions  of  tunnels  and  chambers. 

Four  entrances  wendocated  (A,  B,  (>',  and  D 
in  Fig.  1).  One  of  these  entrances  had  an  asso- 
ciated mound.  Remaining  entrances  opened 
into  semicircular  pits  approximateK  0.6  m  in 
diameter.  No  material  had  been  transported 
from  below  the  surface  or  from  tlie  surrounding 
surface  to  form  a  crater,  as  constructed  bv  black- 
tailed  prairie  dogs  iCi/n())tu/.s  hidoviciaiuis) 
(King  1955,  Cincotta  'l989).  All  entrances. 
except  the  mound,  were  filled  with  loose  soil. 

The  main  entrance  descended  from  one  end 


of  an  oN'al  mound  1.5  m  long,  1.2  m  wide,  and 
0.2  m  high  at  an  angle  of  70°  for  approximately 
0.5  m  and  lexeled  off  at  a  depth  of  0.4-0.5  m. 
Tunnels  connecting  entrances  measured  80- 
220  mm  High  and  80-200  mm  wide  and  were 
approximately  circular  in  cross  section.  These 
connecting  tunnels  were  all  within  0.5  m  of  the 
surface.  A  tunnel  leading  to  the  nest  chamber 
descended  further.  Turning  bavs,  as  described 
by  Scheffer  (1937)  for  black-tailed  prairie  dogs, 
were  found  near  one  entrance,  D  (Fig.  1). 

The  nest  chamber  tunnel  descended  from  an 
entrance  without  a  mound  (D  in  Fig.  1).  A  side 
tunnel  connected  to  the  mound.  After  branch- 
ing, the  tunnel  gradually  descended  to  a  maxi- 
mum depth  of  1.25  m.  Another  branch,  closer 
to  the  nest,  appeared  to  rise  and  was  not  e.xca- 
\ated  due  to  time  constraints.  The  tunnel  lead- 
ing to  the  nest  chamber  was  1 15-150  mm  wide 
and  105-225  mm  high.  In  front  of  the  nest 
chamber  were  three  small  chambers,  190-350 
mm  long  and  100-225  mm  in  diameter.  One  of 
these  chambers,  350  mm  before  the  nest  cham- 
ber, contained  old  fecal  material.  \\'hitehead 
(1927)  reported  a  feces-filled  chamber  in  a 
black-tailed  prairie  dog  burrow  and  suggested 
prairie  dogs  used  it  to  avoid  drowning.  The 
present  burrow  svstem,  however,  had  no  provi- 
sion to  trap  air  if  submerged  (Foster  1924). 
Other  chambers  near  bends  in  the  tunnel  may 
ha\e  permitted  animals  to  pass  one  another.  No 
stored  food  was  found  in  an\'  chambers. 

An  enlarged  chamber  was  located  at  the  end 
of  the  bin-row  svstem.  This  chamber  had  a 
(k)med  ceiling,  a  bowl-shaped  floor,  and  mea- 
sured 2 10  nun  high  bv  210  mm  wide  b\-25()  mm 
long,  (contained  within  the  chamlxM^  was  a  mass 
of  dry;  well-chewed  plant  material,  primarily 


Dfp.irtii)eiitofSv.vt<'nKiticsiimlEcolog\-,  Universih' of  Kansas.  Lawrciice.  Kansas  6604.5-2  KKi  Pn 
Crtifs.  WorccsUi .  M.LSs.iilinsctts  01610-2.39.5. 
•  15410  Helen.  Sontlinalc  .Miclnfrui  -18195. 


lit  address:  Department  of  Biologv.  College  ol  the  llolv 


288 


19921 


NOTKS 


289 


TOP  VIEW 


AcKXow  i.i;i)(;mk\t.s 

This  research  was  supported  in  part  by 
grants  tioiii  tlie  University  of  Kansas  General 
Research  Fund  to  K.  B.  Armitage,  the  IJnixcr- 
sit)'  of  Kansas  Department  of  Systeniatics  and 
Ecolog)',  tlie  Theodore  Roosevelt  Memorial 
Fund,  and  Sigma  Xi.  The  U.S.  Fish  and  Wildlife 
Senice  kindly  permitted  work  on  the  Ara})aho 
National  Wildlife  Refuge.  We  thank  E.  C. 
Patten  and  J.  Solberg  for  assistance  in  locating 
suitable  prairie  dog  stnd\  colonies.  The  manu- 
script was  impro\ed  b\  the  connnents  of  an 
anonymous  reviewer. 


SIDE  VIEW 


Fig.  1.  Structure  of  excavated  white-tailed  prairie  dog 
burrow.  Capital  letters  indicate  entrances  to  the  burrow 
system.  The  nest  chamber  is  indicated  h\-  a  solid  star  The 
location  of  a  feces-filled  chamber  is  indicated  h\  a  solid 
triangle.  Turning  bays  are  indicated  b\  t]>. 


grasses.  This  was  probabK-  a  nest  chamber  and 
not  a  food  storage  area  because  the  plants  found 
were  not  preferred  food  plants  (Kelso  1939, 
personal  obsenation).  Se\eral  small  out- 
pocketings  were  found  off  the  nest  chamber. 
Wliile  the  nest  chamber  and  adjacent  chambers 
and  outpocketings  superficially  resembled  a 
"maternit}^  area"  as  described  by  Burns  et  al. 
(1989),  this  burrow  had  no  known  use  as  a 
maternit)'  burrow  in  three  years  prior  to  e.xcaxa- 
tion.  It  did,  however,  resemble  deep,  permanent 
swstems  described  b\-  Egoscue  and  Frank 
(1984). 

Within  the  nest  materials  were  skeletal 
remains  and  an  eartag  of  a  subadult  female  wlio 
hibernated  in  1987  and  was  not  resighted  in 
1988.  Average  frost  depth  in  this  area  is  betsveen 
500  mm  and  1  m  (X'isher  1945),  just  abo\'e  nest 
chamber  depth.  Ju\enile  males  who  used  this 
burrow  as  a  hibemaculum  in  1988  were  not 
resighted  nor  were  their  remains  found. 


LiTER.ATURE  CiTED 


Bluns.  J.  A..  D.  L.  Ki.ATH  anil  T  W,  Clark  19.S9.  On  the 
stnicture  and  function  of  white-tailed  prairie  dog  bur- 
rows. Great  Basin  Naturalist  49:  .517-524, 

Clahk  T  W.  1971.  Notes  on  white-tailed  prairie  dog  {Ctjti- 
oiiu/s  Icuninis)  burrows.  CIreat  Basin  Naturalist  .31: 
11.5-124. 

.  1977.  EcoIogN'  and  ethologx  ol  the  \\  hitc-tailed  prai- 
rie dog  {Ci/noini/s  laicttnis).  Milwaukee  Public 
Museum  Publications  in  Hiolog\  and  (;eololog\-  .No.  .3. 
97  pp. 

ClNCxrrTA.  R.  P.  1989.  Note  on  mound  architecture  ot  the 
black-tailed  prairie  dog.  (Jreat  Basin  Naturalist  49: 
621-62.3. 

EcoscuE  H.  J.,  and  E.  S.  Fkank  1984.  Burrowing  and 
denning  habits  of  a  captive  colonv  of  the  Utah  prairie 
dog.  Great  Basin  Naturalist  44:  495-498. 

FosTEH,  B.  E.  1924.  Pro\ision  of  prmrie-dog  to  escape 
drowning  when  town  is  submerged.  |ourual  ot  Mam- 
malogN'  5:  266-268. 

Kkiso,  L.'H.  19.39.  Food  habits  of  prairie  dogs.  USD.\ 
Circular  No.  529.  15  pp. 

Kixc.  J.  A.  1955.  Social  behavior  scK'iiil  organization,  and 
population  d\namics  in  a  black-tailed  prairie  dog  towii 
in  the  lilack  Hills  of  South  Dakota.  C^ontribntions  from 
the  L;ib()rator\' o(  Wrtebratc  Biologw  No.  67.  Uni\er- 
sit\' of  Michigan,  -\mi  \rbi)r.  12.3  pp 

.ScHEFFKH.  T.  M.  19.37.  Stu<K  ol  ,1  small  ]iniirie  dog  town. 
Transactions  o(  the  Kansas  .\cadem\  ol  Science  40: 
.391 -.394. 

\|s||i  li  S.  S  1945.  Climatic  maps  of  geologic  interest. 
Bulletin  ol  the  (Jeologiciil  Societv  of  .\merica56:  71.3- 
7:56. 

Will  II  hi:  \l)  1,.  C.  1927.  .Notes  on  priiirie  dogs.  |ournal  of 
.ManmialoiA'  8:  58. 


Received  1  Matj  199} 
Accepted  15  Mot/  1992 


Great  Basin  Naturalist  52(3).  pp.  290-292 

HYBRIDS  OF  WHITE-TAILED  AND  MULE  DEER  IN  WESTERN  WYOMING 


r-        h-2 


Cliarles  E.  Kav  '    and  Edward  Boe 

Ki't/  uonis:  tcliitc-tiiilcd  ihrr.  iniilc  deer.  Odocoileus  \ir(!;inianus,  Odocoileus  hfinionus.  interspecific  hijhridiziition. 
Wi/oiniiif:^. 


Though  .successful  niatiugs  of  captive  mule 
deer  {Odocoileus  Jieinionus)  and  white-tailed 
deer  (O.  virg^inianus)  have  frequently  been  doc- 
umented (Cowan  1962,  Whitehead  1972,  Day 
1980,  Wishart  1980),  interspecific  hvhiidization 
ill  most  natural  populations  appears  to  be  rare. 
Kramer  (1973)  reported  only  10  hybrids  out  of 
()\  cr  1 7,000  deer  killed  in  Nebraska,  2  out  of  983 
deer  from  Kansas,  and  onlv  6  out  of  several 
thousand  obsenations  in  Alberta.  In  34  years  of 
fieldwoi-k  in  Arizona,  Knipe  (1977)  obseived 
onlv  8  definite  hybrids. 

In  recent  years  protein  electrophoresis  of 
serum  albumin  and  restrictive  endonuclease 
anaKsis  of  mitochonchial  deoxyribonucleic  acid 
lia\('  been  uscnl  to  characterize  gene  flow 
between  mule  and  white-tailed  deer  popula- 
tions ( McCK  mont  et  al.  1982).  Based  on  protein 
elctlioplioresis  of  201  deer  from  31  localities, 
maiiiK  in  tlie  .southwestern  states,  Derr  (1991) 
lound  little  exidence  of  nuclear  gene  introffres- 
sion  between  the  two  deer  species.  Cronin  et  al. 
(1988)  reported  that  mitochondrial  DNA  and 
.serum  albumin  appeared  to  be  distinct  between 
umle  deer  and  white-tailed  deer  throughout 
.Montana,  suggesting  that  interspecihc  gene 
flow  was  ver\'  low.  This  was  in  contrast  to  data 
from  Texas  that  showed  a  5.6%  hybridization 
rate  for  319  deer  examined  (Carr  et  al.  1986, 
Stiibbleneld  ct  al.  1986)  and  Alberta  where 
Inbridization  reportetlK  is  increasing  (Lingk^ 
1989). 

Though  whitetail-nmle  deer  hybrids  ha\e 
been  obsened  in  eastern  Wyoming  (Oceanak 
1978),  they  hav-e  not  been  prexiousK  reported 
from  western  Wyoming.  On  .several  occasions 
during  the  winter  and  spring  of  1990-91  we 


obsened  and  photographed  three  female 
h\'brid  deer  west  of  LaBarge,  VWoming,  in  the 
Green  Ri\er  Basin.  The  h\brids  were  always 
associated  with  female  mule  deer  and  fed  with 
the  mule  deer  in  sagel)nish  (Aticniisa  spp.)  hab- 
itats. The  hybrids  were  often  seen  within  a  rel- 
atively short  distance  (0.5  km)  of  willow  {Salix 
spp.)  communities  and  hayfields  along  LaBarge 
Creek,  but  we  never  obsened  the  hvbrids 
kevingon  riparian  areas,  as  whitetails  commonlv 
do  in  the  arid  West  (Wood  et  al.  1989).  Instead, 
the  Inbrids  wintered  in  open  sagebrush  with  the 
mule  deer,  where  there  was  little  hidino;  or  ther- 
mal  cover,  even  though  temperatures  of  -45  C 
or  knver  are  common  in  this  part  of  Wyoming. 

During  the  winter  and  early  spring  of  1991- 
92,  we  made  additional  obsenations  and  photo- 
graphs of  hvbrid  deer  in  the  Green  River  Basin. 
On  two  separate  occasions  we  saw  a  male  h\brid 
8  km  south  of  Big  Piney,  Wvoming,  in  an  alfalfa 
{Medicago  sativa)  field  with  approximateK-  100 
mule  deer  of  both  sexes.  We  also  made  numer- 
ous obsenations  of  hybrids  along  the  section  of 
LaBarge  Creek  where  we  obsen'ed  hvbrids  the 
prexions  xear.  But  in  1991-92  we  saw  more 
hxbrids  including  at  least  two  males,  four 
females,  and  three  fawnis.  The  three  hxbiid 
fawnis  appeared  to  follow  a  single  mule  deer  doe 
and  may  ha\e  been  triplets.  These  deer  were 
usually  obseived  with  mule  deer  and  occupied 
primarih'  nonriparian  areas  as  the  lu'brids  had 
the  prexious  \ear. 

Based  on  published  characteristics  and  mea- 
surements ((^owan  1962,  Oceanak  1978,  Dav 
1 980,  Wishart  1980),  the  deer  that  we  obsened 
appeared  to  be  first-generation  Inbrids.  The 


leuiith  of  the  ridee  on  tlieir  metatarsal  glands 


,l>|)artiiieiUofFislK-ric-saiKl\\'jkllirc.Ulali  State  L.'TiiMMsitw  l.imaij,  i:ialiS1322 
"Present  adclrcs.s:  Institute  of  Political  Kcoiioniv.  Utah 
■  Present  ailelress:  Box  26.  La  Barge.  W'yoniinji  S.3I2;3. 


290 


1992J 


Notes 


291 


was  iiitennediate between  hpical  wliitetails and 
h  pical  mule  deer,  and  the  eolor  of  the  metatar- 
sal tuftwasprimaiiK  w  liite.  Their  tails  appeared 
to  he  slifj;htl\  l()ni:;er  than  normal  whitetail  tails 
and  were  i)ro\\  n  mer<ring  to  black  on  the  dorsal 
side  and  pure  white  on  the  underside.  When 
frightened,  the  h\brids  used  a  bounding  gait 
with  orwithout  tail-flagging  t\pical  ofwhitetails. 
.\s  reported  b\-  Lingle  (1989),  the  Inbrids  did 
not  appear  to  stott  but  used  locomotion  patterns 
intermediate  between  mule  and  white-tailed 
deer.  On  all  occasions  female  Inlands  \\'ere  dom- 
inated b\'  female  mule  deer  the\  associated  with 
and  were  frecjuentK  displaced  from  feeding 
sites  h\  mule  deer. 

Kramer  (1973:298)  po.stulated  that  h\brid- 
ization  between  mule  and  white-tailed  deer  max 
be  more  frequent  where  whitetails  occm-  in  \  eiA 
small  numbers.  This  ma\'  be  true  in  western 
WAoming.  Prior  to  European  settlement,  white- 
tails  were  apparentK"  distributed  throughout 
\\\omin£[,  but  unrestricted  \ear-lon2  meat 
hunting  eliminated  them  from  mo.st  of  western 
\\\oming  b\'  the  tin"n  of  the  centur\'. 

W  hitetiiils  ha\e  been  in  the  process  of  either  reoc- 
cup\ing  fornierk'  occupied  areas  in  western  Wyo- 
ming or  rebuilding  sexerely  depressed  popuhxtions 
for  at  least  30  \ears  (Harrv  Harjii,  \\\oming  Game 
and  Fish  Department,  personal  communication, 
1991). 

Based  on  hunter  sunxns  conducti'd  through  the 
niciil  or  o\er  the  telephone  b\'  the  WVoming 
Game  and  Fish  Department,  85  whitetails  were 
killed  in  all  of  western  Wyoming  in  1974,  while 
159  were  killed  in  1989  (Harju  1991,  personal 
conniiunication).  Since  few  of  these  deer  were 
checked  In  trained  observers,  there  is  no  wav  of 
knowing  how  man\'  deer  reported  b\-  hunters  as 
w  hitetails  were  actually  hybrids. 

bi  contrast,  the  Wvoming  Range  nude  deer 
herd  that  winters  betvyeen  Big  Pine\'  and 
Fontenelle  Resenoir,  including  LaBarge 
Creek,  numbered  approximately  20,000  ani- 
mals after  the  severe  winter  of  1983-84.  Since 
then,  a  series  of  se\en  mild  winters  coupled  with 
limited  doe  hanest  allowed  this  herd  to  increase 
to  55,000  in  1990  (Harju  1991,  personal  com- 
munication). In  fi\e  \ears  of  ol)ser\ation  we  saw 
o\er  40,000  deer  in  the  Big  Pine\-La Barge 
Creek  area,  and  all  but  a  few  were  nnde  deer. 
One  was  a  tvpical  male  whitetail.  and  the  others 
were  the  Inbrids  described  abo\e. 

Though  most  of  these  nuile  deer  svmimer  in 
the  Wyoming  and  Salt  River  mountain  ranges 


60-100  km  to  the  west,  some  reside  year-long  in 
riparian  areas  on  LaBarge  Creek  and  the  (^reen 
River.  Moreover,  bv  the  November  breeding 
season  thousands  of  migrating  mule  ck'er  have 
already  returned  to  their  lower-elevation 
v\intering  areas  and  then  connnonlv  cross  the 
(xreen  River  to  winter  in  the  breaks  to  the  east. 
So  large  numbers  of  nmle  deer  occupy  tvpical 
whitetail  ri])arian  habitats  during  the  nit.  \\ith 
the  marked  chffenMice  in  their  respective  popu- 
lations, it  may  be  difhcnlt  for  white-tailed  deer 
to  find  appropriate  mates  during  the  brecnling 
season.  This  may  lead  to  a  high  hxbridization 
rate  relative  to  the  whitetail  population  as 
appears  to  be  the  case  in  western  Washington, 
where  a  remnant  population  of  (^olumbian 
white-tailed  deer  (O.  v.  Icuciinis)  is  surrounded 
b\  a  nnich  larger  population  of  black-tailed  deer 
(O.  h.  columhiatius)  and  where  18%  of  the 
whitetails  tested  possessed  blacktail  alleles  at 
two  of  three  diagnostic  loci  ((iavin  and  Nhiv 
1988). 

Acknowledgments 

We    tliank    \.    (icist    and    an    anoinnions 
reviewer  for  helpful  comments. 

LiTER.\TURE  Cited 

C:\HK.  S.  .VI.,  ,S.  W.  Bai,i,i\(;kh,  J.  N.  Dkkh  1„  11, 
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white-tailed  deer  and  mule  deer  in  west  Texas.  Pro- 
ceedings of  the  National  Ac  adennof  Science  S3:  9576- 
95S0. 

Chomn,  M.  a.,  E.  R.  \vsi;  and  D.  C.  CwiKHox  19SS. 
Genetic  relationships  between  nuile  deer  and  white- 
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Cow  AN  I.  McT.  1962.  Ihbridi/atiou  between  the  black-tail 
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D\V  C;.  1.  19S().  (Characteristics  and  measurements  of" cap- 
tive liybrid  deer  in  .'\rizona.  Southwestcn  Naturalist  25: 
434-43S. 

Dl  nii  j.  N  1991  (icnctic  interactions  between  white- 
tailed  and  mule  deer  in  the  southwestern  United 
States.  Journal  of  Wildlife  Management  55:  22.5-2.37. 

(;a\I\,  T  a.,  and  B.  M\V  19S8.  Taxonomic  status  and 
genetic  purity  of  Columbian  white-tailed  deer,  an 
endangered  subspecies,  [ounial  of  WikUile  .Manage- 
ment .52:  1-10. 

KmI'K.  T.  1977,  The  Arizona  whitetail  dvvr.  .Arizona  CJanie 
and  Fish  Department  Special  l^eport  6.  lOS  pp. 

Kn WIKR,  .A.  197.3.  Interspecific  behaxior  and  dispersion  of 
two  svmpatric  deer  species.  |oinnal  of  Wildlife  .Miui- 
agenient  37:  2S.S-.300. 

LiNCLE,  S.  19S9.  Limb  coordination  and  bocK  configuration 
in  die  fast  gaits  of  white-tailed  deer,  mule  deer  and 


292 


Great  Basin  Naturalist 


[X'olume  52 


tlieir  hybrids:  adaptix'c  significant  and  nuuiagenient 
implications.  Unpnblishccl  masters  thesis,  University 
of  Calgar);  Calgan,-,  Alberta,  Canada.  289  pp. 

McCly.mont,' R.  A.,  M.  Fenton,  and  J.  R.  Thompson 
1982.  Identification  of  cer\id  tissues  and  hybridization 
bvsenim  albumin.  Journal  of  Wildlife  Management  46: 
54()_544. 

Ot:E.-\N.\K.  C.  1978.  Two  deer  in  one.  WXoming  W'ildliie 
42(3):  24-27. 

Stubblefield.  S.  S..  R.  J.  Wahken  and  B.  R.  Ml  miiv 
1986.  I hbridi/ation  of  free-rimging  white-tailed  and 
mule  deer  in  Texiis.  Journal  of  Wildlife  Management 
.5():  688-690. 


\\i  I ITEIIEAD,  C.  J.,  Jk  1972.  A  preliminary  report  on  white- 
tailed  and  black-tailed  deer  cross-breeding  studies  in 
Tennessee.  Proceedings  of  the  Annual  Conference  o*" 
Southeastern  Association  of  Game  and  Fish  Commis- 
sions 25:  65-69. 

WisilAKT,  W.  D.  1980.  Ihbrids  of  white-tailed  and  mule 
deer  in  Alberta.  Journal  of  Mammalog\-  61:  716-720. 

Wood.  A.  K.,  R.  J.  Mackie,  and  K.  L.  Hamlin  1989.  Ecol- 
ogv'  of  sympatric  populations  of  mule  deer  and  white- 
tailed  deer  in  a  prairie  en\ironment.  Montana 
Department  of  Fish,  Wildlife,  and  Parks,  Helena.  97  pp. 

Received  17  April  1991 
Accepted  17  June  1992 


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(ISSN  0017-3614) 

GREAT  BASIN   NATURALIST  Vol  52  no  3  September  1992 

CONTENTS 

Articles 

Plant  adaptation  in  the  Great  Basin  and  Colorado  Plateau 

Jonathan  P  Comstock  and  James  R.  Ehleringer      195 

Life  history,  abundance,  and  distribution  of  Moapa  dace  {Moapa  coriacea)  .  .  . 

G.  Gary  Scoppettone,  Howard  L.  Burge,  and  Peter  L.  Tuttle      216 

Condition  models  for  wintering  Northern  Pintails  in  the  Southern  High  Plains 

Loren  M.  Smith,  Douglas  G.  Sheeley,  and  David  B.  Wester      226 

Evaluation  of  road  track  surveys  for  cougars  {Felis  concolor) 

Walter  D.  Van  Sickle  and  Frederick  G.  Lindzey      232 

Leaf  area  ratios  for  selected  rangeland  plant  species 

Mark  A.  Weltz,  Wilbert  H.  Blackburn,  and  J.  Roger  Simanton      237 

Ecology  and  management  of  medusahead  {Taeniatherum  caput-medusae  ssp. 

osperum  [Simk.]  Melderis) James  A.  Young      245 

Roost  sites  used  by  Sandhill  Crane  staging  along  the  Platte    River,  Nebraska 

Bradley  S.  Norling,  Stanley  H.  Anderson,  and  Wayne  A.  Hubert      253 

Post-Pleistocene  dispersal  in  the  Mexican  vole  {Microtus  mexicanus):  an  exam- 
ple of  an  apparent  trend  in  the  distribution  of  southwestern  mammals .  . 
Russell  Davis  and  J.  R.  Callalian      262 

Can  Townsend's  ground  squirrels  survive  on  a  diet  of  exotic  annuals? 

Eric  Yensen  and  Dana  L.  Quinney      269 

Notes 

Avifauna  of  central  Tule  Valley,  western  Bonneville  Basin 

Peter  Hovingh      278 

Wildfire  and  soil  organic  carbon  in  sagebrush-bunchgrass  vegetation 

Steven  A.  Acker      284 

Structure  of  a  white-tailed  prairie  dog  burrow 

Lynn  A.  Cooke  and  Steven  R.  Swiecki      288 

Hybrids  of  white-tailed  and  mule  deer  in  western  Wyoming 

Charles  E.  Kay  and  Edward  Boe      290 


H       E 


.     MCZ 

LIBRARY 

FLii   1  6   1/93 
HARVARD 


TjXTFVTriTSrrY 


GREAT  BASIN 


MURAUST 


VOLUME  52  NO  4  -  DECEMBER  1992 


BRIGHAM    YOUNG    UNIVERSITY 


GREAT  BASIN  NATURALIST 

Editor 

James  R.  Barnes 

290MLBM 

Brigham  Young  University 

Provo,  Utah  84602 

Associate  Editors 


Michael  A.  Bowers 

Blandv     Experimental     Farm,     University    of 

Virginia,  Box  175,  Boyce,  Virginia  22620 

JR.  Callahan 

Museum  of  Southwestern  Biology,  University  of 

New  Mexico,  Albuquerque,  New  Mexico 

Mailing  address:    Box   3140,    Hemet,   CaHfornia 

92546 

Jeanne  C.  Chambers 

USDA   Forest   Service    Research,    University   of 

Nevada-Reno,  920  Valley  Road,  Reno,  Nevada  89512 

Jeffrey  R.  Johansen 

Department  of  Biolog)',  John  Carroll  University, 

Universit)'  Heights,  Ohio  44118 

Paul  C.  Marsh 

Center  for  Environmental  Studies,  Arizona  State 

University,  Tempe,  Arizona  85287 


Brian  A.  Maurer 

Department  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 

Robert  C.  Whitmore 

Division  of  Forestry,  Box  6125,  West  Virginia  Uni- 
versity, Morgantovvni,  West  Virginia  26506-6125 


Editorial  Board.  Richard  W.  Baumann,  Chairman,  Zoology;  H.  Duane  Smith,  Zoology;  Clayton  M. 
White,  Zoologv';  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  Biological  and  Agricultural  Sciences;  Norman  A.  Darais,  Director,  University 
Pubhcations;  James  R.  Barnes,  Editor,  Great  Basin  Naturalist. 

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Copyright  ©  1992  by  Brigham  Young  University- 
Official  publication  date:  30  December  1992 


ISSN  0017-3614 
12-92  7503555 


The  Great  Basin  Naturalist 


Published  at  Pucno,  U  iaii,  hv 
Bhicham  Youxc;  Uxaehsity 

ISSN  0017-3614 


Volume  52 


December  1992 


No.  4 


Great  Basin  Naturalist  52i4i,  pp.  293-299 


WINTER  NUTRIENT  CONTENT  AND  DEER  USE  OF 
GAMBEL  OAK  TWIGS  IN  NORTH  CENTRAL  UTAH 


Rosemaiy  L.  Peiidletoii  ,  Fred  J.  WagstaFC',  and  Bruce  L.  Welch' 

Abstract. — We  examined  winter  nutritional  qualitx  ot'current-vear  bud  and  stem  tissues  from  burned  and  unburned 
stands  of  Gambel  oak  (Qiiercu.s  oambclii  Nutt.).  Nutritional  analyses  were  based  on  the  amount  of  forage  consumed  In 
wintering  mule  deer.  Deer  iLse  along  the  UtaJi  \'alle\-  foothills  a\eraged  6.2.5-10.7  cm  of  current-\ear  growth.  Of  the  tissues 
examined,  post-Hre  bud  tissue  had  the  highest  nutrient  content,  with  a  mean  of  9.51%  crude  protein,  0.19%  pliosphonis, 
and  34.0%  in  \itro  digestibilit\'.  Composite  values  (bud  +  stem)  for  unljumed  stands  were  slightK'  higher  in  cnide  protein 
and  phosphorus  and  lower  in  digcstibilitv  than  those  reported  in  pre\ious  studies.  Nutrient  values  from  bumcd  stands  were 
significantly  higher  than  those  of  unbimied  stands  for  all  three  measures.  Tannin  content  ot  the  burned-area  regrowth  was 
also  higher.  OveraO  forage  value  of  Gambel  oak  to  wintering  mule  deer  is  relatix  cl\  low . 


Krij  words:  Quercus  gambelii,  Odocoileus  h 


emionus,  /( 


(xanibel  oak  (Quercus  ^auibelii  Nutt.)  i.s  a 
\  aliiable  \'ear-round  source  of  food  and  cover  for 
many  wildlife  species,  including  deer,  elk,  big- 
horn sheep,  small  mammals,  and  a  variety  of 
birds  (Re\n()lds  et  al.  1970,  Harper  et  al.  1985, 
Tirmenstein  1988).  Because  of  its  abundance 
and  location,  oak  is  an  important  food  source  for 
w  intering  mule  deer,  pro\iding  up  to  75%  of  the 
available  winter  browse  along  the  Wasatch 
Front  (Fern-  1980).  Winter  use  of  oak  \aries 
with  location.  ])nt  it  has  been  report(xl  high  in 
some  areas  along  the  Wasatch  Front,  declining 
in  the  presence  of  more  palatable  rosaceous 
shnibs  (Smith  1952,  julaiuler  1955).  Deer  use 
in  western  North  America  ranges  from  moder- 
ate to  hea\A-  throughout  the  year  (Kufeld  et  al. 
1973  and  references  therein).  In  wint(M"  prefer- 
ence trials.  Smith  (1950)  and  Smith  and  Hub- 
bard (1954)  ranked  oak  as  7t]i  or  lii(j;h(M-  out  of 


iifrieiits.  fora<iiii'^  hcJifivioi:  iitilizafioii.  hnncse,  winter. 

17  browse  species  based  on  time  spent  biowsing 
and  plant  weight  consumed. 

Although  important  to  wintering  mule  deer 
in  terms  ol  forage  a\ailabilit\'  and  jniiatabilitx', 
oak  ranks  among  the  bottom  in  nutritional  \  alue 
(Smith  1957,  Bunderson  et  al.  1986).  Nutri- 
tional studies  report  winter  oak  browse  as  being 
low  in  essential  nutrients  and  digestibilit\' 
(Smidi  1957,  Kufeld  et  al.  1981,  Meneelv  and 
Schenmit/  1981).  Smith  and  Hubbard  (1954) 
described  oak  as  being  well  liked  but  of  low 
forage  (|ualit\-.  CiuTcntlv,  little  information  is 
a\ailal)le  on  the  nutrient  content  of  different 
portions  of  the  plant  stem  or  on  the  selection  ol 
plant  parts  b\  deer. 

The  effect  of  fire  on  the  nutritional  status  ot 
oak  brow^se  is  also  of  some  interest  to  land  man- 
agers. Fire  ma\'  proxide  an  effectixe  manage- 
nuMit  tool  for  opening  the  canopx  of  the  more 


shrub  Science.s  Lahoraton.  Iiitc: 


iiutaiii  Rcscaicli  Station.  L'SD.\  Forest  Senice,  7.3.5  North  .500  East.  Pro\().  Utali  S4606. 


293 


L 


294 


Great  Basin  Naturalist 


[Volume  52 


F'ig.  I.  Location  of  four  oakbrnsh  studv  sites  in  Utiili 
Coimtv,  Ut;ili.  PG  =  Ple;is;uit  Grove;  LB  =  Lindon-lnimed; 
\.\J  =  Lindon-nnhnmed;  IIC  =  Hobble  Creek  Ganvon. 


dense  oak  thickets  to  aHow  greater  herbaceous 
growth  (Anonymous  1966,  Dills  1970,  Hallisey 
and  Wood  1976,  Haiper  et  al.  1985).  Deer  use 
of  browse  species  has  been  found  to  increase 
following  fire  in  some  (Honi  1938,  Hallisey  and 
Wood  1976),  but  not  all  (Kufeld  1983),  cases. 
The  nutrient  content  of  some  oak  species  has 
been  reported  higher  following  burning 
( Hallisey  and  Wood  1976,  Meneely  and 
Schemnitz  1981). 

The  intent  of  this  study  was  to  provide  a 
more  accurate  assessment  of  the  nutrient  con- 
tent of  oak  forage  consumed  by  wintering  nuile 
deer  on  the  Wasatch  Front.  Specific  objectixes 
were  (1)  to  determine  what  portion  of  Gambel 
oak  twigs  was  used  by  wintering  mule  deer  in 
Utiili  Valley,  (2)  to  determine  the  percent  crude 
protein,  phosphorus,  and  in  \itro  digestibilitx  of 
terminal  buds  and  stems  of  C^ambel  oak,  and  (3) 
to  compare  xalues  f)btained  from  adjacent 
burned  and  uiiburned  stands. 


Materials  a.xd  Methods 

Deer  utilization  was  studied  at  locations  near 
Lindon,  Utali,  abo\e  Pleasant  Gro\e,  Utah,  and 
in  the  mouth  of  Hobble  C:rcek  Canyon  (Fig.  1 ). 
Vegetation  at  these  foothill  locations  consists 


primariK'  of  Gaiubel  oak  and  sagebnish  (Arte- 
misia tridentata  ssp.  vuscijana),  with  scattered 
patches  of  cliffrose  {Cowania  stanshitriana)  and 
bitterbmsh  {PumJiia  tridentata).  All  three  loca- 
tions are  heavily  used  by  wintering  mule  deer. 
In  August  1987  a  wildfire  bimied  approximately 
1270  acres  on  the  southwest-facing  slopes  above 
Orem  and  Lindon,  Utali.  Oak  present  on  the 
bum  showed  considerable  regrowth  two 
months  following  the  fire.  Two  study  sites  were 
established  at  the  Lindon  location,  one  on  the 
bum  itself,  the  other  in  the  adjacent  unburned 
vegetation.  Studv  sites  were  also  established  at 
the  Pleasant  Grove  and  Hobble  Creek  Canyon 
locations,  for  a  total  of  four  study  sites. 

Deer  utiUzation  was  determined  by  measur- 
ing the  length  of  marked  twigs  before  and  after 
browsincr.  In  November  of  1987,  679  twio;s  on 
the  Lindon  bum  site  and  660  twigs  on  the  adja- 
cent unburned  site  were  marked  with  colored 
plastic  tape.  Twigs  were  selected  from  around 
the  periphen' of  multiple  clones  to  represent  all 
directiouiil  aspects  and  a  variet}'  of  heights 
accessible  to  deer.  Twig  lengths  were  measured 
from  the  tape  to  the  end  of  the  terminal  bud.  In 
March  1988  the  twigs  were  remeasured  and  the 
number  of  centimeters  browsed  determined  for 
each  tvvig.  The  ratio  of  bud  tissue  and  tvvig  tissue 
consumed  by  deer  was  then  calculated.  The 
procedure  was  repeated  at  the  Pleasant  Grove 
and  Hobble  Creek  sites  the  following  vear, 
where  186  twigs  were  marked  and  measured  at 
each  site. 

Tvvent\-t\vo  samples  for  nutritional  analysis 
were  collected  at  mid-winter  from  12  burned 
and  10  unbumed  oak  clones  at  the  Lindon  loca- 
tion. Portions  of  each  of  the  bumed  clones  were 
fenced  in  early  November  to  ensure  availabilitv 
of  mid-winter  collection  material.  In  late  Janu- 
an'  200-300  stems  were  removed  from  each 
clone,  packed  in  snow,  and  transported  to  the 
laboratoi"v.  Twigs  were  collected  from  all  sides 
of  the  periphen*  of  each  clone  to  eliminate  pos- 
sible differences  due  to  directional  aspect.  At 
the  laboratory,  stems  from  each  clone  sample 
were  divided  into  a  1-cm  terminal  bud  portion 
and  an  adjacent  lO-cm  stem.  The  proportion  of 
current-vear  growth  sampled  (11  cm)  was 
approximately  equal  to  that  removed  bv  winter- 
ing mule  deer.  Where  stem  lengths  measured 
less  than  11  cm,  total  current-vear  growth  was 
used  in  the  analvsis.  Tvvig  diameters  at  1  and  5 
cm  from  the  tip  were  also  recorded. 

The  ensuing  44  bud  and  stem  tissue  samples 


1992]  W'lXTEK  X L  TKIEXT  CUXTENT  (JF  GAMBEL  O.VK  295 

Table  L  Sumnian-  of  deer  utilization  on  marked  twigs  of  Gambel  oak  at  four  study  sites  in  Utah  County,  Utah. 


No.  h\igs 
marked 


No.  twigs 
browsed 


Percent 
browsed 


Mean 
utilization  (cm) 


Lindon-bumed 
Lindon-unburned 
Hobble  Creek 
Pleasant  Grove 


679 
660 
186 

186 


194 
368 
112 
157 


28.6 
5.5.8 
60.2 
83.9 


10.7  ±  0.44^' 

10.7  ±  0.24 

6.3  ±  0.39 

7.7  ±  0.33 


'Mfaii  i  staiuiard  error 


were  ground  u.sing  licjuid  nitrogen  and  stored  at 
—80  C.  In  vitro  digestibility,  cnide  protein,  and 
phosphorus  were  determined  for  both  bud  and 
stem  portions.  These  three  measures  were  con- 
sidered sufficient  to  determine  overall  nutri- 
tional qualit\-  of  oak  as  the\-  are  the  nutrients 
most  commonK'  deficient  in  winter  diets  of 
range  animails  (Welch  et  al.  1986).  In  vitro 
digestibilit\'was  assessed  using  Pearsons  (1970) 
modification  of  the  Tille\-  and  Terr\'  ( 1963)  tech- 
nicjue.  This  technicjue,  while  possibK  oxeresti- 
niating  in  \i\'o  digestion  of  cell  contents  in 
tannin-containing  forages  (Robbins  et  al.  1987, 
Nastis  and  Malechek  1988),  reniiiins  the  easiest 
and  most  accurate  of  the  in  \itro  techniques 
(Nastis  and  Malechek  1988)  and  is  commonly 
employed  in  nutritiontd  studies  of  range  forages. 
Inoculum  for  the  digestion  trial  was  obtained 
irom  a  slaughter-house  steer.  The  CO^-injected 
inoculum  was  processed  within  45  minutes  of 
remoxal  h'om  the  rumen  (Milchunas  and  Baker 
1982).  Studies  have  shown  that  inocula  obtained 
from  domestic  rmuinants  can  successfulK' 
approximate  digestibilitv'  of  range  forages  to 
deer  (Palmer  and  Cowan  1979,  Welch  et  al. 
1983).  Phosphonis  and  cnide  protein  determi- 
nations were  made  at  the  Plant  and  Soil  AnaKsis 
Laboraton  at  Brigham  Young  University'.  Crude 
protein  was  based  on  Kjeldalil  nitrogen  content. 
A  Technicon  Auto  Analw.er  (Technicon  Instru- 
ment CoqD.,  Tarn  towii,  NY)  was  used  to  deter- 
mine phosphoms  content.  To  simplify 
comparisons  with  values  reported  in  the  litera- 
ture, composite  \alues  for  the  complete  11-cm 
sample  were  calculated  as  follows:  composite 
\alue  =  [I0( twig  value)  -I-  bud  value]/ll.  Bulk 
samples  made  up  of  one  twig  from  each  sampled 
clone  were  tested  for  tannin  content.  Twigs 
were  kept  frozen  at  -80  C  until  us(\  then 
ground  under  liquid  nitrogen.  Tannin  content 
for  each  bulk  sample  was  determined  at  the 
Plant  and  Soil  Analysis  Laborator\-  using 
Hagerman's  (1987)  radial  diffusion  method. 


Percentage  data  were  arcsine  transfbrm(>(l 
and  anaK'zed  using  the  General  Linear  Models 
(GLM)  routine  available  on  SAS.  The  mock'l 
used  was  a  2  x  2  factorial  design,  with  burn 
treatment  (burned,  unbumed)  and  tissue  t\pe 
(bud,  twig)  as  main  effects.  Clone  was  used  as 
the  error  term  for  the  bum  treatment  main 
effect.  Tissue  differences  were  also  examined 
separateK'  for  burned  and  imbunied  areas 
because  of  a  significant  burn  treatment  x  tissue 
interaction. 

Results 

Deer  use  at  the  Lin{k)n  sites  averaged  10.7 
cm  for  both  btmied  and  unburned  clones  (Table 
1).  Individual  twig  use  \aried  wideK;  ranging 
from  1.5  to  33  cm.  Although  mean  use  at  the  two 
Lindon  sites  was  the  same,  the  burned  area  had 
a  greater  proportion  of  small  bites  than  the 
unbumed  area  (Fig.  2).  Over  24%  of  the  bites 
were  in  the  1.5-5  cm  category  at  the  bumed  site 
as  compared  to  5.7%  in  this  category'  at  the 
unbumed  site.  Also,  a  smaller  percentage  of 
marked  twi^s  was  browsed  in  the  bumed  area 
(Table  1).  Mean  use  at  the  Pleasant  Grove  and 
Hobble  Creek  sites  during  the  milder  1988-89 
winter  was  somewhat  less  than  at  the  Lindon 
sites,  averaging  7.7  and  6.3  cm,  respecti\el\ 
(Table  1).         ' 

Residts  from  the  nutrient  anaKsis  of  sam- 
pled tissues  arc  gixcn  in  Table  2.  Main  effects 
from  the  anaKsis  of  \ariance  were  all  highly 
significant.  Post-burn  sprouts  contained  more 
crude  protein  and  phosphorus  and  were  more 
digestible  than  unbumed  samples.  Bud  tissue 
exceeded  stem  tissue  in  all  three  measures.  The 
interaction  term  was  also  highly  significant  for 
crude  protein  and  phosphoms  (p  <  .0(X)1  and 
p  =  .0021,  respecti\ely).  Runningseparateanal- 
\scs  for  bumed  and  unburned  areas  revealed 
that  the  difference  between  bud  and  stem 
viilues  was  greatest  for  post-bum  sprouts,  creat- 
ing the  significant  interaction  term.  Bud  and 


296 


Great  Basin  Naturalist 


[\'olume  52 


Lindon  burned 


Lindon  unburned 


2         ♦         6         8        10       12       14       16       18       20       22     >  23 


Hobble  Creek 


8        10       12       14       16       18       20       22     >  23 


2         4         6         8        to       12       14       16       18       20       22     >  23 


Twig  utilization  (cm) 

Fiii;.  2.  I^istrihiition  of  stem  utilization  at  Four  oaklmisli  stutK'  sites  in  Utah  Countw  Utiili. 


TaBLF.  2.  Attained  significance  \alues  from  anakses  of 
variance  for  nutrient  content  of  Cambel  oak. 


Source  of 
variation 


Cnide 
protein 


Phosphonis    Digestibility- 


Bum  treatment 

0.0001 

0.0002 

0.0001 

Tissue  tvpe 

O.OOOl 

O.OOOl 

0.0001 

B\im  X  tissue 

0.0001 

0.0021 

0.3519 

Clone 

0.0001 

0.0228 

0.0015 

twig  values  from  bunied  clones  differed  signif- 
icantly for  all  three  \ariables  (Table  3).  Bud  and 
twig  values  from  unburned  clones  differed  onl\ 
in  nitrogen  content.  Twigs  from  burned  and 
unbunied  clones  also  differed  in  appearance, 
burned  t^vigs  being  more  slender  at  1  cm  (1.<S 
mm  vs.  2.6  mm;  p  =  .0002)  and  at  5  cm  (2. 1  mm 
vs.  2.9  mm; /J  =  .0001). 

Burning  also  had  a  significant  effect  {p  = 
.0001)  on  tannin  content.  The  bud  tissue  sample 
derived  from  burned  clones  had  a  tannin  con- 


tent of  4. 1  mg  per  100  mg  plant  tissue  compared 
with  3.4  mg  for  unburned  clones.  The  stem 
tissue  sample  deri\ed  from  burned  clones  had  a 
tannin  content  of  1 .6  mg  per  100  mg  plant  tissue 
compared  with  0.7  mg  for  unbtmied  clones. 

Discussion 

Previous  reports  on  fall-winter  nutrient  con- 
tent of  Gambel  oak  twigs  from  mature  stands 
range  from  4.6%  to  5.77c  tor  crude  protein,  from 
0.09%  to  0.10%  for  phosphorus  (Smidi  1957, 
Kufeld  et  al.  19S1,  Meneelv  and  Schemnitz 
1981),  and  from  26.6%  to  40.2%  for  in  vitro 
digestibilitN-  (Kufeld  et  al.  1981,  Meneelv  and 
Schemnitz  1981).  Similar  \alues  ha\e  been 
obtained  for  other  oak  species  (MeneeK'  and 
Schenmitz  1981).  Composite  \alues  from 
unbuHK^d  stands  in  our  stiicK"  (Table  2)  are  sim- 
ilar to  prexious  results,  though  slightK"  higher  in 
crude  protein  and  phosphorus  and  slightly 
lower  in  digestibilitv  The  use  of  different  h\ig 


19921 


W'lXTEH  XlTHlENT  CONTFNT  OF  CiAMHKL  OaK 


297 


Tabll  3.  .\kaiis  ami  .staiularil  (.mtois  for  pcicfiit  cIa  matter  iiutriciit  content  oi  hud  and  twig  saniple.s  collected  from 
bunied  and  unhumed  oiik  .stands  growing  near  Lindon,  Utah.  Ix'tters  following  means  indicate  significant  difierences  (/; 
=  .0001)  between  bud  luid  twig \alues  within  bum  treatment.  Composite  \alues  =  [ l()(t\\ig \ alue)  +  bud  value]/l  1. 


Cnide  protein 


Phosphonis 


Digestibility 


Bnmed  stands 
Hud 
Twig 
Composite 

Inhuineil  stand 
Hud 
Twig 
Composite 


9.5  ±  0.36  a 

0.19  ±  ().(X)6<Sa 

34.0  ±  0.59  a 

7.5  ±  0.22  b 

0.13  ±  0.0092  b 

29.8  ±  0.70  b 

7.7  ±  0.23 

0.14-  0.0087 

30.2  ±  0.66 

6.5  :  0.09  a 

0.12  ±  0.0056  a 

26.2  ±  1.68  a 

5.7  ±  0.10  b 

0.11  ±  0.00S6a 

23.5  ±  1.15  a 

5.8  ±0.10 

0.11  ±  0.0081 

23.7  ±  1.16 

leiigth.s  tor  nutrient  anal\  si.s  did  not  affect  the 
overall  results  significantK.  Composite  \alues 
based  on  t\vig  lengths  of  7.7  or  6.3  cm  (mean 
deer  utilization  at  Pleasant  Grove  and  Hobble 
Creek)  differed  \er\'  little  from  those  based  on 
the  11 -cm  sample.  In  the  absence  of  foliage, 
buds  proxide  the  highest  source  of  nutrients 
during  late  fall  and  winter.  X'tilues  for  lea\es  of 
Gambel  oak  are  substantially  higher  in  summer, 
but  comparable  to  bud  tissue  bv  mid-winter 
(Unless  et  al.  1975,  Meneelv  and  Schemnitz 
1981,  Welch  et  al.  1983,  Austin  and  Uniess 
1985). 

Bimiing  had  a  significant  influence  on  the 
nutrient  content  of  oak  forage,  particularK  bud 
tissue,  \alues  obtained  from  bunied  stands 
were  relatively  higher  than  those  obtained  from 
unbunied  stands.  Post-bum  bud  tissue  had  bv 
tar  the  highest  nutritional  \alue  of  anv  tissues 
e.xamined.  Increased  nutrient  content  of  forage 
following  fire  has  been  reported  elsewhere, 
although  indications  are  that  such  an  increase  is 
teniporaiy(  Dills  1970,  Hallisey  and  Wood  1976. 
Meneeh-  and  Schemnitz  1981,  DeB\le  et  al. 
1989).  The  evidence  to  date  suggests  that 
improved  phosphorus  content  of  forage  due  to 
fire  is  fairlv  short-lixcd.  usualK  lasting  onl\  one 
year.  Nitrogen  benefits  nia\  last  longer,  depend- 
ing on  the  species  and  season.  The  higher  values 
reported  for  burned  stands  in  this  stud\-  likelv 
occurred  because  sampling  took  place  less  than 
one  Near  following  the  burn. 

Despite  the  slightK'  highei-  \alucs  lor  ciude 
protein  and  phosphorus  reported  here  lor 
unbunied  stands  and  the  markedl\-  higher 
xalues  for  recenth'  bunied  stands,  the  oxiMall 
winter  nutiitional  \"alue  of  Gambel  oak  remains 
relatively  low.  E\en  post-bm-n  bud  tissue,  which 
had  the  highest  nutrient  content  of  anv  of  the 


tissues  sampled,  had  a  crude  protein  content  of 
less  than  10%.  The  composite  \alue  from 
burned  clones  was  less  than  8%,  whicli  ranks 
below  that  of  sagebrush,  aspen,  and  rosaceous 
.shrubs  (Smith  19.57,  Kufeld  et  al.  1981,  DeByle 
et  al.  1989).  Actual  amounts  of  protein  digested 
ina\-  be  somewhat  less  than  predicted  b\-  the  in 
\  itro  technique.  Tannins  present  in  summer  oak 
and  other  forages  have  been  found  to  increase 
the  feciil  excretion  of  protein  by  domestic  live- 
stock (Robbins  et  al.  1987).  In  mule  deer  and 
other  browsers,  nitrogen  excretion  mav  be 
reduced  1)\'  tannin-binding  proteins  present  in 
the  sali\a  (Robbins  et  al  1987).  \Miiter  digest- 
ibilit\'  of  (xambel  oak  is  also  low  when  compared 
to  other  forages.  Bunderson  et  al.  (1986)  ranked 
digestibilit\'  of  winter  oak  forage  25th  out  of  27 
species  tested.  Our  results  are  similar  in  digest- 
ibilits'  to  that  listed  in  the  ranking,  showing 
sliiihtlv  liidier  di";estiblit\' lor  burned  stands  and 
lower  digestibilit)^  for  unbunied  stands. 

Deer  use  of  oak  depends  on  man\  factors 
including  cover,  exposure,  densitv'  of  oakbnish, 
and  a\-ailabilit\-  of  other  forages  (Smith  1952, 
Inlander  1955,  Kufeld  1983.  .Austin  and  Uniess 
1985).  Fire  affects  the  stnicture  of  xegetation 
and  cover  as  well  as  (juantitx  and  (jualitv  of 
forage  produced  (Horn  1938,  Hallisey  and 
Wood  1976,  Meneely  and  Schemnitz  1981, 
Kufeld  1983).  Whether  or  not  deer  use  an  area 
more  after  bimiing  appears  to  depend  on  struc- 
ture of  the  oak  comnumitx'  and  t\pe  of  vegeta- 
tion present  on  adjacent  areas,  as  well  as 
iutensitx  and  size  of  bum.  W'here  oak  stands 
form  impenetrable  thickets,  or  where  litde 
understoiv  is  available,  burning  has  resulted  in 
increased  use  bv  deer  (Honi  1938.  Hallisev  and 
Wood  1976).  In  contrast,  Kufeld  (1983)  found 
increased  use  b\'  elk  but  not  deer  following 


298 


Great  Basin  Naturalist 


[\  olunie  52 


burning.  Vegetation  at  this  Colorado  location 
consisted  of  a  mixture  of  mature  oak  stands, 
sagebrush,  snowbern-  (Sijiuphoricarpos  alhiis), 
chokecherrv'  {Pntniis  vir^iiiidita),  and  senice- 
bern-  {Aniclancliier  ainifolia).  Burning  elimi- 
nated big  sagebrush  plants  and  decreased 
production  of  several  other  important  browse 
species,  partially  as  a  result  of  abnormally  dr\' 
weather  conditions. 

We  found  no  evidence  for  increased  use  on 
the  Lindon  burned  site  (Table  1).  The  mean 
number  of  centimeters  browsed  at  the  burned 
site  was  identical  to  that  of  adjacent  unburned 
stands,  even  though  twigs  (sprouts)  from  the 
burned  stands  tended  to  be  longer.  Also,  a  lower 
percentage  of  marked  twigs  was  browsed  at  the 
burned  site.  The  apparently  lower  use  of  burned 
twigs  by  deer  despite  higher  nutrient  content 
may  be  due  to  several  factors.  Oak  stands  in  the 
area  form  discrete  clones  rather  than  large 
impenetrable  thickets.  Important  browse  spe- 
cies such  as  sagebrush  and  bitterbrush  present 
on  unburned  areas  were  lost  as  a  result  of  the 
fire.  Also,  a  lack  of  cover  and  increased  tannin 
content  of  forage  on  the  bum  may  have  had 
some  effect  on  deer  preference. 

The  oakbrush  zone  is  critical  to  wintering 
deer  populations  along  the  Wasatch  Front. 
Although  not  the  most  preferred  winter  food,  its 
protective  cover  and  sheer  abvmdance  make  it 
one  of  the  most  widely  used  (Smidi  1949,  Smith 
and  Hubbard  1954.'julander  1955).  Current 
emphasis  in  the  Intermountain  region  is  to 
manage  the  oakbrush  zone  primarily  for  wildlife 
(Winward  1985).  Several  management  tools 
have  been  suggested,  including  fire  (Haiper  et 
al.  1985,  Winward  1985).  Burning  may  result  in 
a  temporary'  improvement  in  nutritional  quality, 
as  well  as  opening  the  canopy  sufficiently  to 
allow  establishment  of  other  shrub  and  forb 
species.  However,  without  some  form  of  ibllow- 
up  treatment,  the  proliferation  of  oak  sprouts 
may  ultimately  result  in  denser,  less  useable  oak 
forage  and  reduction  of  understor\'  species 
(Harper  et  al.  1985,  Stevens  and  Da\is  1985, 
Winward  1985).  Moreover,  the  loss  of  fire- 
susceptible  browse  species  such  as  big  sage- 
brush, mountain  niiiliogany,  and  l)itteri)rush 
may  have  serious  consequences  for  wintering 
mule  deer  (Riggs  et  al.  1990),  outweighing  anv 
possible  benefit. 


ACKNO\yLEDGMEXTS 

Thanks  are  extended  to  Len  Caipenter, 
Steve  Monsen,  and  Art  Tiedemann  for  sugges- 
tions on  research  design;  to  Warren  Claiv,  Philip 
Urness,  Robert  Ferguson,  and  an  anonvmous 
reviewer  for  comments  on  the  manuscript;  and 
to  John  Allen  and  the  Pleasant  Grove  Ranger 
District  for  access  to  the  Lindon  bum  site. 


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Received  9  March  1992 
Accepted  18  October  1992 


Great  Basin  Naturalist  52(4),  pp.  3()()->3().S 

BOTANICAL  CONTENT  OF  BLACK-TAILED  JACKRABBIT 
DIETS  ON  SEMIDESERT  RANGELAND 

Tcliouassi  Wansi',  Rex  D.  Pieper"'  ,  Reklon  F.  Beck",  and  Leigh  W.  Murray 

Abstract. — Botanical  content  of  black-tailed  jackrabbit  diets  was  determined  by  niicrohistological  examination  of  fecal 
samples  collected  from  si.\  different  vegetation  tyjies  in  sontheni  New  Mexico  on  three  dates.  Grasses  comprised  the  largest 
component  of  the  jackrabi)it  diets,  with  dropseed  species  (Sporohohis  spp.)  and  black  grama  (Bontcloua  eriopoda)  die  most 
abundant  grasses  in  tiie  diets.  Leatherweedcroton  {Crotoii  pottsii)  and  siJverleaf  nightshade  (So/c/jn/zu  t'laeagnifolium)  were 
important  lorbs  on  most  vegetation  t)pes.  Diet  composition  vaiied  in  response  to  season  and  vegetation  t\pe.  Grasses  were 
important  during  the  sinnmer  growing  season,  wliile  forbs  were  selected  during  their  growing  season  (summer  or 
winter-spring).  Sln-ubs  were  less  abmidant  in  the  diet  than  grasses  and  forbs. 

Krtj  uonis:  inicroliisloloiiicdl  iniali/sis.  fecal  inuili/sis.  Lepus  califomicus. 


Black-tailed  jackrahbits  (Lepus  califomicus} 
are  widely  distributed  in  western  and  central 
North  America.  Thev  range  from  Canada  sonth- 
ward  to  the  states  of  Sonora  and  (chihuahua, 
Mexico,  and  from  the  Pacific  coast  eastwaixl  to 
the  Great  Plains  (Hansen  and  Flinders  1969). 
Because  of  this  wide  distribution,  jackrabbits 
encounter  a  \ariet\'  of  potential  food  sources 
(McAdoo  and  Young  1980).  Considerable  work 
has  been  conducted  on  food  habits  of  the  black- 
tailed  jackrabbits,  especially  in  Arizona,  Colo- 
rado, and  the  Great  Plains  (Arnold  1942,  Reigel 
1942,  Lechleitner  1958,  Sparks  1968,  Hans-en 
and  Flinders  1969,  Flinders  and  Hansen  1972, 
Uresk  1978,  Fagerstone  et  al.  1980,  Johnson  and 
Anderson  1984).  These  studies  show  that  jack- 
rabbits are  opportunistic  feeders,  varying  their 
diets  depending  on  available  forage. 

In  .spite  of  the  relatively  large  number  of 
pnblications  reporting  the  feeding  habits  of 
black-tailed  jackrabbits,  few  have  been  con- 
ducted in  New  Mexico  and  the  Soutliwest. 
Dabo  et  al.  (1982)  found  jackrabbit  diets  were 
composed  of  many  species,  but  only  a  few  spe- 
cies of  grasses  and  forbs  form(>d  die  bulk  of  die 
diet.  They  found  Uiat  diets,  inlerred  from  fecal 
analysis,  differed  among  habitats  for  jackrabbits 
during  Slimmer  and  fall.  In  c()ntra,st,  Fatchi  et 
al.  (1988)  found  similar  diets  amoue  habitats  on 


similar  raugeland.  The  present  stud\'  represents 
a  continuation  of  earlier  studies  and  should  add 
to  understanding  seasonal  and  \earl\  fluctua- 
tions in  diets  of  black-tailed  jackrabbits. 

Study  Area 

The  study  was  conducted  on  the  New 
Mexico  State  Universitv  College  Ranch  about 
40  km  north  of  Las  Ciiices,  New  Mexico.  The 
ranch  lies  on  the  Joniada  Plain  between  the  San 
Andres  Mountains  and  the  Rio  Grande  at  an 
elevation  of  about  1300  m  (Wood  1969,  Valen- 
tine 1970).  The  climate  of  the  Jornada  Plain  is 
semiarid,  with  a  vearh'  mean  temperature  of 
about  16  C.  Mean  monthly  temperatures  are 
highest  in  June  (35°)  and  lowest  in  Januan-  ( 13°). 
Average  annual  precipitation  is  32  cm  (range 
9.2-36.2  cm),  of  which  about  509f  falls  during 
Jiil\-,  August,  and  September  (Paulsen  and  Ares 
1962). 

Fecal  pellets  from  black-tailed  jackrabbits 
were  collected  from  six  vegetation  t\pes  (habi- 
tats): (1)  mesquite  (Prosopis  glandulosa)  grass, 
(2)  snakeweed  (Guticnvzia  sarothrac),  (3) 
mixed  shrub-grass,  (4)  black  grama,  (5) 
creosotebush  [Larrca  tridoitata),  and  (6)  tar- 
bush  [Flourciisia  ccnuia).  These  \egetation 
t\'pes  are  characteristic  of  destMt  grassland  and 


jDepiutiiK'ntof  Aiiimal;ui(l  K^uii^c  .Sdciiccs,  New  \U-x\m  Slate  rniwiNih,  UusCnurs.  New  McMcdSNllKv  I'lvviil  a^lllr(■s^:  Sec-tor  tor  Ij 
■Departim-nl  of  Animal  and  Kaimc  ScieiK-cs.  New  Mcxic-o  Stale  I'liiversitv,  Las  Cnices.  \e\\  Mevieo  SMK).! 
J  Author  to  wliiiMicorrisponilente  should  l)eati(ln'S.sc(L 
Dcparlim-nt  <>l'  Kxpcrimcntal  Statistics,  New  .Mexico  State  Uni\ersit\ .  Las  Cniccs,  New  .\h\ieo  SSOI):; 


[■stock,  Me/aiii.(iiiiiei 


300 


19921 


Bl.U.KTAlLKD  |A(;K1UBB1  r  DlKTS 


301 


desert  shnihlaiuls  (Huinjilin'x  1958).  Major 
grass  species  include  black  ij;iaiiia  {Boiiteloitd 
criopodd),  mesa  dropseed  (Sporohohis 
ficxuosus),  tliiffgrass  {Ehoiicuro)i  piilchelltini), 
and  threeawais  {Aristida  spp.).  Abundant  torbs 
include  leatlienveed  croton  {Crotoii  pottsii), 
\\()()I\  paperllower  {PsilostropJw  l(i^cti)uie), 
siKerleaf  nightshade  {Solanmii  elaeagiiifoliiim), 
and  other  species.  Shnibs  inchide  mesfjuite, 
creosotebush,  and  tarbusii. 


Methods 


)ackrabbit  lecal  material  was  collected  innn 
each  \egetational  tvpe  in  June,  August,  and 
October'  19SS.  The  sample  consisted  of  15-20 
pellets  collected  randomh'  on  each  date  and  in 
each  ol  t\vo  replications  of  each  vegetational 
tvpe.  Fresh  pellets  were  identified  b\'  their  shim 
appearance.  Field  obsenations  indicated  that 
pellets  lost  their  shim  appearance  within  a  week 
of  deposition.  The  pellets  were  dried  and 
ground  to  pass  through  a  1.0-mm  screen  in  a 
Wiley  mill.  The  gromid  material  was  prepared 
as  described  bv  Bear  and  Hansen  (1966)  and 
Holechek  (1982).  Five  microscopic  slides  were 
prepared  from  each  sample,  and  20  random 
fields  were  read  from  each  slide  (Holechek  and 
Wura  1981 ).  Individual  plant  species  were  iden- 
tified by  comparison  with  known  reference 
slides.  All  identifications  were  made  bv  the 
senior  author  with  an  accuracy  of  94%.  Calcula- 
tions of  percent  composition  bv  weight  were 
made  following  procedures  outlined  b\- 
I  lolechek  and  Gross  ( 1982 ). 

Microhistological  examination  of  fecal  mate- 
rial has  some  limitations  in  diet  evaluations 
(Holechek  et  al.  1982).  Problems  are  related  to 
differential  digestion  of  different  species 
(Sidahmed  et  al.  1981),  differential  detection 
and  recognition  under  a  microscope  (Westobv 
et  al.  1976),  and  differential  particle  siz(^  reduc- 
tion (Crocker  1959).  In  spite  of  thest'  limita- 
tions, fecal  anaKsis  is  one  of  the  main  methods 
for  quantifying  diet  composition  of  w  ide-rang- 
ing  herbivores. 

Statisticiil  anal\ses  of  dietan'  data  were 
based  on  species  counts  using  a  .split-plot,  com- 
pleteK'  randomized  design  with  \egctational 
type  as  the  whole  plot  and  sampling  date  as  the 
split-plot.  Differences  among  tyjDes,  periods,  and 
the  interaction  were  analvzed  using  a  categorical 
modeling  procedure  (Proc  Catmod,  SAS  Insti- 


tute 1985).  Proc  Catmod  is  a  program  for  ana- 
1\ zing  relati\e  frequenc)-  data  by  chi-,s(|uare  tests. 
I  lerbage  standing  crop  (an  estimate  of  herb- 
age availabilit}')  was  det(>rmined  by  clipping 
herbaceous  species  from  ten  0.5  x  1.0-m  (juad- 
rats,  located  randomlv  in  each  of  the  two  repli- 
cations within  each  xegetational  tvpe,  at  the 
time  the  fecal  material  was  collected.  Herbage 
was  separated  bv  species,  oven-dried  (70  C), 
and  weighed.  Shrub  biomass  was  determined 
for  the  major  species  bv  dimension  analvsis  as 
described  bv  Ludvvig  et  al.  (1975).  Preference 
indices  were  calculated  as  the  ratio  between  the 
amount  each  species  contributed  to  the  diet 
div  ided  bv  the  composition  in  the  standing  crop 
(Kiaieger  1972).  Onlv  tho.se  prefenMice  indices 
greater  than  2  are  reported  in  this  papei-  to 
indicate  those  species  with  a  relativt'lv  high 
degree  of  preference. 

Results 

Herbage  Availabilitv 

Grasses  contributed  more  tlian  liallOf  the 
herbaceous  standing  crop  onlv  on  the  black 
grama  tvpe  (Fig.  1).  GeneralK  grass  composi- 
tion increased  from  June  to  .August,  except  on 
the  creosotebush  t>pe.  Summer  is  th(>  major 
growth  period  for  the  C4  perenniiil  grass  species 
in  this  area  (Pieper  and  Herbel  1982).  Forbs 
contributed  more  than  50%  to  the  plant  stand- 
ing crop  on  the  mes(|uite-grass,  black  grama, 
and  snakeweed  t)pes  (Fig.  1).  Shrubs  were 
abundant  (contributing  about  2()9f  of  tlu>  stand- 
ing crop)  on  the  creosotebush,  taibush,  and 
mixed  shrub-grass  tvpes. 

Diet  Composition 

Seasonal  changes  in  jackrabbit  diets 
appeared  to  be  greater  than  standing  crop  a\  ail- 
al)ilit\  for  grasses,  forbs,  and  shrubs  (Fig.  2, 
Table  1).  Generallv,  grass  co)it(Mit  of  the  diet 
peaked  in  August  and  declined  until  October 
(Fig.  2).  Fori)  content  of  the  diet  changed  little 
sea.sonallv  for  jiellets  collected  on  the  tari)ush. 
creosotebush,  and  snakeweed  tvpes.  Forbs 
comprised  a  larger  percentage  of  the  diet  in 
|une  and  October  than  in  .August  on  the  mes- 
(juite-grass  and  black  grama  tvpes.  Shrubs  gen- 
erallv contributed  less  than  25%  of  the  diet, 
except  for  pellets  collected  from  shrubby  tvpes 
at  certain  dates  (e.g.,  October  on  the  mesquite- 
grass  tvpe,  October  on  the  snakeweed  t)pe. 


302 


Great  Basin  Naturalist 


[Volume  52 


Creosotebush 


Snakeweed  Type 


August 


October 


Tarbush  Type 


Mixed  Shrub-Grass  Type 


August 


October 


August 


October 


Mesquite-Grass  Type 


Black  Grama  Type 


August 


October 


June 


June 

Fio;.  1.  Staiuliiiucropori^rasscs.  lorl.s.  and  slinilvs  on  ditTeivnt  \v<j;c-tati()n  types. 


August 


October 


June  aud  October  on  the  creosotebush  and  tar- 
bush t\pes). 

Table  2  shows  the  vegc^tation  t)pe  x  date 
interaction  \Vc\s  significant  (P  <  .05)  for  sexeral 
species.  This  interaction  indicates  these  species 
cUd  not  constitute  a  similar  pcMcentage  of  the 
diet  from  June  to  October  on  the  diflercMit  veg- 
etation types. 

DietaiT  content  of  dropseeds  varied  signili- 


cantly  (P  <  .01)  among  seasons  and  vegetation 
t\pes,  and  the  xegetation  t\pe  x  date  interac- 
tion N\as  also  significant  (Table  2).  Dropseed 
content  of  the  diet  was  highest  in  pellets  col- 
lected from  the  mixed  shrub  t\pe  and  lowest 
from  those  collected  in  the  tarbush  t\pe.  In 
some  t\pes  dropseed  content  of  the  diet  was 
highest  in  |une  (e.g.,  mesquite-grass  andsmike- 
weed  t\pes),  while  in  others  (e.g.,  black  grama 


1992] 


BLACK-'IAH.KD  jACKRAl^Hl  r  Diiri's 


303 


June 


Creosotebush  Type 


August 


October 


June 


Snakeweed  Type 


August 


October 


June 


Tarbush  Type 


August 


October 


June 


Mixed  Shrub-Grass 


August 


October 


Mesquite-Grass  Type 


June 


August 


October 


June 


Black  Grama  Type 


August 


October 


Fig.  2.  Dietan  coiiteiit  of  grasses,  forhs.  and  slinihs  in  |ii'llcts  collected  from  different  \egetation  t\pes. 

and    tarhiisli)     it    was     highest     in     Octohcr.  txpcs  the  (lilTcrcnfc  aniono;  clat(>s  was  rclatixcK 

Dropseed  content  of  pellets  collected  from  the  small  (e.g.,  mi.\ed  shnih  t\pe;  Tahle  1 ).  Th(\se 

creosotebush   t\pe  was  consistent   from   June  inconsistencies  contributed   to  the  significant 

through  October  (Table  1).  vegetational  t\pe  x  date  int(>raction  (P  <  .01; 

Black  grama  content  of  pellets  was  not  dif-  Table  2). 
ferent  (F  >  .10)  among  \egetational  t\pes,  but  13ietar\  content  of  Ihiffgrass  and  threeawn 

was  different  among  dates  (P  <  .10;  Table  2).  In  grasses  was  generalK'  low  (Table  1 ).  I  hmcx  er, 

most  cases  black  grama  content  of  the  diet  Ihiffgrass  contributed  more  than  227f  of  the  diet 

peaked  in  August,  but  for  some  vegetational  in  [une  on  the  black  grama  t\pe  and  more  than 


304 


Cheat  Basin  Naturalist 


[Volume  52 


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Black-tmu'I)  1  ACKiuHHiT  Diets 


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306 


Great  Basin  Naturalist 


[\^olunie  52 


Table  3.  Month  and  xcgetation  t\pe  wIkmi  pivlcrciice  iiulfx  exceeded  2.0  for  all  species  in  hlack-tailetl  jackrahhit  diets 
on  six  vegetational  t\pes. 


\tget; 

itional 

Type 

Species 

Black  grama 

Mesquite-Grass 

Mi.xed  Shi-ul>-C: 

Irass 

Snakeweed 

Creosotebush 

Tarbush 

Dropseed 
Black  grama 

June 

June,  Aug.,  Oct. 

June 
Aug. 

Oct. 

Oct. 

Fluffgrass 

June.  Oct. 

Oct. 

June 

June 

June 

Abert's 
hnckwheat 

Oct. 

|une,  Aug.,  Oct. 

Aug. 

Snakeweed 

June 

Desert  hailexa 

Oct. 

Dcvsert  hollv 

Aug. 

June.  Oct. 

Aug.,  Oct. 

Oct. 

Dwari  dalea 

Oct. 

Fendler's 
hladdei'iiod 

Aug..  Oct. 

[inie 

June 

Gloheniallow 

Aug..  Oct. 

Aug. 

June,  Oct. 

June,  Aug. 

Ihinenopappns 

|une 

Leathenvced 
c  rot  on 

June,  Aug. 

Aug.,  Oct. 

June,  Aug. 

June,  Aug. 

June 

Rattlesnake  wet 

'd     June 

Oct. 

SiKerleaf 
nightshade 

June,  Aug. 

Aug. 

[une,  Aug.,  Oct. 

June,  Aug. 

Speetacle]:)od 

Oct. 

Oct. 

WooK 
])aperno\\'er 

Oct. 

June,  Oct. 

June 

June 

Mcsqnite 

June 

Aug. 

Yucca 

June,  Aug. 

12%  in  June  and  August  on  the  mixed  shrub 
t\pe,  and  in  June  on  the  snakeweed  t\rpe  (Table 
1).  Threeavvns  C()ntri]:)nted  l(\ss  than  9%  of  the 
diet  on  all  dates  and  vegetational  tvpes. 

Other  grass  speeies  made  small  contribu- 
tions to  the  diet.  Plains  bristlegrass  {Setarid 
l('UCO])il(i),  vine  mesfjuite  {Paniaiui  (^hliisin)i). 
and  burrograss  (Sclen)])0(^on  hrevijoliiis)  did 
not  differ  in  diets  {P  >  .10)  among  vegetational 
t\pes  or  dates,  and  the  vegetational  type  x  date 
interaction  was  not  significant  (Table  2). 

Forb  content  of  jackrabbit  diets  varied  over 
time  and  vegetation  t\])e.  For  example,  the  con- 
tent ol  leatlienveed  eroton  differed  significantK 
(P  <  .01 )  among  vegetation  tvpes  and  dates,  and 
the  vegetational  type  x  date  interaction  also  was 
significant  (P  <  .01;  Table  2).  Its  content  \arietl 
from  about  24%  in  pellets  collect(>d  dnring 
Augu.st  in  the  tarbush  tyj^e  to  none  in  the  mixed 
shrub  txpe  at  the  same  tinK\  LeatJKMAveed 
eroton  appeared  to  be  an  important  component 


of  the  diet  on  the  black  grama,  mesquite-grass, 
and  snakeweed  tvpes  during  most  seasons 
(Table  1).  Dietarv  content  of  other  forbs  was 
inconsistent  among  \  egetational  t\pes  and  dates 
(Table  1).  Russian  thistle  {Salsola  ihcrica)  was 
the  onK  forb  species  with  a  nonsignificant  (P  > 
.10)  \egetational  t\pe  x  date  interaction  (Table  2). 
Shrub  content  of  the  jackrabbit  diets  was  also 
inconsistent  among  vegetational  tvpes  and 
dates.  Mescjnite  contributed  substantially  to  the 
diets  on  most  vegetational  txpes  between  June 
and  October.  Mescjuite  constituted  more  than 
24%  of  the  diets  on  the  snakeweed  t\pe  in 
October,  bnt  onI\  1%  on  the  creosotebush  t\pe 
in  August  (Table  1 ).  Yucca  {Yucca  t'/^/fr?)  contrib- 
uted more  than  7%  of  the  diet  from  the 
creosotebush  t\pe  in  June,  but  was  not  found  in 
j)ellets  collected  from  the  snakeweed  t\pe  on 
any  date  (Table  1).  However,  several  shrubby 
species  did  not  show  a  significant  (F  >  .10) 
vegetational  t\pe  x  date  interaction  (cnicifi.xion 


19921 


Black  TAUJ::!)  jACKiivBiiir  Die rs 


30' 


thoni  [Kochcrlinia  spindsa].  creosotcbiish.  zin- 
nia [Zinnia  (icco.sa].  and  ephetlra  [Ephedra  spp.  ]). 

Dit'tan-  Preference 

The  preference  index  was  generalK  helow  2 
for  most  grass  species  (Table  3).  However,  jack- 
labbits  apparentK'  preferred  black  grama  on  all 
dates  in  the  mesquite-grass  tyjje.  Flnffgrass  was 
preferred  dining  some  months  on  all  t)pes, 
except  for  tlie  mixed  shrub-grass  txpe.  The 
preference  index  exceeded  2  for  flnffgrass  in 
June  on  four  of  the  vegetational  t\pes. 

The  preference  index  exceeded  2  for  se\'eral 
forb  species  (Table  3).  Those  with  a  preference 
index  exceeding  2  for  more  than  six  combina- 
tions of  \egetational  t\pe  and  dates  included 
desert  hollv  {Perezia  nana),  fendler  bladdeipod 
[Lcsquerella  femUeri),  gloliemallow  {Sphaer- 
alcea  spp.),  leathen\eed  croton,  and  siherleaf 
nightshade.  I3warf  dalea  [Dalea  nana)  was  pre- 
ferred onlv  in  October  in  the  black  grama  tvpe. 
Dabo  et  al.  (1982)  found  dalea  was  highlv  pre- 
ferred and  comprised  as  much  as  65%  of  the 
diets  in  the  fall  on  grassland  vegetational  t\pes. 
Mes(juite  and  \ucca  showed  a  preference  index 
abo\ t'  2  for  June  and  August  on  three  \egeta- 
tional  txpes  (Table  3). 

Discussion 

I3lack-tailed  jackrabbits  in  southern  New 
Mexico  appear  to  i^e  opportunistic  feeders. 
Although  this  stud\'  and  earlier  ones  indicate 
that  as  man\-  as  30  plant  species  can  be  found  in 
fecal  samples  at  an\'  one  time.  5  or  6  species 
generalK'  made  up  the  l)ulk  of  the  diet.  Forbs 
often  contribute  a  greater  proportion  of  the  diet 
than  grasses,  but  the  important  forb  species  \  ar\ 
considerablx  among  locations,  seasons,  and 
\ears.  Leathenxeed  croton  is  perhaps  tlu^  main- 
sta\'  of  the  diet  auKjng  the  forbs,  although  se\- 
eral  others,  such  as  silverleaf  nightshade  and 
wooK'  paperflower  {Psilostrophc  ta<^ctinae), 
contribute  substantial  amounts  to  the  diets. 

Dropseed,  black  grama,  and  tlulfgrass 
appear  to  he  the  major  grass  species.  (,'ontrar\ 
to  cattle,  which  utilize  black  grama  niaiiiK 
during  the  dormant  season  ( I^osiere  et  al.  1975. 
l^odriguez  et  al.  1978),  jackrabbits  appai"eiill\ 
consume  more  black  grama  during  the  sunnner 
growing  season.  Consecjuentlw  high  jackrabbit 
densities  could  reduce  the  amount  of  black 
grama  a\ailable  for  cattle  later  in  the  \ear. 

Mesquite  appears  to  be  the  main  shiiibb) 


sp(^ci(^s  in  the  diets,  although  preference  for 
mes(juite  was  not  high.  Othei-  important  shiaibs 
xaried  c()nsiderai)l\  o\ci-  time  and  space. 

AcKXow  i,Ki)(;\ii'.\is 

This  is  [ournal  Article  No.  15f")()  of  the  New 
Mexico  Agricultural  E.xperimenl  .Station,  l.as 
Cruces. 

LlTER.\TUHF,  C:iTi;i) 

.Aj^NOLD,  |.  V.  1942.  I'^oraiieconsuiiiption  aiRlprclciciiccol 
experiiiRMitulK  ii'd  .-Xrizoiia  aiul  antelope  jackiahhit.s. 
Uni\ersit\()i  .Arizona  .Agriciiltnral  Experiment  Station 
Bulletin  98.  SB  pp. 

Be.AR.  G.  D..  .\ND  H.  M  IIwsf.N  1966.  Food  haliits. 
growth  and  reprodiietion  ol  white  tailed  jaekrahhit.s  in 
.southern  Colorado.  Colorado  State  Uni\ersit\  .Agrieul- 
tiu-al  E.xperinient  Station  Technical  Bulletin  90.  .59  pp. 

Ciux.KER.  B.  H.  1959.  Coasnniption  of  forage  by  hlack- 
tailed  jackrahhit.s  on  .salt-de.sert  ranges  oi  New  Mexico. 
Journal  of  Wildlife  Management  30:  .3()4-.'31 1. 

D.\BO,  S.  M.,  R.  D.  PiEPER,  R.  F.  Beck,  and  G.  M.  Soltii- 
WARD  1982.  Summer  and  fall  diets  of  black-t;iiled 
jackrabbits  on  semidesert  rangeland.  New  Mexico 
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Fa(;erstone.  K.  A.,  G.  K.  L.\\()iE,  and  K.  E.  Grifkitii, 
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F.-MEHl.  .M..  R.  D.  PlEPER.  AND  R.  F.  Beck.  1988.  SciLsonal 
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Flinders,  f.  T,  and  R.  M.  Hansen.  1972.  Diets  and  hab- 
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I  lolECllECK,  |.  L.  19S2.  Sample  preparation  technitjues  for 
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IIoi.ECHEK,  J.  L.,  AND  B.  D.  Gross  1982.  Evaluation  of 
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IIoi.ECHEK,  J.  L..  ;lnd  .M.  \a\  R\  1981.  The  eiVect  of  slide 
and  irecjuencx  of  obsen  ation  numbers  on  the  precision 
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IIoi.ECHEK,  J.  L..  M.  V.u  ra,  and  R.  D.  Pieper  1982. 
Botanicid  composition  determination  of  range  herbi- 
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iliMiMIREV.  R.  R.  19.58.  The  desert  grassland.  Botanical 
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Johnson.  R.  D.,  and  J.  E.  .Anderson  19S4.  Diets  of 
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Kkiecer  W.  C.  1972.  Exaluating  animal  forage  prefer- 
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the  black-tailed  jackrabbits.  Aniericaii  Midland  Natn- 
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LiiDwic;.  J.  A.,  J.  F.  Reynolds,  and  P.  D.  W'iiitson  1975. 
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P.\ULSEN.  H.  A.,  Jk  .  AND  F.  N.  Ares.  1962.  Grazing  values 
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Bulletin  1270.56  pp. 

PiEi'EH,  R.  D.,AND  C.  H.  Herbei  1982.  Herbage  dynam- 
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Experiment  Station  Bulletin  695.  43  pp. 

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RosiERE,  R.  E.,  R.  F.  Beck,  and  J.  D.  Wallace.  1975. 
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KooNG,  and  S.  R.  R^dosevich,  1981.  Precision  of 
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Sparks.  D'.  R.  1968.  Diet  of  black-tailed  jackrabbits  on 
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Valentine,  K.  A.  1970.  Influence  of  grazing  intensity^  on 
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Mexico  State  Universih'  Agricultural  Experiment  Sta- 
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Westobv,  M.,  G.  R.  Rost,  and  J.  A.  Weis  1976.  Problems 
with  estimating  herbivore  diets  by  microscopic;illv 
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Mammalogy  57:  167-172. 

Wood,  J.  E.  1969.  Rodent  populations  and  their  impact  on 
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Experiment  Station  Bulletin  555. 


Received  9  April  1991 
Accepted  ISJulij  1992 


Crcat  Basin  Naliirdist.  52(4),  pp.  3()^)-ol2 

SPECIES  OF  EIMERIA  FROM  THE  TIIIRTEEN-LINED  GROl  \D  SQUIRREL. 
SPERMOrHlLlS  TIUDECEMLIXEATUS,  FROM  W  YOMIXC; 

Rohcrt  S.  Scxillc   ".  l^ianc  M.  Tlionias  ,  Hiisscll  Pickcriiiij;  .  and  \aiic\  1..  Stanton 

Ans'iHACT. — Fi\e  spt'tirs  ol  tlic  coccidiaii  iiciius  I'.iiurvid  (/■,'.  hceclicyi  [prevalence  =  17.9%],  E.  cdllospcniiopliili- 
iiioraiiicD.sis  [28.6%  ],  E.  lariincrcnsis  [  16. 1%  ].  and E.  hilainclldla  [3.6%] )  were  reco\erecl  from  56, 13-lined ground  squirrels 
(SpcrDiopIjihis  tridicemlimtaUis)  collected  from  t\\()  sites  in  eastern  Wyoming.  Two  s(|uirr<^Is  from  one  site  were  also  passing 
an  unidentified  poKsporocvstic  coccidian.  Infected  scjuirrels  were  found  to  harbor  from  one  to  three  species  simuItaneousK'. 
J'rtviousK  these  sanie  eimerian  species  were  found  infecting  sympatric  populations  of  Wxoming  ground  squirrels  (Spi'r- 
m<>j)liiliis  clc'^diis)  and  \\  hite-tailed  prairie  dogs  (Ci/rioiui/s  Iciininis)  at  one  nl  the  sites;  it  is  suggested  that  the  exchange  of 
these  generalist  parasite  species  among  co-occurring  sciurid  hosts  contrihiites  to  the  consistent  prex'alence  lex'els  reported 
ill  NWoiiiinti  ground  squirrels. 

Kci/  uonl.s:  Eimeria.  Spermophilus  tridecemlineatus,  prevalence,  peihjsporoctjstic  eoceidia. 


Shults  ct  al.  (1990)  reported  the  occurrence 
of  six  .species  of  eimerian  parasites  (Protozoa: 
Apiconiplexa)  in  sympatric  populations  of  Wyo- 
ming ground  squirrels  (Spennopliilii.s  eh'<iaiis 
clegans  Kennicott,  1863)  and  white-tailed  prai- 
rie dogs  (Cynomys  leucunis  Merriam,  1890) 
from  Wyoming.  Stanton  et  al.  (1992)  conducted 
a  studN'  of  eimerian  species  in  four  XWoming 
ground  squirrel  populations  and  found  that 
most  infected  ground  scjuirrels  liarhored  two  or 
more  species  and  that  the  eimerian  assemblage 
was  present  across  populations  and  oxer  >'ears. 

Toft  (1986)  recognized  Iano  classes  of  para- 
sites: micro-  and  macroparasites.  Macropara- 
sites  (e.g.,  helminths)  tend  to  produce 
long-lasting  infections  and  are  endemic  in  host 
populations,  while  microparasites  (protozoa, 
hacteria,  \inises)  prochice  short-lixed  inf(^ctions 
and  long-lasting  inununits',  resulting  in  oscilla- 
tions of  infection  frequence"  (epidemics)  within 
the  host  population.  The  stabilit\'  for  intestinal 
piotozoans  reported  1)\'  Stanton  et  al.  (1992) 
does  not  support  Toft's  prediction  regarding 
microparasites.  While  then"  ha\e  been  no 
mechanisms  propo.sed  for  maintaining  stabilit\ 
in  microparasite  communities.  Stock  and 
Holmes  (1987)  proposed  that  species  richness 
of  intestinal  lielmintli  connnunities  of  grebes 
was  enhanced  b)'  reduced  host  specificit\  which 


allowed  parasite  exchange  among  related  hosts. 
One  important  factor  in  maintaining  the  stabil- 
it\"  of  eimerian  assemblages  is  exchange  of  par- 
asite species  among  closel\-  related  sxnipatric 
host  species. 

The  puipose  ol  (his  stud\  was  to  determine" 
which  eimerian  species  are  present  in  wild  pop- 
ulations of  13-lined  ground  s([uirrels  (Sjut- 
niopltihis  tridecemlineatus  Mitchill,  1821)  and 
to  assess  the  role  these  hosts  pla\'  in  mainte- 
nance of  the  stable  eimerian  guild  obsened  in 
W\()ming  ground  s(|uirrels. 

MKTIIOD.S 

In  1991  we  sampled  13-lined  ground  scjuir- 
nds  from  two  locations:  (1 )  a  natixe  short-grass 
prairie/ha\field  10  km  south  of  Laramie.  W\-o- 
ming  (4LI2'\,  105°33'W).  and  (2)  a  native 
short-grass  prairie/ha\ field  18  km  .south  of  (-il- 
lette,  Wyoming  (44°r7'N.  1()5°31'W). 

At  the  Laramie  site  scjuirrels  wvyc  li\e- 
trapped  using  National  lixe-traps  once  a  month 
from  |uK  to  September.  0\er  the  four-da\  trap- 
ping period  scjuirrels  were  trajijoed  using  three 
60  X  42-m  tra{)ping  grids  with  traj)s  .set  e\en-  6 
m  (162  total  traj)s).  Trai)s  were  set  at  2000  hr 
and  checked  each  morning  by  0800  hr. 

.\t  the  (wllette  site,  six  4()()-m  transc^cts  and 


, Department  of Zoolog)^  and  Ph\ .siology ,  Box  .3166,  University  ofWyoming,  Uiramie,  Wyominj;  USA  8207r 

"Present  address:  Ontario  Ministn'  of  Natural  Resources.  Wildlife  Branch.  Box  5000.  Maple.  Ontario  L6A  IS9,  C:anada 


309 


310 


Great  Basin  Naturalist 


[\  olunie  52 


Table  1.  Total  percent  infected  and  pie\alences  (hosts  infected  vvdth  given  species/liosts  examined)  of  eimerian  species 
in  gronnd-dvvelling  scinrid  hosts  at  Laramie  and  Gillette  collection  sites  in  Wyoming  (%  inf  =  total  percent  infected  with 
FAmcrio:  Elbe  =  £.  hcccluyi;  Eibi  =  K  hihiuuihild:  iMca-mo  =  E.  rallospcnnaphili-inorainciisi.s:  Eila  =  E.  hihincrciisis: 
and  Eisp  =  E.  spermophili). 


Sciurid  host 

%inf 

Eibe 

Eibi 

Eica-mo 

Eila 

Eisp 

Spennophilns 

tridcccinlineatiis 

Laramie  (n  =  41) 

43.9 

7.3 

2.4 

9.6 

14.6 

0 

Gillette  {ii  =  15) 

86.7 

46.7 

6.7 

80.0 

20.0 

0 

Total  (n  =  56) 

51.8 

17.9 

3.6 

28.6 

16.1 

0 

S.  cli-ffnts'^ 

Laramie  (/;  =  1007) 

68.0 

34.0 

11.0 

43.0 

17.0 

5.0 

Ctpiomy.s  Icncurus ' 

Laramie  (n  =  IS) 

94.0 

83.0 

17.0 

22.0 

0 

0 

'Prrceiitagcs  for  S.  clcfiam  ami  C.  Iciin 
''From  Shults  et  al.  1990. 


(U'tcTiTiineil  li\  takine;  liigliest  of  two  sallies  for  E  ivIlnspcnnDpliili  or  E 


f 


^ 


Fig.  1.  Photomicrograph  of  a  polvsporocvstic  coccidiim 
collected  from  a  13-lined  ground  scjuirrel  ( 1250X  Nomarsky 
interference)  showing  t\pical  sporoc\st  with  residuum 
clearK  xisiblc. 


six  50-111  transects  in  \arions  vegetation  t\pes 
were  trapped  dnriiig  the  second  week  in  August 
(3870  trap-nights).  Stations  were  15  m  apait, 
eacli  consisting  oi'one  \ 'ic-tor  rat  trap,  two  Victor 
mouse  traps,  and  one  Sliernian  H\e-trap.  Sher- 
man traps  remained  closed  during  daxHght 
hours,  and  traps  were  c-hecked  and  reset  at  dawn 
and  (hisk. 

All  fecal  samples  collected  from  animals  at 
both  sites  were  placed  in  2%  potassium  dicliro- 
mate  solution  at  room  temperature  (25  C)  for  at 
least  three  weeks  to  allow  oocyst  sporulation  for 
species  identification.  Ooc\sts  were  isolated  1)\ 
flotation  in  saturated  sucrose  solution  (specific 
gra\it)'  =  1.2)  and  identified  at  lOOX  objective 


with  an  Olvmpus  (CH)  microscope.  Identifica- 
tion to  species  in  most  cases  could  be  accom- 
plished based  on  oocyst  size  and  external  and 
internal  moiphologv.  Howexer,  for  Eimeria  callo- 
spennophili  Henn;  1932  and  E.  inorainencsis 
Torbett  et  al.  1982,  the  respective  size  ranges 
overlap  making  identification  dependent  on 
internal  moipholog)'.  Unfortunately,  rarely  do 
all  oocvsts  in  a  fecal  sample  sponilate.  Therefore, 
although  both  species  were  identified,  the  two 
are  combined  into  a  single  species  complex,  E. 
callospcnnopluli-inorainoisis. 

Comparisons  of  total  percent  infected  and 
prevalences  of  each  species  between  the  two 
sites  were  made  using  chi-square  tests  (Number 
Cruncher  Statistical  System  \ersion  5.03; 
Hintze  1990). 

Results 

ForK'-one  13-liiied  groiuid  scjuirrels  were 
sampled  at  the  Laramie  site  and  15  at  the  Gil- 
lette site.  Five  species  of  Eimeria  were  found 
infecting  scjuirrels  in  both  populations.  Overall, 
51.8%  of  all  squirrels  examined  were  infected 
with  at  least  one  species  of  Eimeria.  The  total 
percent  infected  was  significantK  higher  at  the 
(Gillette  (86.7%)  than  at  the  Laramie  site 
(43.9%;  P  <  .05).  Infected  squirrels  at  Gillette 
also  had  higher  parasite  species  richness  (1.77 
species/infected  squirrel)  than  at  Laramie 
(1.17).  Total  percent  infected  and  pre\'alences 
by  species  at  each  site  are  presented  in  Table  1. 

Overall,  the  Eimeria  eallospermophiU- 
moraineiisis  complex  was  the  most  prexalent 
species  found,  infecting  28.6%  of  the  56  hosts 
examined.     SisnificantK'     more     hosts     were 


19921 


EiMEiUA  FKUM  13-Lim-:d  Groum)  Sgi  iukkls 


311 


infected  with  this  species  complex  at  the  CTillette 
than  the  Laramie  site  (8()9f  \s.  9.67r ;  P  <  .05). 

Ehiwrid  bccchciji  HemA;  1932  was  tlie 
second  most  pre\alent  species  fonncl.  inlectintj; 
1 7.9%  ot  the  hosts  examined.  SignificantK'  more 
hosts  were  infected  at  the  Gillette  site  (46.7% 
vs.  7.3%;  F<. 05). 

Eimeria  larinwrcnsis  N'etterling,  1964  was 
fonnd  infecting  16.1%  of  the  scjuirrels  exam- 
ined. Prexaleiice  was  higher  at  the  Gillette  site 
(20%  \s.  14.6%),  but  the  difference  was  not 
significant  (P<. 05). 

Eiinerio  bilamellata  was  the  least  common 
species  found  dming  the  stnd\'  (3.6%).  Again, 
prevalence  was  higher  at  the  Gillette  site  (6.7% 
\s.  2.4%),  but  the  difference  was  not  significant 
(F<.05). 

Two  squirrels  at  the  Laramie  site  were  also 
infected  with  a  subspherical  poKsporocwstic 
coccidian  (Fig.  1)  with  10-12  sporoc\sts.  The 
number  ot  sporozoites  could  not  be  determined 
due  to  the  large  amount  of  residuum  present  in 
the  sporocvsts.  Mean  size  for  15  measured 
oocNsts  was  38.62  x  30.20  |jl.  Sporocvsts  w^ere 
spherical  and  measured  10.65  x  10.65  |jl  (/j  = 
15)  and  had  no  steida  bod\'.  Both  oocvsts  and 
sporocwsts  contained  numerous  residual  bodies. 
Attempts  to  infect  t\v()  captix  e  WVoming  ground 
S(juirrels  {SpcrDtophilus  dedans)  were  unsuc- 
cessful. 

DISCUS.SION 

The  occurrence  of  E.  beccJiet/i,  E.  hihi- 
iiwllata,  and  E.  morainciisis  in  13-lined  ground 
scjuirrels  constitutes  new  host  records  for  these 
species  in  this  host.  Polysporocystic  oocysts  have 
not  been  pre\iouslv  reported  from  sciurid 
rodents.  Levine  et  al.  ( 1955)  identified  two poK- 
sporocystic  species,  Klossia  perf)Iexens  fiom 
deer  mice  (Peromysais  maniadatiis)  and  K. 
variabilis  from  the  western  big  eared  bat 
iConjiiorliiiuis  rafinesc/iiii)  collected  at  the 
Grand  CauNon,  Arizona.  Becau.se  all  species  of 
Klossia  previousK'  described  were  found  in 
inxertebrates,  Levine  et  al.  (1955)  postulated 
tliat  the  t^vo  species  were  parasites  of  in\erte- 
brates  eaten  b\-  the  deer  mouse  and  bat.  Dornev 
(1965)  reported  finding  tvvo  poKsporocwstic 
oocysts  in  feces  from  a  woodchuck  (Mannota 
inonax)  from  Pennsylvania  that  resembled  the 
descriptions  of  the  two  species  in  the  genus 
Klossia  reported  b\  Le\  ine  et  al.  ( 1955).  Donie\' 
speculated  that  the  two  oocysts  might  represent 


spurious  infections  of  in\  (Mtebrate  origin.  Based 
on  thes(^  reports,  it  is  likcK  that  the  poK- 
sj)()roc\stic  coccidian  ol)ser\-ecl  in  13-lined 
scjuirrels  is  a  member  of  the  genus  Klossia  and 
possibly  of  invertebrate  origin.  Howexer.  iden- 
tification to  species  recjuires  further  woik. 
including  the  identification  of  the  priman  host. 

The  results  of  this  study  indicatc_'  that  while 
the  eimerian  fauna  of  13-lined  ground  scjuirrels 
is  \eiy  similar  to  that  of  \V\oming  ground  scjuir- 
rels and  white-tailed  prairie  dogs,  at  the  Lara- 
mie site  there  were  some  differences  in  the 
prevalences  of  the  different  parasites.  Of  the 
fi\e  species  found  infecting  13-lined  scjuirrels. 
all  ha\e  been  reported  previousK  from  svmj)at- 
ric  ground  squirrels  (Shults  et  al.  1990,  Stanton 
et  al.  1992),  and  all  have  been  reported  from 
w  liite-tailed  prairie  dogs  in  WVoming  (Todd  and 
Hammond  1968a,  1968b,  Todd  et  al.  1968, 
Shults  et  al.  1990).  However,  at  the  Laramie  site 
13-lined  scjuirrels  were  not  as  frecjuentlv 
infected  and  had  lower  prevalences  than  Wvo- 
ming  ground  squirrels  for  all  species  and  lower 
prevalences  than  white-tailed  j)rairie  dogs  for  E. 
beeclieyi,  E.  callospcrniopliili-inorainciisis,  and 
E.  bilamellata.  Values  for  13-lined  squirrels  at 
the  Gillette  site  (where  no  other  species  of 
sciinids  were  present)  were  more  similar  to 
those  forWvoming  grovmd  scjuirrels  at  the  Lar- 
amie site  (Table  1).  Additionallv.  W'voming 
ground  squirrels  had  greater  species  richness 
than  13-lined  squirrels  (Stanton  et  al.  1992). 
Species  richness  for  prairie  dogs  has  not  been 
reported. 

Results  indicate  that  related  sv  nipatric  hosts 
can  be  infected  by  the  same  species  of  Eimeria, 
which  mav  contribute  to  the  stabilitA   of  the 


eimerian  guild. 


ACKNOW  Li:i)(;.\lENTS 

This  resc^arcli  was  suj)j)c)rted  in  part  bv  the 
Dej)artnient  of  Zoologv  and  Phvsiologv  and  the 
Office  of  Research,  Universit\ of  W'voming,  and 
NSF  Grant  #BSR-8909887.  ' 

LiTKHATlHK  GlTED 

DoKM  V  H.  S.  1965.  Eimeria  tuscarorensis  n.  sp.  (Protozoa: 
EiiiK-iiiilae)  and  rede.scriptions  of  other  c-occidia  of  the 
w(K)dchuck,  Marmota  monax.  Journal  of  Protozoologv' 
12:  42.3-426. 

I.I  \  INE,  N.  D.,  V.  I\'KNs.  .WD  F.J.  Kkuidkmkh  1955.  Two 
new  species  of  Klossia  (Spirozoa:  Adeleidae)  from  a 
deer  mouse  and  a  hat.  Journal  of  Parasitology-  41: 
623-629. 


312 


Great  Basin  Naturalist 


[Volume  52 


SiiULTS.  L.  M.,  R.  S.  Skvillk.  N.  L.  Stanton,  and  G.  E. 
Mf.nckf.ns.  Jr  1990.  Einwria  sp.  (Apicomple.va: 
Eiineriidae)  from  Wyoming  ground  .s(juirrel.s  {Spcr- 
tnophilus  clegam)  and  white-tailed  prairii>  dogs  {Cijn- 
oinijsleiiciint.s)  inWVoniing.  Great  Basin  Naturalist5(): 
327-331. 

Stanton.  N.  L..  L.  M.  SiiiiLTs,  M.  R\hkek.  and  R.  S. 
Sf.\ti,lk  1992.  Coccidian  assemblages  in  the  Wyo- 
mingground  squirrel,  S})enn()])liiliis  elc'^diis.  Journal  of 
Par;Lsitol()_g\- 78:323-,328. 

Stock,  T.  M.,  and  J.  C.  Holmks,  1988.  Functional  rela- 
tionships ;uid  microhabitat  distributions  of  enteric  hel- 
minths of  grebes  (Podicipedidae):  the  exidence  for 
interactive  communities,  journal  of  Parasitologx  74: 
214-227. 

Todd.  K.  S.,  Jr..  and  D.  M.  Hammond  1968a.  Life  cycle 
and  host  specificity- oiEiiueiia  callospennoph'di  Heniy, 


1932  from  the  Uinta  ground  squirrel,  SjU'niiopliilns 
annatiis.  Joimial  of  Protozoology-  15:  1-.8. 

.  1968b.  Life  cycle  and  host  specificiK-  of  Ehueria 

lariiiwrcnsi.s  Vetterling,  1964  from  the  Uinta  ground 
squirrel,  Spennophilu.s  (iiiiuiiiis.  fournal  of  Protozool- 
o,g\- 15:  26.8-275. 

Todd,  K.  S.,  Jk  .  D.  M.  Ha.mmond  and  L.  C.  .Anderson 
1968.  Observations  of  the  life  cvcle  of  Eiuwria 
hilanu'llafa.  Henn-  1932  in  the  Llinta  ground  squirrel 
Speniutphiliis  (innatus.  (ourual  of  Protozoology'  15: 
732-740. 

Toft,  C.  A.  1986.  (xjuununities  of  species  with  parasitic 
life-styles.  Pages  445—463 /»  J.  Diamond  and  T  |.  C^ase. 
eds..  Community- ecology-.  Harper  and  Row.  New  York. 


Received  12  Fehnuin/  1992 
Aceeptecl  IS  Septeuiher  1992 


(iivat  Basin  Xatiiralist  52(4).  pp.  313-.320 

PLANT  AGE/SIZE  DISTRIBUTIONS  IN  BLACK  SAGEBRUSH 
{ARTEMISIA  NOVA):  EFFEGTS  ON  COMMUNITY  STRUCTURE 

James  A.  Youiiy;   and  nei)ra  E.  Palmqiiist 

Abstiuct. — The  demographv  of  black  sagebrush  (Ai-tciitisia  nova  Nelson)  wius  iTivestigated  in  the  Buckskin  Mountains 
ofwesteni  Nexacla  to  determine  patterns  of  stand  renewal  in  sagebrush  communities  currentK'  tree  Ironi  wildiires.  Biouiass 
sampling  was  conducted  to  de\eIop  growth  ckisses  that  reflected  apparent  age  of  the  shrubs.  The  densit\  of  black  sagi'l)nish 
plants  was  twice  that  of  basin  big  sagebrush  (A.  tiidcntata  ssp.  trklcntata  Nutt.)  in  adjacent  comunmities  on  contrasting 
soils  (2.2  \ersus  1.1  plants  per  m~).  Black  sagebmsh  accumulated  only  759^  as  much  woodv  biomass  as  big  sagebnish. 
f-legression  equations  were  de\eloped  and  tested  for  predicting  total  wood\  biomass,  current  annual  growth  (CAG),  and 
leaf  weight  of  black  sagebnish  plants.  Apparent  age  classes  were  de\elopetl  both  lor  the  black  sagi'brush  plants  and  die 
sub-canop\'  mounds  on  which  thev  grew.  Discriminant  lUUiKsis  was  used  to  test  this  classification  system.  Plant  succession, 
apparentk'  controlled  b\  nitrate  content  of  the  surface  soil,  appealed  to  eliminate  the  successful  establishment  of  black 
sagebnish  seedlings  on  the  mounds.  After  the  shnibs  die,  the  mounds  eventiuilK  deflate.  We  projiose  tliat  mounds  reform 
aroiuid  shrub  seedlings;  but  because  seedling  establishment  is  so  rare  in  these  coiiimunities.  this  could  not  be  xcritied. 

Kci/  iiords:  hioinass.  shnth  .succession,  dcscii  sail  fonnnlidii.  soil  nil  rale,  black  sY/gcAn/.v//.  Artemisia  nox  a. 


Black  .sagt'hru.sh  [Aiicniisid  uolci  Xclson)  is 
one  of  tlie  dwarf  sagebnish  species  which  col- 
lectiveK'  constitute  about  half  the  sagebitish 
\egetation  in  Ne\ada  (Beetle  1960).  Black  sage- 
1  irush  plays  a  dominant  role  in  a  number  of  plant 
communities  in  the  Great  Basin  (Zamora  and 
Tueller  1973).  Rarel\-  does  black  sagebrush 
share  dominance  with  another  species  of  Artc- 
inisia.  In  the  section  Tridentate  of  the  genus 
Aiicniisid.  black  sagebrush  is  perhaps  the  spe- 
cies most  adapted  to  arid  en\ironments.  Black 
sagebnish  is  closeK'  associated  with  shadscale 
[Atriphw  conjciiijolid  (Torr.  &  Frem.)  Wats.] 
dominated  landscapes  (Blaisdell  and  Holmgren 
19S4).  The  browse  of  black  sagebnish  is  higliK 
prcdcrrcd  b\'  domestic  sheep  {Ovis  aries). 
pronglioni  {Antilocaiya  anwiicana),  and  Sage 
Grouse  {Centrocerens  orophasianus) .  From  the 
189()s  until  the  late  195()s,  black  sagebni.sh  plant 
communities  in  the  Carson  Desert  of  Ne\ada 
werea\ital  part  ol  winter  range  for  tlu^  domestic 
range  sheep  indnstn".  Years  of  e\cessi\  e  brows- 
ing b\' sheep  actually  shaped  the  outline  of  black 
sagebnish  shrub  canopies;  Zamora  and  Tueller 
(1973)  reported  the\-  had  difficult\'  in  finding 
relic  communities  in  high  range  condition. 

Vetretation  of  the   Buckskin   Mountains  of 


west  central  \e\ada  is  characterized  In  black 
sagebruslVdesert  needlegrass  {Sfipa  spcciosa 
Trin.  &  Rupr.)  plant  communities.  The  Buck- 
skin Mountains  are  located  100  km  southeast  of 
Reno,  Nexada,  in  the  rain  shadow  of  both  the 
Sierra  Ne\ada  and  Pinenut  Mountains.  This  is  a 
portion  of  the  Canson  Desert  in  which  Billings 
(1945)  suggested  that  Afn/^/c.v-dominated  salt 
desert  shnib  \egetation  occurred  because  of 
atmospheric  drought  rather  than  occurreuc("  of 
soluble  salts  in  the  soil.  Ifwe  compare  the  black 
sagebrush  comniunities  of  the  ISuckskin  Nh)un- 
tains  with  those  describetl  in  the  regional  stutK 
conducted  b\  Zamora  and  Tueller  (197.3),  we 
find  that  the  highest-elexation,  north-facing 
slope  communities  of  the  Buckskin  Mountains 
correspond  to  the  most  arid  communities  pre- 
viouslv  described.  P'rom  this  we  assume  the 
black  sagei)riish  communities  in  this  stiuK  rep- 
resent an  arid  (extension  of  this  t\pe. 

Only  rec(^iitl\  haxc  occasional  wiUllires  of 
any  extent  occurred  in  black  .sagebrush  commu- 
nities in  western  Nexada.  The  fires  that  ha\e 
occurred  ha\e  been  associated  with  the  recent 
si)read  of  the  ali(Mi  annual  cheatgrass  {Broinus 
tcctoiiim  L.i  into  these  arid  emironments 
(Young  and  Tijiton  1990).  ApparentK  for  much 


USDA,  .Agricultural  Ht-.si'arcli  Senile.  920\'allc\  Kciad.  Heuo.  Ne\aila  S9.512 


313 


314 


Great  Basin  Naturalist 


[X'olume  52 


of  the  tAventietli  centun-  these  communities 
have  not  been  subject  to  wildfires  because  of 
lack  of  herbaceous  vegetation  to  cany  the  fire. 
Because  of  the  lack  of  trees  to  produce  fire  scars, 
it  is  difficult  to  determine  whether  these  sites 
were  subject  to  periodic  burning  under  pristine 
conditions.  This  is  in  sharp  contrast  to  basin  big 
sagebnish  communities  where  periodic  cata- 
strophic stand  renewal  by  burning  from  wild- 
fires has  been  common.  The  lack  of  catastrophic 
stand  renewal  in  black  sagebrush  communities 
should  be  reflected  in  the  age/size  class  struc- 
ture of  the  communities. 

Our  puipose  was  to  determine  the  age/size 
distribution  of  black  sagebrush  plants  to  deter- 
mine community  structure. 

Materials  and  Methods 

Studies  were  conducted  from  1984  through 
1988  in  the  Buckskin  Mountains  located  about 
100  km  southeast  of  Reno,  Nexada.  The  geo- 
logic features  of  this  moimtain  range  have  been 
described  in  detail  bv  Hudson  and  Oriel  ( 1979). 
Vegetation  and  soils  of  the  range  ha\e  been 
mapped  and  related  to  the  geologic  map  of  the 
area  (Lugaski  and  Young  1988).  The  plant  com- 
munities used  in  this  study  were  located  on  the 
Guild  Mine  member  of  the  Mickey  Pass  tuff 
This  geologic  unit  consists  of  crvstal-rich,  mod- 
erately to  poorlv  welded  ash  flow  tuff  (Proffett 
and  Proffett  1976).  It  has  been  proposed  that 
the  soils  (a)  developed  in  place,  (b)  developed 
from  subaeriallv  deposited  material  from  long- 
distance transportation,  or  (c)  dexeloped  from  a 
combination  of  residual  and  subaerialh'  depos- 
ited material  (unpublished  research,  ARS- 
USDA).  The  bulk  of  the  profile  is  an  argillic 
horizon,  about  50  cm  thick,  which  consists  of 
50%  or  more  clay-te.xtured  material.  It  is  pro- 
posed that  this  clay  horizon  is  a  reHc  of  a  soil  that 
de\'eloped  on  the  site  and  whose  original  surface 
horizon  lias  been  removed  by  erosion.  The 
important  point  is  that  the  clay  horizon,  which 
is  interniittentl)  exposed  on  the  soil  surface, 
developed  under  different  environmental  con- 
ditions from  the  current  surfiice  horizon.  The 
current  surfiice  soil  consists  of  a  relatively 
recenth'  deposited  layer,  apparently  from  sub- 
aerial  deposition,  that  is  largely  confined  to 
mini-mounds  beneath  the  canopies  of  the  black 
sagebrush  plants.  The  soil  is  classified  as  a  fine, 
iridic,  montmorillonitic,  Typic  Paleargid. 

S})atial  structure  of  the  black  sagebnish  com- 
nuinities  was  detennined  by  .sampling  five  .stands 


located  along  the  western  flank  of  the  Buckskin 
Mountains.  The  five  stands,  located  on  the  same 
outcropping  of  Micke\'  Pass  tuff,  were  separated 
bv  small  canvons  where  the  westerK^  tilted  ash 
flows  were  broken  bv  faulting.  All  sites  were 
west  facing  and  located  in  a  band  alono;  the 
mountainside  at  1720-1780-ni  elevation. 

A  starting  point  was  located  on  aerial  photo- 
graphs in  each  stand,  and  10  plots,  each  10  m" 
in  area,  were  located  random Iv  along  Line  tran- 
sects parallel  to  the  slope.  A  total  of  50  plots 
were  established  (5  stands  X  10  plots  per  stand). 
In  each  plot  the  following  were  determined:  (a) 
shnib  densit)'  by  species,  (b)  crowii  coxer  of 
shnibs  (ocular  estimate),  (c)  shnib  height,  (d) 
area  of  mound  and  interspaces,  and  (e)  herba- 
ceous cover  (ocular  estimate).  Mound  co\'er 
refers  to  the  slightly  raised  areas  beneath  shrub 
canopies  where  subaerialh'  deposited  soil  and 
saltation  deposits  accumulate. 

At  each  plot  location  the  herbaceous  \egeta- 
tion  frequencv  was  sampled  with  100  step  points 
arranged  in  4  lines  of  25  points  each  following 
the  procedures  of  Evans  and  Love  (1957).  The 
herbaceous  xegetation  was  resampled  annually. 

Using  the  same  starting  point,  but  bv  placing 
the  transects  up  and  down  the  slope,  25  black 
sagebnish  mounds  were  located  in  each  stand. 
The  shrubs  rooted  on  each  mound  were  mea- 
sured for  (a)  height,  (b)  ma.ximum  and  mini- 
mum crowni  diameter,  (c)  stem  number  (as  black 
sagebnish  ages  the  cambium  splits,  forming 
multiple-stemmed  plants),  and  (d)  stem  diame- 
ter at  the  soil  surface  (diameter  of  the  group  of 
split  stems).  The  aerial  portion  of  the  plant  was 
subdixided  by  clipping  into  the  following  cate- 
gories: (a)  coanse  stems,  2.5  cm  or  larger  in 
diameter;  (b)  fine  stems,  0.25  to  2.4  cm  in  diam- 
eter; (c)  current  annual  growth;  and  (d)  leaves. 
The  material  was  dried  at  80  degrees  C  for  24 
hours  and  weighed. 

After  the  aerial  portion  of  the  shrub  was 
remoxed,  the  litter  beneath  the  canopx' xx^as  col- 
lected and  screened  through  a  2-mm  screen. 
The  material  too  coarse  to  pass  through  the 
screen  xxas  saxed,  dried,  andxxeighed.  The  max- 
imum and  minimum  diameters  of  the  mound 
xvere  measured,  and  the  height  of  the  mound 
xx'as  determined  bx'  digging  to  the  clax'  horizon. 
The  number  of  perennial  grasses  rooted  on  the 
mound  xvas  counted  bv  species,  and  the  cover  of 
cheatgrass  xx^as  estimated  ocularlx'  per  mound. 

A  series  of  age/size  classes  xvas  established 
fcjr  the  black  sagebrush  plants  sampled.  These 


1992] 


Black  SACKiiiasii  Dkmocivm'Iiv 


315 


Table  1.  Mean  plus  standard  error  (SE)  for  shrub  densitv-  per  m".  percent  ])rojeeted  eanop\  cover,  fre(juenc\-  (lO-m" 
iloti  within  stands  (.V  =  lOl,  and  constanc\-  among  sttinds  (A'  =  5). 


Species 

Densih^ 

SE 

Cover 

(%) 

SE 

Frequency 

(%) 

SE 

Constancv 

(%) 

SE 

Artemisia  nova 

2.2 

0.40 

22 

2.4 

100 

0 

100 

0 

Chn/sotliainniis  riscidijloni.s 

0.7 

0.10 

o 

0.4 

40 

S 

SO 

8 

Ephnda  iiciiitlcnsi.s 

0.3 

O.OS 

1 

0.4 

64 

10 

100 

0 

Tc'tni(lt/mi(i  <H(ihmtti 

0.2 

0.04 

-■' 

- 

32 

5 

60 

4 

EriD^onuiii  iiuvrothccnin 

0.1 

0.04 

_■' 

- 

5 

1 

20 

2 

Eriogonum  UDibcUdtiim 

0.2 

0.00 

_■' 

- 

8 

1 

20 

1 

"Imlicatfs  less  tliaii  T 


classes  were  based  on  the  size,  growth  form, 
percentage  dead  canopy,  and  apparent  age  of 
the  plants.  The  classes  were  (a)  seedling,  (b) 
\oung  plant,  (c)  mature  plant,  (d)  patriarch,  (e) 
senescent,  and  (f)  dead. 

Soil  samples  from  the  sin"face  5  cm  were 
taken  (a)  ne.xt  to  the  shiaib  stem,  (b)  at  the 
can()p\-  edge,  and  (c)  10  cm  Ixnond  the  edge  of 
the  shnib  canop\.  These  samples  were  dried, 
screened,  and  shipped  to  a  connnercial  labora- 
toiA  ior  nitrate  nitrogen  anahsis. 

A  tAvo-wa\  anal\ sis  oi  xariance  and  post  hoc 
Duncans  Multiple  Range  test  were  performed 
to  analwe  differences  in  soil  nitrate  concentra- 
tion between  sagebnish  age/size  classes  and 
sample  location.  A  series  of  stepwise  regressions 
was  performed,  utilizing  the  general  linear 
model,  wherein  a  subset  of  \ariables  was  chosen 
that  would  best  predict  plant  weight,  annual 
growth  of  plant  (weight),  and  leaf  weight  of 
black  sagebnish.  Separate  step-up  regressions 
were  performed  for  plant  and  mound  character- 
istics (Neter  and  Wasserman  1974).  The  inde- 
pendent variables  that  were  significant 
contributors  to  predicting  age/size  classes  for 
these  two  groups  were  chosen  as  discriminant 
\ariables  to  be  used  in  two  separate  discriminant 
anal\  ses.  The  age/size  classes  of  the  sagebrush 
plants  were  used  as  the  grouping  stnicture  in 
the  discriminant  analysis.  The  plant  character- 
istic variables  .selected  as  significant  contribu- 
tors to  classification  into  age/size  classes  were 
(a)  weiglit  of  coarse  stems,  (b)  nimiber  of  stems, 
(c)  plant  height,  and  (d)  plant  diameter.  The 
mound  characteristics  (ranked  in  order  of 
importance)  used  were  (a)  litter  cover,  (b)  litter 
weight,  (c)  soil  nitrate  concentration,  and  (d) 
cheatgrass  cover  on  the  mound. 


Results  and  Dlscussiox 

C>ommimit\  ("ompetition 

The  plant  comnumities  of  the  Buck.skin 
Mountains  dominated  bv  black  sagebnish  are 
low  ill  diversitv  (Table  1).  Green  rabbitbnish 
[Chnjsothaminis  liscklijlonis  (Hook.)  Nutt.] 
occurs  in  patches  in  the  community  Nevada 
ephedra  {Ephedra  ncvadciisis  Wats.)  is  rather 
evenly  distributed  through  the  black  sagebnish 
communities,  but  at  a  low  density  Littleleaf 
horsebnish  (Tctrachpiiia  <j^labrata  Gra>")  is  a  rel- 
ativeh  infrequent  component  of  the  communi- 
ties. The  two  species  of  Eiio<io)}iim  are 
semiwoodv  species  that  also  occurred  in  tlie 
most  arid  black  sasebrush  communities  that 
Zamora  and  Tueller  (1973)  reported. 

S(jnirreltail  [Ehjmus  lujsfhx  Scribn.)  and 
cheatgrass  are  the  most  frequent  herbaceous 
species  (Table  2).  The  relative  fr(H|iiencvof  the 
two  species  reverses  from  \ear  to  xcar  depend- 
ing on  available  moisture  for  plant  growth. 
Cheatgrass  is  abundant  only  in  years  with  ade- 
({iiate  moi.sture  during  the  spring.  The  densits' 
of  squirreltail  plants  remmns  relatively  constant. 
In  dn'  years,  squirreltail  is  virtually  the  only 
herbaceous  species  in  these  communities. 

Biomass 

Along  the  western  margin  of  the  Buck.skin 
Mountains,  black  and  basin  big  sagebrush  com- 
niiiiiiti(\s  occur  side  by  side  on  shaqily  contrast- 
ing .soils.  The  basin  big  sagebnish  communities 
have  been  burned  in  wildfires,  ba.sed  on  historic 
records  and  fire  scar  aiialv  sis  (Young  et  al.  1 9S9). 
Our  analysis  of  black  sagebrush  communities  is 
essentially  based  on  abovegroimd  woody  mate- 
rial accumulation  of  the  dominant  shrub.  We 
had  previouslv'  conducted  a  stud)'  of  the  biomass 


316 


CiHKAT  Basin  Naturalist 


[\  olunie  52 


T.\BI,K  2.  Mean  plus  staiitliuxl  error  (SE)  lor  lrL'(|nL-ncv  ol  herhaeeou.s  species  for  an  a\erage  of  four  years'  sampling 
(a\erage  precipitation  175  nnn),  tor  a  dn'  spring  (1989,  no  April  precipitation),  and  a  year  with  above-a\erage  moistine 
av;iilal)le  for  plant  growth  (1986,  225  mm  precipitation).  Based  on  5000  sample  points  per  year. 


Fr 

equency 

Av( 

;^rage 

Diy(1989) 

Wet  (1986) 

Foiu"  ye 

ars 

SE 

Sprir 

ig 

SE 

Spring           SE 

GKOWTIIKHOMSI'IX.IKS 

..-%- 

PKHIONNIALCaU.SS 

Elijiiius  lu/stiix 

39 

4.1 

70 

6.9 

6              0.5 

Stipa  spcckmi 

3 

0.3 

8 

0.7 

- 

Siipa  ihurhciidiKi 

- 

- 

- 

- 

Poo  scamdd 

- 

- 

1 

O.S 

- 

On/Z(>j)si.s  liiiinciiuklcs 

- 

- 

1 

0.2 

-                 - 

.\N\r\i.(;HASS 

Broiiiiis  Icctoniin 

44 

6.6 

14 

2.8 

76             3.8 

PKHKNNI.\LF()Hli 


Cast i Ih'ja  cliromo.sa 

1 

0.2 

1 

0.2 

- 

- 

Spluicmlcai  pairifolid 

- 

- 

2 

0.3 

- 

- 

Phlox  luHxIii 

- 

- 

3 

0.3 

- 

- 

AWIALKOKB 

Enxliiiin  cinitariinn 

5 

0.8 

- 

- 

2 

0.8 

Dcsairaiiiiti  piitiiata 

- 

- 

- 

- 

5 

0.7 

Sistjmhriiiin  altissiitiiiin 

5 

0.9 

- 

- 

10 

0.8 

■'Inclicatt's  li's.s  than  19r  avcraiit- 


ol  l)a.sin  big  sagebrush  adjacent  to  the  western 
edge  of  the  Buckskin  Mountains  (Young  et  ah 
1989).  This  allowed  comparison  of  the  produc- 
tion of  bioniass  of  basin  and  l)lack  sagebmsh 
from  the  same  area.  The  basin  big  sagebrush 
communit)'  had  a  .sandy  loam  surface  soil  and  a 
greater  soil  depth  (Haplargids  deri\ed  from  meta- 
N'olcanic  sources).  Big  sagebmsh  ages  were 
clumpt>d  at  5.5-60,  40^5,  and  10-15  years  old. 

The  general  aspect  of  the  t\vo  communities 
is  strikingK'  different,  with  the  maximum  height 
of  th(^  black  .sagebrush  being  60  cm  and  that  of 
the  big  sagebmsh  over  1  m.  In  contrast  to  the 
central  wood\'  stems  of  the  big  sagebmsh  plants, 
black  sagebrush  plants  appear  multi-stemmed. 
Despite  the  difference  in  height,  the  two  com- 
munities have  .similar  biomass  because  of  the 
higher  detisit)  of  plants  in  the  black  sagebrush 
connnunit).  There  is  more  coarse  and  fine 
woody  material  in  the  basin  big  sagebrush  com- 
munit)' (Table  3). 

If  we  assume  both  populations  are  the  same 
age  (assumption  is  necessaiy  becau.se  actual  age 
of  black  sagebmsh  plants  could  not  be  esti- 
mated), the  rate  of  woody  biomass  accumula- 
tion was  13.2  g/nr/year  and  64.5  g/m"/vear  for 
black  and  basin  big  sagebmsli,  respectively.  The 


wide  difference  between  the  two  connnunities 
is  apparently  due  to  the  higher  woody  biomass 
of  more  mature  basin  big  sagebmsh  plants. 

Woody  bioniass  of  black  sagebaish  was  best 
predicted  b\'  the  ecjuation: 

Y  =  9.87  +  1.21  "  XI  +  1.12  "  X2  +  0.88  °  X3 

where  Y  =  total  woody  biomass  (grams),  XI  = 
fine  stems,  X2  =  coarse  stems,  and  X3  =  root 
crown.  R~  -  .96  for  this  determination.  YearK' 
growth  hicrement  was  predicted  b\  the  e( {nation: 

Y  =  16.96  +  0.26  "  XI  +  0.16  ^  X2  - 
0.73  °X3- 0.14  "X4 

where  Y  =  current  growth,  XI  =  fine  stems,  X2 
=  coarse  stems,  X3  =  iilant  lieidit,  and  X4  -  root 
crown.  R~  =  .57  for  this  determination,  despite 
the  inclusion  of  a  fourth  \ariable.  Our  third 
ecjuation  predicted  leaf  weight: 

Y  =  23.53  +  0.29  "  XI  -  0.93  °  X2  + 
0.2  "  X3  -  0.35  °  X4 

where  Y  =  leaf  weight,  XI  =  coarse  stems,  X2  = 
height  of  plant,  X3  -  fine  stems,  and  X4  =  plant 
density.  These  four  \arial)les  in  the  eejuation 
accounted  for  64%  of  the  variability'  in  the  data. 


19921 


Bi^u;iv  S.u;kbiu;.sii  Di;.\iuc;haimiv 


31' 


30 


20 


10- 


0- 


■10 


-20- 


-4 


A 
A  A 


A 
A  A     A     A 


® 
®         ®®  ® 

®®  ®s)®®8e6®  ®  ®®(S®S®  ®®  ®  ®  ® 
®^  ®®s®  ®®s®    ®®®  ®  ®  ®       ® 

®    ®  ® 


— I 1 r 

-1  0  1 

Canonical  Fnc.  2 


Fig.  1.  Plot  of  l)Iatk  sagebrush  group  membership  based  on  plant  characteristic  (liscrimiuaut  ecjnationswliereO  =  \oung. 
®  =  mature.    ▲  =  patriarch.  ■  =  senescent,  and  □  =  dead. 


Age/Size  Classes 

The  selected  variables  for  both  plant  and 
mound  characteristics  were  important  contrib- 
utors in  distinguishing  between  age/size  classes 
and  were  good  indicators  of  group  composition 
(Fig.  1).  Ver\-  few  niisclassifications  occurred  1)\ 
use  of  the  resulting  discriminant  functions. 

The  bulk  of  the  black  sagebrush  stands  was 
composed  of  mature  plants  20-60  cm  tall  with 
canopies  20-50  cm  in  dianu^er  (Table  4).  This 
is  a  wide  range  in  height  and  canop)'  size,  but 
tl le  1 1  uitu re  age/size  class  was  distinguished  f  r( )n  i 
young  plants  b\  the  presence  of  up  to  10%  dead 
material  in  the  canop\-  and  the  beginning  of  the 
separation  of  the  stem  into  individual  cambium 
bundles.  The  patriarch  class  was  distinguished 
from  the  mature  class  b\-  an  increase  in  dead 


material  in  the  canopv  (to  307c)  and  complete 
separation  of  the  stems.  The  separated  stems 
fbnned  U-shaped  flutes  with  the  open  end  of 
the  U  toward  the  former  center  of  the  stem.  It 
was  not  possible  to  establish  tlie  maximum  age 
of  the  class  because  the  center  of  the  stem  was 
missinji.  The  indixidual  .section  had  at  least  40 
growth  rings. 

Senescent  plants  formed  the  next,  appar- 
cntK  older,  age/size  class.  In  this  cUiss  at  least 
50%  of  the  canopv  was  dead.  Older  black  sage- 
brush plants  do  not  get  taller,  prol)abl\  because 
tl  wv  ha\e  no  central  stem  to  support  the  canopy. 
The  diameter  of  the  crowns  does  increase. 
There  is  a  marked  increase  in  wood\-  biomass 
between  the  patriarch  and  senescent  classes. 

Seedlings  and  \oung  plants  constituted  only 
6%  of  the  black  sagebnish  populations  (Table  4), 


318 


Great  Basin  Naturalist 


[Volume  52 


TaBLK  3.  Mean  density  (stem.s/ni  ,)  plus  staiiclard  error  (SE)  and  oxcn-iln  hioniass  (g/ni")  oi  Aiicinisid  nma  and  A. 
tridentata  subsp.  tridentata.  Data  for  A.  tridentata  suhsp.  tridcntata  from  a  previous  study  (Young  et  al.  1989). 


Biomass  per 

■-> 
m" 

Species 

Deni 

5it\- 

Co; 

u"se 

Fine 

GAG 

Le; 

.ives 

Total 

m"" 

SE 

g 

SE 

g 

SE 

g 

SE 

g 

SE 

g         SE 

Artemisia  nova 
Artemisia  tridentata 
subsp.  tridentata 

2.2 
1.1 

0.4 
0.3 

750 
,S50 

90 
KM) 

.520 
970 

fiO 
110 

ISO 
170 

4.5 
40 

1.30 
130 

40 
50 

15.50      240 
2120      420 

T\HLK  4.  Artemisia  luna  crown  and  hioniass  characteristics  for  indixidual  age/size  classes.  Demographic  breakdown  of 
black  sagebnish  communities  In  growtli  classes.  Glasses  are  related  to  age  tor  younger  plants,  but  once  stems  separate,  ages 
are  not  based  on  luinud  rings. 


Grown  characteristics 

BiomiLss  char 

acteristics 

Dead 

Goarse 

Fine 

Age/size 

Height 

Diametei 

■  Densit\-  bi 

•anchlets 

stems 

stems 

GAG 

Lea\e 

Stem 

Percentage 

Age 

class 

(cm) 

(cm) 

{%) 

(%) 

(g) 

(g) 

(g) 

(g) 

number 

of  stand 

(vears) 

Seedling 

5 

5 

30 

0 

0 

15 

5 

10 

1 

>1 

2-5 

Young  plant 

10-20 

.5-10 

.50 

0 

2S 

64 

.30 

40 

1 

5 

5^30 

Mature 

20-fi() 

10-.5() 

SO 

10 

140 

120 

SO 

60 

Multiple 

60 

30-50 

Patriarch 

20-60 

20-S() 

60 

.30 

,560 

420 

140 

100 

Multiple 

17 

40+ 

Senescent 

20-60 

20-100 

30 

60 

9S() 

640 

60 

.30 

Multiple 

12 

•p 

Dead 

20-60 

20-100 

0 

100 

910 

320 

0 

0 

,Multiple 

5 

p 

with  seedling.s  being  very  rare.  The  separation 
between  seecUing  and  young  plants  was  based 
on  the  occurrence  of  coarse,  woody  biomass  in 
the  latter  class.  Young  plants  had  entire  stems 
with  no  exidence  of  division  of  the  cambium. 

Mound  Tvpes 

Eacli  black  sagebnish  age/size  class  had  a 
corresponding  t)pe  of  sub-canopy  mound.  The 
only  seedling  found  in  the  entire  study  was 
located  in  an  interspace  between  mounds.  Obvi- 
ously, one  seedling  is  not  a xalid  sample,  but  tlie 
lack  of  seedlings  is  a  critical  factor  in  the  d\  nam- 
ics  of  the  communities  studied.  The  first  detect- 
able mound  occurred  imder  young  plants.  Only 
5-10%  of  the  sub-canopy  area  imder  black  sage- 
brush plants  in  the  voimg  plant  age/size  class 
was  covered  with  litter  (Table  5).  The  litter  was 
coTnpo.sed  of  fragnu^nts  of  black  sagebnish  leaxes. 

In  the  mature  plant  age/size  class  the  co\er 
of  litter  and  the  weight  of  litter  increa.sed  (Table 
5).  The  mounds  were  easily  distinguished  by 
both  height  and  surface  soil  color  and  te.xture. 
The  surface  of  the  mounds  appeared  darker  in 
color,  and  the  reddish  tinge  to  the  cla\'  surface 
soils  of  the  interspace  was  not  apparent.  If  the 


surface  of  the  mound  was  disturbed,  the  dark 
color  was  replaced  bv  a  gra\ish  shade.  Mounds 
appear  to  reacli  their  maximum  height  with  this 
growth  stage  of  black  sagebnish.  Mounds  of 
mature  plants  had  perennial  grasses  associated 
with  the  sub-canopy  area.  The  most  frecjuent 
perennial  grass  was  squirreltail. 

Litter  accumulations  increased  witli  the 
patriarch  age/size  class,  but  height  oi  the  mound 
did  not  increase.  ApparentK;  trapping  of  sub- 
aerial  deposition  material  and  saltation  particles 
must  be  related  to  gro\\'th  stage  of  black  sage- 
brush plants  in  terms  of  crowii  architecture. 
SubaerialK'  deposited  particles  are  ob\iousl\' 
\'er\'  unstable  and  subject  to  redeposition  if  the\' 
fall  in  the  largelv  bare  interspace  among  shrub 
mounds  (Young  and  E\ans  1986).  If  litter  accu- 
mulation increases  on  patriarch  mounds,  wliy 
do  the\  not  trap  these  secondan'  erosion  prod- 
ucts and  the  mound  keep  growing  in  height? 
Canopv  stnicture  changes  with  the  patriarch 
aee/size  class,  with  increasino;  bare  stems  and 
spreading,  but  not  taller,  plants.  It  would  appear 
that  aerial  d\iiamics  of  the  crown  of  black  sage- 
brush plants  influence  mound  height. 

W'itli  the  senescent  age/size  class,  adivergence 


19921 


Black  Sa(;ki51u  sii  Dkmockapiiv 


319 


Table  5.  Mouiul  characteristics  in  relation  to  a<j;<Vsi/.(>  classes  of  Aiicinisia  noia.  Illustrati's  tliat 
liuige  with  age/size  classes  of  shrubs. 


ound  characteristic; 


Ml 

ound 

Litter 

1 

\'renni; 

il 

Cl 

leatgrass 

Black  saijelirush 

DiaiiK 

■tcr 

gnwtli  classes 

Heigiit 

Max 

Mm 

Co\er 

Depth 

Weight 

g' 

iuss  deusitv 

cover 

Number 

(cm) 

(cm) 

(cm) 

{%) 

(cm) 

(g) 

(per  mound) 

(%) 

Samples 

Seedling 

0 

0 

0 

0 

0 

0 

0 

0 

1 

Young  plant 

2-5 

fiO 

30 

5-10 

0.5 

40 

0 

2 

6 

Nhiture 

5-15 

SO 

40 

40-60 

1-1.5 

4S() 

2.0 

15 

76 

Patriarch 

10-15 

100 

60 

SO 

2-3 

690 

2.S 

12 

21 

Senescent 

10-15 

100 

60 

SO 

2-3 

720 

2.1 

60 

15 

Dead 

10-15 

100 

60 

SO 

2.5-5 

970 

6.4 

5 

6 

T.\BLF.  6.  Mean  nitrate  level  (mg/kg)  of  soil  at  the  stem,  canop\'  edge,  and  onts 
in  relation  to  maturit\-  classes.  Buckskin  Mountains,  Ne\ada.'' 


the  c;uu)p\  ol  black  sagebrush  plants 


Age/size  class 


Location ' 

Stem 

Canopv 

Outside 

Age/size 

(ppnv) 

(ppm) 

(ppm) 

class  mean' 

4.7  h 

4.3  hi 

4.1  hi 

4.4  d 

6.6  g 

5.5  g 

4.0  1 

5.4  c 

10.5(1 

12.0  b 

S.Oe 

10.2  a 

13.2  a 

11.3c 

7.0  f 

10.5  a 

S.4  c 

7.0  f 

7.1  f 

7.5  b 

8.7  a 

S.Ob 

6.0  c 

Young  plant 

Mature 

Patriarch 

Senescent 

Dead 

Mean  location 


'Means  followed  by  the  same  letter  are  not  significantly  different  at  the  .01  le\el  of  probability  as  detennined  by  Unncans  .Multiple  Kange  test. 
'■-Means  of  location  followed  b\'  the  same  letter  are  not  significantK'  ilifTerenI  at  the  .01  le\el  of  probability  as  determined  by  Duncan's  Multiple  Hange  test. 
Mi-ans  of  age/size  cliisses  followed  bv  the  same  letter  are  not  significantly  different  at  the  01  le\cl  ol  prnbabilit\  as  delermin.-d  b\  Duncans  Multiple  Range  lest. 


ill  lu'ii);iceous  species  composition  on  the 
mounds  occurs  (Table  5).  Some  mounds 
become  densely  covered  with  cheatgrass  as 
i)lack  sagebrush  plants  become  senescent  antl 
others  support  colonies  of  squirreltail. 

After  the  black  sagebmsh  plants  die,  litter 
weight  continues  to  increase  and  litter  changes 
in  appearance.  Litter  under  dead  plants  is  com- 
posed of  stringN'  bark  fragments,  and  indixidual 
black  sagebrush  lea\es  cannot  be  distinguished 
in  the  litter. 

Soil  Nitrate  Le\els 

Surface  soil  nitrate  le\els  were  higher 
beneath  shrub  canopies  than  in  the  interspace 
(Table  6).  Levels  were  highest  next  to  shrul) 
stems.  Nitrate  le\e]s  beneath  the  canop\'  rose  as 
age/size  classes  of  black  sagebmsh  indicated 
older  plants  and  moimds.  This  is  in  itself  an 
indication  that  age/size  classes  actual]\  do 
reflect  increasing  age.  The  development  of  xer- 
tical  and  horizontal  patterns  in  soil  nitrogen, 
attributed    to    the    localization    of   litter    fall 


beneath  the  canopies  of  desert  shrubs,  has  been 
documented  by  the  research  of  N.  E.  West  and 
co-workers  (Charley  and  West  1977,  West  and 
Skujins  1977,  West  1979).  Nitrate  lexels  of  sur- 
face soils  dropped  significantK  (F  <  .01)  once 
the  black  sagebmsh  plants  died.  Nitrate  le\els 
in  surface  soils  at  the  edge  ol  slimb  iiiouiids 
incn\ised  with  apparent  increasing  age  of  black 
sagebrush  j^lants  and  mounds.  These  areas  cor- 
respond to  the  micro-topoedaphic  situation 
described  as  c()[)pice  benches  by  Eckert  et  al. 
(1989)  for  shmb  mounds  in  big  sagebru.sh  com- 
munities. ApparentK  the  increase  in  soil  nitrate 
n^sults  from  leaching  (rom  the  mounds.  Once 
black  sagebrush  ])lants  are  dcatl  and  grasses 
doiiiinate  the  iiiomid.  soil  nitrate  le\els  decrea.se. 

Mounds  and  Black  Sagebrush 
CommunitN  Structure 

We  did  not  find  grass-dominated  inounds  or 
grass-dominated  mounds  with  black  sagebmsh 
seedlings.  We  did  note  the  remains  of  mounds 
that  appeared  to  be  eroding  awa)'.  Apparently, 


320 


Great  Basin  Nathhallst 


[Volume  52 


mounds  are  dvnamically  formed  and  eroded  in 
relation  to  the  establishment  and  eventual  death 
of  black  sagebrush  plants.  The  failure  to  find 
grass-dominated  mounds  may  be  a  function  ol 
herbivoiy-  by  domestic  livestock  [sheep,  feral 
horses  {E(j{ius  cahallus),  and  black-tailed  jack- 
rabbits  {Lcpiis  californicus)].  Grass-dominated 
mounds  nia\-  fail  to  persist  since  grasses  cannot 
maintain  mounds  because  of  leaf  fall  andcanop)' 
structure  differences  compared  with  black  sage- 
brush plants.  The  onK'  patchy  vegetation 
encountered  in  the  communities  was  groups  of 
rabbitbrush  plants.  Perhaps  rabbitbinish 
increases  after  relatively  short-lived  squirreltail 
plants  die  or  are  reduced  h\  grazing.  In  an 
adjacent  liig  sagebmsh  commimit)'  we  pre- 
\iously  determined  three  episodes  of  seedling 
establishment  at  12,  42,  and  57  years  before 
1985  (Young  et  al.  1989).  Plant  ages  were  clus- 
tered around  these  apparent  establishment 
dates.  The  clusters  mav  represent  periods  of 
desirable  climate  for  seedling  estabhshment  or 
a  single  season  when  establishment  occurred; 
they  may  also  represent  variabilitv  in  growth 
ring  deposition  or  recognition.  The  classes  we 
constnicted  in  this  study  are  much  too  broad  to 
pinpoint  this  t)pe  of  epi.sodic  stand  establisli- 
ment  for  black  sagebrush.  Perhaps  black  sage- 
bmsh conununities  not  renewed  catastrophicalK 
b)'  wildfires  onh  require  stand  renewal  at  such 
low  levels  (5%  of  the  stand,  standing  dead 
plants)  that  our  one  seedling  sampled  is  suffi- 
cient for  conmnmit^'  regeneration. 

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ZvMORA,  B.,  AND  P.  T  TiEi.i.ER  1973.  Artciiiisifi 
(irhiisnild.  A.  loii'^ilolxi,  and  .A.  mnd  habitat  tvpes  in 
northern  Nevada.  (Jreat  Basin  Naturalist  33:  22.5-242. 


Rrniiid  21  Chiohcr  Ih)yi 
Accepted  lU  September  1992 


Cireat  Basin  Naturalist  52(4 K  pp.  321-52' 


MUSHROOM  CONSUMPTION  (MYCOPIIAGY) 
BY  NORTH  AMERICAN  CER\  IDS 


Karen  L.  Lai 


iiR'hhaii'jii 


and  I^ln'Iip  j.  Unless 


Abstiuct. — Nati\c  miishrooms  pla\  iui  iiriportaiit.  tli()uu;li  olteii  uiulerc'stiinatetl.  role  in  deer  elk.  and  Ciiribon  diets  in 
Nortli  America.  Mushrooms  are  often  noted  as  an  unusual  or  anomalous  food  in  the  diets  of'eenids;  \et  the\-  often  dominate 
diets  in  the  late  summer  and  tall  in  forested  areas  of  western  North  America  and  throusrhout  the  \'ear  in  tlie  southeiLsteni 
U.S.  Mushrooms  are  particularh'  high  in  protein  ( 16-19% ).  phosphorus  (a\  erage  0.759f ).  and  potassium  (a\erage  2*^ ).  Also, 
mushroom  production  is  generalK'  greatest  in  tall  Tlieic^toic.  th('\  are  a  liigliK  nutritious  lood  in  late  se;LSon  when  other 
nati\e  forages  ma\'  marginallv  meet  basal  nutrii'ut  recjuirements  of  ungulates. 

Kci/  words:  ctirihoii.  aTiid.  deer.  diet.  dk.  iin/<(ij)liii!^i/.  iitiislirooiii.  iiiihitidii.  nnniiKint. 


\\'ildlife  scientists  ha\e  lon^  recognized  that 
certiiin  higliK'  nntritious,  "bonus"  foods  fre- 
cjuently  contribute  significanth-  to  animal  wel- 
fare though  their  contribution  (%)  to  the  diet 
nia\'  be  small  (e.g.,  acorns,  mushrooms,  and 
mestjuite  beans).  By  seeking  out  these  high- 
( [ualits'  but  generalK-  scarce  or  ephemeral  foods, 
li(n-bi\ores  can  balance  nutrients  against  lower- 
(jualit\'  forages  that  are  more  abundant.  Natixe 
nuishrooms  ha\'e  often  been  recorded  as  a 
"bonus  "  food  in  the  diets  of  deer,  elk,  and  cari- 
bou in  North  America.  However,  their  contribu- 
tion to  cenid  nutrition  is  not  commonh' 
miderstood. 

The  term  "mushroom"  refers  to  the  flesh\ 
fruiting  bod\  (sporocarp)  of  mam'  species  of 
fungi.  Mushrooms  are  technicalK"  not  "plants." 
The\'  belong  to  tlie  kingdom  Mxcetae  under  the 
fi\e-kingdom  classification  system  (Whittaker 
1969).  The  priman'  mushroom-producing  fungi 
are  in  the  group  called  Basidionncetes,  but 
man\  mushrooms  eaten  b\'  wildlife,  including 
morels,  are  Asconncetes.  Mushroom  produc- 
tion is  triggered  when  species-specific  rec^uire- 
nients  of  minimum  temperature  and  moisture 
conditions  are  met  (Smith  and  Weber  1980). 

Mushroom  consumption  (mvcophag))  has 
been  recorded  for  man\'  wildlife  species  in 
North  America.  Mushrooms  are  eaten  b\  ungu- 
lates (e.g.,  deer  and  elk),  small  manunals  (e.g., 
squirrels  and  armadillos),  as  w(>ll  as  birds,  tur- 


tles, and  insects  (Miller  and  Halls  1969,  Fogel 
and  Trappe  1978,  Martin  1979).  Mushrooms 
ha\e  long  been  recognized  as  an  important  com- 
ponent of  small  mammal  diets  (Fogel  and 
Trappe  1978).  Howexer,  nuishrooms  are  seldom 
considered  a  significant  component  of  cerxid 
diets  even  though  the\  ha\e  been  anecdotalK 
recorded  as  a  "preferred"  food  item,  l^iscount- 
ing  mushrooms  as  an  important  dietan conijx)- 
nent  ma\stem  from  a  misunderstandingol  their 
nutriti\-e  \alue.  The  piuposes  of  this  re\iew  are 
to  (1)  assess  the  contribution  of  mushrooms  to 
cenid  diets,  (2)  summarize  the  known  literature 
on  the  nutritixe  \alue  of  nuishrooms  to  ungu- 
lates, and  (3)  assess  the  im[)lications  of  nncoph- 
ag\' to  liabitat  selection  and  iiuli-itional  ecologv 

contriihition  of  mush1u)()ms  to 
Deer.  Elk.  ani3  Cai^ibol  Diets 

Mushroom  (lonsumjition  b\  Deer 

Mam  stu(li(^s  haxc  recorded  mushrooms  in 
diets  of  both  mule  {Odoroilciis  hcinioiuis)  and 
white-taikxl  (Odocoilciis  vir^inianii.s)  deer 
(Table  1).  Diet  composition  estimates  range 
from  a  trace  to  a  majoritx'  of  the  diet.  On  the 
up[)('i-  limit.  71.2%  mushrooms,  on  a  fresh- 
weight  basis,  were  recorded  in  fall  deer  diets  in 
.Mai)ama  (Kirkixitrick  et  al.  1969),  65.8%  in 
Augu.st  diets  in  Arizona  (Hungerford  1970),  and 
59.59f  in  .August  diets  in  Montana  (Loxiuis  1958). 


^Range  Science  Department.  Utah  State  University',  Logan.  Utah  S4.322-.5230. 

"Present  address:  Range  and  WildHfe  Management  Department.  Te,\a.s  Tech  University  l.nhbock.  'Pi-xas  79409-212.5. 


321 


322 


Great  Basin  Naturalist 


[Volume  52 


Table  I.  Proportion  of  mushrooms  in  deer,  elk,  luid  caribou  diets  in  North  America  a\eraged  o\er  season''. 


Species 
State  or  Province  (Vegetation  t>pe)  ^ 


%  of  diet 


Spring      Summer        Fall  \\ 'inter  Kind  of  data'       Source' 


Mule  deer  (Odocoileus  hemionus) 

Colorado  ( spruce/fir/pine  forest) 
Montana  (spruce/fn/pine  forest) 
Utiili  (dn'  mountain  meadow) 
Utiili  (mature  conifer  forest) 
Utah  (stagnated  conifer  forest) 
Utah  (conifer  forest/oak  woodland) 
Arizona  (mixed-conifer  forest) 
California  (chaparral-oak  woodland) 
British  C'olumbia  (conifer  forest) 

White-tailed  deer  (Odocoileun  virginianiis) 

.\ew  Brunswick  (conifer/deciduous  forest) 
Maine  (pine-hemlock  forest) 
Penn.svKania  (clear-cut  forest) 
Southeastern  U.S.  (oak-hickoiy-pine  forest) 
Southeastern  U.S.  (mixed-pine  forest) 
Southeastern  U.S.  (southern  evergreen  forest) 
X'irginia  (eastern  deciduous  forest) 
North  Carolina  (oiik-hickor\'-pine  forest) 
South  Carolina  (mLxed  pine  forest) 
CJeorgia  (southern  evergreen  forest) 
Florida  (southern  evergreen  forest) 
Florida  (southern  evergreen  forest) 
Florida  (pine-scmb  oak  forest) 
Alabama  (southern  pine-hiirdwood  forest) 
Alabama  (southern  pine-hardwood  forest) 
Louisiana  (pine-bluestem  range) 
Louisi;ma  (pine-hardwood  forest) 
Louisiana  (clear-cut  forest) 
Texas  (pine-mixed  hardwood  forest) 
Oklahoma  (oak  savannah) 
Wisconsin  (northern  hiirdwood  forest) 
Miiniesota  (northern  hardvvood  forest) 
South  Dakota  (pine  forest) 
South  Dakota  (pine  forest) 

Elk  (Cervus  eUiphus) 

X'irginia  (eastern  tleciduous  forest) 
Saskatchewan  (pine  forest) 
Saskatchewan  (mixed  forest) 
Utah  (diy  niountiiin  meadow) 
Utah  (mature  conifer  forest) 
Utah  (stagnated  forest) 
California  (Pacific  rain  forest) 

Caribou  (Ratif>ifer  tarandus) 

Newfoundland  (conifer  forest) 
Northern  Canada  (conifer  forest) 
Northern  Canada  (boreal  forest) 
Alaska  (spnice  forest/tundra) 
Alaska  (spruce  forest) 


- 

0.3 

- 

- 

Obs.(%  bites) 

31 

0.0 

12.1 

0.0 

0.0 

Rum.  (%  vol.) 

21 

- 

7.0 

- 

- 

Obs.  (%  mass) 

10 

- 

15.0 

- 

- 

Obs.  (%  mass) 

10 

- 

14.0 

- 

- 

Obs.  (%  mass) 

10 

- 

5.4 

9.3 

- 

Obs.  (%  ma.ss) 

4 

- 

16.4 

- 

- 

Obs.  (%  time) 

16 

- 

- 

- 

<1.0 

Rum.  (%vol.) 

20 

0.0 

0.0 

13.0 

4.0 

Rum.  (7c  vol.) 

8 

13.7 

6.7 

9.1 

Rum.  (%  m;rss) 

26 

0.0 

0.0 

45.0 

0.0 

Obs.  (%  mass) 

9 

1.6 

0.2 

0.8 

4.5 

Obs.  (%  time) 

14 

2.1 

19.8 

8.4 

6.2 

Rum.  (%  vol.) 

12 

0.4 

15.6 

8.6 

4.9 

Rum.  (%  vol.) 

12 

0.6 

16.4 

5.4 

3.2 

Rum.  (^c  vol.) 

12 

0.0 

40.0 

2.5 

0.0 

Rum.  (%  vol.) 

19 

0.0 

10.6 

7.0 

0.0 

Rum.  (%  vol.) 

19 

0.2 

33.4 

2.6 

10.7 

Rum.  (%  vol.) 

19 

0.0 

9.7 

9.0 

13.8 

Rum.  {'7c  vol.) 

19 

1.4 

10.4 

26.7 

13.2 

Rum.  (%  vol.) 

19 

- 

- 

- 

9.2 

Rum.  (%  vol.) 

11 

- 

- 

- 

25.2 

Rum.  {7c  vol.) 

11 

0.0 

71.2 

0.5 

17.4 

Rum.  {9c  vol.) 

19 

7.3 

- 

4.8 

0.8 

Rum.  {%  vol.) 

1 

0.5 

1.5 

3.5 

<0.5 

Obs.  (%■  bites) 

28 

- 

0.4 

1.9 

0.7 

Obs.  (%  bites) 

29 

- 

<0.1 

2.1 

0.2 

Obs.  (%  bites) 

29 

3.0 

34.0 

1.0 

7.0 

Rum.  (%  mass) 

25 

0.0 

0.0 

4.3 

1.0 

Rum.  (rel.  freq.) 

30 

- 

2.0 

- 

- 

Rum.  (%  vol.) 

22 

- 

- 

<1.0 

0.0 

Rum.  (%  vol.) 

2 

0.0 

4.0 

2.1 

0.0 

Rum.  (9c  vol.) 

15 

- 

0.7 

0.5 

<0.5 

Rum.  [9c  vol.) 

23 

1.0 

Rum.  (%vol.) 

3 

- 

5.3 

- 

- 

Rum.  (%  mass) 

17 

- 

4.2 

- 

- 

Rum.  (%  mass) 

17 

- 

4.2 

8.3 

- 

Obs.  (%  mass) 

"" 

- 

18.7 

55.7 

- 

Obs.  (%  mass) 

"" 

- 

18.4 

55.4 

- 

Obs.  (%  mass) 

"* 

- 

- 

0.3 

- 

Obs.  (%  time) 

13 

0.0 

25.0 

12.0 

0.0 

Rum.  {9c  vol.) 

5 

- 

- 

- 

0.4 

Rum.  (%  vol.) 

24 

- 

1.2 

- 

- 

Rum.  {9c  vol.) 

18 

0.0 

12.0 

10.0 

2.0 

Obs.  (%  vol.) 

6 

- 

- 

45.0 

- 

Rum  (%  vol.) 

27 

\\  cliLsli  (-)  listed  a.s  %  in  diet  me:m.s  no  data  were  availablf . 

'General  description.s  given  by  authors  or  vejretation  area  according  li>  Aldruli  19(i:). 

'Obs.=  observational  data,  Hnni.=  rumen  contents. 

"Key  to  references:  (l)AdaiMS  1959;  (2)Aldous  and  Sniitb  19.%;  i3IHalduin  and  I'alton  193,S;  (4U3eale  and  Darbv  1991;  (.5)Bergenid  1972:  Ui'Boertje  19S4; 
(7)Collins  1977;  (.S)Cowan  1945;  (9)Crawford  19S2;  ( l{))Descbamp  et  al.  1979;  ( 1 1  )Ilarl()\v  J9fil ;  ( 12)IIarlow  and  Iloojx-r  1971;  (1.3)Harper  1962;  (14)Healv  1971; 
(15)Hillandriarris  194.3;  (16)IIungcrford  1970:  (17)IIunlI979;(lS)Kels;illH)6S(19*Kirkpatricket;il.  1969;(2())I^opoldetal.l95l;(21)Lovaas  195S;(22)Mc<;;affen- 
et  al.  1974;  (23)Scliencket  al  1972;  (24)Sc()tter  19R7;  (25).Sliort  1971 ;  (2R)Skinn<T  and  Teller  1974:  ,27iSkooii  196.S:  i28)Tliill  .md  Martin  19.Sfi:  l29n-liill  et  ;il.  199(): 
(3())Van  Vreede  19S7;  (31)\Vallmo  et  al.  1972 


1992] 


MVCOIMIACY  BY  CEKN'IDS 


323 


Late  summer  mid  fall  are  o;enerall\'  the  sea- 
sons of  greatest  imisliroom  consumption,  prob- 
ably because  mushroom  production  is  general  K 
greatest  then.  Though  mushroom  biomass  pro- 
duction is  seldoiu  recorded  in  diet  studies,  se\  - 
eral  authors  note  that  mushroom  production  is 
triggered  b\  tall  rains  (Te\is  1952,  Hungerford 
1970,  Umess  1985). 

The  mushroom  species  most  consumed  b\ 
deer  are  not  precisely  knowai  because  species 
are  seldom  recorded  in  diet  sunews  and  prefer- 
ence studies  ha\"e  not  been  conducted.  In  addi- 
tion, species  identification  is  rare  because  most 
wildlife  researchers  are  not  ac(juainted  with 
common  mushroom  species  and  professional 
taxonomic  help  is  difficult  to  obtain  (Cowan 
1945).  In  most  field  studies,  nuishrooms  are 
categorized  into  groups  such  as  "field  mush- 
rooms," "mixed-mushrooms,"  or  simpK  "fungi." 
Howe\er,  when  listed,  species  of  the  following 
genera  are  consistently  taken  by  deer:  Amanita 
(Hungerford  1970),  Annillaria  (Healv  1971, 
Miller  and  Halls  1969),  Boletus  (Cowan  1945, 
Hungerford  1970,  Beale  and  Darb\-  1991), 
Chivaria  (Dixon  1934),  Clitocybe  (Cowan  1945, 
Beale  and  Darby  1991),  Cortinarius  (Hunger- 
ford 1970),  Morchella  (Cowan  1945),  Lactarius 
(Millerand  Halls  1969),  Lentinm  (Dixon  1934). 
Pohjponis  (Skinner  and  Teller  1974),  Rus.siila 
(Cowan  1945.  Millerand  Halls  1969.  Hunger- 
ford 1970).  and  .S/////,/,s  (Miller  and  Halls  1969). 

Mushroom  Consumption  In  Elk 

Elk  {Ceixiis  claplius)  diet  studies  rarely 
record  fungi  as  a  component.  An  extensi\e  liter- 
ature rexiew  of  elk  food  habits  in  1973  did  not 
mention  mushrooms  as  a  recorded  food  item 
I  Kufeld  1973).  However,  at  least  foiu"  studies 
lia\e  recorded  mushrooms  as  a  component  of 
t'lk  diets  (Table  1).  Composition  estimates  range 
from  a  trace  to  as  high  as  757c  on  a  diy-weiglit 
basis  (Collins  et  al.  1978).  As  with  deer,  mush- 
room consiunption  is  greatest  during  s(\i,sons  ol 
greatest  axailabilitv — late  summer  and  fall. 

It  seems  reasonable  to  assume  that  nnisfi- 
room  species  sought  bv  deer  would  also  Ix' 
acceptable  to  elk,  though  evidence  is  lacking. 
Collins  ( 1977)  listed  species  oi'Alciiria,  Boletus, 
and  Russula  as  important  and  "highly  preferred" 
dietar\-  components. 

Mushroom  ('onsumption  In  ( Caribou 

Mushrooms  lia\e  often  l)een  recoixled  as 
\er\-  palatable  and  highl\-  sought  dietan'  items 


in  caribou  {Ran^ifcr  tarandus)  diets.  When 
nuishrooius  are  axiiilable,  the\'  mav  constitute 
l()-259f  of  caribou  diets,  but  tlicx  inava\'erage 
as  nuicli  as  45%  (Table  1)  and  ha\e  been 
recorded  as  liigh  as  84%  in  one  individual 
(Skoog  1968).  Even  in  winter,  reindeer  "unerr- 
ingly" detect  snow-co\ered  frozen  mushrooms, 
"consuming  them  greedily"  (Karae\-  1968). 
Boertje  (1981)  reported  that  most  genera  of 
nuishrooms  are  taken  without  hesitation  b\-  car- 
ibou. Mushrooms  of  the  genera  Boletus,  Coph- 
luis,  Laciarius,  Li/coperdoii.  Morchella,  and 
Russula  ha\e  been  listed  as  major  dietaiT  com- 
ponents ( Karaex- 1968,  Skoog  1 968.  Boertje  1981). 

NUTHITINE  VaLUP:  OF  MU.SIIH{ )( )MS 

Man\  authors  state  that  deer,  elk.  and  cari- 
bou "stronglv  prefer"  mushrooms  and  in  some 
cases  actually  traxel  from  site  to  site  seeking 
mushrooms.  The  obvious  question  is.  why? 
What  nutritional  benefits  do  cenids  gain  from 
fungi?  Some  authors  consider  nuishrooms 
nearly  devoid  of  nutrition,  while  others  suggest 
they  compare  favorabK  with  sovbeans  or  spin- 
ach (C^iisan  and  Sands  1978). 

Little  is  knowai  about  the  tnie  nutritiv  e  \  alue 
of  mushrooms  since  few  comprehensive  studies 
have  been  conducted.  Crisan  and  Sands  (1978) 
conducted  a  thorough  literature  rexiew  on  the 
uutiitixe  xalue  of  xxild  luushrooms  to 
monogastrics  (e.g.,  humans).  Sexeral  range  and 
xxildlife  scientists  haxe  collected  and  analxyed 
mushrooms  prominent  in  ruminant  diets.  But, 
the  nutritional  procedures  used  bx  most  range 
and  w  ildlif(^  scicMitists  xx'ere  designed  to  analx"ze 
grass(^s  and  forbs.  W'licu  these  procedures  are 
applied  to  mushrooms,  the  results  are  often 
incorrectlx  interpreted  because  mushrooms  are 
much  different  from  xasciilar  plants  in  their 
chemical  composition.  Further  information  on 
the  nutritixc  \alue  of  inushrooms  can  be  gained 
Iroiu  research  on  mxcophagx'  by  insects  and 
small  mammals.  The  folloxxing  discussion  is  a 
sumnuuA'  and  inteipretation  of  nutrition  studies 
to  assess  the  \alue  of  inushrooms  to  ruminant 
animals. 

Moisture  Content  of  Nbishrooiiis 

Over  80%  of  the  fresli  xxeight  of  most  mush- 
rooms is  water  (Table  2).  This  large  x\ater  pro- 
portion re(juir(\s  that  the  consumer  eat  large 
xojumes  to  obtain  nutritional  benefit,  although 
hitih  water  content  rarelx  restricts  intake.  The 


324 


Great  Basin  Naturalist 


[Volume  52 


Table  2.  Nutritive  value  and  digestibility  of  wild  nuishroonis"'. 


Initial      (Jnule  N-free 

Composite  samples  based  on:       moisture  protein     Ash       Fat     extract  Fiber 


Cal 


Plios-    Digesti- 
pliorus      l)ilit\'    Source' 


Species  a\'ailable 

Species  a\'ailable 

Species  available 

Species  in  cattle  diets  (summer) 

Species  in  cattle  diets  (fall) 

Species  available  (winter) 

Species  available  (spring) 

Species  available  (summer) 

Species  available  (fall) 

Species  in  deer  diets 

Species  in  elk  diets 

Species  in  caribou  diets  (summer) 

Species  in  caribou  diets  (fall) 

Species  in  caribou  diets  (winter) 


34.8 


8.1        4.8      31,6 


23.0        9.0       5.0      48.0 


21.5 


(S.6 


3.9      54.2 


- 

22.0 

- 

25.0 

89.4 

22.1 

87.6 

23.1 

87.2 

29.0 

85.9 

24.8 

88.9 

21.3 

89.5 

24.1 

- 

34.7 

- 

35.3 

- 

40.0 

20.8  (cmde) 

- 

- 

- 

( 

]5.0(cnide) 

- 

- 

- 

5 

]3.8(cnide) 

0.09 

0.56 

- 

4 

- 

<0.10 

0.42 

- 

2 

- 

<0.10 

0.55 

- 

2 

- 

0.08 

0.46 

58.8 

1 

-         - 

0.07 

0.47 

64.7 

1 

- 

0.05 

0.53 

56.6 

1 

- 

0.04 

0.53 

59.9 

1 

- 

- 

- 

80.8 

6 
6 

3 

31.7  (NDF) 

0.03 

0.70 

90.0 

31.5  (NDF) 

0.03 

0.71 

9t).() 

3 

29.9  (NDF) 

0.03 

0.79 

91.0 

3 

■'.^11  (lata  expressed  as  a  '7i  (it  (In  matter  except  initial  moist  lire,  wliieli  is  expressed  as  5i  of'fresli  weiylil. 

''Kev  to  referenees:  (HBlair  et  al.   19S4:  <2iB|iii;stad  and  l),ilr\mple  1968:  (.3)Hoertje  1981;  (4)Cns,in  and  Sands   1978:  i5iKelsall  1968;  (6)Pallesen  1979; 
(7)SvTJala-Qvist  1986. 


addition  of  water  to  the  Rimen  per  se  has  little 
effect  on  intake  because  it  is  easily  absorbed  or 
removed  (Van  Soest  1982).  Mushrooms  ma\'  in 
fact  be  an  important  somx-e  of  water  for  some 
mammals  (Fogel  and  Trappe  1978). 

Mushrooms  as  an  Energy-  Source 

Mushrooms,  like  tnie  plants,  contain  lipids 
(or  fats),  nonstructural  carbohydrates,  and  fiber 
that  are  all  used  as  energ\'  sources  b\-  nmiinants. 
The  average  gross  energ)'  of  mushrooms  ranges 
from  300  to  400  kcals  per  100  grams  diy  weight. 
Fleshv  fungal  tissue  compares  flivorably  ^^^th 
many  fruits  and  vegetables  but  is  less  rich  in 
energ\than  seeds  or  nuts  (Martin  1979). 

The  fat  content  of  edible  imishrooms  ranges 
from  <1%  to  as  high  as  20%  (Crisan  and  Sands 
1978).  On  average,  however,  mushrooms  con- 
tain 2-6%  fat.  The  fat  component  of  fungal 
tissue  includes  free  fatt^■  acids,  mono-,  di-,  and 
trighcerides,  sterols,  sterol  esters,  and  phos- 
phoHpids. 

On  a  diy-weight  basis,  nuishroonis  are  pri- 
marily composed  of  nonstructural  carboln- 
drates  (nitrogen-free  extract  [Table  2]).  A  large 
\ariet\of  compounds  make  up  the  carbohydrate 
components,  including  pentoses,  medivl  pen- 
toses, hex(xses,  disacchaiides,  amino  sugars, 
sugar  alcohols,  and  sugar  acids  (C^risan  and 
Sands  1978).  By  compaii.son,  the  most  promi- 
nent nonstnictural  cad)oh\clrates  in  green 
plants  are  fnictosans,  sugars,  de.xtrin,  and  starch 
(Trlica  1977). 

In  plants  most  energ\-  available  to  nuninants 


comes  from  the  microbial  degradation  of 
fibrous  cell  walls.  However,  fungal  cell  walls  are 
much  different  from  those  of  higher  plants.  The 
primaiy  component  of  fungal  cell  walls  is 
chitin, whereas  plant  cell  walls  are  mostly  cellu- 
lose (Crisan  and  Sands  1978,  Martin'  1979). 
Chitin  is  a  N-acet)lglucosamine  polymer  linked 
with  |3-1,4  bonds  similar  to  cellulose.  Unlike  the 
fiber  of  higher  plants,  chitin  contains  a  signifi- 
cant proportion  of  nonprotein  nitrogen  as  an 
amino  sugar.  A  |3-glucan,  with  (3-1,3  linkages  and 
3-1,6  branches,  also  forms  a  part  of  the  cell  wall 
(Martin  1979).  AdditionalK;  lignin  and  pectin 
are  not  known  to  occur  in  fmigi. 

Protein  Content  of  Mushrooms 

Early  investigators  used  the  term  "\egetable 
meat"  to  describe  nnishrooms  because  anaK'sis 
rexealed  that  nati\e  mushrooms  contain  20- 
50%  of  their  dn-  matter  as  protein  (Peck  1895). 
More  recent  studies  on  nnishroom  protein  con- 
tent suggest  tliat  muslu-ooms  probabK'  rarely 
reach  50%  protein  \)\  dn'  weight.  Howexer, 
relati\eK' speaking,  mushrooms  are  an  excellent 
protein  soiux-e.  There  is  extreme  \ariation  in 
protein  content  from  a  \o\\  of  about  4%  to  as 
high  as  44%  depending  on  species,  stage  of 
growth,  and  emironmental  conditions  (Crisan 
and  Sands  1978).  B\-  comparison,  fresh-cut 
alfalfa  {Mcdicago  safiva)  is  generalK  16-19% 
protci)!  (jurgens  1978). 

('rude  protein  is  iisualK"  calculated  b\  nnilti- 
j)l\  ing  total  nitrogen,  determined  by  Kjeldahl 
anaKsis,  bv6.25.  This  correction  factor  is  based 


19921 


MVCOl'II.UiY  hvCkhmds 


325 


on  the  assumptions  tliat  most  proteins  contain 
\6%  nitrogen,  that  these  proteins  are  eom- 
pletely  cligestil)le,  and  that  amounts  of  non[)ro- 
tcMU  nitrogen  in  the  cell  are  negHgihle.  Since  a 
suhstantial  aiiioimt  ol  nitrogen  in  mushrooms  is 
in  chitin  and  other  nonprotein  compounds,  such 
as  urea  and  micleic  acids,  ('lisan  and  Sands 
(1978)  suggested  a  correction  term  hased  on  the 
assumption  that  onK  liV/r  of  the  nitrogen  in 
nnishrooms  is  in  the  lorni  ol  (Ugestible  protein 
{7()7cN  °  6.25  =  4.38).  This  correction  tcM-m  of 
4.38%  ma\'  he  consenatixe  wiien  considering 
tlie  use  ol  mushrooms  b\  niminants  and  com- 
paring nnishrooms  to  other  forage  eaten  l)\' 
ruminants.  Onlx*  60-70%  of  the  nitrogen  in 
fungal  tissue  is  in  the  form  ol  protein  (Moore- 
I  .andecl<er  1982).  Howexer,  tliis  estimate  is  sim- 
ilar to  the  proportion  of  nitrogen  in  proteins  in 
forage  plants  (60-80%;  \'an  Soest  1982).  Fur- 
tliermore,  nonprotein  nitrogen,  such  as  urea,  is 
leadih'  con\erted  to  ammonia  h\  rumen 
microbes  and  is  either  used  for  microbial  growth 
or  absorbed  across  the  rumen  wall.  The  nitrogen 
fraction  of  chitin  is  iniaxailable  to  monogastrics 
but  is  probabh'  con\erted  to  microbial  protein 
in  the  mmen.  In  fact,  chitinous  nitrogen  may  be 
more  available  to  ruminants  than  the  cell-wall 
nitrogen  of  higher  plants  due  to  the  lack  of  lignin 
in  fungi. 

\'itamin  and  Mineral  Composition 
of  Mushrooms 

.\bishrooms  are  a  good  scjurce  of  several 
vitamins  including  the  B  comple.x  and  vitamin  C 
(Change  1980,  Crisan  and  Sands  1978).  How- 
ever, these  are  not  essential  vitamins  for  rumi- 
nants l)ecause  thev  can  be  sviithesized  bv  nunen 
microbes  (\an  S()(\st  1982).  Additionallv,  nursh- 
rooms  are  basicalK  devoid  of  \  itamins  A  and  1^, 
which  are  es.sc^ntial  dietan  comj)onents  for 
ruminants. 

Mushrooms  accumulate  minerals  Irom  the 
soil  and  plant  material  on  wliich  thev  grow. 
Therefore,  mushrooms  probablv  contain  all  the 
minerals  present  in  their  growth  substrate 
(Crisan  and  Sands  1978).  Stating  average  min- 
eral concentrations  mav  l)e  misleading  becan.se 
mineral  concentration  \  aries  greatlv  depending 
on  species  and  .soil  feitilitv.  For  example,  though 
potassium  level  averages  2%  (in  24  species  from 
.several  locations),  it  varies  from  0.18  to  4.8% 
(Crisan  and  Sands  1978). 

The  most  abundant  minerals  in  mushrooms 
are  potassium,  averaging  about  2%  diy  weight. 


and  pliosj)liorus,  averaging  about  0.75% 
(Change^  1980,  C^risan  and  Sands  1978,  Martin 
1979).  Both  mineral  levels  exceed  maintenance 
re(|uirements  of  most  weaned  ungulates  (ba.sed 
on  sheep  and  cattle  recjuireinents;  jurgens 
1978).  Mushrooms  also  contain  calcium  l)ut  at 
lower  concentration  than  [)ho.sphorus  or  potas- 
sium. H ( )\\ ever,  calciu m  c( )ncentratioi i  averages 
0.14%,  v\hich  would  not  meet  calcium  recjuire- 
ments  ol  weaned  dew  (  Ullrev  et  al.  1973).  (-al- 
ciinn  is  often  in  excess  of  ruminant  needs  in 
otliei-  forages,  wliiie  phosphorus  is  more  com- 
mouK  inade(|uate. 

Dig(^stibilitv  ol  Musfirooms 

The  degradation  ol  lungal  cell  walls  requires 
chitinase  and  p-1.3  and  (3-1,6  glucanases 
(Martin  1979).  Cliitin  is  degrachible  in  tlu> 
rumen  becan.se  of  chitina,se  activitv  bv  rumen 
microbes,  although  there  mav  be  an  adaptation 
period  necessaiy  to  obtain  adecjuate  levels  of 
chitinase  activitv  (Clieeke  1991).  The  abilitv  of 
rumen  microbes  to  degrade  the  p-glncans  in 
lungal  cell  vxalls  is  unkiumn. 

Tlie  in  vitro  digestibilitv  of  nnishrooms  is 
ven  high  r(4ative  to  other  ungulate  forages 
(Table  2)  and  mav  exceed  90%  in  some  cases. 
Consequentlv,  identification  of  nnishrooms  in 
fecal  analvsis  is  rare  (Boertje  1981). 

iMl'LICATIOXS  OF  MY(  :()IM  lACV  liV 

Di:kh  AM)  Ei.K 

To  conchid(^  this  discussion  it  is  lair  to  ask. 
What  difference  does  it  make  if  dcc\\  elk.  or 
caribou  eat  nnishrooms  or  not?  Mvcophagv  bv 
cenids  mav  be  important  for  several  reasons. 
First,  nnishrooms  niKk)ubtedlv  make  an 
im|:)ortant,  though  s])oradic.  contribution  to 
cenid  nutrition  in  musli room-rich  environ- 
ments. Mushrooms  are  highlv  preferred  and 
nutritions  foods  for  cervids,  particularlv  in  late 
snminer  and  fall  in  forested  areas  of  western 
North  .'\mei  ica  and  throughout  the  vear  in  the 
Southeast.  .Mushrooms  mav  be  a  particularlv 
im[)ortant  protein  and phosphonis  source  in  late 
season  when  main  forages  vield  onlv  enough 
digestible  d\y  matter  to  meet  basal  energv 
re(iuirements  (Short  1975,  Blair  et  al.  1984). 
Therefore,  even  a  fev\-  bites  of  mnshrooins  bv  an 
herbivore  may  contribute  substantiallv  to  meet- 
ing the  nutritional  requirements  and  helping  to 
balance  nutrients  obtained  from  other  forages 
of  (^uite  different  composition. 


326 


Great  Basin  Natuhai.ist 


\\<>\ 


iiiiic  o2 


Second,  mushrooms  may  attract  herbivores 
to  mature  and  stagnated  forest  areas  that  might 
otherwise  go  unused  as  foraging  areas  (Rasirius- 
sen  1941,  Collins  et  al.  1978,  Warren  and  Mys- 
terud  1991).  Additionally,  mushrooms  may 
become  an  important  dietaiy  supplement  when 
herbi\"ores  are  forced  to  seek  densely  forested 
areas  for  protection  from  biting  insects  or  pred- 
ators (Bergemd  1972).  Mushroom  production 
is  usucilK"  greatest  in  dense  forested  areas,  in 
part  because  mushrooms  do  not  require  sun- 
light for  o;ro\\th. 

Finallw  fungi  pla\'  an  important  s\nibiotic 
role  in  m\corrhizal  relationships  with  several 
conifer  species,  including  ponderosa  pine 
(Kotter  1984).  Since  the  spores  of  fungi  are 
apparentK'  not  destroved  in  the  nmien,  herbi- 
\"ores  ma\'  sene  as  \ectors  for  fungal  spores  to 
initiate  nncorrhizal  associations  (Fogel  and 
Trappel978). 

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Ullrev,  D.  E.,  ET  AL.  1973.  Calcium  re(jnirements  of 
wetined  white-tailed  deer  fawns.  Journal  of  Wildlife 
Mtmagement  37:  187-194. 

Urness,  p.  J.  1985.  Managing  lodgepole  pine  ecosystems 
for  game  and  rtuige  values.  In:  Lcxlgepole  pine:  the 
species  and  its  management.  Symposium  Proceedings. 
Cooperative  E.xtension  Senice,  Washington  State  Uni- 
versity, Pullman. 

Van  Soest.  P.  J.  1982.  Nutritional  ecolog\  of  the  ruminant. 
Cornell  University  Press,  Ithaca,  New  York. 

Van  Vreede.  G.  T.  1987.  Seasonal  diets  of  white-tailed  deer 
in  south-central  Oklalioma.  Unpublished  master's 
thesis,  Texas  Tech  UniversiK,  Lubbock. 

Wali.mo.  O.  C,  W.  L.  RJ'^celin.  and  D.  W.  Rf:iciiEin". 
1972.  Forage  use  b\'  mule  deer  relative  to  logging  in 
Colorado.  |onrnal  ofWildlile  Management  .36:  1025- 
10.33. 

Warren,  J.  T,  and  1.  Mvsterud  1991.  P'migi  in  the  diet 
of  domestic  sheep.  Rangelands  13:  168-171. 

Willii\Ki:n  R.  II.  1969.  New  concepts  of  king(k)ms  of 
organisms.  Science  163;  1.50-160. 


Received  15  June  1992 
Accepted  25  September  1992 


Great  Basin  Naturalist  52(4).  pp.  328-334 


TERRESTRIAL  VERTEBRATES  OF  THE 
MONO  LAKE  ISLANDS,  CALIFORNIA 


Michat'l  L.  Mcirrison',  William  M.  Block-,  Joseph  H.  Jchl,  Jr. '.  and  Liiinea  S.  Hall'^ 

.ABSTHAcr. — \\\'  compared  \ertebrate  populations  between  the  two  major  islands  (Paoha  and  Nes^it)  in  Mono  Lake, 
Calirornia.  and  tlie  atljac(^nt  mainland  to  further  elucidate  the  mechanisms  underlying  island  colonization.  Deer  mice 
{Peroini/sais  maiticuhitn.s)  iuid  montane  voles  (Microtus  iin>nt(iniis)  were  captured  on  Paoha,  but  only  deer  mice  were 
captnred  on  Negit.  In  contrast,  eight  .species  of  rodents  were  captured  on  the  mainland.  Oxerall  rock^nt  abundance  on  Paoha 
and  the  mainland  was  similar,  but  on  Negit  it  was  about  three  times  greater  than  on  Paoha  or  die  mainland.  Adult  deer  mice 
from  Faoiiawi're  significanth'  (F  <  .05)  smaller  in  most  e.xternal  bodv characteristics  than  m;unland  mice.  Coyotes  (Canis 
latniiis)  and  one  or  two  species  of  lagomoiphs  were  observed  on  the  islands  and  the  nuiinland.  No  amphibians  or  reptiles 
wt-re  found  on  the  islands;  both  occmred  in  low  numbers  on  the  nnrinland.  Ratting  and  hinnan  transport  are  probable  means 
of  colonization  for  mice  and  voles.  Tile  occiu'rence  ot  co\otes  on  the  islands  max  have  moditied  liistoric  predator-pre\' 
relationshi|is.  and  thus  the  population  of  rodents  and  lagomoiphs. 

Kcij  iLonls:  Moiiii  Lake,  islaiids.  coloniziilidii.  Peromyscus  maniciilatns,  Microtus  montanns. /c//if/ /i/iV/gc. 


Lslancl  animal  popukitions  ha\c  attracted 
much  scientific  interest  because  they  sene  as 
natural  experiments  for  the  study  of  cok)niza- 
tion,  dispersal,  extinction,  competition,  and 
otlier  biological  processes  (MacArthur  and 
Wilson  1967).  Because  islands  are  stnall  and 
isolated,  populations  inhabiting  them  are  more 
N'ulnerable  to  stochastic  ewnts  than  their  main- 
land counteiparts. 

Most  previous  studies  of  island  zoogeogra- 
phy ha\e  emphasized  patterns  of  island  occu- 
pancN',  morpholog\',  and  genetics  of  restricted 
subsets  of  the  islands'  fauna  (reviewed  b\' 
]^'lt()iK>n  and  Ilanski  1991 ).  Our  goals  were  to 
couipare  island  and  mainland  \eii:ebrates  of 
Mono  Lake  and  the  surrounding  Mono  Basin, 
California,  in  light  of  natural  and  human-infhi- 
enced  processes.  This  area  was  of  interest 
because  no  thorough  sunevs  had  been  con- 
ducted on  the  islands  of  this  large  saline  lake, 
and  because  of  possible  changes  in  local  ecolog\- 
as.sociatcnl  with  falling  lake  levels  from  water 
diversion  lor  human  consumption. 


Mono  Basin  and  Islands 

Mono  Basin  is  the  In  drolotjic  drainage  basin 
for  Mono  Lake.  The  l^asin  is  surrounded  by  the 
Sierra  Ne\'ada  to  the  west  and  the  Great  Basin 
ranges  to  the  north,  east,  and  south.  Mono  Lake, 
estimated  at  500, 000  vears  of  age,  is  one  of  the 
oldest  lakes  in  North  America.  Because  no 
water  naturalK  flows  out  of  the  basin,  and 
because  of  long-term  evaporation  coupled  with 
water  diversion,  the  lake's  salinitv'  is  aliout  2.5 
times  that  of  the  ocean.  In  October  1986  the 
surface  area  of  the  lake  was  about  177  knr 
(Mono  Basin  Ecosvstem  Stud\  (xnnmittee 
1987). 

There  are  t^\■o  major  islands  in  the  lake: 
Paoha  Lsland  at  about  7.7  km'  and  Negit  Island 
at  onK  about  1.3  knr  (Fig.  1).  Paoha  formed 
liom  volcanic  activitv  and  an  uplift  of  lake  sedi- 
ment some  time  between  1723  and  1850  A.D. 
Negit  formed  as  a  result  of  a  series  of  eruptions 
beginning  about  200  A.D.  (Mono  Basin  Ecosys- 
tem Studv  (>'oiMmittee  1987,  U.S.  Forest  Senice 
1989). 


ill72(l  ,uhI  Willi. ■  \1. mill. nil  Ki 


n<-|iartiiiriil.,rK(m'sti-\^UKl  Hc-soiucc  Maiui<;riiic-iil,  Unu<isil\  ,il  ( :.ilil,,iiii,i   B,  lit  lr\    Calil. 
Lin^'Shvrl,  liisliop,  (:alil(inii.i9:V51  I, 

-USIM  Kciivst  St-nuc,  HmU  Mcuiilain  Kiiicst  and  Hangc  K\|..i  iinrnl  Si.iIi.h,    Inu^l  S,  1,11,  ,  s  l,.il,    l\ni|.,     \n/.,ii.i  S-i2S7. 

'^Sca  World  Hcsca.vli  InsliluU-,  llulihs  Marine  HcscMi-chCfiilri    ITOIISmilli  Si s  ILnmI   San  I  )i,  00  (  .ilil.ania  l)i!l(W 

■  I'reseiit  address:  Sc-li(ii.l  i)(  Henewahle  Nadual  Uesouree.s.  Uni\.  imI\  nl  Ari/oii.i,  ria.s.iii,  Ai  1/011,1  s"-)721 . 


328 


1992] 


Mono  Laxe  Islands  N'ehtebrates 


329 


Fig.  1.  Mono  Lake  and  the  two  major  islands,  Paoha  and  Negit;  small  islets  are  not  shown.  The  Ixwes  intlicate  the  generd 
location  of  the  1991  stud\' plots;  stippling  indicates  the  marsh  on  Paoha.  Redrawn  from  \arions  U.S.  Forest  Service  maps. 


Beginning  in  1941  the  major  streams  enter- 
ing Mono  Lake  were  dixerted  and  tlieir  water 
was  transported  to  Los  Angeles,  California.  This 
dixersion  lowered  the  lake  lexel  abont  15  m  In" 
19S1,  to  the  modem  historic  low.  and  also 
decreased  the  lake  volnme  b\-  about  50%  ( Mono 
Basin  Ecos\stem  Stud\'  Committee  1987, 
Botkin  et  al.  1988).  Although  dixersions  ha\e 
been  halted,  a  continuing  drought  (through  at 
least  1992)  that  began  in  1986  has  pre\  ented  an\ 
significant  rise  in  the  lake  le\"el. 

Paoha  and  Negit  islands  are  located  along  an 
a.\is  ninning  peipendicidar  from  the  northern 
shore  of  Mono  Lake,  with  Paoha  the  farthest 
away  and  about  1  km  from  Negit.  Since  its 
formation,  Negit  has  been  separated  from  the 
mainland  bv  0  to  >3  km  (Mono  Basin  Ecos\s- 
tem  Stud)  Committee  1987:  Figs.  1.3  and  6.1). 
Howexer,  no  mainland  coiniection  with  Negit 
existed  since  the  formation  of  Paoha  until  the 
late  197()s;  the  next  most  recent  laud  bridge 
apparenth-  occurretl  about  500  xcars  before 
present.  During  our  stud\-  in  1990-91,  Negit 
was  separated  from  the  mainland  In'  sexi'ral 
lumdred  meters  of  nuidflats  and  a  fex\'  meters  of 
shalloxx'  xx'ater;  this  area  is  referred  to  herein  as 
the  land  bridge. 

^^'eknox\■of  onlxtxx'opnnious  small  mauunal 
trapping  efforts  on  the  islands.  In  1975  W.  M. 


Hoffmann  (unpublished  report)  captured  no 
small  nuunmals  on  Paoha  in  one  night  of  effort. 
|.  IL  Harris  (personal  conununication)  cap- 
tured deer  mice  {Peromyscus  maniculatus)  on 
Negit  during  sexeral  dax's  of  trapping  in  the  earlx' 
198()s.  One  of  us  (fRp  has  nuuU^  repeated  xisits 
annuallx  to  the  islands  since  1980,  making xisual 
obsenations,  but  not  trapping.  .\1I  otlicr 
accounts  of  the  islands"  manunal  hiunaare  troui 
recollections  of  earlv  settlers  and  local  residents 
(e.g.,  Fletcher  1987.  personal  connnunication 
xxith  JRJ). 

Study  Akkas 

I'aoha  Island  can  be  dixided  into  txxo  general 
X  egetatixe  /ones:  a  small  (about  2  ha)  spring-fed 
marsh  along  the  southeastern  shore,  and  the 
remaining  nonmarsh  xegetation.  \egetation  in 
tlu^  marsh  is  composed  of  rush  (Junciis  effusus), 
bullrush  {Scii~})iis  americanus),  saltgrass  {Dis- 
ticltlisspicata),  foxtiiil  {Hoirleiunjtibatum),  and 
bassia  {Bas-sia  Iii/ssopifolia).  Nonmarsh  areas 
aic  dominated  bx  greasexx'ood  (Sarcohatiis  vcr- 
inicuhitus)  and  hopsage  (Grai/ia  spinosa):  sage- 
brush {Aiiemi-sid  tridentaia)  is  present  but  rare. 
(brasses  and  herbaceous  plants  are  scarce  and 
concentrated  in  the  marsh  and  one  small  (about 
0.3  ha.)  gnisslaud  site  located  upslop-  about  300  m 


330 


Great  Basin  Naturalist 


[X'olume  52 


from  the  marsh.  The  grassland  area  is  domi- 
nated bv  exotic  clieatgrass  [Bronms  tectoniin). 
Negit  Island  lacks  any  marsh  \egetation  and  has 
no  permanent  freshwater.  The  upland  is  similar 
to  Paoha  e.xcept  for  more  cover  by  sagebmsh. 
Dominant  \egetation  on  the  mainland  plots  is 
sagebmsh,  rabbitbrush  {CJinjsotlunnnus  nausc- 
osus),  bitterbmsh  {Purshia  triclentata),  and  scat- 
tered individuals  of  greasewood,  curlleat 
maliogany  {Cercocatyus  k'difolitis),  and  desert 
peach  (Pninns  ai}ch'rso)iii).  Vegetation  in  the 
basin  was  detailed  b\-  Burch  et  al.  (1977).  Soils 
are  a  loose  mixture  of  sand,  gra\'el,  ash,  and  silt 
(Loeffler  1977). 

In  1990  trap  lines  were  established  to  deter- 
mine species  composition  and  approximate  dis- 
tributions of  small  mammals  on  Paoha  and 
Negit.  Specific  trap  locations  were  based  on 
ease  of  boat  landing  and  proximit\'  to  the  next 
nearest  trapping  location;  adjacent  trap  lines 
were  at  least  200  m  apart. 

In  1991  we  systematicalK*  established  10 
fixed  study  plots  (50  x  20  m)  on  Paoha  Island 
and  5  on  the  adjacent  mainland  to  compare 
mammals  on  the  island  and  mainland;  island 
plots  were  placed  in  the  marsh  (3  plots)  and  diy 
shrub  vegetation  (7  plots).  All  mainland  plots 
were  located  to  the  north  and  northeast  of  Black 
Point  on  the  northwest  shore  of  Mono  Lake. 
This  location  was  selected  because  its  vegeta- 
tion resembles  the  dominant  \egetation  on 
Paoha  Island  and  represents  a  likeK'  sovux-e  for 
terrestrial  animals. 

Methods 

Small  Mammal  Lixe-Trapping 

All  traps  used  during  this  stud\'  were  large 
(7.6  X  8.9  X  22.9  cm"  [3  x  3.5  x  9  inch]) 
Sherman  live-traps.  In  1990  trapping  was  done 
on  Paoha  Island  on  27-29  April  and  23-25 
August,  Negit  Island  on  27-29  April,  and  the 
mainland  on  4-7  September.  Trap  spacings 
ranged  fnjm  10  to  20  m  and  were  based  on 
axailabilit)-  of  vegetative  coxer.  Traps  were 
baited  with  rolled  oats  and  peanut  butter  and 
checked  each  morning  for  1-3  davs  depending 
upon  weather  conditions  and  thus  access  to  the 
islands.  Mainland  trapping  in  1990  was 
restricted  to  a  marsh  on  the  northern  shore  of 
the  lake.  Captures  were  identified  to  species, 
sex,  and  age  and  were  measured,  iriarked,  and 
released  at  the  trap  location.  Measurements 
between  sexes  and  betAveen  island  and  main- 


land populations  of  deer  mice  were  compared 
using/  tests  (Zar  1984:126-131). 

In  199 1 ,  within  each  plot  on  Paoha  described 
abo\'e,  18  large  Sherman  live-traps  were  placed 
at  lO-m  spacings  (1  row  of  6  traps  along  each 
long  axis  of  a  pk)t).  Each  plot  was  trapped  for  a 
total  of  54  trap-nights  and  da\s  (i.e.,  traps  were 
left  open  constantly  for  3  days  and  checked  both 
during  the  morning  and  in  late  afternoon) .  Traps 
were  baited  and  animals  handled  as  in  1990. 
Mainland  and  Paoha  traps  were  run  7  Mav-24 
June.  Trap  lines  were  rim  on  Negit  4-5  August, 
as  described  for  1990.  Data  are  reported  here  as 
the  number  of  new  individuals  (i.e.,  excluding 
recaptures)  captured  per  100  trap-nights;  we 
assume  that  this  measure  of  capture  success  is 
an  adequate  index  of  actual  population  abun- 
dance. Indices  of  abundance  were  compared 
using  chi-square  goodness  of  fit  (Ziu- 1984:40-43). 

Other  Sune\'s 

During  1991,  one  4.2-L  (1-gal)  can  was 
placed  near  the  center  of  each  trapping  plot. 
Cans  were  placed  on  all  mainland  plots  and  on 
six  Paoha  Island  plots.  Each  was  coxered  with  a 
wooden  board  raised  2-3  cm  abo\e  the  can. 
Traps  were  am  4-17  da\s.  Three  additional 
traps  were  placed  in  the  marsh  on  the  southeast 
side  of  Paoha  Island,  this  being  the  most  likeK' 
location  for  shrews  (Soricidae).  Thus,  six  traps 
were  placed  in  the  marsh.  All  mainland  pitfalls 
were  opened  9-12  June;  island  traps  were 
opened  7  Ma\'-4  June. 

A  1-m"  area  in  an  open  location  near  the 
center  of  each  plot  was  selected  to  determine 
the  presence  of  mediiuii-  to  larger-sized  mam- 
mals traxeling  across  the  plot.  The  soil  in  a  track 
plot  was  smoothed  bx'  hand  and  moistened  xxith 
xxater;  fine-grained  sand  or  soil  xxas  added  as 
needed.  A  can  of  chicken-flaxored  cat  foodxx'as 
secured  at  the  center  of  each  track  plot.  Each 
plot  xxas  checked  dailv  for  three  days  for  evi- 
dence ofxxildlife  use.  One-half  of  the  studx' plots 
on  Paoha  and  three  mainland  studx  plots  xxere 
used. 

Time-constraint  sunexs  of  one-person-hour 
duiation  each  were  conducted  in  all  study  plots. 
The  species,  date,  time,  location,  and  general 
xegetation  tvpe  for  each  obserxation  xvere 
recordctl. 

Museum  Records 

\\e  obtained  records  for  all  xertebrates  col- 
lected in  Mono  Basin  from  the  Los  Angeles 


1992] 


Mono  Lake  Islands  Vertebrates 


331 


T\iil.K  1 .  Index  ofabuiulance  (iio./KX)  trap-niijhts)  for  small  niaimnals  captured  on  stiicK  plots  on  Paolia  Island  (n  =  10 
plcits    and  .idjaeent  mainland  (n  =  5  plots).  ;uid  on  Negit  Island  (trap  lini's).  Mono  Basin,  (-'alifornia.  1991.'' 


Species 


Paoha  Island  (trap-nights) 


Totd 

Total 

Total 

marsh 

nonmarsh 

island 

(108) 

(324) 

(432) 

17.6 

13.0 

14.1 

8.3 

9.0 

-S.8 

9.3 

4.0 

5.3 

5.6 

0.9 

2.1 

Negit 
Island 

( 120) 


Total 
mainland 

(.■342) 


Pcroini/sni.s  uumicnlatiis 

Male 

Female 
Micmtiis  iiioiitanus 
Pcmi^)i(itliiis  panti.s 
Dipodoiiuis  pauainintinits 
D.  microps 
Pcroiiii/.srus  hoi/lii 
Eutamiits  minbntis 
Sp('n)to)>hilus  hccclwyi 
Total 


62.5 
32.5 
30.0 


23.2 


13.9 


16.2 


62.5 


6.4 
4.1 
2.3 
0.3 
6.7 
5.3 
1.8 
0.3 
0.3 
0.3 
21.3 


■'Chi-srjiiare  anaKsis:  all  comparisons  between  total  marsli  and  total  nonmarsli  cm  Paoli.i  /'  >  .0.5;  between  P.iolia  total  island  and  \e0t  Is 
Paoha  and  total  mainland  P  >,0.5;  and  between  total  mainland  and  Ne^il  I'  ■    1)1 


1  P  <  .(X)l;  l)et%veen  total 


Counh'  MiLseuni  of  Natural  Iliston 
(LACMNH)  and  the  Museum  of  Wrtebrate 
Zoolog)',  UniversitS'  of  California,  Berkele\ 
(MVZ).  Although  no  records  were  available  for 
the  islands,  data  from  the  basin  were  sununa- 
rized  to  supplement  published  accounts  of 
mainland  \ertebrate  sun'eys.  X'oucher  speci- 
mens were  deposited  at  the  M\"Z. 

Results 

Small  \himmal  Trapping 

Only  deer  mice  and  montane  xoles 
{Microttis  montanus)  were  captured  on  Paoha 
Island.  Most  voles  were  captured  in  the  marsh; 
deer  mice  were  also  slightK'  more  abundant 
there  than  in  dn'  sliRib  plots,  but  these  differ- 
ences were  not  significant  (F  >  .  1).  The  sex  ratio 
of  deer  mice  was  sk'ewed  toward  males  in  the 
dr\'  sliRib,  but  was  about  e\(Mi  in  the  marsh 
(Table  1). 

Onlv  deer  mice  were  captured  on  N(^git 
Island.  Nh)us('  abundance  was  ai)out  4.5  times 
higher  on  Negit  than  on  Paoha  (F  <  .05),  and 
sex  ratios  were  about  e\en  (Table  1). 

Eight  species  of  small  mammals  wen^  cajv 
tured  on  the  mainland  plots  in  1991.  Cireat 
Basin  pocket  mice  {Perognathus  parvus),  deer 
mice,  and  Panamint  kangaroo  rats  (Dipodoinijs 
))(i)ia)nintiiius)  had  similar  relatixe  abundanc(\s 
and  were  the  onl\-  species  with  abundances  >5 
indi\iduals/100  trap-nights).  Except  for  the 
Great  Basin  kangaroo  rat  {Dipodomi/s  microps), 
all  species  were  captured  rareK'  (all  at  0.3  ani- 
mals/100 trap-nights).   Oxerall  abimdance  of 


small  mammals  on  Paoha  was  similar  to  that  on 
the  mainland,  but  on  Negit  it  was  almo.st  three 
times  greater  than  that  on  Paoha  (F  <  .001)  or 
the  mainland  (F  <  .01;  Table  1). 

Abundance  of  deer  mice  approximateK' dou- 
bled (F  <  .01)  on  Paoha  betvxeen  April  (earl\ 
breeding)  and  August  (end  of  breeding)  1990. 
Subadult  males  accounted  for  fiT'^'f  of  this 
increase  (Table  2),  wiiile  subadult  females 
accounted  for  onlv  67c.  Total  male  and  female 
abimdance  was  about  etjual  in  April;  the 
number  of  males  caught  increased  b)'  63%  and 
females  onlv  by  35%  in  Augvist,  although  the 
difference  was  not  significant  (F  >  .1). 

Nhile  and  female  abundances  of  deer  mice 
were  similar  on  Negit  in  April  1990;  no  compa- 
rable August  data  were  a\  ailable.  Total  abun- 
dance on  Negit  in  April  was  48%  higher  (F  < 
.05)  dum  that  on  Paoha  (Table  2). 

Adult  male  deer  mice  from  Paoha  weighed 
significantly  less  and  had  significantK  shorter 
tails,  feet,  and  tail :bodv-length  ratios  than  main- 
land animals;  bod\  and  ear  lengths  were  not 
different  (Table  3).  .Adult  females  from  Paoha 
were  significantK  less  hea\A-  than  mainland  ani- 
mals and  had  smaller  but  not  significantly  dif- 
ferent a\(M-age  measurements  for  other 
characters.  Comparisons  with  Negit  mice  were 
not  j)ossil)le  because  an  insufficient  number  of 
animals  were  measured. 

Other  Sun e\s 

Islands. — The  six  pitfalls  in  the  Paoha 
marsh  were  nm  for  13  da\s  (78  trap-da\s)  and 
captured  7.7  xoles/lOO  pitfall-days;  the  three 


332 


Great  Basin  Naturalist 


[N'olume  52 


TAIU.K2.Al.un(lancv(noyi()()trapMn,uhts)<.i7Vr<<mv,sr^^^^^^^        ^,^j,^   ^^^^   ^j^^   ^^^^^   adjacent   tO   the    plots   Oil 
»N«»/r»wA».v  on  Paolia  and  Nc-trit  islaiuis.  Mono  Lake,  Call-  ,  .  ■'  ^ 

se\  eral  occasions. 


fornia,  1990. 


Paohii 

I  Island 

Negit  Island 

April 

August 

April 

Trap-nis^lits 

290 

160 

74 

Mae 

Adult 

7.6 

S.S 

13.5 

Suhadult 

1.0 

15.0°°° 

0.0 

Juwnile 

0.7 

1.3 

0.0 

Total 

9.3 

25.0°° 

13,5 

Feniiilc 

Adult 

.5.9 

8,1 

1(),S 

Suhadnit 

.3.8 

5.0 

4,1 

Juvenile 

0.0 

1.9 

0.0 

Total 

9.7 

15.0 

14.9 

Oxerall 

19,0 

40.0°° 

36.5°° 

Clii-square  analv.si.s:  ^"P 
April  vs.  Negit  April. 


"/'<    (101.  I'.iolia  Apnl' 


sI,,i,mI  Pa„l.; 


pitfalls  in  the  chA'  shnil)  were  iiin  for  17  da\'s  (51 
pitiall-da\s);  no  animals  were  captnred.  Track 
plots  were  run  tor  3  davs  on  Paolia,  resulting  in 
a  total  effort  of  15  track-pl()t-da\s.  One  set  of 
coyote  {Canis  latrans)  tracks  was  found  on  a  plot 
in  the  marsh,  and  one  set  of  unidentified  rodent 
tracks  (likely  deer  mouse)  was  found  on  a  dn' 
shrub  plot.  Coyote  tracks  and  scat  were  seen 
throughout  botli  islands;  the\'  were  especialK 
evident  on  the  southeast  end  of  Paoha,  includ- 
ing the  marsh.  Black-tailed  jackrahbits  {Lcpiis 
calijonticiis)  were  uncommon  but  were  seen 
occasionally  on  both  islands.  Cottontails 
(Sijlvilaffis  spp.)  were  seen  rareh'  on  Negit  but 
were  not  exident  on  Paoha.  Rabbit  pellets  were 
conspicuous  on  the  islands,  indicating  that  the 
populations  had  been  greater  at  a  previous  time. 
No  herps  were  obsened  on  either  island  during 
any  sunev,  or  in  anv  vearK  island  \  isit  b\  jRf 
since  1980.  Scattered  individuals  of  sagebrush 
lizard  (Scclo})onis  gmcioiis)  were  seen  wliile 
walking  on  and  near  the  mainland  study  plots. 

Mainland. — The  five  pitfalls  w/ere  run  for 
4  days  (20  pitfiill-days).  Four  sagebrush  li/.aids 
were  captured  (20  li/ard.s/lOO  pitfall-davs). 
Track  plots  were  run  for  a  total  of  3  dav  s  on  three 
study  plots,  vvitii  one  set  of  black-tailed  jackrab- 
bit,  two  sets  of  kangaroo  rat  (species  unknown), 
and  one  set  of  unidentified  small  rodent  tracks 
obsened.  Thus,  there  were  four  separate  ani- 
mals in  9  track-plot-days.  Covote  tracks  were 
seen  on  the  plots  and  coyotes  were  heard  calling 
adjacent  to  plots.  Numerous  rabbit  and  kanga- 
roo rat  tracks  were  present  on  all  plots;  cotton- 


DlSC;USSION 

Only  tAvo  species  of  small  mannnal  (deer 
mouse  and  montane  vole)  were  trapped  on 
Paoha,  and  one  species  (deer  mouse)  on  Negit, 
compared  with  eight  species — including  deer 
mice  and  niontane  voles — on  the  adjacent 
mainland.  \''isual  and  track  sunevs  found  the 
jackrabbit,  cottontail,  and  coyote  on  Negit 
Island  and  the  mainland;  all  but  the  cottontail 
were  evident  on  Paoha.  In  contrast,  at  least  20 
species  of  small  manimals  have  been  obsened 
around  the  shores  of  Mono  Lake  (Harris  1982, 
1984).  In  addition,  weasels  {Miistcla  spp.), 
badger  {TaxkJea  taxiis),  bolicat  {Lij)ix  nifiis), 
mountain  lion  {Fclis  concolor),  black  bear 
{Ursus  a))UTicaitus}.  and  mule  deer  {OdocoiJeus 
hcinioiiiis)  occur  around  Mono  Lake  (Harris 
1982).  Furthermore,  lion  remains  have  been 
reported  from  an  islet  near  Negit  and  from  the 
vicinitA'  of  the  Negit-mainland  land  biidge 
(Mono  Lake  Conimittee,  unpublished  obsena- 
tion).  The  presence  of  niontane  voles  on  Paoha 
was  associated  vvitli  the  marsh  and  grass  vegeta- 
tion that  is  absent  on  Negit.  The  current  lake 
level  has  allowed  the  Paoha  marsh  to  expand 
onto  an  exposed  lake  shelf,  thus  increasing 
potential  vole  habitat.  The  environment  maybe 
unsuitable  on  the  islands  for  persistence  of  the 
larger  carnivores  and  deer  but  appears  suitable 
on  Paoha  (because  of  water  and  rodents)  for 
weasels  and  possibly  badgers. 

Animals  can  colonize  islands  by  svvimnung, 
rafting,  using  ice  bridges,  being  inadvertent  pas- 
sengers on  watercnift  (C\illioun  and  Greenl^aum 
1991),  intentional  or  unintentional  releases,  or 
by  flving;  all  but  flving  may  apply  to  the  animals 
discussed  herein.  The  lack  of  historic,  (juantita- 
tive  data,  however,  prevents  determination  of 
the  method(s)  and  date(s)  of  arrival  of  animals 
on  tlu^  Mono  Lake  islands.  However,  Hoffman 
(unpublished  report)  set  76  Sherman  traps  for 
one  night  (24  Mav )  in  1974  in  various  locations, 
including  in  ami  around  the  same  marsh  and 
grassland  an^as  we  trapjied.  He  caught  no  ani- 
mals but  did  locate  a  rodent  faex.  Although 
I  lof  fmans  efforts  were  minimal,  his  data  at  least 
indicate  the  presence  of  rodents  prior  to  1974. 

Although  the  earliest  historic  accounts  of 
local  Native  Americans  date  to  the  early  186()s 
(Jelil  et  al.    1984,   1988),  various  peoples  are 


19921 


Mono  LakI';  IsiandsXhrtebiuths 


io6 


T \Hl.i:  3.  Characteristics  or;ulu]tP('r(>Hi(/.sr(/.s;;i^/;i/r((/r////.s  captiiicdon  Faolia  Island.  .Mono  Lake,  and  atljaccnt  mainland 
dnrinii  1990  iuid  1991. 


Adult 

male" 

Adult  f( 

i-mde'^ 

Paoha 

Mainland 

Paoha 

Mi 

iiinli 

uid 

Characteristic 

X 

SD 

X                SD 

X 

SD 

X 

SD 

Mass  (gf 

17.2 

2.11 

IS.S               1.29°° 

18.1 

2.29 

19.9 

2.80"" 

Hod\- length  (mnO 

SI. 4 

5.13 

SI. 4              2..S7 

79.4 

6.50 

Sl.l 

3.18 

Tail  length  (nnn) 

64.5 

4.32 

67.1              5.;34° 

66.4 

6.85 

69.1 

6.34 

Foot (nnn) 

20.0 

1.07 

20.9              0.95°° 

20.0 

0.91 

20.6 

1.08 

Ivir  (mm) 

17.4 

J.OS 

17.4              1.45 

17.4 

1.04 

17.9 

1.38 

Tail/I)c)d\ 

0.79 

0.06 

0.S2            0.07° 

0..S4 

0.07 

0.85 

0.07 

■■Saiiiple  size  =  50  indi\i<liuils  eacli  urea,  cxcfpt  lor  Tiiass. 
'Sample  sizi"  =  15  inili\idiials  each  area.  e.«ept  for  mass. 
'Excliules  pregnant  teniale.s:  Paoha  ii  =  12.  mainland  n  =  1:5 
°P<  .0.5.  "P<  .01,/ test. 


thoiitilit  to  lia\(.'  \isitc>(l  tlu^  ba.sin  lor  a  iiiucli 
longer  period  (Fletcher  1987).  Western 
immigrants  began  making  trips  to  the  islands  1)\ 
the  1860s  (Jehl  etal.  1984, 1988,  Fletcher  1987). 
A  cliicken  [GaUu.s  galliis)  and  domestic  lago- 
moiph  ranch  was  established  on  Paoha  in  tlie 
late  187()s,  a  domestic  goat  {Capra  sp. )  ranch 
was  initiated  in  the  1896s  (Fletcher  1987).  and 
a  mineral  salts  and  health  spa  \entnre  was 
attempttnl  in  the  194()s.  Lagomoiphs  raised 
c()mmerciall\  werc^  apparentK  European  hares 
(Lej)us  sp.),  but  there  is  no  exidence  that  these 
hares  remained  on  Paoha  after  the  earlv  192()s 
when  the  commercial  operation  ceased.  A  few 
goats  suni\ed  on  Paoha  until  at  least  1975 
(Hoffman,  unpublislied  report)  but  were  extir- 
pated b\  1980. 

Thus,  human  moxxMueuts  onto  the  ishinds 
\\(M"e  li"e(|ucnt.  and  rodents,  such  as  deer  mice 
and  \ oles,  couki  ha\e  been  inadxertentK  traus- 
portcxl  in  the  grain,  ha\.  and  other  it(^nis  taken 
to  support  acti\iti(\s  on  the  islands.  We  do  no! 
know  il  natixc  lagomoiphs  were  traiiportcd  to 
th(^  islands  In  liumans. 

There  is  debate  in  the  literature  oxer  the 
abilities  of  PcrojiKjscus.  Microfiis.  and  other 
small  mammals  to  coloni/e  iskmds  b\  s\\  iinming 
or  rafting  because  the\  are  not  well  adapted  lor 
exposure  to  water  (Redfield  1976,  (^alhoiiu  and 
Greenbaum  1991,  Peltoneu  and  Hanski  1991). 
We  haw  no  direct  wa\-  of  (juantifxing  the  rela- 
tixe  prol)abilities  of  inadxertent  liuman  trans- 
port xersus  rafting.  Ih)x\exer,  the  knoxxn  and 
fre(juent  histonof  liuman  xisitation  and  habita- 
tion tor  commercial  puq3oses  during  this  cen- 
tun-  results  in  a  higher  frecjuencx'  of  occurrence 
and  less  harsh  means  of  possible  transport  than 


(k)es  raiting  due  to  Hooding  (ncnts.  (.'onfonnd- 
ing  the  present  situation  is  the  laud  bridge  or 
near  land  britlge.  .Moxement  across  the  land 
bridge  to  Negit,  folloxx'cd  bx  swimming  or  raft- 
ing to  I-*a()ha,  is  likelx  more  probable  nox\-  than 
historicalK'. 

The  ab.sence  of  lizards  on  the  islands  is  [)er- 
plexing.  Iioxxexer,  as  there  appears  to  be  ample 
habitat  on  the  islands,  and  species  on  the  main- 
land are  potentiallx  good  colonizers  (sensit  ("ase 
1975,  1983).  Hoxx'ex'er,  mainland  [)()puIations 
are  small,  as  the  elexatiou  of  the  .Mono  Basin  is 
at  th(Mipperendof  the  normal  range  for  reptiles 
in  the  Sierra  Nex'ada  (summarized  from  Storer 
and  Usinger  1968).  Therefore,  their  chance  of 
arrixal  and  persistence  is  loxx*. 

Snakes  {Fitiiopliis  iitclanolciiciis  and 
Tluniniopliis  clc^^atis)  and  amphibians  [Biifo 
horcds.  llifla  re^^illa,  ScapJiiojnts  Juiinmoiulii,  S. 
intcniiontauus)  are  found  around  Mono  Lake 
M\'Z  specimens,  personal  obsenation ),  but 
tliex-  are  scarce  locallx'  (personal  obsenation). 
There  are  no  historic  records  of  snakes  or 
amphibians  on  either  island,  and  x\e  .sax\-  no 
ex  idence  of  either  during  our  xisits.  .\s  discussed 
abox(^  for  lizards,  it  api)ears  that  the  chance  of 
arrixal  and  persistence  of  snakes  and  amphibi- 
ans is  loxx'. 

The  Mono  Fake  islands  parallel  other  islands 
in  haxing  a  greater  population  al)uiKlance(espe- 
ciallx  Xegit)  and  a  simple  species  comp(Jsition 
relatix(^  to  the  mainland.  Larger  relatixe  abun- 
dances max  be  because  fexx- predators  are  pres- 
ent and  tlie  lack  of  nonaxian  food  competitors, 
as  has  !)een  postulated  for  other  island  rodent 
populations  (e.g.,  Halpin  and  Sullixan  1978). 
The  fexx-  rodent  species,  absence  of  lizards,  and 


334 


Great  Basin  Naturalist 


[Volume  52 


reduced  bird-species  richness  (Hall  et  al.,  in 
preparation)  on  the  islands  may  result  in  density 
compensation  (semn  MacArthur  et  al.  1972, 
Case  1975)  by  the  islands'  Fewmijsais  popula- 
tions. 

In  contrast  to  island  biogeographic  theory 
(Redfield  1976,  Sullivan  1977),  deer  mice  are 
smaller  on  the  islands  than  on  the  mainland. 
Although  a  founder  effect  (sen.sti  Kilpatrick 
1981,  Calhoun  and  Greenbaum  1991)  could 
ha\e  resulted  in  smaller  individuals  on  the 
islands  than  on  the  mainland,  there  is  likely 
some  combination  of  ecological  factors  on  the 
Mono  Lake  islands  that  has  either  resulted  in 
maintenance  of  small  body  size  or  has  directed 
selection  toward  smaller  body  size.  In  our  study 
the  sex  ratio  of  deer  mice  appears  to  be  male 
biased,  although  more  intensive  trapping,  both 
within  and  betsveen  years,  would  be  necessan' 
for  confirmation  because  of  potential  trapping 
biases  associated  with  dispersing  young  males. 

ACKNO\\'LEDGMENTS 

We  thank  Paul  Aigner,  fean  Block,  Martin 
Morton,  and  Paul  Stapp  for  assistance  with 
fieldwork;  Edward  Beedy  for  helping  with  stud\' 
design;  John  Harris  and  several  anonymous  ref- 
erees for  reviewing  earlier  dratts;  and  Lorraine 
Merkle  for  manuscript  preparation.  Data  col- 
lection was  supported  by  subconsultant  contract 
to  Jones  and  Stokes  Associates,  consultant  to 
Los  Angeles  Department  of  Water  and  Power 
and  C^alifoniia  State  Water  Resources  Control 
Board;  and  the  White  Mountain  Research  Sta- 
tion, Uni\ersit\()f  California,  Los  Angeles. 

Literature  Cited 

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AND  J.  A.  WiKNS  1988.  The  future  of  Mono  Lake. 
University  of  Califomia,  Water  Re.source.s  Center, 
Report  No.  68.  Berkeley,  California.  29  pp. 

BUIK.II.  J.  B.,  J.  RoiiHINS.  AND  T.  Wainwhiciit  1977. 
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Calhol'n.  S.  W,  and  I.  F.  Chkknbai  \i  1991^  Evolution- 
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Case,  T  j.  1975.  Species  numbers,  dcnsit\  compensation, 

and  colonizing  abilitv  of  Lizards  on  islands  in  the  (iulf 

of  California.  Ecology- .56:  .3-18. 
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tow  of  the  Bodie-Mono  Lake  area  in  the  nineteenth 

centniY.  Artemisia  Press,  Lee  Vining,  Ctilifomia.  123  pp. 
IIaepin.  Z.  T.  and  T.  P.  Sullivan  1978.  Social  interactions 

in  island  populations  of  the  deer  mouse,  Pcwmy.sai.s 

maniailatus.  Journal  of  Mammalogv' 59:  .39.5-401. 
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foraging  ecology.  Ecology  65:  1579-1584. 
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reference  to  the  California  Gull  colony  at  Mono  Lake. 

CaLifomia.  Colonial  Waterbirds  11:  322-.327. 
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Mammalian  population  genetics.  Uni\ersit\- of  Georgia 

Press,  Athens. 
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in  D.  W.  Winkler,  ed..  An  ecological  stnd\  of  Mono 

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of  island  biogeography.  Princeton  Uni\ersit\   Press. 

Princeton,  New  Jersey.  20.3  pp. 
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1972.  Densit\' compensation  in  island  launas.  Ecolog\' 

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occupancy  explained  1)\'  colonization  and  extinction 

rates  in  shrews.  Ecolog\-  72:  169<S-1708. 
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Received  19  Fehnianj  1992 
Accepted  1  October  1992 


Great  Basin  NatiuiJist  52(41,  pp  335^'543 


\'\SCULAR  FLORA  OF  KANE  LAKE  CIRQUE, 
PIONEER  MOUNTAINS,  IDAHO 


Rfihcrt  K,  MosclcN 


Susan  Bcruatas 


Abstract — Kane  Lake  Cirque  lies  in  the  western  Pioneer  Monntains  of  south  central  Ichilio.  An  inventor\-  of  the 
liii;;h-ele\ation  flora  of  the  cirque  revealed  the  presence  of  ISO  \ascniar  taxa  representiu'j;  95  genera  and  30  families.  Fi\e 
alpine  taxa  are  here  documented  from  Idaho  for  the  first  time:  Carcx  inainiforntis  Mack.,  Draha  flachiizcnsis  Wilfen., 
Polentilh  nivca  L.,  Rannunilus  i^clidus  Kar.  &  Kir.,  and  Ranniicuhts  pijgmacus  Wdilenl).  Kane  Lake  Cirtjne  also  contains 
populations  of  four  additional  alpine  taxa  considered  to  be  of  consi-rvation  concern  in  hialio:  Eiii^croit  limiiilis  CiriJiam. 
Panui.ssiti  kotzi'huei  Cham.,  Saxifra^a  adscendens  L.,  and  Saxifra^ti  cennia  L. 

Kcij  words:  Idahti.  rioiiccr  Moiintdiiis.  K/iuc  Ijikc  (Injuc.  alpine  idscidar  flora,  state  reeords.  rare  flora. 


Studies  of  alpine  flora  have  been  numerous 
throughout  the  North  American  Cordillera,  but 
onl\'  recenth"  haxe  inxestigations  of  this  kiiul 
been  undertaken  in  Idaho.  Floristic  stuches  ini- 
tiated b\  Douglass  Henderson  of  the  Uni\ersit\' 
of  Idaho  Herbarium  in  the  mid-197()s  were  the 
first  to  swstematicalK'  explore  the  alpine  zone  of 
Idaho.  Through  numerous  collections,  he  and 
co-workers  documented  Idaho's  alpine  flora  to 
be  unique  in  man\'  respects  (Henderson  197S, 
Bmnsfeld  1981,  Henderson  et  al.  1981.  Bruns- 
feld  et  al.  1983,  Caicco  et  al.  1983,  Lackshewitz 
et  al.  1983,  Hartman  and  Constance  1985). 
NearK"  all  these  inxestigations  took  place  in  the 
large  Basin  and  Ranre-like  massifs  in  east  cen- 
tral  Idaho,  with  fe\\'  extending  into  the  western 
Pioneer  Mountains  of  .south  central  Idaho. 

Rare  plant  imentories  initiated  In  the  U.S. 
Forest  Senice  were  the  first  to  point  out  tlu^ 
phxtogeographic  importance  of  the  circjues  in 
tlie  western  Pioneer  Mountains,  in  general,  and 
Kane  Lake  Cirque,  in  particular  (Caicco  and 
Henderson  198L  Bmnsfeld  et  al.  1983\  Bar- 
bara Ertters  collections  in  1977  and  our  collections 
in  1987  further  highlighted  the  significance  of 
Kane  Lake  Cirque.  Because  of  increasing  recre- 
ational use  of  the  Kane  Lake  area.  sensiti\it\ of 
the  habitats,  and  preliminar\  nature^  of  the  flo- 
ristic inxenton;  we  undertook  this  stucK  in 
cooperation  with  the  Challis  National  Forest  to 
provide  them  with  adequate  data  on  the  distri- 


bution and  abundance  of  rare  plants  and  habi- 
tats in  the  basin  for  future  manaiiement. 


Stiioy  a  hi:  a 

The  Pioneer  Mountains  rise  abruplK  from 
tlie  northern  edge  of  the  Snake  \\\\vr  Plain  in 
central  Idaho  between  the  Big  Lost  and  Big 
Wood  ri\ers.  These  mountains  form  a  large, 
complex  block  about  60  km  long  and  50  km 
w  ide.  oriented  northwest  to  soutlunist.  Topogra- 
pln  \aries  bom  shaip  horns,  serrate  ridges,  and 
broad  upland  surfac(\s  in  the  al})ine  zone  to 
steep-sided  \alle\s  and  rounded  ridges  in  the 
foothills.  Elexations  range  from  1900  m  in  the 
\alle\s  on  the  western  slope  to  3658  m  al  the 
summit  of  IIvTidman  Peak. 

The  PionecM-  Moimtains  arc  composed  of 
TtMtiaiA  Challis  \'olcanics  consisting  of  inter- 
bedded  la\a  and  tuffaceous  miits.  which  lie 
uncomformabK  oxer  a  core  of  Precambiian 
metamoq^hic  and  Paleozoic  sedimentar\  units. 
During  the  formation  of  the  Cretaceous  Idaho 
Batholith.  small  ■'satelliti'"intnisi\e  bodies  were 
emplaced  in  the  western  PionetM"  Mountains. 
TertiatA  and  Quatenian'  block  faulting  is 
beliexcd  to  be  the  cause  of  the  subsequent  upHft 
aiul  present  relief  (Doxer  1981).  The  geomor- 
phologic  setting  has  been  greatly  influenced  by 
(,)naternan  glacial  and  flu\ial  acti\it\'.  Most 
streams  in  glaciated  \alleys  are  underfit.  and 


^Conservation  Data  Center.  Ichiho  Department  ot  Fisli  and  Game,  Box  2-5,  Boise.  Idaho  83707. 

"Present  address:  Science  Application  International  Corporation.  40.5  S.  8th.  Suite  201.  Boise.  Idaho  8.3702. 


335 


336 


Great  Basin  Naturalist 


[\  oluiiie  52 


uplands  display  classic  alpine-tyjoe  glaciated 
features  including  cols,  aretes,  honis,  and 
cirques  (Evenson  et  al.  1982). 

The  Pioneer  Mountains  lie  in  a  transition 
zone  between  the  maritime  climate  of  northern 
and  western  Idaho  and  the  continental  climate 
of  southeastern  Idaho,  and  are  affected  by  two 
basic  storm  patterns.  From  Noyember  through 
March  most  precipitation  comes  from  low- 
altitude  cyclonic  storms  that  moye  east\vaid  from 
the  Pacific  Ocean.  During  May  and  fune  most 
precipitation  results  from  high-altitude  ccjuyec- 
tional  storms  moving  northward  from  the  Gulf 
of  Mexico  and  California  coast.  The  combina- 
tion of  maritime  and  continental  influence  cre- 
ates t\yo  wet  seasons,  winter  and  late  spring, 
respectively  (Caicco  1983).  No  climatic  data  are 
available  from  high  elexations  in  the  Pioneer 
Mountains;  hovx^ever,  Moseley  (1985)  estimates 
mean  annual  precipitation  at  2835  m  in  the 
southern  part  of  the  range  to  be  813  nun. 
Throughout  the  mountainous  regions  of  the 
world,  the  altitude  of  upper  treeline  has  long 
been  obsen'ed  to  coincide  with  the  10  C  iso- 
therm of  the  warmest  month  (Griggs  1937, 
Daubenmire  1954,Wardle  1974).  Extrapolation 
of  temperatures  from  valley  stations  in  the  vicin- 
ity to  timberline  (3000  m)  using  an  adiabatic 
lapse  rate  of  0.62  C/100  m  (Baker  1944)  sub- 
stantiates this  obsenation  for  the  Pioneer 
Mountains. 


DEst:KiPTU)i\  OF  Kane  Lake  Cirque 

Kane  Lake  Cirque  encompasses  approxi- 
iiiatelv  567  ha  at  the  head  of  Kane  Creek  in  the 
western  Pioneer  Mountains  21  km  northeast  of 
Ketchum,  Custer  Countv,  Idaho  (43°44'N 
114°6'W;  T5N  R20E,  Boise  Meridian).  The 
circjue  is  characterized  bv'  permanent  snow- 
fields.  glaciall)-  .scoured  bedrock  (gneiss  antl 
(juartz  diorite),  and  unstable  talus  and  morainal 
(k'posits.  Although  several  small  ponds  are  scat- 
tered throughout  the  basin,  5.3-ha  Kane  Lake  is 
the  only  large  body  of  water.  Elevations  of  the 
.study  area  range  from  2800  m  to  3648  iii.  The 
only  appreciable  soil  dexclopnient  is  in  (k-posi- 
tional  areas  such  as  along  streams  and  rixulets, 
around  ponds  and  lakes,  and  in  the  coniferous 
woodland  on  the  north  side  of  Kane  Lak(\ 

\'egetation  in  the  cinjue  reflects  a  inoisttM- 
regime  than  has  been  noted  at  high  elevations 
elsewhere  in  Idalio  (Brunsfeld  1981,  Caicco 
1983,  Moseley  1985).  This  mesic  en\iromnent 


can  be  attributed  to  several  factors,  including 
the  north-facing  orientation  of  the  cirque,  a 
massive  headvxall  on  the  south,  and  high  peaks 
on  the  east,  south,  and  west  sides  of  the  basin. 
These  features  contribute  collectixely  to  a  heavy 
snov\'  accunnilation  in  the  v\inter  and  its  reten- 
tion throuiihout  the  summer.  Late-lvinci  snovy 
and  an  impermeable  substrate,  augmented  by 
siunmer  thundershowers,  appear  to  provide 
plentiful  moisture  to  nearly  all  habitats  through- 
out the  growing  season,  except  the  south-facing 
slopes  north  of  Kane  Lake. 

Habitats  in  the  circjue  can  be  divided  into 
two  distinct  groups:  snbalpine  and  alpine.  Sub- 
alpine  vegetation  is  restricted  to  areas 
innnediatelv  adjacent  to  Kane  Lake  and  gener- 
all)'  does  not  exceed  2850  m.  Alpine  habitats 
cover  most  of  the  area  and  are  generallv  sparsely 
vegetated,  although  small  areas  with  continuous 
vegetative  cover  occur  along  streams  and  rivu- 
lets throughout  the  basin  and  contain  much  of 
the  plant  species  diversity  of  the  alpine  zone. 
Plant  associations  of  the  study  area  are  not 
included  in  published  vegetation  classifications 
of  the  region  and  are  subjectiveK  characterized 
below. 

Snbalpine  Connmmities 

Coniferous  woodland.  Open  stands  of 
Piiiiis  (lU)ic(iuJis  Engelm.,  vvith  lesser  amounts 
of  Ahics  hi.siocaiya  (Hook.)  Nutt.  and  Picea 
cii^chiuiiiiui  Parn,  occupy  the  level  bench  north 
of  Kane  Lake.  The  relatively  xeric  understoiy  is 
characterized  by  Vacciniuin  scopariuin  Leiberg, 
Pod  lu'iTosa  (Hook.)  Wisev,  and  Seiwcio 
strfpidiifUlfoJius  Greene. 

Upland  meadows.  Interspersed  in  low- 
King  areas  w ithin  the  coniferous  woodland  are 
meadows  ck)minated  bv  graminoids  and  forbs. 
Species  characteristic  of  these  sites  include 
Fcstiicd  iddhoensis  Elmer,  DanfhonUi  iiifcniw- 
(lia  \'ase\,  Erig^croii  simplex  Greene,  Ccircx 
inirroptcra  Mack.,  antl  Potciitilla  diccrsifolia 
Lehm. 

Tree  islands  and  krunimolz.  Isolatetl  tree 
islands,  consisting  ot  small,  upright  Ahics 
hisiocarpa  with  an  underston  dominated  b\ 
Plu/Ihxiocc  spp.,  occur  as  high  as  3050  m  on 
benches  south  of  the  lake.  These  relatively  moist 
sites  are  surrounded  by  bedrock  or  alpine 
meadow  c-onnnnuities.  Areas  of  kruniinholz  are 
ran^  in  the  study  area.  Small  isolated  patches 
consisting  of  low-growing  A.   lasiocaiya  and 


19921 


K.WK  Laki-;  CiHoL  e  Xascl  l.\k  Flora 


337 


Fimis  (ilhirauVis  occur  as  high  as  3230  in  on  tlie 
stccj)  soiith-lacing  slopes  iioith  of  Kane  Lake. 

Lakeside  meadows.  Snrronndinij;  Kane 
l^ake  and  radiating  out  along  inlet  and  ontlet 
streams  are  mc^ulows  that  ha\e  soil  saturated  to 
the  sni'laee  and  are  high  in  organic  matter. 
Carcxscopuloniin  1  h)lni  is  dominant  here  along 
with  scattered  iorhs.  such  as  Ehiici'oii  pcr- 
('ilritiiis  (Pursh)  Greene,  Sowrio  ci/inba- 
larioidcs  Buck.,  and  Gcntiaiia  cali/cosa  Griseh. 
Low  sln-iihs.  iiiclnding  Salix  phinifolia  Pursh 
and  Ledum  ^Idiuliilosiini  Xutt..  occur  occasion- 
all\  in  these  meadow  s. 

Alpine  (lommnnities 

Meadows.  This  mesic  conuuunit\"  is  limited 
in  extent  and  generalK  occurs  in  isolated 
patches  around  seeps  or  as  stringers  along 
streams  and  rixiilets.  Dcscliamjisia  ce.sj)it<)s(i 
(L.)  Beam',  is  1)\'  far  the  dominant  species  her(\ 
with  a  high  di\ersit\'  of  forbs  and  othei" 
graminoids  occurring  in  low  co\er 

Cliffs  and  ledges.  This  is  the  most  common 
conniiunitx  in  the  cirque.  Most  cliffs  and  ledges 
are  north-facing  and  wet  to  mesic,  with  Draha 
loncJiocaiya  R\db.  being  the  most  constant  spe- 
cies, alongwith  several  species  of  S/ixifra^a.  The 
xeric  counteiparts  occur  ouK'  on  the  south- 
facing  slopes  northeast  of  Kane  Lake  and  ha\e 
few  \ascular  plants. 

Talus  and  scree.  Common  at  upper  ele\"a- 
tions  in  tlu^  cirque,  this  relatix  el\'  mesic  conmiu- 
nit\  is  characterized  b\  a  uni(|ut>  suite  ol  species 
able  to  with.stand  constantK  shifting  substrates. 
Specic^s  characteristic  ol  material  greater  than 
5  cm  in  diameter  (talus)  include  Hulscci  al0(l(i 
Gra\  and  Sciwcio  frcDioiitii  T  &:  (i.,  while 
Saxifra<ia  cerniia  L..  Liiziila  spicdta  (L.)  IX^. 
and  Andwsacc  scptcnirioiuilis  L.  characterize 
small-diameter  material  (scree). 

Fellfield.  On  fellfield  habitats  are  rare. 
occiuringonK  in  small  pockets  on  bedrock  slabs 
on  the  cirque  floor  east  of  Kane  Lake.  Species 
t\pical  of  this  poorK  dexcloped  c'ommunitx 
include  Potoit'dhi  hrcrifolia  Xutt.,  jiniciis 
dnimntoitdii  E.  Me\'er,  and  Sihhaldia  ))n)- 
cuinhcns  L.  Carcx  chjiioidcs  Holm  also  occ-urs 
in  this  comnnmitx  but  does  not  de\elop  into  the 
ext(Misi\e  turfs  that  are  found  elsewhere  in  cen- 
tral Idaho  (Caicco  19S3).  .\11  ridges  surrounding 
the  basin,  t\pical  sites  for  fellfield  connriunities 
elsewhere  in  the  state,  are  shaq)  aretes  witli  no 
well-de\ eloped  \egetation . 


Mktiiods 

The  checklist  is  based  on  263  collections, 
made  mostK  b\  tlu^  authors  in  |ul\'  and  .\ugu.st 
19(S7  and  July  1991.  Other  collectors  include 
Barbara  Ertter,  who  \isited  Kane  Lake  in  |ul\ 
1977,  and  Ste\en  C^aicco.  who  collected  in  the 
cii(|ue  during  juK  I9S1  and  .\ngust  I9S2.  A 
nearK  complete  set  of  sp(>cimens  is  deposited  at 
the  Uni\ersit\  of  Idaho  IIed)ariuni  (ID),  with 
duplicates  distributed  wid(^l\.  Ertters  collec- 
tions are  deposited  at  the  Albertson  (College  of 
Idaho  (CIC). 

(Collections  thought  to  l)e  new  records  for 
Idaho  were  confirmed  b\  exjxMts  in  a  particular 
taxon  and/or  fiom  a  search  of  up  to  59  k)cal, 
regional,  and  national  herbaria  ii\Iosele\'  1989). 
Range-extension  data  for  these  s  tate- record  ta.xa 
were  determined  from  herbarium  records  and 
from  the  atlases  and  data  bases  maintained  b\ 
tli{^  Idaho  (>)nsenati()n  Data  (Center  and  Mon- 
tana Natural  Heritage  Program  on  the  location, 
distribution,  numbers,  and  condition  of  rare 
plant  populations  in  their  respectix'c  states 
(Jenkins  19S6).  The  Ickiho  (Consenation  Data 
Center  data  base  was  also  consulted  concerning 
the  current  distribution  of  additional  rarc^  spe- 
cies in  Idaho. 

Results  axd  Discission 

The  \ascular  flora  of  Kane  Lake  (Circjue  con- 
sists of  180  species  representing  95  genera  in  30 
families  of  pteridophxtes,  g\nino.sperms.  and 
angiosperms.  Of  these,  53  species  (29%)  are 
ic'stricted  to  subalpine  conununities  in  the 
c  injue,  while  58  species  (33%)  are  re.stricted  to 
al])ine  habitats.  The  remaining  69  species  (38%) 
transcend  the  subalpine-alpine  boundaiA'  and 
occur  in  both  l\  pes  of  comnumities.  Our  collec- 
tions of  fi\e  species  from  the  stucK  area  repre- 
sent their  first  documented  occurrence  in 
Idaho.  In  addition,  four  otlier  arctic-alpine  spe- 
cies are  known  from  Idaho  from  ouK  a  few 
occurrence's  and  are  considerc-d  rare  in  the  state 
(.\Iosele\  and  (inncs  1990).  OnK  one  alien 
taxon.  Taraxacum  officiualc  Weber  was  found 
in  tlie  stud\  area. 

Taxa  New  to  Idaho 

Carex  incurviformis  Mack.  Fhis  species 
occurs  in  tx\()  areas  of  the  North  American  Cor- 
dillera: \ar.  incuniformis,  known  from  the 
HockA  Mountiiins  of  British  Columbia.  Alberta, 


338 


Great  Basin  Naturalist 


[Volume  52 


Montana,  and  now  Idalio;  and  \ar.  danaensis 
(Stacey)  Hermann,  occurring  in  the  southern 
Rocky  Mountains  of  Colorado  and  the  Sierra 
Nevada  and  White  Mountains  of  California  (J. 
Mastrogiuseppe,  Washington  State  Universitv; 
personal  connnunication,  1991).  The  popula- 
tion in  Kane  Creek  is  disjunct  south  from  the 
next  closest  knowii  population  in  Deer  Lodge 
Count\-,  Montana,  b\'  about  260  km  {Lack- 
scliew'itz  393S  MONTU;  Lesica  and  Shelly 
1991).  We  found  one  small  population  in  the 
Kane  Lake  Cirque,  occurring  in  a  steeply  slop- 
ing meadow  on  seepy  ledges  at  3350  m  at  the 
southern  end  of  the  cinjue. 

Draba  fladnizensis  Wilfen.  A  widespread 
circmnpolar  species,  Draba  fladnizensis  is 
sparselv  distributed  in  North  America,  from  the 
arctic  south  through  the  Rock-v  Mountains  to 
Utah  and  Colorado  (Hitchcock  1941).  As 
withC<7/Y'.v  incitniforniis,  the  Kane  Lake  Cirque 
population  is  disjunct  south  from  the  next  ck)s- 
est  known  population  in  the  Storm  Lake  area  of 
the  Pintlar  Range,  Deer  Lodge  Counts',  Mon- 
tana, bv  about  260  km  [Lackschewitz  6120 
MONTU).  Several  verv  small  populations  occur 
on  ledges  and  in  rocky  areas  south  of  Kane  Lake, 
including  sprav  zones  of  waterfalls,  bare  stream 
gravels,  and  on  steep,  rock^'  slopes  near  seeps. 

Potentilla  nivea  L.  This  circumpolar  spe- 
cies occurs  in  arctic  and  alpine  regions  of  North 
America,  being  pre\iousl\'  knowai  in  western 
North  America  from  Alaska  south  along  the 
mmn  crest  of  the  Rock-\'  Mountains  to  Montana, 
Wyoming,  Colorado,  and  Utah  and  east  to 
Nevada  (Hitchcock  and  Cronquist  1973).  The 
Kane  Lake  Cirque  population  is  disjunct  from 
the  nearest  Montana  populations  by  perhaps 
280  km.  A  small  population  of  about  a  dozen 
[)lants  was  seen  in  a  moist,  sloping  meadow  at 
the  top  ()(  the  waterfalls  south  of  Kane  Lake  at 
2950  m. 

Ranunculus  gelidus  Kar.  &  Kir.  A  North 
American  endemic,  this  species  is  distributctl 
across  the  arctic,  southward  in  the  Rock\  Moun- 
tains to  Colorado  (Benson  1 948).  The  very  small 
population  in  Kane  Creek  Cirque  represents  a 
disjunction  southwestward  of  about  350  m  froui 
the  Beartooth  Plateau,  Stillwater  Countw  Mon- 
tana {Stickney  4  MRC;  Lesica  and  Shellv  1991 ). 
In  the  study  area  it  occurs  in  a  stringer  of 
Deschanipsia  cespitosa  along  the  northeastern 
tributaPy-  of  Kane  Lake  at  about  3170  m. 

Ranunculus  pygmaeus  Wahlenb.  This 
buttercup  is  circumpolar,  occurring  south  along 


the  Rock-x-  Mountain  crest  to  Colorado  (Benson 
1948).  Its  presence  in  the  Kane  Lake  Cirque 
represents  a  disjunction  of  about  200  km  south- 
west from  the  next  nearest  known  populations 
in  the  Pioneer  Mountains,  Beaverhead  Countv, 
Montana  (Hitchcock  and  Midilick  12899  WS'). 
Raniincidus  pi/(iniacus  is  relati\el\-  connnon  in 
the  Kane  Lake  Cirque,  occurring  in  moist, 
exposed  soil  along  creeks,  on  ledges  and  slopes, 
and  occasionally  in  cracks  in  cliffs. 

Additional  Rare  Species 

Erigeron  humilis  Grahm.  This  circmnpo- 
lar species  was  not  known  from  Idaho  until 
Henderson  et  al.  (1981)  reported  it  from  the 
Lemhi  and  Lost  Ri\er  ranges.  Eight  occur- 
rences are  now  knowii  from  the  state,  with  the 
Kane  Lake  Cirque  populations  being  the  only 
ones  known  outside  the  two  ranges  mentioned 
above  (unpublished  data  on  file  at  the  Idaho 
Conseivation  Data  Center,  Boise).  Erigeron 
liuniilis  is  relatixeh'  common  in  moist 
Dcscliainpsia  cespitosa  meadows  throughout 
the  lower  portion  of  the  cirque. 

Paruassia  kotzebuei  Cham.  This  species 
was  also  not  knowni  from  Idaho  until  recently 
when  Brunsfeld  et  al.  (1983)  reported  it  from 
the  Lost  River  Range  and  Pioneer  Mountains. 
Foui-  occurrences  are  now^  known  from  the  state 
(unpublished  data  on  file  at  the  Idaho  Conser- 
\ation  Data  Center,  Boise).  It  is  relativelv 
common  on  moist  ledges  and  in  sloping 
Descliampsia  cespitosa  meadows  throughout 
the  lower  portion  of  the  circjue. 

Saxifraga  adscendens  L.  The  North  Amer- 
ican representatixe  of  this  wide-ranging  species, 
\ar  orcf^onoisis  (Raf.)  Breit.,  occurs  throughout 
the  RockA  Mountains  and  northern  Cascade 
Range  (Hitchcock  and  Cronquist  1973).  In 
lelaho  it  is  known  from  nine  sites  in  the  White 
('loud  Peaks,  Pioneer  Mountains,  and  Lost 
\\\\vr  Range  (unpublished  data  on  file  at  the 
klaho  (]onsenation  Data  Center,  Boise).  Kane 
Lake  (>ir(jue  populations  occur  throughout  the 
area  on  moist  scree,  sand,  and  graxel.  often 
along  streams. 

Saxifraga  cernua  L.  Se\en  small  popula- 
tions of  this  circumboreal  species  are  known 
from  Idaho  (unpublished  data  on  file  at  the 
klaho  Consenation  Data  Center,  Boise).  At 
Kane  Lake  (>irque  it  is  wideK'  scattered  in  small 
[)ojiulations  from  moist  subalpine  ledges  north 
of  Kane  Lake  at  2S00  m  to  ledges  and  cracks  on 
the  headwall  at  3400  in. 


19921 


Kane  Lake  Cikole  \'asli l.\k  Flora 


339 


Annotated  Checklist  of 
Vascular  Plants 

The  checklist  is  arranged  b\  dixisioii  and 
class  (in  Magnohoph\ta),  then  alphaheticalK  h\ 
hmiiK;  genus,  and  species  within  these  major 
groupings.  Nomenclature  generalK  follows 
Hitchcock  and  Cronquist  (1973),  exceptions 
heing  Salix  (Bnmsfeld  and  |ohnson  1985), 
Carex  incuniformis  and  C.  sco})iil()rum  \ar. 
hracteosa  (Hermann  1970),  antl  Eno<i(>iiuiii 
(■(ipisfrafii))i  (Re\eal  1989).  Unless  othen\ise 
noted,  th(^  collection  numbers  are  the  authors". 

Di\  isiON  Lvcopodkm'uvta 

Srlasjiiirllaceac" 

Selaginella  densa  Rvdb.  Conuiion  in  suhalpinc  and 
alpine  zones;  moist  to  ili"\  slopes  antl  lecl<j;es  and  staiiilized 
scree.  2245. 

Dl\  IsloN  Pol.YI'oDlOI'IlVTA 

PoKpodiaeeae 

C ryptogramma  crispa  (L.)  R.  Br.  Unconinion  in  moist 
suhalpine  and  alpine  tains;  eircnnilioreal.  2292. 

Cystopteris  fragilis  (L.)  Bemh.  Common  among 
rocks  in  moist  alpini.'  sites;  circnnihoreal.  2294. 

Pellaea  breweri  D.C.  Eat.  Uncommon  in  snbalpine 
and  alpine  zones;  stabilized  scree,  rockv  ledges,  and  bonlder 
fiekls.  2293. 

Woodsia  scopulina  D.C.  Eat.  Common  on  rocks  in 
snbalpine  zone.  2252. 

Dl\  ISION  Pl\()I'llVT,\ 

Cupressaceae 

Juniperus  communis  L.  var.  montana  Ait.  Rare  on  diA 
ledges  of  lower  alpine  zone  and  in  krnnnniiolz.  2355. 

Pinaceae 

Aliies  lasiocarpa  (Hook.)  Nutt.  Common  in  woodland 
and  krnmmliol/  lommnnities.  2.392. 

Picea  engelmanuii  Parr\'.  Connnon  in  woodland  and 
krnmmliol/  connnnnties.  239L 

PinuN  albicaulin  Engelm.  Common  in  wootlland  ;uid 
krnmmliolz  conminnities.  22fi.3. 

Dl\  ISION  .Maonoi.iopiiv'ia 

Class  Macnoi.iopsida 

Apiaceac 

Lomatiiim  idahoense  Math.  &  C^onsl.  K;u('  in  dis- 
turbed microsites  in  moist  snbalpine  meadows  north  ol 
Kane  Lake.  2256. 

Osmorhiza  chilensis  H.  &  A.  Uncommon  in  deep  soil 
of  forest  understor\.  237fiB. 

.\steraceae 

Achillea  miUefolium  L.  ssp.  lanulosa  (Nutt.)  Piper 
^■ar.  alpicola  (Rvdb.)  GaiTt'tt.  Connnon  on  dn  snbalpine 
and  alpine  slopes.  2262. 

AgoHerift  aurantiaca  (Hook.)  Greene.  Unconunon  in 
snbalpine  meadows  north  of  Kane  Lake.  23S7. 

Antennaria  alpina  (L.)  Gaertn.  van  media  (Greene) 


Jeps.  Common  in  moist.  sand\  soil  in  alpine  zone.  ILSG, 
2336. 

.A»i/t'»i»ian«  dimorphu  (Nutt.)  T.  &  G.  Rare  in  dr\ 
forest  opcniiiii  north  ol  Kane  l,ake.  2393. 

Antennaria  microphylla  Rvdb.  Common  on  iln  snb- 
alpine ;in(l  ;il])inc  slojies,  2244, 

.4»i/e»iii«riV/  umbrinella  Rvdb.  (lonnnon  in  dn  to 
moist  snbalpine  and  alpine  meadows.  230S.  2312. 

Arnica  latifolia  Bong.  var.  gracilis  (Rvdb.)  Cronq. 
("ommon  in  dt'ep  soil  ol  snbalpine  and  alpine  slopes  and 
bonlder  fiekls.  2246. 

Arnica  mollis  Plook.  ('onnnon  iTi  moist  subalpine  and 
lower  alpine  meack>ws  and  bonlder  fields.  1177,  2319,  2.343. 

Artemisia  michauxiana  Bess.  Uncommon  in  moist, 
unstable,  rock'^•  drainage  bottoms;  snbalpine  and  lower 
alpine  zones.  2.363. 

Artemisia  tridentata  Nutt.  Rare  on  dr\  snbalpine 
slopes  north  ol  Kane  i^ake.  2261. 

Aster  alpigenus  (T.  &  G.)  Gray  var.  haydenii 
(Porter)  Cronq.  Di-\-  op'-nings  in  forest  north  of  Kane 
Lake.  2419. 

Aster  foliaceus  Lindl.  var.  apricus  Gray.  (Connnon  in 
moist  alpini'  meadows  east  ol  Kane  Lake.  1175. 

Aster  stenomeres  Grav.  Dn,  rock)'  ledges  in  forest 
openings  nortli  of  Kane  L;ike.  22.35. 

Chaenactis  alpina  (Gray)  Jones.  Uncommon  in  snb- 
alpine and  alpine  d?\.  saiuK  scree.  2.361. 

Cirsium  tweedyi  (Rvdb.)  Petr.  (Connnon  in  moist 
meadows  and  on  ledges  in  idpine  zone.  2.37S. 

Erigeron  acris  L.  var.  debilis  Gray.  Connnon  in 
moist,  sand\'  soil;  snbalpine  and  aljMne  zones.  11S.3,  22S7. 
2.344. 

Erigeron  asperuginus  (Eat.)  Grav.  Dv\  slopes  and 
ledges;  common  in  snb;ilpine  and  nnconnnon  in  lower 
alpine  zones.  22.50.  2.350. 

Erigeron  compositus  Pursh  \av.  glabratns  Macoun. 
("omnion  on  tin  snbalpine  antl  alpine  letl^es.  22(')5. 

Erigeron  coulteri  Porter.  Rare  in  alpine  meatk)ws 
along  creek  east  ol  Kane  Lake.  1 174. 

Erigeron  humilis  Graham.  L>ocall\  connnon  in  moist 
;il|)ine  meadows.  2274.  24  10:  C.iicco  2S-1, 

Erigeron  percgrinus  (Pursh)  Cireene  ssp. 
calliunthemtis  (Greene)  Cronq.  \  ar.  scaposus  (T.  &  G.) 
Cronq.  Connnon  in  moist  to  wet  snbalpine  meadows 
aronntl  Kane  Lake.  2.306.  2.367. 

Erigeron  simplex  Greene.  C^ommon  in  snbalpine  and 
al[inie  zones;  moist  meatlows  anil  slopes.  IISS.  2270.  2.3.38; 
Caicco  476:  I'a-tter  21()S, 

Haplopappus  Itfallii  Gra\.  Unconnnon  oti  dn  alpine 
letlues.  2402. 

Haplopappus  macronema  (iray.  Unconnnon  on  dr\ 
subalpine  knoll,  within  forest  Tiorth  of  Kane  Liike.  Not 
collectetl. 

Haplopappus  suffruticosus  (Nutt.)  Gray.  Uncom- 
mon on  (lr\  sub;ilpine  knoll,  within  forest  nortli  ol  KiUie 
L;ike.  Not  collected. 

Ilieracium  gracile  Hook.  Uncommon  in  dn  forest 
openint^s  north  ol  Kane  Lake.  2.30.5. 

llulsea  algida  Gray.  Common  in  alpine  t;ilns.  2403. 

Microseris  nutans  (Geyer)  Shultz-Bip.  Uncommon 
in  snbalpine  meadows  north  ol  Kane  L;ike.  2.3S5. 

Senecio  cymbalarioides  Buek.  Connnon  in  moist  sub- 
alpine and  alpine  me;ido\ss.  !  17.3. 

Senecio  fremontii  T.  &  G.  var./rp»io»i/ii.  Common  in 
alpine  talus.  24(K). 

Senecio  streptanthifolius  Greene.  Common  in  sub- 
alpine zone;  dn  slopes  and  Itjrest  understor)'.  22.55. 


340 


Great  Basin  Naturalist 


[Volume  52 


Solida^o  multiracUata  .\it.  \ar.  scoptilorum  Gray. 

DiT.  roc'k\'  siihalpiiic  and  alpiiu'  Icdiies.  225S. 

Taraxacum  lyrutum  (Ledeb.)  DC.  (^omiiion  in  alpine 
zone;  moist  meadows  and  slopes.  I  185,  22(S9. 

Taraxacum  officinale  Weber.  Alien:  rare  in  snhalpine 
meadows  north  of  Kane  Lake.  23S(S. 

Hora<j;inaeeae 

Mertensia  ciliata  (Torr.)  G.  Don.  (xinimon  alon'j; 
snhalpine  and  lower  alpine  rixniets.  226S. 

Brassieaceae 

Arabis  .sp.  Innnatnr(>  and  nnidentifiable  to  species. 
Uncommon  in  dn  to  moist  forest  openings  north  of  Kane 
Lake.  2375. 

Arabis  lemmonii  Wats.  var.  lemmonii.  (Common  on 
(h-v.  mistahle  iiipiiie  slopes.  2313.  2356. 

Arabia  microphylla  Nutt.  ^ar.  microphyUa. 
Connnon  on  su!)alpine  ledges  and  slopes  north  ol  Kane 
Lake.  224S. 

Arabis  microphyUa  Nutt.  \ar.  saximontana  Rollins. 
Uncommon  in  moist  soil  of  alpine  zone.  2374. 

Draba  sp.  Unable  to  identifv;  possibK'  a  new  taxon. 
Rare;  .seen  onl\-  in  one  small,  steepK'  sloping,  moist  meadow 
at  3353  m.  east  ol  Kane  lake.  2412. 

Draba  fladnizensis  Wilfen.  ("ircnmpolar:  rare  on  tlis- 
tnri)ed,  bare-soil  microsites  of  steep  alpine  slopes  and  along 
rivulets.  1 107. 

Draba  lonchocarf)a  Rydb.  var.  lonchocarpa. 
Clonmion  througliont  cinjue  on  moist  ledges  and  slopes; 
alpine  zone.  1  lOtS.  2:U4;  Lrtter  2106. 

Draba  oligosperma  Hook.  var.  oligosperma.  Haic  on 
di"\  alpine  slopes  and  letlges.  2357,  2362;  Ertter  2102. 

Draba  paysonii  Macbr.  var.  treleasii  (Schulz) 
Hitchc.  Unconnnon  in  dr\.  sand\  alpine  soil.  2405. 

Erysimum  asperum  (Nutt.)  DC.  Rai"e  in  dn'  subalpine 
tains  noi'tli  ol  Kane  Lake.  2377. 

Smelowskia  cahjcina  (Steph.)  C.A.  Mey.  var.  amer- 
icana  (Rej^el  &  Herd)  Drurv  &  Rollins.  Unconnnon  on 
dn',  exposed  alpine  slopes  2340;  Ertter  2107. 

(Janoplnllaceae 

Arenaria  aculeata  Wats.  Dw.  sandv  slopes;  connnon 
in  subalpine  and  rare  in  alpine  zone.  2236,  2351. 

Arenaria  congesta  Nutt.  I'ncornmon  on  dn  alpine- 
slopes  east  of  Kane  Lake.  1  1S7. 

Arenaria  obtusiloba  (Rydb)  Fern.  Dn,  exposed 
slopes  and  ledges;  connnon  in  alpine  and  uni'ommon  in 
suba!pin<'  zone.  2354. 

Arenaria  rubella  (Wahlenb.)  J.E.  Smith,  ('ir- 
cnuiboreal;  nncouunon  on  moist  to  dr\  alpine  ledges.  2424. 

Cerastium  berringianum  C^iain.  &  .Schlechl. 
(lonmiou  in  alpine  zone  throughout  cinjue;  moist  slopes, 
meadows,  and  Ii'dges.  2321,  2413. 

Sagina  saginoides  (L.)  Britt.  C>'ircnmborial:  uncom- 
mon in  moist  al|)ine  uu^adows.  I  179,  2260. 

Silene  douglasii  Hook.  var.  douglasii.  i)r\,  rot  k\ 
ledges;  uncommoTi  in  subalpine  and  lower  aliiine  zone. 
2364. 

Silene  repens  Pers.  var.  australe  Hitchc.  &  Maj^. 
Rare  among  rocks  ol'boulder  field  east  of  Kane  I  .ake.  (  laicco 
2S6. 

Stellaria  longipes  Goldic  \ar.  altocauUs  (Hulten) 
Hitchc.  Uncommon  in  moist,  sandv  sites  and  scree  in  alpine 
meadows.  1180,2327. 

Stellaria  umbellata  Turcz.  Rare  in  wet  to  moist  grawls 
along  alpine  rivulets.  2326,  2347. 


("rassnlaceae 

Sedum      lanceolatum      Torr.      \ar.      lanceolatum. 

(iommon  on  moist  to  df\  snl)al[iinc  and  alpine  slopt^s  and 
ledges.  2240. 

Ericaceae 

Kalmia  microphylla  (Hook.)  Heller,  ('onunon  in 
moist  to  wt't  sub;ilpine  ;md  alpine  meadows.  2304. 

Ledum     glandulosum     Nutt.     var.     glandulosum. 

(Common  in  moist  subalpine^  forest  and  meadows  around 
Kane  Lake.  2303. 

Phyllodoce  empetriformis  (Sw.)  D.  Don.  (Connnon  on 
moist  s\ibalpine  and  alpini'  slopi's.  2300. 

Phyllodoce  glandulifera  (Hook.)  Co\.  Common  on 
moist  subalpine  and  alpine  slopes.  2302. 

\Phyllodoce  intermedia  (Hook.)  (Jamp.  ( .'ommon  on 
moist  subalpine  and  alpine  slopes.  2301. 

Vaccinium  scoparium  Leiberg.  (Jonunon  in  dn  sites 
in  un(lerstor\  of  forest  and  knnnmholz.  2237. 

Fabaccnie 

Astragalus  alpinus  L.  ('ircumboreal.  ('ommon  in 
moist  meadows  diroughont  cir(|ue;  subalpine  and  alpine 
zones.  2318.  2373;  C^aicco  474. 

Astragalus  eucosmus  Robins.  Rare  in  cracks  of  moist 
cliff  u("ar  stream;  alpine  zone.  2396. 

Astragalus  kentrophyta  Gra'v  var.  implexus  (Canby) 
Baniebv.  Common  on  exposed,  dn'  alpine  slopes  and 
letlges.  2352;  Ertter  2102. 

Trifolium  longipes  Nutt.  var.  pedunculatum  (Rydb.) 
Hitchc.  I'ncommon  in  deep  soil  alf)ng  subalpine 
strcambank  north  ot  Kane  Lake.  2380. 

Gentianaceae 

Frasera  speciosa  Dougl.  Uncoiumon  in  dr\  subalpine 
talus  north  of  Kane  Lake.  2415. 

Gentiana  calycosa  Griseb.  \'ar.  asepala  (Maguire) 

Hitchc.  (Connnon  in  moist  subalpine  and  low  alpine  iriead- 
ows.  1172. 

Gentiana  prostrata  Haenke.  Rare;  seen  onl\  in  moist, 
steepK'  sloping  meadow  above  ponds  t'ast  of  Kane  Lake; 
;ilpine  zone.  240S. 

(ii'ossnlariaceae 

Ribes  ceniitum  Dougl.  var.  inebrians  (Lindl.) 
Hitchc.  Uncommon  in  subalpine  and  alpine  zones;  dw 
h'dges  and  boulder  fields.  2400. 

Ribes  hendersonii  Hitchc.  Rare  anil  local  in  dr\  boul- 
der field  (.'ast  of  K;me  L;ike;  alpine  zone.  2416. 

Ribes  lacustre  (Pers.)  Poir.  L'ncommon  along  snb- 
;ilpine  creek  near  outlet  of  Kani'  Lake.  2398. 

Ribes  montigenum  McClatchie.  Common  in  boulder 
fields  and  dv\  forest  nud(-rstor\;  subalpine  zone.  2310. 

i  hdrojiliv  llaceae 

Phacelia  hastata  Dougl.  \ar.  alpina  (Rydb.)  Cronq. 

Unconnnon  in  moist  to  dr\  alpine  tains.  2406. 

Ouagraceae 

Epilobium  alpinum  L.  var.  alpinum.  Conmiou  on 
moist,  unstable  subalpine-  ;iud  alpine  slopi-s;  eircnmboi"i-al. 

23:5;5. 

Epilobium  angustifolium  L.  Rare  in  i.\y\  forest  open- 
ing nortli  ol  K;me  Lake.  2418. 

Epilobium  glaberrimum  Barbey  var.  fastigiatum 


1992] 


Kaxk  Laki-;  CiHyiiE  VAScuLy^  fYoiu 


341 


(Nutt.)  Trel.  Uncoiiinion  in  moist  meadow  al()ii<j;  stifain 
east  ol  Kane  L;ikc:  alpine.  225S. 

Oenothera  andina  Nutt.  Hare  in  disturbed  microsites 
in  dr\  snhalpine  meadows  north  of  Kane  Lake.  2264. 

Polemoniaceae 

Phlox  piilvinata  {\VhciT\)  C^nnitj.  (Common  on  dn, 
e\]X)sed  alpine  slopes.  23-tS. 

Polemonium  viscosuin  Nutt.  Connnon  tlirous^Iiout 
cirque  in  talus  and  unstable  sites  on  ledges;  alpine.  2311; 
I-:rtter21()4. 

PoK'gonaceae 

Eriogonum  caespitostim  Nutt.  Uncommon  on  ih\ 
sul)alpine  knoll  north  ot  Kane  Lake.  2359. 

Eriogonum    capistratum    Rev.    \ar.    capintratum. 

LocalK  eomiiion  in  subalpine  and  ;ilpine  zones:  dn,;  rock^' 
slopes  antl  led<j;es.  23fi(). 

Eriogonum  ovalifolium  Nutt.  var.  depressum  Blank. 
Connnon  in  sul)ali)iue  and  alpine  zones;  dn.  unstable  slopes 
and  ledges.  2249. 

Oxyria  digtjna  (L.)  Hill,  ('ircumboreal.  Common 
thronijhoiit  cin  jue  on  moist.  rock\  slopes;  alpine  zone.  22S6. 

Polygonum  histortoides  Pursh.  Common  in  moist  to 
wet  snl);iljiine  iind  ;ilpine  meadows.  2267. 

Polygonum  kelloggii  Greene.  I'nconnnon  in  dn 
forest  openings.  2390. 

Polygonum  viriparum  L.  (."omnion  in  moist  al[)ine 
meadows.  2395;  Caicco  477. 


Port 


niacaceae 


Claytonia  megarhiza  (Gray)  Pariy  var.  megarhiza. 

Uncommon  in  ;ilpine  t;ilus.  2404. 

Lewisia  pygmaea    (Gray)    Robins,   var.   pygmaea. 

Common  in  dn'  snb;ilpine  and  ;ilpine  sites.  2271,  2369. 

Priumlaceae 

Androsace  septentriomdis  L.  Common  on  dr\.  sand\ 
iilpine  slopes;  eiri-nmbore;iI.  2353,  2422. 

Dodecatheon  pulchellum  (Raf.)  Merrill  var. 
watsonii  (Tide.stroni)  Hitchc.  Common  in  moist  subal- 
pine iuul  aljiine  me;idows.  2341. 


H;u 


mneniaceae 


Anemone  parviflora  Michx.  Hare  in  moist  alpine 
me;idow  south  ol  Kane  L;ike.  2339. 

Aquilegia  formosa  Fisch.  Connnon  in  moist,  slopiuii 
meadows:  snbiilpine  ;ind  lower  alpine  zones.  2269:  C;ueeo 
472. 

Caltha  leptosepala  DC.  var.  leptosepala.  Connnon 
throuiihont  cir(jue  in  alpine  ;uid  sulnilpine  zones;  wet  mead- 
ows ;ilong  stre;nns  ;ind  ;n()und  kikes  ;ind  ponds.  IISI,  229S. 

Delphinium  depauperatum  Nutt.  Unconnnon  in  dn 
subiilpint'  in(M(lows  noith  ol  K:nie  L;ike.  2257. 

Ranunculus  eschscholtzii  Schlecht.  var.  esch- 
scholtzii.  ("ommon  throughout  cinjue  on  moist  subaljiine 
and  ;ilpine  slopes.  22S4:  Litter  2109. 

Ranunculus  gelidus  Kar.  &  Kir.  Hare;  seen  onK  in 
moist  iilpine  nie;idow  at  ;il)ont  -3170  in.  iilonij  stream  e;ist  ol 
Kane  Lake.  11 S2. 

Ranunculus  pygmaeus  W'aiilenb.  Cireunipol;ir. 
LocalK  connuon  in  moist  to  wet  sites  in  alpine  zone;  iilong 
iiNulets,  ledges,  and  cracks  on  rock  face.  1110.  2315,  2:346. 

Ranunctdus  verecundus  Robin.s.  Rare  in  moist  alpine 
boulder  field  at  3050  in.  southwest  of  Kane  Lake.  2.345. 


Rosaceae 

Polcniilla  hrciijolia  Nutt.  L(K-all\  connnon  on  dn 
iilpine  outcrops  iit  .3050  ni,  southwest  ol  Kane  Lake.  2399. 

Potentilla  diversifolia  Lehm.  var.  diiersifoUa. 
C^oimnon  throughout  cinjue  in  moist  subalpine  and  alpine 
uK'adows.  22S;5.  23)07,  2372. 

Potentilla  fruticosa  L.  [Pentaphylloides  floribunda 
(Pursli)  Lo\e|.  Connnon  on  moist  ledges  iind  in  boukler 
liekLs  ol  snbiilpine  iind  iil])ine  zones;  circnmborciil.  1171, 
2317. 

Potentilla  glandulosa  Lincll.  var.  pseudorupestris 
(Rydb.)  Brcil.  Local  on  i\\\  snbiilpine  lediies.  2366. 

Potentilla  nivea  L.  Circnnipolar.  Hare  in  iilpine  zone; 
.seen  onl\  in  moist,  sloping  meadow  at  head  of  waterfall 
south  ol  Kiiiie  Liikc-.  2379. 

Ruhus  idaeuH  L.  var.  grucilipes  Jones,  ('ommon  in 
snbiilpine  l)onlder  lields.  2114. 

Sihhaldia  procumbens  L.  Connnon  in  iilpine  ;ind  sub- 
iil])ine  zones  on  moist,  SiincK  slopes  ;ind  ledges;  cir- 
cnmboreal.  22SS. 

Salicaceae 

Salix  arctica  Pall.  \ar.  petraea  -\ndress.  Common 

throni^hont  circjiie  in  moist  snbiilpine  iind  alpine  sites.  2276, 
2342,  2376A,  2397;  Ertter2100. 

Salix  sp.  OnK'  \("getati\'e  specimens  obtained,  but 
iippears  to  be  S.  casticoodiae  Cockerell  ex  Heller  (S. 
Bnmsfeld,  Uni\c'rsit\  ol  Idaho,  personal  coinmnnication, 
1991),  Unconnnonin  wet  snbiilpine  ineiidowiidjacent  to  the 
north  shore  ol  Kiuie  Lake.  23S6 

Salix  nivalis  Hook.  var.  nivalis.  Wmv  on  moist  slopes 
in  snbiilpine  iind  iilpine-  zones.  2337. 

Salix  planifolia  Pursh.  Unconnnon  in  snbiilpine 
ineiidow  west  ol  Kiine  Lilke.  2266. 

Salix  tweedyi  (Bebb)  Ball.  Hare;  onK  one  robust  plant 
seen  iit  biise  ot  sniiill  cascade  at  2865  m,  west  of  Kane  Lake; 
snbiilpine  zone.  242  1. 

Sii\ilrii<iiiceiie 

Heuchera    cylindrica    Doug!.    \ar.    alpina    \\ats. 

C^oimnon  in  subalpine  ;ind  lower  alpine  zones  on  dn  ledges 
iind  outcrops  and  moderateK  stiibilized  scree.  2239. 

Lithophragma  hulhifera  Rydb.  [L.  glabra  Nutt.]. 
Uncommon  on  moist  subalpine  slopes.  2370, 

Mifella  pentandra  Hook.  I  iiconmion  in  lower  iilpine 
iiiid  sul)iil])ine  zones:  moist  nu'iidows  and  slopes.  2330. 

Parnassia  fimhriata  Common,  var.  fimbriate. 
l.ociilK  common  in  moist  snbiilpine  iind  iilpine  meadows. 
1  176:  Ciiicco2S3. 

Parnassia  kotzebuei  Cham.  \ar.  kotzebuei.  L'ncom- 
mon  iind  lociil  in  gentK  to  steepK  sloping  iilpine  mciidows 
iind  on  ledges.  22S5,  232S;  Caicco  2S(). 

Saxifraga  adscendens  L.  \ar.  oregonensis  (Raf.) 
Breit.  Uncommon  in  moist,  sloping  meadows  and  t;ilns  and 
alon^  ri\iilets  in  iilpine  zone.  233 L  C'aicco  2.S2. 

Saxifraga  arguta  D.  Don  [S.  odontoloma  Piper]. 
( Connnon  along  strciims  iind  n\  nlets  in  snbiilpine  iind  lower 
iiljiine  zones.  1  I  7S.  2.335. 

Saxifraga  cemua  L.  Uncommon  ;ind  widely  scattered 
in  moist  scree  luid  sloping  meiulows  and  on  ledges  oi  iilpine 
iind  subalpine  zones;  circuinboreal.  2420, 

Saxifraga  debilis  Engelm.  Common  on  moist  and 
|)rotected  .ilpiiie  ledi^es  iind  slojies.  1  lOS,  1 109, 2.324;  Caicco 
2S1:  Lrtter  210.5. 

Saxifraga  occidentalis  \\als.  \ar.  occidentalis. 
(Common  in  moist  subalpine  ;ind  al[)ine  meatlows.  2242, 
2277,2316,2.329.2401. 


342 


Great  Basin  Naturalist 


[Volume  52 


Saxifraga  oppositifolia  L.  Commcm  on  moist  alpinr 
cliff  fLices;  circ'umhorcal.  235S. 


Scmj 


iliiuanaceae 


Castilleja  miniata  Dougl.  Common  in  moist  to  wvt 
siil)al[iinc  aiul  low  alpine  nuMtlows.  2299. 

Mimultis  tilingii  Regel  var.  caespitosiis  (Greene) 
Grant,  (.'onnnon  alon^  alpine  streams  and  ri\nlets.  1 169. 

Penstemon  procerus  Dougl.  var. /ormo.sus  (A.  Nels.) 
Cronq.  ("onnnon  in  snbalpine  zone  and  micommon  in 
alpine  zone  on  dr\.  roek\  ledges.  2259. 

Veronica  tcormskjoldii  Roem.  &  Schult.  (Jonmion  in 
moist  t()\\(4  alpine  and  snbalpine  meadows.  1184,  2295. 

Violaceae 

Viola  adiinca  Sm.  var.  bellidifolia  (Greene)  HaiT. 

Common  in  suhdpine  and  alpine  zones  on  moist  meadows 
and  .slopes.  2290.  2371. 

Viola  macloskeiji  Lloyd  var.  macloskeyi.  C'onmion  in 
wet  snbalpine  meadow  adjacent  to  Kane  L;ike.  2309. 

CLAS.S  LiLIOPSIDA 

Cvperaceae 

Carex  atrata  L.  var.  erecta  Boott.  Rare  in  moist  soil 
oFbonlder  field  north  of  Kane  Lake;  snbalpine.  Ertter  21 10. 

Carex  capillaris  L.  Circnmboreal.  Uncommon  in 
moist,  steeplv  sloping  meadow  sonth  of  Kane  Lake;  lower 
alpine  zone.  2332. 

Carex  elynoides  Holm.  Unconnnon  on  exposed  alpine 
ledges  east  oi  Kane  Lake.  2425. 

Carex  haydeniana  Olney.  C'onnnon  in  moist  snbalpine 
anil  alpine  meadows.  227S,  2431. 

Carex  incurviforinis  Mack.  cf.  var.  inctirviformis. 
Rare;  seen  only  in  one  small,  steeplv  sloping,  moist  meadow 
at  3353  m,  ea.st  of  Kane  Lake.  241 1. 

Carex  microptera  Mack.  Uncommon  in  moist  snb- 
alpine meadows  north  of  Kani-  Lake.  23S3. 

Carex  nova  Bailey.  Conmion  in  moist  alpini'  meadows. 
2291,  242S;Caicco  475. 

Carex  phaeocephala  Piper.  W'ideK  scatteretl  in  tin 
alpine  sites.  1 190,  2430;  Ertter  21  lOA. 

Carex  proposita  Mack.  C^ommon  on  moist  snbalpine 
and  alpine  slopes.  2279. 

Carex  rossii  Boott.  Uncommon  in  dn'  areas  f)f  forest 
imdcrston.  2247. 

Carex  scirpoidea  Michx.  var.  psetidoscirpoidea 
(Rydb.)  Oonq.  Common  on  moist,  sand\-  snbalpine  nud 
iilpine  slopes.  1 1S9.  22.54,  2432;  C;ucco  47.1. 

Carex  scopulorum  Holm  var.  bracteosa  Hermann. 
Common  in  wet  meadows  tJong  creeks  and  inonnd  K;ine 
Liike  in  snbalpine  and  alpine  zones.  22(S2. 

Carex  suhnigricans  Stacey.  Uncommon  in  moist 
alpine  and  snbalpine  meadows.  2429. 

Inncaccae 

JuncuH  drummondii  E.  Meyer  var.  dnimmondii. 

Common  in  moist  to  dn,  sand\  soil  of  snluiipinc  and  alpine 
slopes.  2253,  2297. 

JuncuH  mertensiattus  Bong.  Common  m  moist  :ilpine 
meadows.  1195. 

Luztila  parviflora  (Ehrh.)  Desv.  Common  in  moist 
snbalpine  and  low  alpine  meadows.  2320. 

Luzula  spicata  (L.)  DC.  Connnon  on  moist,  nnstable 
slopes  of  snbalpine  and  ;ilpine  zones;  circnmbonal.  1  197, 
2323;  Caicco  480;  Ertter  2103. 


Liliaceae 

Allium  brandegei  VVat.s.  Rare  in  dn'  forest  opening 
north  of  K:ine  L;ike.  2394. 

Allium  brevistylum  Wats,  (^onnnon  in  moist,  steeply 
sloping  nu-ackjws  in  snbalpine  and  lower  alpine  zones.  2334. 

Calochortus  etirycarpiis  Wat.s.  Rare  in  dr\'  areas  of 
snbalpine  meadow  north  ol  Kane  Lake.  2384. 

Zigadenus  elegans  Pur.sh.  Common  in  moist,  sloping 
alpine  meatlows.  2340;  (Jaieco  478. 

Poaceae 

Agropyron  scribneri  Vasey  [Elymus  scribneri 
(Vasey)  Jones].  Unconnnon  on  dn',  nnstable  alpine  and 
snbalpine  slopes.  2272. 

Agrostis  humilis  Vasey.  Uncommon  on  moist,  sand\' 
alpine  ledges.  1196,  2.3fi8. 

Agrostis  variabilis  Rvdb.  Uncommon  in  moist  alpine 
meadows.  1194. 

Calamagrostis  puqjurascens  R.  Br.  Unconnnon  on 
dn',  rock^  snbalpine  and  alpine  ledges.  2365. 

Dantlionia  intermedia  Vasey.  Locally  connniMi  in  snb- 
alpine meadows  nortli  ol  Kane  Lake.  2389. 

Deschampsia  cespitosa  (L.)  Bcauv.  var.  cespitosa. 
Common  thn)nghont  cir(|ne  in  snbalpine  and  alpine  moist 
meadows  where  it  is  often  dominant;  circnmboreiJ.  1192, 
2325;  Caicco  481. 

Festuca  idahoensis  Elmer  var.  idahoensis.  Uncom- 
mon in  dn'  forest  openings  north  of  Kane  Lake.  2241. 

Festuca  ovina  L.  var.  brevifolia  (R.  Br.)  Wats.  [F. 
brachyphylla  Schult.  &  Schult.].  Unconnnon  in  alpine 
zone;  moist  to  dn'  meadows  ;nid  ledges.  2273,  2407. 

Oryzopsis  exigua  Thurb.  Common  in  dry  snbalpine 
siti'S  north  of  Kant-  Lake.  2251. 

Phleum  alpinum  L.  Common  in  wet  to  moist  snbalpine 
and  iilpine  meadows;  circnmboreal.  1191,  2281. 

Poa  alpina  L.  Common  in  wet  to  moist  snbalpine  and 
alpine  meadows.  2296. 

Poa  cusickii  Vasey  var.  cusickii.  Uncommon  in  moist 
to  tin  snbalpine  meadows.  2296. 

Poa  cusickii  Vasey  var.  epilis  (Scribn.)  Hitchc. 
Unconnnon  in  moist  snbalpine  meadows.  2382. 

Poa  gracillima  \'a.sev.  Unconnnon  on  dn  ledges  in 
forest  openings.  22.'58. 

Poa  incurva  .Scribn.  &  \\'iU.  Unconnnon  in  iln  snb- 
alpine meadows.  2381. 

Poa  interior  Rvdb.  Ihicommon  on  dn  alpine  slopes 
;md  in  scree.  2426. 

Poa  nervosa  (Hook.)  Vasey  var.  wheeleri  (Vasey) 
Hitchc.  Connnon  on  dn'  ledges  and  in  forest  nnderston' 
nortli  oi  Kane  Lake.  2234. 

Poa  rupicola  Nash.  Uncommon  on  dn.  rock^'  alpine 
slopes.  2427. 

Sitanion  hystrix  (Nutt.)  Smith  var.  hystrix  [Elymus 
eh/moides  (Raf.)  Swezev].  Unconmion  in  snbalpine  and 
alpine  zones  on  dn.  rock\  ledges  and  slopes.  224.3. 

Trisetum  spicatum  (L.)  Richter.  Circnmbore;il. 
( lommon  in  alpine  and  snbalpine  zones;  moist  meadows  and 
Ic-tlees.  2280. 


ACKNOW'LEDCMENTS 

Tlii.s  study  was  supported  hx  the  Challis 
National  Forest,  and  we  greatK  appreciate  the 
help  of  forest  personnel,  especially  Cindy 
Haggas,  Dave  Reeder,  and  Stexe  Spencer.  We 


1992] 


Kane  Lakk  (:iiu,)UE  X'asculah  Flora 


343 


thank  Joy  Mastrogiuseppe,  Wasliiiigton  Stat(^ 
Uni\ersit\-,  for  rapiclK  iclentiF\ing our  Carcx  tol- 
K^ctioiis;  Ste\'e  Brunsfeld,  Uni\-ersit\-  of  Idaho, 
tor  identifxing  our  Salix;  Arthur  Cronquist,  New 
York  Botanical  Garden,  for  identif)ingAn^t'/(/;r/- 
ha  dimoiyha:  and  Barbara  Ertter,  Universit\()i 
("ahtornia.  ior 'K\v\\\.'\[\in^^  Pot  cut  ilia  nivea.  Bar- 
bara Ertter  and  Ste\en  Caicco  graciously 
opened  their  collecting  books  for  our  examina- 
tion. The  manuscript  benefited  greatly  from 
rexiews  b\  Douglass  Henderson,  Barbara 
Ertter,  and  an  anon\nious  reviewer. 

Literature  Cited 


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Benson,  L.  1948.  A  treatise  on  the  North  Anu'rican 
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Bhunsff.ld  S.  J.  1981.  .\lpine  flora  of  east-central  Idaho. 
Unpublished  thesis,  Uni\ersit\- of  Ickilio,  Moscow.  205  pp. 

Bhunsfeld.  S.,  S.  C.\k:co.  and  D.  Hendekson  1983. 
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Madrono  30:64. 

Bkunsfeld.  S.  J..  AND  F.  D.  Johnson  1985.  Field  guide 
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Caicco.  S.  L.  1983.  Alpine  \egetation  of  the  Copper  Basin 
area,  south-central  kUilio.  Unpuhlislied  thesis,  Univer- 
sit)'  of  Idaho,  Moscow.  99  pp. 

Caicco.  S.,and  D.  M.  Hendekson  1981.  A  survey  of  the 
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District-West  Side,  with  recommendations  and  man- 
agement implications.  University  of  Idaho  Herbarium. 
University  of  Idaho,  Moscow.  45  pp. 

Caicco,  S.,  J.  Civille,  and  D,  Henderson  1983,  Note- 
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Pensfemon  procerus  Vhxr.fonnosiis.  Madroiio  30:64. 

Dm  HENMiHE  R.  19.54.  .-Mpine  timberlines  in  the  Americas 
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DcnER.  J.  H.  1981.  Geolog)  of  the  Boulilcr-Pionecr  Wilder- 
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Pages  16-75  in  Mineral  resources  of  die  Boulder-Pio- 
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EvENsoN,  E,  B..  J.  F  R  Cotter,  and  ).  M.  Clinch  1982. 
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model  for  Idaho.  Pages  653-665  in  B.  Bonnichsen  and 
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trends.  Science  85:  251-2.55. 

1  Iartman.  R.  L.,  and  L.  Constance  1985.  Two  new  spe- 
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America.  Brittonia  37:88-95. 

Henderson,  D,  1978,  Notes  on  the  flora  of  ea.st-central 
Idaho,  Madrono  25:172-174, 

Henderson,  D„  S.  Brlnsfeld.  and  i'  Brunsfeld 
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Kansas. 

Lackshewitz,  K.,  D.  Henderson,  and  S.  Brunsfeld 
1983,  Noteworthy  collections  of  Idaho  and  Montana: 
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Fish  and  Game,  Boise.  33  pp. 

Re\  EAL,  J.  L.  1989.  New  combinations  antl  novelties  in 
Eriogonuni  ( Polygon  aceae:  Eriogoniodeae).  Phv- 
tologia  66:251-265. 

Wardle.  P,  1974,  Alpine  timberlines.  Pages  371-402  in 
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Received  3  March  1992 
Accepted  29  September  1992 


Great  Basin  Naturalist  52(4).  pp.  344-.351 

LAKEWARD  AND  DOWNSTREAM  MOVEMENTS  OF 

AGE-0  ARCTIC  GRAYLING  {THYMALLUS  ARCTICUS) 

ORIGINATING  BETWEEN  A  LAKE  AND  A  WATERFALL 

Mark  A.  Delera\    and  CaKin  M.  Ka\a 

Abstiucf, — Arctic  ti;ra\  liiit^  in  Deer  Lake,  Montana,  spawn  onK  in  the  35()-ni  segment  ol  outk't  stream  between  the  kike 
and  a  waterfall.  The  puipose  of  this  study  wtxs  to  e.xamine  consequences  of  and  possible  adaptations  bv  this  population  to 
spawning  abo\'e  the  falls,  bv  determining  the  extent  of  loss  over  the  ftJls  of  age-0  young,  the  daily  and  seasonal  patterns  of 
such  losses,  and  the  .seasond  pattern  of  movement  upstream  into  the  lake  by  the  remtuning  young.  We  measiued  fish 
movements  during  1989  ;uid  1990  with  traps  placed  at  the  outlet  and  at  the  falls,  from  fiv  .swimup  in  July  until  October  or 
November.  Young  went  over  the  falls  predominantly  as  newly  svxTmming  fi-y  at  night.  In  1989  about  5()()()-90()()  were  lost 
downstream,  representing  an  estimated  4-7%  or  less  of  young  produced.  Most  young  thus  appetu"  adapted  to  maintaining 
their  position  above  the  falls.  A  few  started  entering  the  lake  in  August  and  September,  but  only  95  in  19S9  and  23  in  1990 
had  done  so  bv  the  time  obsenations  were  ended  by  the  onset  of  winten'  conditions.  Most  movement  into  the  lake  appeared 
to  occur  sometime  during  the  six  to  seven  months  of  annual  ice  cover.  This  extended  period  of  sti^eam  residence  contrasts 
with  eiirlv  lakeward  movements  reported  for  other  inlet-spawiiing,  lacustrine  gra\'ling  populations  and  ma\'  be  an  adaptation 
for  a\  oiding  predation  by  ku'ge  conspecifics  in  Deer  Lake. 

Kit/  iLiu'd.s:  iiiitj^riitidii.  fish.  jS.r(n/!iiiii.  Thvmallus  arcticus,  .sy//)/i(i//(V/.s,  ndtcijail.  strctim.  hike. 


Limited  infonnation  is  available  on  iiioxe- 
ments  of  young  fish  from  populations  inhabiting 
or  spawniing  in  small  headwater  streams  above 
waterfalls.  An  innate  tendencv  of  voimg  fish 
from  such  populations  to  hold  position  or  move 
upstream  in  water  current  (positixe  rheotaxis) 
would  be  highly  adxantageous  in  preventing 
their  irretrievable  loss  over  the  falls.  Such  loss 
should  be  limited  to  enable  the  population  to 
maintain  itself,  and  appropriate  beha\aoral 
adaptation  would  be  promoted  through  remo\al 
from  the  gene  pool  of  voung  hsh  that  did  go 
downstream.  Evidence  for  such  adaptation  is 
provided  by  studies  reporting  little  or  no  loss 
over  waterfalls  of  young  fish  from  long-estab- 
lished, native  ])opulations  of  rainbow  trout 
{Oncorhijnchus  mijkis.s)  and  cutthroat  ti'out  (O. 
clarki)  in  North  America  (Northcote  1969. 
Northcote  and  Hartman  1988)  and  brown  trout 
{Saliuoirntta}  in  Europe  (Jonssou  1982).  Exper- 
imental studies  haw  provided  evidence  for  a 
genetic  basis  of  such  rheotactic  adaptation  in 
rainbow  trout  andbrowai  trout  (Northcote  198 1 , 
Northcote  and  Kelso  1981,  jons.son  1982). 

Although  there  is  evidence  for  geneticalK 


based,  positi\e  rheotaxis  b\'  Noung  Arctic  gray- 
ling {ThifmaUus arcticus)  m  streams  (Kaya  1989, 
1991),  there  have  been  no  prexious  studies  on 
their  possible  loss  over  waterfalls.  Yoinig  gra\- 
ling  may  be  more  susceptible  to  such  loss  than 
voung  trout,  since  voung  gravling  are  much 
smallei-  and  appear  to  be  weaker  s\\dmmers.  At 
swimup  (initiation  of  swimming),  yoimg  grav- 
ling are  about  9-11  nun  in  length  (Ka\a  1991), 
compared  to  20  mm  or  more  for  rainbow  trout 
(Northcote  1962).  The  present  obsenations 
were  conducted  (^n  a  population  of  gravling  that 
li\es  in  a  lake  near  the  head  of  a  m()initain\alle\' 
and  spawns  onK'  in  a  short  stream  section 
betvveen  the  lake  outlet  and  a  waterfall.  Objec- 
tixes  of  the  stucK  were  to  determine  whether 
age-0  (first-\ear)  voimg  are  lost  downstream 
o\er  the  falls,  the  daih'  anil  seasonal  patterns  of 
such  losses,  and  the  seasonal  patterns  of  their 
upstream  moxcment  into  tlic^  lak(\  Peipetuation 
ol  such  a  population  would  depend  on  limited 
ilow  nstr(\un  loss  of  their  progeux,  and  resitleuce 
in  the  lake  would  require  upstream  migration  by 
the  voung.  The  stud\'  was  desiiined  to  include 
movements  of  the  earliest  mobile  lanae,  an 


^BiologN-Di'parti 
■Aiitliortowlioiii 


III.  Montana  Statf  University,  Bozcn 
incspondiMKc  sliould  lif  acklresscd. 


,  M(iTitana59Ti; 


344 


19921 


Stream  Mu\  emexts  oi'  Ace-u  L.\ke  Giuvling 


345 


aspect  that  appears  lacking  from  most  past  stud- 
ies iiixoKiinj;  tlownistream  iiunenieiits  from  sal- 
nioiiid  populations  al)o\('  waterfalls. 

Study  Site  and  Fopulatiox 

Obsenations  were  conducted  in  1989  and 
f  990  on  the  35()-m  long  section  of  Deer  Creek 
tliat  flows  from  Deer  Lake  to  a  3-m,  \ertical 
w  aterfall.  Tlie  lake  is  located  at  2780  m  altitude 
near  the  head  of  a  mountain  \  alle\  in  the  Mad- 
ison Range  of  southwest  Montana.  Dimensions 
of  the  stream  on  19  August  1990,  measured 
hank-to-hank  at  fi\e  locations  along  each  of  34 
transects  between  the  lake  outlet  and  the  water- 
fall (Deleray  1991),  were  mean  width  of  5.88  m 
(range  1.08-21.0),  mean  depth  of  0.10  m  (range 
0.0-0.41),  and  mean  water  \'elocit\-  (measured 
at  0.6  X  depth  at  each  location)  of  0.05  m/sec 
(range  0.0-0.48).  Estimated  discharge  \olume 
ranged  from  about  0.02  to  0.05  m'Vsec  between 
2  JuK-  and  9  September  1990. 

Prexious  obsenations  had  indicated  that 
Arctic  grayling,  the  onl\'  fish  in  the  lake,  spawn 
onlyin  the  outlet  stream  (Kaya  1989).  The  outlet 
stream  is  inhabited  by  gravUng  fiT  (age-0  hsli 
smaller  than  about  2.5  cm  in  length;  Piper  et  al. 
1982)  and  other  young  up  to  about  14  cm  in 
length.  Larger  fish  are  rare,  except  when  spawn- 
ing adults  are  present  during  early  summer. 
Numbers  of  adults  spawiiing  in  the  stream  were 
estimated  b\"  electrofishing  mark-and-recap- 
ture  methods  at  803  (95%  CL  ±104)  in  1989  and 
1 109(95%CL±124)in  1990,  with  similar  num- 
bers of  males  and  females  (Delera\'  1991).  The 
350-m  segment  between  the  lake  and  the  water- 
fall is  the  onlv  part  of  Deer  Creek  inhabited  b\ 
gra)ling.  Near  the  base  of  the  waterfall  the 
stream  disappears  beneath  the  svnface  of  a  steep 
tahis  slope  before  reemerging  about  200  m 
(k)wnslope.  Gra\ling  are  not  present  in  the  10 
km  of  stream  between  the  lake  and  tlie  (Tallalin 
Ki\er,  perhaps  because  of  the  streams  steej) 
gradient  (about  1000  m/lO  km)  and  munerous 
cascades.  Fish  habitat  is  absent  u[)stream  from 
the  lake,  and  the  population  is  thus  plnsicalK' 
isolated  within  the  lake  and  the  short  .section  of 
stream  abo\e  the  waterfall. 

Methods 

Methods  and  obsei'xatiou  schedules  were 
influenced  b\-  the  relatixeK  remote  location  of 
the  study  site.  The  lake  is  located  within  a  des- 


ignated wild(M-ness  area  and  is  reached  \ia  a  trail 
that  extends  about  10  km  from  and  climbs  about 
10(10  III  al)()\('  the  nearest  motor \ehicle  access. 
Loss  of  ice  co\er  from  tlu^  lake  and  stream  and 
spawning  actixities  b\-  gra\ling  were  monitored 
through  weekK'  hikes  to  the  site  starting  in  late 
Ma\ .  Obsenations  of  fish  beha\ior  started iis  the 
ice  thawed  and  adults  began  entering  the 
stream,  mid-June  in  1989  and  late  juikmu  1990. 
and  ended  as  ice  started  forming  on  the  lake  and 
stream  margins  (10  No\ember  1989)  or  as  snow 
accumulations  on  the  trail  made  access  difficult 
(11  October  1990i.  Stream  temperature  was 
continuousK  recorded  throughout  both  obser- 
\ation  seasons  with  a  PeabocK  H\an  Motlel  D 
tliermograph  placed  about  30  m  dowTistream 
from  the  lake.  Daily  mean  temperature  was 
calculated  as  the  ayerage  of  (lail\-  maxiiuuiii  and 
minimum. 

Traps  with  1-m i ii-n lesh  n( 'tting  were  placed  to 
detenuine  the  dates  fn  became  free-swimming, 
and  to  monitor  their  moxement  downstream 
and  upstream  out  of  the  outlet  stream.  In  1989 
three  fry  emergence  traps  (45  X  45  cm)  of  the 
txpe  described  by  Fraley  et  al.  (1986)  were 
placetl  oxer  the  substrate  after  most  spawning 
had  ceased,  oyer  areas  where  fish  had  been  seen 
spawning  and  where  concentrations  of  eggs 
were  \isible.  Because  Arctic  gra\'ling  spawn 
o\'er  the  substrate  without  excaxating  redds, 
eggs  were  readily  \isible  among  the  substrate 
[)articl(\s.  One  emergence  trap  was  placed  in  a 
spawning  area  about  30  m  below  the  outlet,  and 
the  other  two  were  placed  in  the  printipal 
spawning  area  about  niidwax  through  the 
stream  length.  Swimup  fn  in  the  traps  were 
renuned  and  tabulated  daih  or  on  alternate 
da\s  until  emergence  ceased. 

One-wa\'  traps  wen^  placed  across  the  lake 
outlet  and  at  the  top  of  the  waterfall  to  monitor 
iiioxement  of  xoung  out  of  the  stream.  The 
upstream  trap  had  \-shaped,  screened  barriers 
extending compl(4el\- across  theoutlet  and  \vm\- 
ing  upstream  into  a  holding  box.  This  traji 
retained  fish  as  the\  entered  the  lake.  The  trap 
was  installed  after  most  adult  spawners  had  left 
tlu^  stream  but  before  the  xoung  became  free- 
swinnning.  After  installation,  the  trap  was  in 
contiimous  operation  through  both  obsenation 
seasons.  It  was  inspected  at  intenals  \aning 
from  seyeral  da\s  to  about  one  week;  young 
were  removed,  measured,  and  released 
upstream  into  the  lake. 

The  downstream  trap  was  a  drift  net  with  its 


346 


Great  Basin  Naturalist 


[Volume  52 


260 

220 

180 

S    140 
CD 

s: 

Z) 

^    100 

60 

20- 


—  Emerge 
D    Falls 


6      8     10    12    14    16    18    20    22    24    26    28    30     1      3 

JULY  AUGUST 

Fig.  1.  Total  nuinberof  voung  Arctic  crravlingCr/; ;///(rt//».v 
arctiais)  in  three  emergence  traps  placed  over  the  sub- 
strate, and  in  the  waterfdl  trap,  Deer  Creek,  Montana,  1989. 

opening  positioned  at  the  lip  of  the  waterfall;  it 
collected  \'(ning  that  were  going  over  the  ftxlls. 
In  1989  this  trap  sampled  about  0.3-0.5  of  the 
stream  \olume,  as  estimated  b\' comparing  flow 
rate  into  a  plastic  sack  attached  to  the  trap  versus 
estimated  stream  discharge  \'olume.  In  1990 
V-shaped  barriers  were  added  to  direct  all  flow 
through  the  net.  In  1989  the  trap  was  installed 
on  sampling  days  and  left  in  place  for  about  24  h 
before  the  young  within  were  tabulated  and 
measured.  The  trap  was  deployed  on  6  Julv, 
before  fry  became  free-swimming,  and  oper- 
ated at  intervals  of  one  to  two  days  until  num- 
bers in  the  trap  declined  shaiplv.  Thereafter,  the 
trap  was  operated  at  intenals  of  several  days  to 
two  weeks  until  19  October.  In  1990  this  trap 
was  operated  less  frequently,  at  intervals  rang- 
ing from  five  days  during  the  swimup  period  to 
about  four  weeks  in  September  and  October,  to 
determine  diel  patterns  of  movement  o\er  the 
falls  of  young  at  different  ages  post-swimup. 
Sampling  began  on  23  Julv  as  fiy  started  to 
swim.  On  sampling  dates  the  trap  was  deployed 
at  1000  or  1100  h  (Mountain  Standard  Time); 
the  trap  was  emptied  of  \()ung  at  1400  h,  and 
thereafter  e\-er\-  4  h  until  1000  h  the  next  da\-. 

Results 

Spawaiing  occurred  through  much  of  the 
350-m  length  of  the  stream,  from  about  10  m 
below  the  lake  outlet  to  within  15-20  m  of  the 
falls.  The  most  heavily  used  area  was  a  10-in 
reach  about  130-140  m  above  the  falls.  In  1989 
spawning  occurred  during  the  last  week  of  June, 
and  swimup  of  fr)-  in  the  emergence  traps  began 


about  1 1  }ul\-,  peaked  in  mid-month,  and  con- 
tinued until  about  25  Jul\'  (Fig.  1).  Spawning  in 
1990  occurred  during  the  first  week  in  |ul\',  and 
swimup  of  fiy  began  in  mid-month  and  contin- 
ued to  the  end  of  the  month. 

In  1989  fn'  started  appearing  in  the  falls  trap 
as  they  became  free-swimming  (Fig.  1).  Highest 
daih'  totals  of  frv  in  the  falls  trap,  generaUv  over 
200  per  dav,  occurred  15-22  Julv  as  mnnbers  of 
frv  becoming  free-swimming  in  the  emergence 
traps  peaked,  and  then  declined.  The  swimup 
period  ended  about  25  July;  thereafter,  within  a 
week,  numbers  of  young  in  the  falls  trap 
declined  to  0-6  per  dav.  No  voung  entered  the 
falls  trap  after  20  September. 

Movement  of  fry  over  the  falls  was  concen- 
trated within  a  19-clav  period,  13-31  July.  The 
falls  trap  was  operated  for  13  of  these  days,  and 
the  mean  number  of  frv  per  24-h  sample  was 
127.3.  Extrapolation  from  the  estimated  30- 
50%  of  total  stream  volume  that  passed  through 
the  net,  and  application  of  the  13-da\'  mean  to 
19  days,  yielded  a  crude  estimate  of  4837-8062 
young  grayling  lost  over  the  falls  13-31  July. 
Numbers  in  the  falls  trap  axeraged  onl)'  2.7  per 
day  during  the  1 1  davs  sampled  from  1  August 
to  20  September,  the  last  da)  voung  entered  the 
trap.  Similar  extrapolation  to  this  51-da\'  period 
yielded  a  cnide  estimate  of  an  additional  275- 
459  voung  lost.  Thus,  the  number  of  \oung  lost 
downstream  over  the  falls  in  1989  during  the 
period  from  swimup  of  fry  to  onset  of  ice  cover 
over  the  stream  was  roughly  estimated  at  5000- 
9000. 

Frv  were  alreadv  becoming  free-swimming 
w4ien  the  falls  trap  was  installed  on  23  Jul\'  1990. 
Numbers  of  voung  per  da\  in  the  falls  trap 
peaked  at  561  on  28  }ul\,  diminished  to  49  ten 
da\s  later  on  6  August,  and  to  5  bv 8  September. 
No  v'oung  entered  the  trap  on  12  October,  the 
last  dav  sampled  in  1990.  During  the  swimup 
period  fiy  went  over  the  falls  predominantly  at 
night  (23  and  28  Juh';  Fig.  2).  However,  there 
was  no  consistent  pattern  of  diurnal  \s.  noctur- 
nal movement  among  the  fewer  young  fish  that 
went  over  the  falls  on  later  dates  (6  and  17 
August;  Fig.  2).  Too  tew  daws  wert^  sampled  at 
the  falls  in  1990  to  estimate  total  numbers  lost. 

In  contrast  to  earK'  losses  over  the  falls, 
upstream  movement  of  voung  grayling  into  the 
lake  did  not  begin  until  late  summer,  when  the 
fish  were  larger  and  w^ater  temperatures  were 
cooling  (Fig.  3).  Small  numbers  of  voung  were 
trapped  at  the  lake  outlet  starting  in  mid-August 


1992] 


Stream  Monements  oe  Ac;e-()  Lake  (;iuvei\g 


341 


90- 

July   23 

80- 

70- 

i 

1 

cr      60- 

LU 

^      50- 

Z) 

z      40- 

30- 

20- 

10- 

1 

1 

July   28 

260- 

220- 

cr    180- 

LU 

m 

i 

1 

^    140-1 

z 

100- 

60- 

1 

20- 

1 

40- 

August   6 

cc 

LiJ     30- 

i    ao- 

i                            ♦ 

lOi 

1 

40- 

August    17 

n 

t^      30- 

1     20- 

i 

1 

10- 

1 

I 

0-^ 

1 

^ , 

12 


16  20  0 

HOUR   OF   DAY 


Fig.  2.  Diel  pattern  of  Noting  Arctic  gra\ling  (Thi/iiiallus 
arcticiis)  accumulated  in  waterfall  trap  during  3-  or  4-li 
sampling  periods  (100()  or  1100  h  to  1400  li,  then  at  4-li 
intenals  thereafter).  Deer  Creek,  Montiuia,  1990.  Note 
cluuige  of  \-a\is  on  Jnl\'  28.  Mean  sizes  of  \()ung  on  these 
sampling  dates  are  in  Figure  3.  Arrow  s  indicate  sunset  and 
sunrise. 


10     20     30       10     20     30       10     20     30      10     20 
July  August  SeptemDer      October 


Fig.  3.  Mean  dail\  temperature  i  (.'  ol  Deer  Creek, 
Montiuia,  mean  total  lengths  oi  age-0  gra\ling  [ThijmaUns 
iircticris)  in  waterfall  and  lake  traps,  and  numbers  of  age-0 
grayling  in  the  kike  trap,  1989  and  1990.  Aitows  indicate 
dates  when  numbers  of  \()ung  becoming  free-sun nmiing 
(collected  in  emergence  traps)  peaked  during  I9S9  and 
1990. 


1989  and  earK-  September  1990.  Total  nniiibers 
of  )oung  trapped  per  3-da\-  to  l-week  periods  in 
September  and  October  were  0-26  in  1989  and 
0-14  in  1990  (Fig.  3).  Onlv  95  age-0  vonng  had 
ino\ed  np  into  the  lake  in'l989  and  23  in  1990. 
before  obsei^ation.s  were  terminated  by  the 
()n,set  (A  winterlike  conditions  in  November 
1 989  and  October  1990.  While  age-0  grayling  in 
the  falls  trap  were  mostK  newl\-  swimming  fr\ 
that  a\eraged  12-14  mm  in  length,  the  smallest 
mo\ing  upstream  into  the  lake  trap  a\eraged 
52-54  mm  in  length  (Fig.  3). 


Although  numbers  of  resident  \-oung  in  thc^ 
stream  were  not  estimated,  \isnal  observations 
indicated  that  age-0  fish  were  abundant  in 
\o\  ember  1989  as  ice  was  starting  to  form  on 
the  stream,  but  were  present  in  much  fewer 
numbers  (as  age-1  fish)  when  ice  co\er  melted 
the  following  June.  The  age-1  fish  still  in  the 
stream  in  June  1989  and  1990  had  upstream 
{)atterns  of  movement  similar  to  those  of  the 
age-O  fish;  \en'  few  entered  the  lake  during  the 
[une-No\ember  .stud\  period,  and  these  lim- 
ited  upstream    movements    occurred   mostl)' 


348 


Great  Basin  Naturalist 


[N'olunie  52 


between  earl\  September  and  the  end  of  obser- 
vations in  October  or  November.  In  1989  onl\ 
se\en  age-1  fisli  were  in  tlie  lake  trap  from  June 
to  the  end  of  August,  and  38  more  from  Septem- 
ber to  the  end  of  obseivations  in  No\  ember.  In 
1990  onlv  two  age-1  fish  were  trapped,  both  in 
September.  Age-1  fish  were  nearl)'  absent  from 
the  falls  ti-ap;  three  were  trapped  in  1989  and 
two  in  1990.  Fish  older  than  age-1  were  rare  in 
tlu^  stream  when  ice  cover  thawed  in  June  of 
both  vears. 

louring  the  summer  of  1990,  six  adults 
remained  in  the  outlet  stream.  These  fish  were 
seen  in  shallow  water  (5-10  cm  deep)  chasing 
groups  of  young  in  late  July.  One  was  captured 
with  a  dip  net  and  had  12  age-0  gra\ling  in  its 
stomach. 

Discussion 

Since  we  did  not  estimate  the  number  of 
young  produced  in  the  stream,  we  do  not  know 
the  percentage  of  total  \'oung  lost  over  the  falls 
between  swimup  and  the  end  of  obsei"vations  in 
Octoloer  and  Noxember.  Two  considerations 
suggest  that  the  losses  represented  a  relatively 
small  percentage  of  voung  produced.  First,  it 
was  visually  apparent  that  age-O  young 
remained  abundant  and  wideK"  distributed 
throughout  the  stream  until  the  end  of  each 
obsenation  season.  Second,  we  estimated  that 
the  number  of  eggs  that  could  ha\e  been 
spawned  by  this  population  dining  1989  was 
about  1.3  million.  This  was  based  on  the  esti- 
mated average  of  2988  eggs  in  each  of  seven 
females  sampled  (range  2459-3674)  and  the 
estimated  number  of  426  adult  females  in  1989 
(Deleray  1991).  If  we  assume,  as  an  example, 
that  swimup  fry  resulted  from  10%  of  this  poten- 
tial egg  deposition,  then  the  estimated  loss  of 
young'over  the  falls  (500()-9( )()())  would  be 
about  4-7%  of  fry  produced  in  1989.  We  do  not 
know  of  any  estimates  of  the  relationship 
between  potential  egg  deposition  and  actual  fn 
production  b\-  gravling.  Howe\er,  a  figure  of 
10%  seems  consenative  compared  with  recent 
estimates  of  11. ,5-22.2%  for  chum  salmon 
iO)icoHu/nc]ius  kcta)  and  16.4-29. 17r  for  colio 
salmon  (O.  kisittcJi)  in  a  Canadian  stream,  with 
the  lower  percentages  associated  with  poor  sub- 
strate qualih-  (Scrivener  and  Brownlee  1989). 

The  grayling  lost  ckmiistream  were  predom- 
inantly small,  newlv  swimming  fW  that  went 
over  tlie  falls  at  niiiht.  The  nocturnal  dowii- 


stream  movement  of  the  young  was  similar  to 
those  of  young  from  inlet-spawning  populations 
of  grayling  (Knise  1959,  Lund  1974,  Wells  1976) 
and  other  salmonids  (McCart  1967,  Northcote 
1969,  Brannon  1972).  These  obseivations  were 
also  consistent  with  results  of  experiments  in  an 
artificial  stream  (Kaya  1989),  which  indicated 
that  although  young  Deer  Lake  gravling  had  an 
innately  greater  tendency  to  swim  upstream 
thim  tliose  of  an  inlet-spawiiing  population,  man\- 
mo\ed  downstream,  especial!)'  in  darkness. 

If  loss  o\er  the  falls  results  from  deliberate 
downistream  migration  by  the  yoimg,  then  this 
mav  indicate  that  the  Deer  Lake  population  has 
not  \et  completelv  adapted  to  outlet  spawning. 
If  so,  then  the  waterfall  is  continuing  to  act  as  a 
selectixe  factor  remoxing  those  voung  with 
inappropriate  responses.  Incomplete  adapta- 
tion has  also  been  suggested  as  an  explanation 
for  downstream  movement  bv  man\'  swimup  fiy 
of  rainbow-cutthroat  hvbrid  trout  that  spawn  in 
the  outlet  of  a  Colorado  lake  (Lentsch  1985). 
The  lake  had  first  been  planted  with  trout  about 
100  years  earlier.  Little  or  no  downstream  loss 
has  been  reported  from  populations  of  brown 
and  rainbow  trout  natixe  to  waters  abo\e  falls 
(Northcote  1969,  1981,  Jonsson  1982, 
Northcote  and  Hartman  1988),  in  contrast  to 
downstream  movement  oxer  cascades  of  an  esti- 
mated 22%  of  marked  rainboxv  trout  in  a  stream 
that  had  been  stocked  repeatedly  in  preceding 
years  xxith  nonnatix  e  rainboxx'  trout  (Chapman 
and  Max  1986).  The  Deer  Lake  population 
almost  certainlv  originated  through  a  transplant 
of  )()ung  from  an  inlet-spaxxaiing  population 
sometime  during  the  present  centun'.  In  Mon- 
tana, graxling  xx'ere  not  present  al:)oxe  natiu'al 
barriers  to  upstream  moxement,  and  the  onlx" 
lakes  xxithin  the  original  range  that  xx'ere  natu- 
ralh'  accessible  to  fish  and  knoxxn  to  haxe  con- 
tained natixe  graxling  xvere  Upper  and  Loxx'er 
Red  Rock  lakes  and  perhaps  Elk  Lake,  of  the 
Red  Rock  Rixer  drainage  (Nelson  1954,  \incent 
1 962).  Another  lacustrine  population  originated 
xxith  the  creation  of  Funis  Resenoir  on  the 
Madison  Rixer,  xvhich  contained  natixe  graxling. 
The  Red  Rock,  Elk,  and  Funis  populations  are 
inlet-spawning.  Populations  in  other  lakes  orig- 
inated through  stockings  that  began  after  artifi- 
cial culture  of  the  species  xxas  initiated  in  1898 
(Ilenshall  1906).  Unpublished  records  of 
regional,  state,  and  federal  hatcheries  inx'olxed 
in  these  stocking  programs  indicate  that  fertil- 
ized eggs  xx'ere  obtained  from  Upper  Red  Rock 


1992] 


S THKAM  M(  )\  IvVlENTS  OF  ACK-O  L\Kt:  GlUVLlNG 


349 


LakeorEiinis  Resen oiror other inlet-sjxiwniiitj; 
populiitions  established  thr()U!j;h  transplants 
from  these  two  sources  (Ka\a  19S9,  1990). 
Outlet-spaw  uing  populations  are  known  to  have 
exoKed  elsewhere  from  transplants  of  inlet- 
spawning  gra\linti;  (Kriise  1959)  and  lainhow 
trout  (Northeote  1969). 

It  is  possible  that  downstream  loss  ol  inanx 
\oung  fish  occurs  e\eu  from  populations  well 
adapted  to  spawning  above  a  waterfall.  With 
iiati\e,  abo\e-falls  populations  that  ha\e  been 
studied,  the  \()img  sampled  were  browni  trout 
from  about  10  cm  to  over  20  cm  in  length 
(lonsson  1982),  or  rainbow  and  cutthroat  trout 
whose  sizes  were  not  stated  (Noithcote  1969, 
19S1,  Northcote  and  Hartman  1988).  Given  the 
rapid  post-swlmup  decline  of  dowmstream 
movement  observed  in  the  present  stucK,  con- 
clusions on  magnitude  of  such  losses  would  have 
been  \en'  different  if  the  sampling  had  begun 
one  or  two  weeks  after  the  end  of  the  swimup 
period,  or  if  the  onK'  fish  sampled  were  lai'ger 
than  1.5-2.0  cm. 

Factors  other  than  deliberate  downstream 
moxement  could  ha\e  produced  losses  oxer  tlie 
falls,  including  passixe  drift  or  local  dispersal. 
Those  \c)img  that  were  lost  could  have  origi- 
nated from  eggs  either  spawiied  within  or 
drifted  to  locations  close  to  the  falls.  Adults 
spawned  within  15-20  m  above  the  falls,  and  we 
confirmed  visuallv  that  many  eggs  drift  down- 
stream from  spawning  areas  after  being  broad- 
cast oxer  the  sub!>trate.  Fn-  originating  from 
eggs  near  the  falls  could  be  lost  through  passixe 
drift  if  thev  became  free-sxximming  at  night  and 
xxere  consequentlx  displaced  doxxnstream  in  the 
darkness,  as  has  been  described  of  European 
graxling  (7'  tiiyinallus;  Bardonnet  and  Gaudin 
1990).  Doxxnstream  losses  could  also  represent 
passive  drift  of  dead  or  vmhealthx'  fish,  as  sug- 
ge.sted  by  a  report  that  819f  of  xoung  broxxn 
trout  produced  in  a  section  of  stream  did  not 
surxixe  and  drifted  downstream,  mostly  at  night 
(Elliott  1986).  We  did  not  attempt  to  determine 
the  health  of  xoung  graxling  in  the  falls  trap. 

Loss  oxer  the  falls  could  be  an  indirect  con- 
sequence of  local  dispersal  of  young  xvithin  the 
stream  as  thex'  became  free-.sxximming.  Young 
sockexe  salmon  {OncoHit/ncliiis  iicrka)  ol 
outlet-spaxxning  populations  liaxe  been 
reported  to  temporarilx  disperse  doxxnstream 
before  holding  position  or  sxximming  upstream 
into  lakes  (McCart  1967,  Brannon  1972).  Younu 
graxling  in  Deer  Dreek  iilso  disperse  localK 


from  th(^  immediate  spaxxiiing  areas,  some  of 
them  apparentlx'  doxxnstream.  For  tho.se 
becoming  free-sxximming  near  the  falls,  exen 
localized  doxxnstream  dispersal  coukl  result  in 
some  being  carried  oxer,  especiallx  under  con- 
ditions of  poor  xisibilitx'  at  night. 

The  results  indicate  that  Deer  Lake  grax  ling 
spend  at  least  the  first,  and  possibly  also  their 
second,  sunnnerand  earlx' to  mid-autunm  in  tlie 
outlet  stream,  lloxxever,  the  results  ditl  not 
permit  us  to  determine  the  exact  timing  of  most 
moxement  by  young  into  the  lake,  or  xx'hether 
they  move  upstream  predominantlx'  as  age-0  or 
as  age-1  fish.  The  x-ery  fexv  xoung  that  moxed 
into  the  lake  during  both  obsenation  seasons 
coidd  not  account  for  the  numbers  of  spaxxning 
adults  produced  in  the  population.  Since  there 
is  no  other  source  of  xoung,  and  since  the  1989 
obserxation  season  extended  oxer  the  entire  ice- 
free  period  on  the  stream,  maintenance  of  the 
Deer  Lake  pojiuiation  must  depend  on 
upstream  moxement  of  xoung  .sometime  during 
the  six  to  sexen  months  of  annual  ice  coxer. 
Althouo;h  age-O  xoung  greatlv  diminished  in 
numbers  and  age-1  fish  xirtuallx'  disappeared 
from  the  stream  betxxeen  the  on.set  of  ice  coxer 
in  Nox  ember  1989anditsthax\ingin  fune  1990. 
xxe  do  not  knoxv  the  proportions  ol  these  reduc- 
tions in  numbers  attributable  to  moxement  into 
the  lake,  death,  or  loss  oxer  the  falls.  The  greatlx' 
diminished  numbers  of  xoung  in  the  falls  trap 
during  late  sununer  and  their  absence  in  the 
trap  bx  October  of  both  xears  suggest  that  doxxn- 
stream losses  during  winter  max'  be  small.  The 
chronologx  of  major  moxement  bx  xoung  grax- 
ling into  the  lake  and  the  numbers  and  ages  of 
fish  inxolxed  xxould  need  to  be  resoKod  bx 
obserx  ations  during  xxinter. 

Little  is  knoxxn  about  duration  ol  stream 
residence  lor  outlet-spaxxniing  populations  ol 
.\rctic- grax  ling.  Younglrom  inlet-spaxxning  pop- 
ulations of  the  .species  txpicallx-  haxe  an  early 
descent  to  the  lake,  ranging  from  immediately 
after  sxximup  (Knise  1959,  Lund  1974,  Wells 
1976)  to  xxithin  .sexeral  xxeeks  (Nelson  1954). 
We  are  not  axxare  of  other  studies  on  stream 
residence  times  of  voung  grax  ling  from  outlet- 
spaxx'uing  populations  and  so  do  not  knoxx' 
w  hether  extended  period  of  stream  residence  is 
txpical  for  such  populations.  Young  rainboxx' 
trout  of  outlet-,spax\ning  populations  tend  to 
remain  for  extended  periods  of  at  least  a  month 
to  a  xear  or  more  before  migrating  upstream  to 
lakes,  xx'hile  those  of  inlet-spaxxning  populations 


350 


Great  Basin  Naturalist 


[Volume  52 


mio;rate  when  newlv  swimiuing  in  some  popula- 
tions and  after  extended  periods  of  stream  resi- 
dence in  others  (Northcote  1969).  The 
extended  stream  residence  of  voung  Deer  Lake 
grayling  is  also  consistent  with  their  lesser  ten- 
dencv  to  swim  upstream  in  an  artificial  stream 
as  early  fiy  (from  swimup  to  three  weeks),  com- 
pared with  their  responses  when  older,  within  a 
study  period  of  up  to  10  weeks  post-swimup 
(Kayal989,  1991). 

It  may  be  that  young  of  an  outlet-spawning 
population  need  to  attain  larger  sizes  and 
thereby  become  stronger  swimmers  before  they 
can  swim  upstream  into  the  lake.  However,  this 
possibilit)'  appears  contradicted  by  oiu"  casual 
obsei-vations  that  age-0  grayling  of  all  sizes  in 
Deer  Creek,  starting  from  those  newly  swim- 
ming, were  capable  of  swimming  upstream 
when  they  w^ere  disturbed  by  our  presence. 
Those  young  originating  from  spawaiing  areas 
within  a  few  meters  of  the  lake  outlet  could  ha\e 
entered  the  lake  by  mo\ing  onl\'  a  short  distance 
upstream. 

Another  possible  factor,  (jualitv  of  rearing 
habitat,  also  does  not  appear  to  favor  extended 
residence  in  Deer  Creek.  Deer  Lake  grayling 
grow  slower  during  their  first  two  years  than 
those  of  other  lacustrine  populations  studied 
thus  far  in  Montana,  but  thereafter  they  grow  at 
similar  or  faster  rates  (Deleray  1991).  Unlike 
yovmg  Deer  Lake  grayling,  those  from  inlet- 
spawning  populations  in  Montana  spend  their 
first  summer  and  autumn  growing  season  in 
lakes.  The  slower  early  growth  of  Deer  Lake 
gra)'ling  thus  appears  related  to  their  spending 
their  first  growing  seasons  in  the  stream  rather 
than  in  the  lake. 

We  speculate  that  young  Deer  Lake  grayling 
may  remain  in  the  outlet  stream  to  avoid  intra- 
specific  predation  in  the  lake.  Eriksen  (1975) 
obsen'etl  that  age-0  grayhng  in  several  Montana 
lakes  occupied  shallow,  near-shore  areas  among 
rooted  aquatic  vegetation,  and  suggested  that 
their  distribution  pro\ided  protection  against 
predation  by  the  adults.  Behavior  of  the  few 
post-spawning  adults  that  remained  in  Deer 
Creek  during  the  suuuner  of  1990  confirmed 
that  adults  will  prey  on  the  voung.  Young  gray- 
ling would  likely  be  susceptible  to  predation  by 
larger  conspecifics  in  Deer  Lake  because  of  its 
high  water  clarity-  throughout  the  summer  and 
the  lack  of  rooted  macrophvtes.  In  the  outlet 
stream  the  only  potential  predators  of  x'oung 
gra\ling  that  we  saw  were  the  relatively  few 


residual  adult  and  age-1  grayling  remaining 
through  the  summer,  and  an  occasional  belted 
kingfisher  (Axes,  Ccnjlc  alci/oii).  Thus,  the 
movements  of  age-0  Deer  Lake  grayling  that 
remain  in  the  outlet  stream  appear  adapted  both 
to  beginning  their  existence  a  short  distance 
aboxe  a  waterfall  and  to  a\oidance  of  predation 
b\-  larger  conspecifics  in  the  lake. 

Acknowledgments 

These  observations  were  part  of  a  stud)-  sup- 
ported by  a  grant  from  the  Montana  Depart- 
ment of  Fish,  Wildlife  and  Parks. 

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Received  7  May  1991 
Accepted  15  August  1992 


Givat  Basin  Naturalist  52(4).  pp.  352-356 

EFFECTS  OF  BROWSING  BY  MULE  DEER  ON  TREE  GROWTH 
AND  FRUIT  PRODUCTION  IN  JUVENILE  ORCHARDS 

Dennis  D.  AiLstin    and  Philip  J.  Unless 

Abstiuct. — The  effects  of  big  game  depredation  on  jnvenile  fruit  trees  were  studied  in  northern  Utah.  Utilization  of 
trees  was  determined  by  counts  of  nipped  tuid  intact  buds  in  spring.  Heiglit,  width,  l)asal  diameter,  number  of  l:)uds,  and 
initial  fruit  production  of  peach  and  apple  trees  were  determined  from  trees  protected  from  or  bi'owsed  b\'  mule  deer  in 
winter.  Results  from  the  10  orchards  studied  indicated  that  remov;il  of  buds  at  the  observed  browsing  levels  had  no  effect 
on  tree  growth  or  initial  truit  jirotluction. 


brow.' 


Ki'i)  tLortls:  (Icprcchitiou.  mule  deer,  orrluirds.  fruit  trcc\.  deer  dtiina^c  crdluiitinii.  <ipplc  trees,  peach  tree.s.  whiter 


^\'hene^•er  depi'edation  occurs  in  commer- 
cial orchards,  potential  crop  losses  due  to  big 
game  browsing  become  a  major  concern  to 
growers.  Bro\\'sing  of  juvenile  fitiit  trees  has 
important  economic  conse(jiiences  because  the 
effects  ma\"  limit  future  crop  production  and 
increase  tree  mortalit);  Research  has  clearK' 
shown  that  browsing  bv  big  game  on  mature 
apple  trees  causes  significant  crop  loss  within 
the  browsing  zone  (Katsma  and  Rusch  1979, 
1980,  Austin  and  Umess  1989).  However,  lim- 
ited information  on  the  effects  of  browsing  on 
jmenile  fniit  trees  is  extant. 

Westwood  (1978)  suggested  deer  browsing 
may  be  especialK  damaging  to  young  trees,  but 
rarely  would  browsing  be  expected  to  cause 
niortalitA.  Harder  (1970)  reported  no  differ- 
ences in  trunk  diameter  growth  between  pro- 
tected and  unprotected  apple  trees  with  one 
wint{M-  of  l)ud-remo\al  browsing  b\  mule  deer. 
In  this  ('olorado  stud\  of  160  trees,  no  mortality 
was  attributed  to  bud-reuKjxal  browsing, 
although  8  trees  died  as  a  result  of  bark  damage 
caused  by  antler  rubbing.  Similarly,  McAninch 
et  al.  (1985)  in  a  New  York  study  reported  9  of 
]()  growth  parameters  measured  between  pro- 
tected and  browsed  trees  showed  no  significant 
differences.  One  parameter,  basal  diameter,  was 
smalk^r  on  browsed  trees.  However,  this  studv 
with  white-tailed  deer  also  showed  that  axerage 
diameters  of  brow.sed  limbs  appeared  greater 


than  protected  limbs,  suggesting  possible 
growth  stimulation  as  a  result  of  deer  browsing. 
In  our  project  onK  bud-remo\al  browsing 
was  studied,  and  since  browsing  dunng  summer 
was  negligible,  we  considered  onl)  o\en\inter 
depredation.  The  puipose  of  this  study  con- 
ducted in  northern  Utah  was  to  measiu'e  the 
degree  of  browsing  in  xoung  fruit  trees  and  to 
assess  the  browsing  effects  on  tree  growth  and 
initial  crop  production. 

Methoi^s 

The  percentage  of  buds  browsed  b\-  mule 
deer  was  determined  in  March,  dunng  late  dor- 
mancy, after  deer  .switched  diets  from  winter 
browse  to  herbaceous  spring  growth  (Kufeld  et 
al.  1973,  Austin  and  Unless  1983).  Percent  bud 
remoxal  was  determined  b\'  counting  all  intact 
and  nipped  buds  and  then  dividing  nipped  buds 
bv  the  total  nipped  plus  intact  buds.  Nipped 
buds  are  easih'  identified  b\  the  exposed  and 
broken  woody  twigs  (Katsma  and  Rusch  1979). 
Counted  intact  buds  were  restricted  to  terminal 
buds  of  the  previous  summers  annual  growth, 
and  all  protruded  buds  along  second-vear  and 
older  stems  >1  cm  in  length  (Austin  and  Umess 
1987).  Protnided  was  defined  by  visualizing  a 
perpendicular  line  from  the  twig  to  the  tip  of  the 
bud,  and  an  obsenable  space  was  re(juired 
between  the  line  and  the  bud-twig  intersection. 

Tree  growth  measurements  were  taken  after 


Department  ol'Raiiile  Seience,  Utali  State  Univer.si(\ ,  Lxigaii,  Utali  84:322-.523(). 


352 


19921 


Dekh  Bhowsinc;  in  |r\  iaii.i:  Okcii ahds 


353 


tlie  end  of  the  growing  season  ImU  before  winter 
browsing  occurred.  Tree  height  was  measured 
to  tlie  nearest  1.0  em  from  ground  le\('l.  tree 
width  to  (he  nearest  1.0  cm  at  the  height  where 
maximum  width  occnn-(nh  Width  was  measured 
in  north-south  and  east-west  directions  and  the 
mean  recorded.  Basal  trunk  diameter  was  mea- 
sured to  the  nearest  0. 1  cm  using  dial  calipers  at 
10  cm  ah()\e  the  graft  scion.  Diameter  was  sim- 
ilarh  measiu-ed  on  north-south  and  east-west 
directions  antl  the  mean  recorded.  The  number 
of  intact  buds,  using  tlie  same  definition  as  tliat 
for  bud-remo\al  determinations,  was  counted 
using  hand-tall\'  registers.  W'liere  hanestable 
crops  were  produced,  all  fniits  were  hand- 
picked  and  counted.  Specific  methods  are 
reported  in  the  results  for  each  orchard. 

Data  were  anal\"zed  between  prottx'ted  and 
browsed  trees  and  bet\\'een  trees  with  \arious 
intensities  of  browsing,  using  the  standard  t  test 
of  the  means.  Confidence  lexel  was  .set  at  P  <  .05. 

Results 

Orchard  1 

A  4  X  6  block  of  24  ec^ual  age  and  size  Elberta 
peach  trees,  planted  in  spring  1986,  was 
selected  for  study.  Alternating  trees,  deter- 
mined b\  coin  toss,  were  fenced  during  three 
winters,  1986-89.  During  the  fourth  winter, 
1989-90,  all  trees  were  fenced.  Because  within- 
vear  browsing  effects  decrease  fniit  production 
(Katsnia  and  Rusch  1980,  Austin  and  Umess 
1989),  trees  were  protected  from  browsing  to 
compare  production  between  prexionsK 
browsed  and  protected  trees.  Tree  measuic- 
inents  were  taken,  and  peaches  were  hand- 
picked  and  counted  in  late  summer  1990,  the 
lirst  year  of  commercial  harvest. 

Percent  bud  remoxal  as  measured  in  spring 
1987,  1988,  and  1989  was  35.6,  76.6  and  73.57^. 
respectively.  Even  with  (his  high  degree  of 
brow.sing  by  deer,  trees  fulK  recoxcred  during 
(lie  summer  groxxing  seasons.  No  differences 
between  protected  and  browsed  trees  were 
found  for  anx-  tree  measurements  or  fruit  pro- 
duction (Table  1  ). 

Orchard  2 

A  small  commercial  orchard  comprising  210 
Elberta  peach  trees  x\as  planted  in  spring  1986. 
Percent  oxenxinter  bud  remoxal  xvas  deter- 
mined in  earlx-  spring  1987.  Since  9  trees 
shox\ed    bark    scraping    damage,    they    xxere 


deleted  from  the  sample.  Trees  were  placed  into 
three  ecjnal  groups  of  67  bx'  the  percentage  of 
bud-remoxal  browsing  damage:  heaxy  61- 
100%,  moderate  34-60%,  and  light  0-33%. 
Tree  measurements  xvere  made  folloxxing  the 
1987  summer  growing  period.  No  differences  in 
tree  measurementsxx'ere  found  aniongthe  three 
intensities  of  browsing  bx  mule  decM^  (Table  1). 

Orchaicl  3 

TweKc  [xuvs  of  ecjual  age  and  size  Yellow 
I^elicious  aj)ple  tr(H\s  w(m'(^  carefully  .selected  bx' 
( )ci  1  lar  ( )1  )seiA  at  ion  wi(  hi  1 1  a  commercial  orchard 
planted  during  spring  1984.  One  tree  of  each 
pair,  determined  bx  coin  toss,  xx'as  protected 
liom  broxvsing  bx'  fencing  dming  fixe  xxinters, 
1984-89.  During  the  .sixth  winter.  19S9-90.  for 
tlu^  same  reason  as  described  for  orchard  1.  all 
trees  were  fenced. 

Percent  bud  remoxal  from  browsing  was 
76.4, 60.5, 41.7. 23.6.  and  63.2%  foryears  198.5- 
89,  respectixelx'.  No  differences  betxx'een  pro- 
tected and  broxvsed  trees  were  found  for  anx' 
tree  measurements  or  I  ruit  production  Table  1) 

Orchard  4 

Twelxe  pairs  of  equal  age  and  size  Red  Deli- 
cious apple  trees  xx'ere  carefullx'  selected  b\ 
ocular  ob.seiA ation  xxithin  a  connnercial  orchard 
planted  in  spring  1983.  One  tree  of  each  pair, 
determiiu^d  b\  coin  toss.  x\as  protected  from 
broxvsing  bx  fencing  during  three  x\inters, 
1984-87.  During  winter  1986-87  a  deer-proof 
fence  xvas  constructed  around  the  orchard,  and, 
cf)ns(H|uentlx,  deer  use  was  close  to  zero  (0.4% ). 
During  the  txx'o  prexious  winters  (1984—86)  per- 
cent bud  remoxal  xx'as  71.0  and  17.0%,  respec- 
ti\(4x.  No  differences  between  protected  and 
browsed  trees  xxere  found  for  either  tree  niea- 
surcMuents  or  number  of  fruits  (Table  1).  Also, 
flow(>r  cluster  counts.  x\hich  were  collected  in 
spring  1987  as  part  of  an  ongoing  jiarallel  stud\- 
(Austin  and  Unless  1987),  showed  no  difference 
between  protected  (x  =  166)  and  broxxsed  (x  = 
169)  trees. 

Orchard  5 

Txx'elxe  pairs  of  equal  age  and  size  Red  Deli- 
cious apple  trees  xx'ere  selected  xxithin  a  com- 
mercial orchard  planted  in  spring  1985.  One 
tree  of  each  pair,  determined  bx'  coin  toss,  x\as 
protected  from  broxxsing  during  four  xxinters, 
1985-1989.  During  the  fifth  winter.  1989-90,  all 
trees  xx^ere  fenced. 


354 


Great  Basin  Naturalist 


[Volume  52 


Table  1.  Mean  growth  incasurement.s  and  initial  fruit  production  from  juvenile  peach  and  apple  trees  protected  from 
or  browsed  bv  mule  deer  in  winter 


Mean  tree  measurements 


Orcluird 
No.  Fruit  tree 


Treatment 


Years 


Basal 
%  buds      Height         Width     diameter      No.  of       No.  of 
removed      (cm)  (cm)         (mm)  buds         fniits 


1 

Elberta  peach 

Browsed 

12 

1986-90 

62 

225 

257 

5.6 

104 

Protected 

12 

230 

247 

5.7 

— 

103 

2 

Elberta  peach 

Hea\il\ 

browsed 

67 

1986^87 

61-100 

120 

88 

2.6 

61 

— 

Mfxlerately 

browsed 

67 

34-60 

124 

92 

2.7 

67 

— 

Lightly 

browsed 

67 

0-33 

122 

91 

2.7 

65 

— 

3 

Yellow  Delicious 

Browsed 

12 

1984-90 

53 

192 

1.36 

5.1 

250 

72 

apple 

Protected 

12 

193 

149 

5.2 

238 

70 

4 

Red  Delicious 

Browsed 

12 

1984-87 

44 

569 

248 

4.4 

.349 

75 

apple 

Protected 

12 

588 

262 

4.4 

375 

59 

5 

Red  Delicious 

Browsed 

12 

1985-90 

24 

259 

163 

5.4 

.577 

3 

apple 

Protected 

12 

250 

158 

5.4 

570 

3 

6 

Golden  Delicous 

Heaxily 

apple 

browsed 
Moderatelv 

20 

1987 

6.5-92 

198'' 

9:1' 

3.5 

96 

— 

browsed 

20 

2<8-64 

192'' 

88 

3.5 

93 

— 

Lightly 

browsed 

20 

0-27 

175'' 

8& 

3.5 

92 

— 

7 

Red  Delicious 

Hea\ily 

apple 

browsed 
Moderatelv 

8 

1985-.S6 

49 

88 

22 

1.7 

11 

— 

browsed 

S 

21 

98 

30 

1.8 

10 

— 

Protected 

8 

92 

21 

1.6 

" 

— 

8 

Mcintosh  apple 

Hea\ily 

browsed 

8 

1985^86 

50 

132 

62 

2.4 

31 

— 

Moderately 

browsed 

8 

35 

126 

47 

2.1 

22 

— 

Protected 

8 

129 

44 

2.6 

17 

— 

9 

Jonathan  apple 

Ileavilv 

browsed 

8 

1985-^86 

28 

147 

69 

2.4 

26 

— 

Moderatelv 

browsed 

8 

22 

123 

48 

2.0 

22 

— 

Protected 

8 

131 

69 

2.0 

45 

— 

10 

Red  Delicious 

Browsed 

12 

1985-87 

39.4 

167 

67 

5.1 

90 



apple 

Protected 

12 

159 

63 

5.0 

107 

— 

'  'F"igiires  with  tltffcrt'iit  .supi.Tscriptfii  nnnihi-rs  uitliiii  tolii 


vere  .signiHcaTitK  (lilferent,  P  <  .0.5. 


Percent  bud  renunal   hoiu  browsing  was  Orcliard  6 
16.7,  ().(),   16.7,  and  61.0  for  years   1985-89, 

respecti\-ely.  No  differences  behveen  protected  A  2  x  .30  block  of  60  two-year-old  Golden 

and  browsed  trees  were  found  for  any  tree  mea-  Delicious  apple  trees  was  measured  for  over- 

surenients    or    fmit    production,    which    was  winter   bud-reni()\al    browsing   use   in   spring 

greatly  reduced  in  1990  dut^  to  cold  temptMa-  1987.  Utilization  during  the  pre\ious  winter  was 

tures  in  spring  (Table  1).  unknowai,  but  was  probably  similar  to  the  use 


1992] 


Dkkh  Hi^()\\si\(;  i\  ji  \  kmlk  Ohcilvj^ds 


355 


ineasunHl  in  1987.  Percent  lnul  renunal  ranged 
from  0  to  927f .  with  a  mean  of  46.79f  (Table  1  ). 
Trees  were  plactnl  into  three  groups  of  20  hv 
l)ud-renio\-al  classes:  0-27,  28-64,  and  64-929f . 
SinprisingK",  heaxilv  and  moderateK'  browsed 
trees  had  significanth'  greater  height  at  the  end 
of  the  growing  season  than  lightK  browsed 
trees,  and  hea\il\  browsed  trees  also  had  greater 
width  than  lightK'  browsed  trees  (Table  1). 
Although  other  factors,  such  as  pRuiing,  could 
ha\e  accounted  for  these  increases,  height  and 
width  ma\  ha\e been  increased b\  browsing.  No 
differences  were  found  in  basal  diameters  oi- 
number  of  buds. 

Orchards  7.  S.  9 

Twentv-four  ecjual  age  and  size  trees  of  Red 
Delicious,  Mcintosh,  and  Jonathan  apples  were 
planted  in  spring  1985  for  this  stud\'.  In  winter 
1985-86,  one-third  (8  of  each  species)  of  the 
trees,  randoniK'  selected,  were  protected;  one- 
third  receixed  moderate  browsing  by  tame  mule 
deer  as  modified  by  temporary  fencing;  and 
one-third  recei\ed  hea\A'  browsing.  Mean  bud 
remo\al  \aried  from  21  to  35%  under  moderate 
browsing,  and  28  to  50%  under  heavy  browsing 
(Table    1).    Following    the    summer    growing 

season  in   1986,  no  significant  srowth  differ- 
ed o 

ences  in  tree  measurements  were  found 
betx\een  protected,  moderately  browsed,  or 
hea\il\  browsed  trees  (Table  1). 

Orchard  10 

TweKe  pairs  of  equal  age  and  size  Red  Deli- 
cious apple  trees  were  selected  within  a  com- 
mercial orchard  planted  in  spring  1983.  One 
tree  of  each  pair,  determined  b\-  coin  toss,  was 
protected  from  browsing  during  winters  1985- 
87.  Percent  bud  removal  from  browsing  was 
76.6,  37.4,  and  4.1%,  respectixelw  No  differ- 
ences between  protected  and  browsed  trees 
were  found  (Table  1). 

Discussion 

Percentages  of  bud  remcnal  measured  Irom 
these  10  orchards  were  mostk'  less  than  65%. 
Browsing  by  mule  deer  during  winter  dormancv' 
at  this  level  of  use  was  not  sufficient  to  cau.se  a 
decrease  in  tree  growth  parameters  measured. 
From  the  view  of  carboh\drate  resenes, 
decreased  producti\it\  would  not  be  expected 
if  the  total  number  of^  intact  buds  axailable  for 
spring    growth    were    sufficient    to    maintain 


balance  with  the  root  swstem.  This  was  the 
obsened  case. 

In  this  stiuK  trees  were  not  browsed 
sexerely.  As  a  suggestcnl  dehnition,  severely 
browsed  trees  would  include  browsing  of  >90% 
of  the  axailable  protruded  buds,  removal  of 
>70%  of  the  current  animal  growth,  scraped 
bark  on  the  central  leader  and/or  scraped  bark- 
on  two  or  more  priman-  branches,  or  limb 
breakage.  C-'eitaiuK,  as  the  level  of  browsing 
increases  toward  severe  levels,  the  potential  for 
permanent  daiuage  and  reduced  growth  also 
increases.  The  level  of  l)r()wsing  intensitv' 
needed  to  damage  juxenile  fruit  trees  is 
unknowii,  but  it  is  apparenth  higher  than  that 
w  hich  occurs  in  most  depreciation  situations  in 
northern  Utah  and  elsewhere  (Harder  1970, 
McAninch  et  al.  1985). 

The  intensitv  of  browsing  needed  to  cause 
measurable  damage  would  also  be  expected  to 
\"an-  with  the  qualitv*  of  the  horticultural  prac- 
tices inx'olved  in  managing  the  orchard.  In  this 
stud\  all  orchards  received  high-intensit\'  care, 
including  adequate  irrigation,  periodic  spra\- 
ing,  weed  control,  etc.  Orchard  trees  receixing 
lower  intensities  of  care  and  increased  emiron- 
mental  stress  from  pests,  or  competition  from 
weeds,  may  respond  negativelv  to  similar  levels 
of  deer  browsing. 

In  conclusion,  the  results  from  this  stud\  of 
juvenile  apple  and  peach  fruit  trees  were  con- 
sistent with  pre\ious  research  (Harder  1970, 
McAninch  et  al.  1985).  Browsing  bv  mule  deer 
at  the  intensities  observed  had  no  negatixe 
effects  on  tree  height,  width,  basal  diameter, 
number  of  buds,  or  initial  fruit  production. 

ACk'XOWLF.nCMF.XTS 

This  report  is  a  contribution  ot  the  Itali 
State  Dixision  of  Wildlife  R(\s{)urces,  Federal 
Aid  Project  \V-105-R. 

Liti:k.\tl'hk  Citkd 


.'Vl  sri\  I).  D..  AM)  P.  J.  Uhnkss  198.3.  Overwinter  forage 
.selection  bv  mule  deer  on  seeded  big  sagebnisli-grass 
range.  Journal  oiWildlife  Management  47: 1203-1207. 

.  1987.  Guidelines  for  evaluating  crop  los.ses  due  to 

depredating   big    game.    Utidi    Di\ision    of    Wildlife 
Resources  Publication  87-5.  42  pp. 

1989.  E\'iJuating  production  losses  from  mule  deer 


deprecLition  in  apple  orcluirds.  Wildlife  Societv  Bulle- 
tin 17:  161-16.5. 


L 


356 


Great  Basin  Natuhallst 


[N'oluiiie  52 


riMiDEH,  J.  D.  1970.  Evdiiatingvvinterdeeruse  of  orchards 
in  western  Colorado.  Transactions  of  the  Nordi  Amer- 
ican Wildlife  Conference  35:  35^7. 

IGvTSMA.  D.  E.  AND  D.  H.  Ruscil  1979.  Evaluation  of  deer 
damage  in  mature  apple  orchards.  Pages  123-142  in 
J.  R.  Beck,  ed.,  \'ertel)rate  pest  control  tuid  manage- 
ment materids,  ASTM  STP  680,  American  Societ\-  for 
Testing  Material. 

.    1980.    Effects   of  simulated   deer   browsing   on 

branches  of  apple  trees.  Journal  of  W'ildlile  Manage- 
ment 44:  603-612. 

KuFELD.  R.  C,  O.  C.  Wallmo,  and  C.  Ff.ddema  1973. 
Foods  of  the  Rockv  Mountain  mule  deer.  United  States 


Department  of  Agriculture,  Forest  Service  Research 

PaperRM-111.31pp, 
Mc'^NiNcii.  J.  B.,  M.  R.  Ellingwood  M.  J.  Farcjione. 

AND  p.  PicoNE  1985.  Assessing  deer  damage  in  young 

fruit  orchards.  Proceedings  of  the  Wildlife  Damage 

Control  Conference  2;  215-223. 
Westwood.  M.  N.  1978.  Temperate-zone pomolog\-.\\'.  H. 

Freeman  and  Companx;  San  Franscisco.  Calitomia. 

428  pp. 


Received  20  April  1992 
Accepted  18  September  1992 


(Jrt-at  Basin  Naturalist  52i4i.  pp.  o57-.')fi3 

chanc;es  in  riparian  \'E(;etati()n  along  the  Colorado 
rl\  er  and  rio  grande,  colorado 

\\  ancii  D.  SiiNclci"    aiul  (ianCJ.  Miller" 

Abstract — Clianges  in  vegetation  inchicling  area  oteiipied.  canopx  co\er,  ;uk1  niatiirit)'  class  of  cottonwoods  (Poptihts 
spp.)  within  lower-clcxation  zones  of  the  (Colorado  Hiverand  Rio  (Grande  in  (Colorado  were  monitored  o\er  25-  and37-\ear 
inten als,  respectively,  nsing  photo-inteipretatixc  nietliocLs.  l"",stiniated  loss  oi  cottonwoods  along  the  C^olorado  Ri\er  was 
2  lui/kni  ( — 17.5% ),  and  remaining  stands  had  become  more  open  and  older.  ( "ottonwoods  along  the  Rio  (Grande  increa.sed 
1  .f)  h;i/km  (9.3% )  with  minor  canopx'  cover  and  maturitx  class  changes.  Area  occnpied  1)\'  shrnhs  and  ri\er  channel  changed 
little  along  the  ("olorado  Ri\er,  hut  declined  along  the  Rio  Grande.  Loss  of  ha\  meadow  occurred  along  both  ri\ers.  whereas 
dexeloped  land  increasi'd  along  the  Colorado  River  and  iarmland  increased  along  the  Rio  CJrande.  W'ildlile  habitats  alon<4 
the  Colorado  deteriorated  nuich  more  rapidl\  than  diose  along  the  Rio  Grande  tluring  mcjiiitored  intervals. 

K('i/  ircnis:  riparian.  Colorado,  iiiroiton/.  coiiomvood.  Populus  ,v^)^j..  wihllijc  liahitaf. 


Ri\-eriiie  .swstems  in  the  Great  Basin  and 
southwestern  United  States  are  important  hab- 
itats for  resident  and  niigraton  wildlife  (Ander- 
son and  Ohniart  19S(),  Ilnnter  et  al.  1985).  Two 
major  ri\er  swstems  (Colorado  and  Rio  Grande) 
in  the  southwestern  United  States  originate 
within  Colorado.  While  substantial  work  has 
been  conducted  to  identify  wildlife  use  and  to 
manage  riparian  habitats  in  lower  reaches  of 
these  ri\er  swstems  (Stexens  et  al.  1977,  Ander- 
-son  et  al.  1978,  Anderson  and  Oh  mart  1980, 
1985,  Swenson  and  Mullins  1985),  little  infor- 
mation has  been  published  from  studies  con- 
ducted near  the  headwaters  of  these  ri\ers. 

The  cottonwood-willow  {Popidus-Salix) 
riparian  ecosystem  along  Colorado's  major 
rixers  has  the  highest  wildlife  species  richness 
and  densits'  in  the  state  ( Beidleman  1 978,  Fitz- 
gerald 1978,  Hoover  and  Wills  1984)  andisu.sed 
In  283  species  of  \ertebrate  wildlife.  Howe\-er, 
most  studies  ha\e  centered  on  the  South  Platt(" 
Rixer  in  northeasteni  Colorado  (Graul  and 
Bissell  1978).  Wildlife  \alues  of  riparian  habitats 
along  streams  and  rivers  in  the  mountainous 
western  two-thirds  of  (Colorado  ha\ e  receixed 
little  stud\.  Among  ecosx'.stems  in  mountainous 
areas,  cottonwood-willow  rixerbottoms  nsnalK' 
possess  higli  \alues  for  resident  and  migratorx' 
wildlife  (Schnipp  1978,  Thomas  et  al.  1979, 


Melton  et  al.  1984).  Awareness  of  these  x'alues 
has  increascnl  in  recent  \ears  along  with  concern 
for  increasing  actixities  in,  and  degradation  of. 
these  critical  xxildlife  zones  (Windell  1980). 
These  habitats  are  of  .special  concern  in  moun- 
tainous areas  because  xallevs  are  frequentlx 
narroxx'  and  centers  of  himian  actixih'. 

Before  attempting  to  manage  riparian  xegeta- 
tion  for  xxildlife,  it  is  necessarx'  to  leani  xx'hether 
these  habitats  are  declining  in  ability  to  sustain 
species  richness  and  abundance.  This  paper 
assesses  recent  changes  and  status  of  riparian 
xegetation  along  the  Rio  (irande  and  Colorado 
Rixer  in  southern  andxx'estem  C'olorado. 


Study  a  hi;  a 

Lf)xx'er-elexation  zones  of  the  Rio  Grande 
and  (Colorado  Rixer  in  C>olorado  xxere  selected 
for  study  (Fig.  1,  Table  1).  The  Colorado  Rixer 
and  its  tributaries  drain  about  46,196  km"  of 
western  Colorado  (Ugland  et  al.  1984,  \V)I.  2). 
The  Colorado  Rixc-r  is  confined  to  relatixelx' 
narroxx- xallex s  until  it  is  joined  bx  the  (immison 
Rixer  near  CJrand  function  xxhere  the  xallex 
broadens  xxith  reduced  .stream  gradient.  It 
leaxes  the  state  xxith  floxxs  approximatelx  75% 
greater  tlian  at  the  upstream  end  of  the  studx 
area  (Table  1). 


^Colorado  Dhi.sion  ol Wildlifi-.  .306  Cottonwood  Liine,  Sterling,  Colorado  807.51. 
"Colorado  Dhiskm  of\\ildlif"f,.317X\'.  Prospect  Hoad.  Fort  Collins.  Colorado  80.526. 


357 


358 


Great  Basin  Naturalist 


[\ blume  52 


WYOMING 


NEW  MEXICO 


Fig.  1.  C^olorado  River  and  Rio  Grande  with  iinentoried  portions  ( — )  and  segments  (  |  )  in  western  and  south  centr 
Colorado. 


Tablk  1 .  Characteristics  of  variables  measured  along  th 
Colorado  River  and  Rio  Grande,  Colorado. 


\ariable 


Colorado  Ri\er      Rio  (Traude 


.V  sampling  intervixl,  \t.s''  25.0 

Distance  sampled,  km  '  1 67.3 

Sample  units  21 

.V  hii/sample  unit  ST.O 

Sampling  intensiU',  %  20 
Elc\ati()n,  m 

upper  1829 

lower  i;372 
X  daily  stream  flow,  m  Vs 

ii])per  1005 

lower  175. .5 


36,8 
117.4 

20 
163.2 


2438 
2286 


25.3 
7.0 


■'.Vi-rial  plioto.s  were"  fn 

to  19.S()(C;olora<l()Kiverl, 

Liticar  di.staiKc  wa,s  ri 


1941  to  197.3-S:?  iKio  CraiKlrl  .uul  Ironi  19.51 -.57 
■siireilal  llu- cfiitc-r  ol  tin-  n\rr  tlianiu'l. 


Tlie  Rio  Grande  drain,s  appro  .xiniately 
1 9, 1 94  knr,  of  which  7612  knr  i,s  within  a  closed 
l)a,sin  in  sonth  central  (Colorado  (llglanel  et  al. 
1984,  Vol.  1).  River  flow  originates  priinariK  in 
the  San  jnan  Range  with  lesser  anionnts  from 
the  Sano;re  de  Gristo  Ran<ie.  The  ri\  er  enters  the 
western  part  of  the  San  Luis  \  alley,  a  high- 
elevation  (2286-2438  m)  park,  and  travels 
throngh  farmed  areas  for  approxiniateK  100  km 
(where  most  stream  flows  are  used  for  irrigation 
[Table  1  ] )  before  entering  a  cam  on  that  extends 
into  New  Mexico. 

Harrington  (1954)  noted  that  luirrowleaf 
cottonwoods  {P.  angtisiijoJui)  dominate  along 
the  Rio  Grande  and  upper  pc^lions  of  the  Col- 
orado   Ri\er,   whereas   lanceleaf  cottonwoods 


1992] 


CoLOHAi:)()  Hii'AHiAN  \'Kc;t:rAri()\ 


359 


(P.  acuminata)  occur  sparselv  over  a  slightK' 
broader  ele\ation  range.  Rio  Grande  cotton- 
woods  [P.  wislizeni)  dominate  at  lower  ele\  a- 
tions  along  the  Colorado  Ri\er.  Willows  are  tlie 
priiiiaiA' shrubs  along  the  Rio  Grande  and  uppei' 
portion  of  the  Colorado  Ri\"er  gi\ing  wa\  to 
tamarisk  (Tanuirix  (^allica)  at  lower  elevations 
along  the  latter  iplant  names  follow  I  hirrington 
11954]). 

Methods 

ApproximateK  167  km  ot  the  Colorado  Ri\er 
and  1 1 7  km  of  the  Rio  Grande  were  selected  for 
stndv  and  respectixeh  stratified  into  four  and 
three  segments  (strata)  based  on  empirical 
assessments  of  \egetation  (area  occupied  b\'  cot- 
tonwoods,  plot  width,  etc.;  Fig.  1).  Segments 
(numbered  from  upstream  to  downstream;  Fig. 
1)  were  used  to  distribute  random  sample  units 
(linear  1.61-km  river  tracts)  more  uniformK 
along  the  ri\ers.  Twent\'  sample  units  were  dis- 
tributed along  the  Rio  Grande,  whereas  the 
Colorado  River  stud\-  area  contained  21.  An 
electronic  planimeter,  positioned  at  mid-chan- 
nel on  U.S.  Cicological  Sune\  topographic 
maps,  was  used  to  delineate  the  randomh 
selected  1.61-km  (ri\er  mile)  sample  units. 
Width  of  sample  imits  varied  and  was  based  on 
flood  plain  width,  primariK'  encompassing  nat- 
ural riparian  xegetation  readiK  discerned  on 
aerial  photos  (some  adjacent  cropland  and 
grassland  were  included). 

The  earliest  (scale  1:20, ()()())  and  most  recent 
(.scale  1:40,000)  aerial  photos  axailable  (U.S. 
Department  of  Agriculture)  were  acquired  for 
each  sample  unit  to  \ield  changes  over  time. 
The  same  area  was  inxentoried  within  each 
sample  iniit  duiing  both  earlv  and  recent  inter- 
\als  to  assess  changes.  Earliest  aerial  photos 
were  from  1941  and  the  most  recent  photos 
were  from  1973  through  19S3  for  the  Rio 
C^rande.  Those  for  the  CJolorado  Rixcr  were 
from  1951-57  (early)  and  1980  (recent  >. 

Inteipretati\e  anaKses  of  aerial  photos  were 
contracted  to  the  C>'olorado  State  Forest  Senice. 
Vegetation  t\pes,  including  trees  (primariK  tot- 
t(^nwoods),  shmbs  (tamarisk  [Colorado  Rixcr] 
and  willow),  hav  meadows,  grasslands,  agricul- 
ture (farmland),  de\"eloped  (roads,  towns,  etc.), 
ri\en  standing  water,  and  umegetated  (sand- 
bars) were  delineated  on  acetate  overlaxs  using 
a  stereoscope.  Ri\er  and  unvegetated  wei(> 
combined  as  ri\er  channel.  Minor  vegetation 


tynpes  {<!%  of  total  area)  were  omitted.  The  area 
per  \egetation  t) pe  was  recorded  to  0. 1  ha  using 
an  electronic  planimeter.  On-site  inspections 
w(^re  conducted  within  se\-eral  plots  along  both 
risers  to  \ciil\  that  [ilioto  interpretation  was 
accurately  assessing  cottonwood  stand  matnritA, 
canopy  co\er,  and  \egetation  types.  Photo  inter- 
pretation accuiacv  approximated  95%. 

Maturit)  classes  (tmnk  diameter)  were  esti- 
mated from  tree  crown  si/.e  using  photo  inter- 
pretation. The  relationship  between  trunk 
diameter  and  tree  crown  si/.e  was  basetl  on 
pre\i()us  sampling  of  cottonwoods  along  tlie 
South  Platte  Ri\er  in  Nhjrgan  ('ount\-,  Colorado 
(Getter  1977).  A  close  relationship  (r  =  .SI) 
between  tree  crown  size  and  trunk  diameter  at 
breast  height  (dm  dbh)  was  indicated.  Howexer, 
data  lelating  dbh  to  tree  age  were  lacking,  as 
increment  boring  to  estimate  age  of  cotton- 
woods  did  not  \ield  satisfacton-  age  data.  Matu- 
rit\'  classes  included  stands  dominated  b\  trees 
<1.5,  1..5-4.0,  4.1-7.6,  and  >7.6dm  dbh.  Stands 
of  trees  were  classified  b\  canopy  co\er  as  open 
(10-35%),  intermediate' (36-55%).  and  clo.sed 
(>55%.). 

Changes  in  stands  of  cottonwoods  from  earl\' 
to  recent  photos  were  anah'zed  using  paired 
t  tests  appropriate  for  stratified  (segment)  sam- 
ples based  on  the  Inpothesis  that  mean  change 
was  zero.  Initial  tests  included  anaKses  of  indi- 
\  idual  maturitx/canopv-cover  classes;  howexer, 
sample  sizes  were  inadecjuate  to  \ield  meaning- 
ful results.  Therefore,  maturitx-class  data  for 
pooled  canop\-  cox'er  classes  and  canop\-co\er 
data  for  pooled  maturit\'  classes  are  presented, 
hi  addition.  carK  to  recent  changes  were  pre- 
s(Mited,  wluMc  cauop\'  cover  and  maturit\' 
classes  were  [)artitioned.  Changes  for  other 
cover  t\p)es  were  anaKzed  using  paired  t  tests; 
ANON'A  was  u.sed  to  detect  differences  among 
segments.  Mean  conipaiisoiis  were  considered 
significant  at  P  <  .05. 

RE.su  LTS 

("oloiado  l^iNcr 

Estimated  loss  of  cottonwood  stands  along 
the  Colorado  River  was  1.9  hii/km  sample  unit 
( 17.5%:  Table  2).  Losses  in  the  upper  segment 
(Fig.  1),  where  cottonwoods  initiallv  averaged 
only  2.2  hii/km,  were  >90%  (Table  3).  Area 
occupied  by  cottonwoods  was  highest  in  seg- 
ment 2  where  they  declined  4.4  ha/km.  Within 
downstream  segments,  cottonwoods  axeraged 


360 


Great  Basin  Naturalist 


[\  olume  52 


TaULK  2.  Area  occupied  (.v  lia-'kiii )  In  \  egetation/Iand-iise  h  pe  during  eaiK  and  recent  inten  ids 
and  Rio  Grande,  Colorado. 


■  the  C'olorado  Ri\'er 


G 

olorado  River 

Ri()( 

Grande 

Early 

Recent 

P 

Etu-K- 

Recent 

Type 

X 

SE 

X 

SE 

.V 

SE 

.V 

SE 

P 

Cottonwoods 

11.2 

2.1 

9.2 

1.7 

NS 

17.4 

2.9 

19.0 

3.3 

NS 

Shrnhs 

9.5 

l.S 

10. 1 

2.1 

NS 

6.5 

0.9 

4.9 

0.7 

<.05 

Hav  Meadow 

14.7 

2.9 

11.2 

3. 1 

NS 

68.6 

7.0 

54.5 

6.3 

<.03 

(^nissland 

3.1 

O.S 

4.1 

1.0 

NS 

0.9 

0.6 

3. 1 

1.4 

<.05 

Agricultiu'e 

.5.5 

1.6 

5.1 

2.6 

NS 

0.1 

0.1 

13.5 

5.3 

<.03 

De\(>loped 

0.7 

0.3 

3.2 

0.9 

<.()1 

0.7 

0.2 

1.0 

0.3 

NS 

Ri\er  cliannel 

9. .3 

0.7 

S.S 

O.S 

NS 

6.2 

0.4 

3.9 

0.3 

<.01 

Standing  water 

0.1 

0.05 

2.3 

0.8 

<.03 

1.0 

0.3 

1.2 

0.3 

NS 

T.\BLE  3.  Area  occupied/segment  (.v  lui/kni)  In  cottonwoods  troin  earK  to  recent  sampling  intenals  along  the  (-olorado 
Rixer  and  Rio  Grande,  Colorado. 


Segment 


Colorado  River 


Early 


SE 


Recent 


SE 


Rio  Grande 


Earlv 


SE 


Recent 


SE 


Upper 
Middle 
L<n\er 
L(n\est 


2.3 

0.6 

0.2 

0.1 

.02 

14.7 

1.2 

18.4 

2.3 

NS 

14. 0 

4.2 

19.6 

2.3 

NS 

29.9 

3.0 

.32.2 

3.5 

NS 

7.8 

3.1 

7.4 

•1  ■-> 

NS 

4.9 

2.9 

4.3 

2.9 

NS 

9.3 

1.3 

8.2 

1.8 

NS 

about  7.5-9.3  lui/kin  and  ck-cliiied  at  more 
modest  rates. 

Fith-eight  percent  ol  the  cottonwoods  along 
the  Colorado  River  were  in  the  two  xounger 
matim't)'  classes  (Fig.  2).  The  percentage  of 
young  trees  (  dm-dbh)  declined  almost  50% 
(F  <  .01)  during  the  25-vear  intenal.  Numbers 
of  large  trees  (>7.6  dm)  also  declined  dramati- 
cally (/'<  .02). 

Hectares  of  cottonwoods  were  similar 
among  all  canop\ -coxer  classes  during  the  earK 
sampling  intenal.  However,  In  thc^  recent 
sample  intenal,  open  stands  increased  11%, 
whereas  intermediate  and  closed  stands 
declined  42%r  {P  <  .01)  and  27%  {P  =  .05), 
respectiveK-  (Fig.  2). 

Hay  meadow,  the  most  abundant  xegetation 
t\pe  along  the  Colorado  Hiver,  declined  23.7%o 
during  the  sample  inten-al  (Table  2)  with  the 
primaiy  decrca.se  occurring  in  the  lower  seg- 
ment. (Grassland  occupied  5.7%  of  the  area 
during  early-year  .sampling  but  increased  31%. 
About  10%  of  the  sampled  area  was  in  agricul- 
ture during  both  snnxns.  Developed  land  and 
standing     water    were     initiallx'     minor     but 


increased  to  10%  of  the  total.  Oxerall,  rixer 
chaiuiel  changed  little,  but  xariance  among  seg- 
ments was  exident;  the  channel  xxidened  in  the 
t\x'o  upstream  segments  and  narroxxed  doxxni- 
stream. 

Shrubs,  primanlx'  tamarisk,  occupied  17- 
18%'  of  the  sampled  rix'erbottom  and  increased 
slightlx;  primarilx'  in  the  second  segment. 
Shnibs  occupied  onlx  1.9-2.5  hii/km  xxithin  the 
upper  segment,  >12.4  hii/km  xxithin  the  second 
and  third  segments,  and  9.3  hii/km  xxdthin  the 
loxx'er  segment. 

Rio  (rrande 

Cottonxx'oods  xxere  moderatelx  abundant 
x\ ithin  the  uppcM-  segment  of  the  Rio  Grande, 
increasing  3.7  h;i/km  (24.9%),  and  xx'ere  most 
abundant  xxithin  the  middle  segment  xxhere 
they  increased2.3  lu»/km  (7.7%;  Table  3,  Fig.  1). 
They  xx'ere  absent  xxithin  sexeral  doxxiistream 
sample  units,  and  estimated  loss  xxas  0.7  hii/km 
(13.8%).  biitiallx;  cottonxx'oods  occupied  17. f%' 
of  the  sampled  area,  increasing  to  18.8%  bx'  the 
sec-ond  suncx'  (Table  2). 

Small  trees  (<1.5  dm)  represented  10.4%'  of 


19921 


(]()I,()H\I)()  HiI'Mll  W  \'1':CKT.\TI()\ 


361 


TRUNK  DIAMETER 
(dm-dbh)r]..6 

□ 


EARLY         RECENT  EARLY  RECENT  EARLY  RECENT 

CANOPY  COVER 

FitT.  2.  Earlv  to  recent  clianges/sample  in  niatnritx  class, 
aiul  canop\  c()\ei"  of  cottonwoods  along  the  lower  Colorado 
Hi\er,  western  Colorado. 


the  coiiipo.sitiou  duiing  hotli  saiiipk\s  and 
increased  9.3%  in  occupied  area  (Fig.  3).  Trees 
of  intermediate  size  (1.5-4.0  dm)  declined  (F  = 
.13)  over  the  36.7-year  interval,  giving  way  to  the 
next  larger  (4.1-7.6  dm)  maturitv  class  that 
increased  27.2%  (P  =  .16)  (Fig.  3)'.  This  latter 
group  dominated  among  inatnritv"  classes 
dnringboth  sunevs.  Large  trees  (>7.6  dm)  rep- 
resented onlv  3%  of  the  total  during  both  sur- 
veys and  showed  little  evidence  of  increasing  in 
occupied  area. 

Open  stands  initiallv  occupied  31%  of  the 
timbered  area  and  declined  (P  =  .25)  to  259^ 
I  Fig.  3).  In  contrast,  stands  of  intermediate  clo- 
sure increased  {F  =  .02)  from  33  to  40%.  Closed 
stands  increased  modestlv  {P  =  .49,  9%),  repre- 
senting 359f  of  the  total  during  both  sui"ve\s 
(Fig.  3). 

Hav  meadows  dominated  among  v  egetation 
tvpes  (Table  2),  decreasing  from  68  to  54%  of 
tli(^  sampled  area.  Declines  occurred  primariK 
\\  itliin  the  two  nj^per  segments.  Initiallv.  grass- 
land was  minor  but  it  increased.  primariK 
w  ithin  the  upper  segment.  Onlv  2  of  20  samples 
originallv  contained  cropland,  but  the  propor- 
tion increased  to  9  of  20  samples  (0.1  to  13.4%). 

Developed  land  and  standing  water  were 
minor  components  in  both  earlv    and   recent 


I 

LU 


TRUNK  DIAMETER  (dm-dbh) 


EARLY  RECENT   EARLY  RECENT  EARLY  RECENT 

CANOPY  COVER 

Fig.  3.  FarK' to  recent  clianges/saniple  in  niatnrit\' cUiss. 
and  eanop\'  co\er  oi  cottonwoods  along  the  lower  Hio 
Grande,  southern  Colorado. 

suiA'evs.  Ri\er  channel  decreased  (36.7%) 
throughout  the  studv  area.  .\rea  occupied  by 
shrubs  was  minor  and  estimated  loss  v\as  25% 
(Table  2). 


Dl.SCUSSION 

('omparison  of  clianges  along  the  two  rivers 
leads  to  greatest  concern  for  habitats  along  the 
(Colorado  River,  the  much  larger  of  the  two 
(Table  1 ).  The  25-vear  interval  along  the  C^oio- 
rado  River  was  considerablv  less  than  that  for 
the  Rio  Grande,  but  a  17.5%  decline  occurred 
in  area  occupied  bv  trees.  Development  along 
the  y'wcr  increased  dramaticalK  and  replaced 
manv  stands  of  trees. 

Lack  of  natuial  reproduction  and/or  liigli 
uiorialitA of  voung  trees  v\  as  indicated  by  a  50% 
reduction  in  stands  of  voung  trees  along  the 
(Colorado  River.  Reduction  of  stands  dominated 
b\  old  trees,  which  provide  primaiy  habitat  for 
ca\itv  nesting  wildlife,  was  also  evident.  IIov\- 
evcr  ra])id  shifts  toward  more  open  stands, 
which  indicated  excessive  mortalit)-  within 
stands,  were  more  discouraging  than  changes  in 
luaturitv  structure.  Thus,  there  were  fewer  and 
smaller  stands  and  those  remaining  v\ere  more 


362 


Great  Basin  Naturalist 


[\ bliime  52 


open  and  occupied  In'  iritennediate  maturih 
classes. 

Losses  of  cottonwoods  were  especially  dra- 
matic (>90%)  in  the  upper  segment  where 
occurrence  was  initiallv'  low.  Expansion  of  urhan 
areas,  highway  construction,  and  other  de\elop- 
ments  were  responsible  for  much  of  the  riparian 
habitat  loss  in  a  relativeK'  narrow  valley  that 
initially  possessed  limited  riparian  habitat  and 
relati\el\-  rapid  stream  flows.  Loss  of  trees  to 
beaver  {Castor  canadensis)  was  noted  and  ma\' 
be  important,  especially  in  the  upper  segments, 
since  many  stands  of  cottonwoods  were  con- 
fined to  streamsides  by  valley  relief 

Expansion  of  tamarisk  was  evident  along 
lower  reaches  of  the  Colorado  River  wdthin  a 
broadened  floodplain  and  slower  stream  flows. 
Increasing  expansion  of  tamarisk  severely  limits 
opportunities  for  natural  regeneration  of  cot- 
tonwoods and  willo\\'S.  Russian  olive  {Elaeagnus 
angustifolia)  also  is  pioneering  along  the  Colo- 
rado Ri\'er.  This  species  possesses  a  growth  form 
of  intermediate  height  and,  like  tamarisk,  may 
form  monocultures  (Knopf  and  Olson  1984). 

Stream  flows  along;  the  Colorado  River  haxe 
not  shown  major  declines  in  recent  decades. 
Large  impoundments  and  high-elevation  diver- 
sions, primarih-  occurring  during  the  last  50 
years,  ha\e  altered  and  reduced  peak  flow 
sequences  on  the  Colorado  and  Gunnison 
rivers. 

Extensive  flooding  occurred  along  the  Colo- 
rado River  in  1983-84,  resulting  in  considerable 
natural  reproduction  of  seedlings.  However, 
infrerjuent  flooding  is  not  likely  to  offset  the 
impacts  of  stream  flow  regulati(5n,  streamside 
developments,  and  invasions  of  exotic  species. 
Vegetation  conditions  and  changes  along  the 
(Colorado  River  appear  to  be  following  the  pat- 
tern of  disrupted  recruitment  of  native  riparian 
phreatoplntes  occurring  along  many  western 
rivers  (Howe  and  Knopf  1991). 

In  contra.st  to  changes  documented  along  the 
Colorado  River,  riparian  habitats  along  the  Rio 
Grande  were  relatively  stable  during  the  sample 
intenal,  with  an  increase  in  area  occupied  by 
cottonwoods.  However,  several  of  the  sample 
units  within  the  lower  segment  contained  few  or 
no  cottonwoods.  Little  evidence  of  seedling 
establishment  was  noted  subsequent  to 
increased  stream  flows  during  1983-84,  which 
raises  concern  for  future  trends.  Stream  flow  s 
averaged  over  10-year  intenals  since  1890 
showed  little  evidence  of  decline  at  Del  Norte 


in  the  west  central  portion  of  the  San  Luis  Valley 
(Ugland  et  al.  1984,  \bl.  1).  However,  upstream 
impoundments  have  reduced  peak  flows  and 
altered  patterns  with  stabilized  increased  vol- 
umes into  late  summer  for  irrigation.  Flows 
downstream  at  Alamosa  (Fig.  1)  averaged  about 
30%  of  those  at  Del  Norte,  and  average  flows 
since  1930  have  been  about  one-half  of  those 
from  1913  to  1930.  Reduction  in  channel  width 
was  indicative  of  reduced  and  stabilized  stream 
flows.  Streamsides  were  dominated  bv  peren- 
nial herbaceous  vegetation,  which  provides  lim- 
ited opportunity  for  establishment  of 
pioneering  species  such  as  cottonwoods  and  is 
indicative  of  moderately  stable  and  slow  stream 
flows  through  the  relatively  flat  San  Luis  \  alley. 
Increased  farmland  was  the  most  pronounced 
land-use  change  along  the  Rio  Grande,  whereas 
little  development  occurred. 

Shnibs  (primarily  willows)  have  not  been 
major  components  along  the  Rio  Grande  in 
recent  decades.  Severe  cold  winters,  due  to  high 
elevations  (Table  1),  ma\"  prevent  invasions  of 
tamarisk,  which  has  developed  as  a  streamside 
monoculture  at  lower  elevations  elsewhere 
along  riparian  systems  in  the  Southwest.  Rus- 
sian olive  was  not  v'et  invading  the  inventoried 
Rio  Grande  riverbottom. 

Similar  inventories  of  riparian  vegetation 
changes  and  status  were  conducted  along  the 
South  Platte  and  Arkansas  rivers  in  the  High 
Plains  of  eastern  Colorado  (Snvder  and  Vliller 
1991 ).  Deterioration  of  habitat  along  the  Arkan- 
sas River  was  much  greater  than  along  western 
rivers  in  Colorado.  However,  conditions  along 
the  Colorado  River  seemed  to  be  deteriorating 
more  rapidly  than  along  the  South  Platte  River. 
There  was  also  much  less  riparian  habitat  along 
western  rivers,  making  that  wliicli  remained  of 
greater  importance.  Sampling  of  changes 
between  two  points  in  time  may  not  give  an 
accurate  assessment  of  long-term  trends.  A 
third  inventory  of  these  same  sample  units  is 
recommended  in  the  near  future. 


ACKNOW  LEDCMENTS 

T  E.  Owens  and  D.  Teska  of  the  Colorado 
State  Forest  Senice  performed  aerial  photo 
inteqoretation.  Assistance  in  sampling  design 
and  statistical  analysis  was  provided  by  D.  C. 
Bowden.  W.  D.  Graul  and  G.  R.  Craig  assisted 
in  study  design  and  implementation.  A.  E. 
Anderson,  C.  E.  Braun,  R.  W.  Hoffman,  and 


1992] 


COLOIUDO  Kll'AKlAX  NKCKTATIOX 


363 


R.  C.  Kiifeld  proxided  constnicth'e  re\ie\\'  of 
tilt'  inaiiustript.  This  project  was  supported  b\- 
Federal  Aid  to  Wildlife  Restoration  Project 
\\'-152-R  and  the  Nongame  Checkoff  Program 
of  the  C-olorado  Dixision  of  W  ildlife. 

LiTERATURK  CiTED 

.\.NDKKS()N.   B.  \\.,  AND   R.   D.  OlIMAKT     lySO.    Desit^Ilillg 

and  de\eloping  a  predictive  model  and  testing  a  re\eg- 
etated  riparian  community  for  southwestern  birds. 
Pages  4.34-449  in  R.  M.  DeGraff,  technical  coordina- 
tor. Management  of  western  forests  luid  grassliuids  for 
nongame  birds.  U.S.  Department  of  Agriculture, 
Forest  Serxice  Generd  Technical  Report  I\T-S6. 

.  19S5.  M;uiaging  riparian  vegetation  and  wildlife 

along  the  Colorado  Ri\er:  SMithesis  of  data,  predictixe 
models,  and  management.  Pages  123-127  /»j  R.  R. 
Johnson.  C.  D.  Ziebell,  D.  R.  Patton,  P  F.  Ffolliott,  and 
R.  H.  Hanire,  technical  coordinators.  Riparian  ecosys- 
tems and  their  niiinagement.  First  North  American 
Riparian  Conference,  U.S.  Department  of  .Agriculture. 
Forest  Ser\ice  Generd  Technical  Report  R.\l-12(). 

Anderson,  B.  W.,  R.  D.  Oiimakt.  and  J.  Di.sano  1978. 
Revegetating  the  rip;uian  floodplain  for  wildlife.  Pages 
SlS-S'ai  in  R.  R.  Johnson,  and  J.  F.  McCormick,  tech- 
nical coordinators.  Strategies  for  protection  and  man- 
agement of  floodplain  wetlands  and  other  ripari;ui 
ecosystems.  U.S.  Depiirtment  of  Agriculture,  Forest 
Senice  General  Technical  Report  \\'-0-12. 

Beidleman.  R.  G.  1978.  The  cottonwood-willow  riparian 
ecosystem  us  a  vertebrate  habitat,  with  particular  ref- 
erence to  birds.  Pages  192-195 ;'/!  \\'.  D.  Graul  and  S.  J. 
Bissell,  technical  coordinators.  Lowland  ri\er  and 
stream  habitat  in  Colorado:  a  sviiiposium.  Colorado 
Chapter,  The  Wildlife  Societs-  luid  Colorado  .Audubon 
Council,  Greeley. 

Fit/,c;krald,  J.  P.  1978.  Vertebrate  associations  in  plant 
comnumities  along  the  South  Platte  Rixerin  northe;ist- 
em  Colorado.  Pages  73-88  in  W.  D.  Graul  and  S.  J. 
Bissell,  technical  coordinators.  Lowland  river  and 
stream  habitat  in  Colorado:  a  sxinposium.  Colorado 
Chapter.  The  Wildlife  Societ\'  ;uk]  Colorado  .Audubon 
Council,  Greeley. 

Getter,  J.  R.  1977.  Procedures  for  inventor\ing  plains 
cottonwoods,  Morgan  Count);  Colorado.  Colorado 
State  Forest  Service  Report.  44  pp. 

Graul.  W.  D.,a\d  S.J.  Bissell, TECHNK.wLcoouDiN.vroas. 
1978.  Lowland  river  and  stream  habitat  in  Colorado:  a 
s\-mposium.  Colorado  Chapter,  The  Wildlife  SocieU' 
and  (Colorado  Audubon  Council.  Greelev. 

ll\HHiNt:T()N.  11.  D.  1954.  Manual  of  the  pkuits  of  Colo- 
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II()()\  KR.  R.  L.,  AND  D.  L.  Wills,  eds.  1984.  Managing 
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Knoi'F  F.  L.,  and  T  E.  Olson.  1984.  Naturalization  of 
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Melton,  B.  L.,  R.  L.  Hoover.  R.  L.  Mooke.  and  D.  J. 
Pfankucii  1984.  Aquatic  and  riparian  wildlife.  Pages 
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Greeley. 

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Received  28  January  1992 
Accepted  20  September  1992 


Great  Basin  Naturalist  52(4),  pp.  .■3fi4-)72 

RESIDENT  UTAH  DEER  HUNTERS"  PREFERENCES 
FOR  MANAGEMENT  OPTIONS 

Dennis  D.  Austin  ,  Philip  ].  Uniess  ,  and  Wes  Sliields" 

Abstract. — A  total  of  3291  resident  deer  liunters  returned  questionnaires  distributed  at  eheei<ing  stations  in  fail  1989 
and  1990  pro\  iding  opinions  and  management  data  concerning  tlie  Utah  rifle  Inmt.  Hunters  reported  hunter  crowding  and 
too  few  big  bucks  as  critical  reasons  for  possibly  choosing  to  quit  deer  hunting  in  Utah.  Indeed,  hunter  age  stmcture  and 
measured  satisfaction  suggested  a  negative  future  trend  in  hunter  participation.  Results  suggested  the  adoption  of  several 
huntcr-prclrrrcd  management  options  would  increase  satisfaction,  motivation,  and  success. 

Kci/  uords:  mule  ilcci:  (jucstioiiiidircs.  cltcckinf^.stdtions.  deer  iitinta^iutcitt .  hunter  Dpiiiions.  uihilife  methiuls.  wildlife 
teelinujues. 


Competition  for  wildlife  recreation  in  the 
Rocky  Monntain  region  will  increase  in  the 
future,  while  projected  populations  of  niajor 
wildlife  species  will  show  little  change.  In  the 
ne.xt  30  \'ears  the  number  of  big  game  hunters 
is  expected  to  slowly  increase  from  about  1.5  to 
1.7  million,  compared  with  the  rapid  inci'ease  in 
nonconsumptive  users  of  3.9  to  7.1  million 
(Flatherand  Iloekstra  1989).  Certainh;  the  per- 
centage of  hunters  in  the  total  population  will 
decHne,  w^hile  the  percentage  of  nonconsmnp- 
tive  users  will  increase.  Conse(jueutlv,  to  bal- 
ance resource  use,  wildlife  managers  must 
obtain  a  clear  understanding  of  user  prefer- 
ences, particularK  aiuoug  those  users  who  his- 
toiicalK  and  curreutK  ha\e  paid  most 
managenuMit  costs  \ia  license  permit  lees  and 
excise  taxes  on  spotting  equipiuent. 

In  Utah,  mule  deer  are  preeminent  among 
hunted  wildlife  species  in  terms  of  income 
received  for  wildlife  manageiuent  and  hunter 
days  afield.  Ilowexer,  compared  with  the  197()s 
and  in  contrast  to  past  regional  trends  (Flather 
and  Iloekstra  19(S9),  total  big  game  licenst^  sales 
(k^creased  slightly  (().<S%)  in  tlie  f98()s  whik^ 
total  lifle  hunters  afield  declined  3.1%-  (jen.se 
and  Shields  1990).  These  figures  warn  of  possi- 
ble negatixe  trends  for  deer  hunter  participation 
and,  ak)ng  with  uncertain  hmiter  satisfaction, 
strongK  suggest  a  need  for  constant  and  (effec- 


tive communication  betvveen  state  wildlife  offi- 
cials and  Utah  hunters. 

One  means  of  conuuiuiicating  information  is 
through  hunter  opinion  questionnaires,  which 
ha\e  become  an  important  data  source  for  game 
management  decisions.  In  Utah  during  the 
19(S0s,  six  questionnaire  sunexs  were  con- 
ducted, and  that  mmiber  will  likely  double  in 
the  1990s  (Bunnell  and  Austin  1990).  The  use 
of  postcard  questionnaire  surxevs  distiibuted  to 
hoiueward  looinid  hunters  at  deer  checking  sta- 
tions is  one  method.  This  simple  technique, 
de\eloped  in  Utah  during  tlie  late  1980s,  is 
inexpensive,  deiuographicalK  iml)iased,  and 
accurately  representative  of  hunters"  opinions 
concerning  deer  luanagement  (Austin  and 
Jordan  1989,  Austin  et  al.  1990). 

Methods 

Questionnaires'  were  piinted  on  4  1/4  X 
6-inch  postage-paid  cards.  Dming  opening 
weekend  of  the  1989  Utah  lifle  deer  himt,  7()4{) 
(jnestionnaires  were  distril)uted  to  hunters  at  f  1 
checking  stations,  and  in  1990,  8750  question- 
naires were  distiibuted  at  16  locations.  One 
(juestioimaire  was  given  to  each  licensed  hunter 
checked  until  the  suppK*  was  exfiausted. 

Data  vx'ere  analyzed  w ithin  xears  using  the 
l\nus()n  chi-square  statistic.  Tlu^  cross-tabula- 
tion method  from  the  SPSS  program  on  a  \'AX 


^Dcpartiiu-iit  orHantit'  Scii-iici-,  IJlali  Stak-  L;iii\<'rsitv,  l^)s;an.  L't.ili  S4:;22-.52:?(l. 

-  Utah  Division  orWildlilc.  H,-s()niXf,s,  1.596  UVsl  .\ortli  Temple,  Sail  I  M-  ( :il\ .  ll.ili  S-1 1  Ui, 

■  Copies  olllii-  ()iu'sti()iinaiie  cards  are  a\ailal)le  I'roiii  llie  senior  aiitlior. 


364 


1992]  1  k  XTKH  Ol'IMOXS  ON  Deeh  Ma\a(;eme.\t 

Taislk  1.  Questionnaire  rehini  rates. 


365 


19S9  Sur\'ey 

1990  Sur\ey 

No. 

No. 

% 

No. 

No. 

% 

Region 

Lt)eation 

distributed 

retimied 

returned 

distributed 

returned 

returned 

Nortlu'rn 

SnowA  illc 

.500 

107 

21.4 

500 

109 

21.S 

Blacksmith 

.500 

95 

U).0 

.500 

83 

16.6 

Ogden 

1000 

151 

15.1 

.500 

126 

25.2 

Kaiuas 

— 

— 

— 

500 

91 

18.2 

Wellsville 

72 

20 

27.8 

— 

— 

— 

Salt  Eake 

(.'an\()us 

— 

— 

— 

250 

48 

19.2 

Northeastern 

\ernal 

666 

44 

6.6 

.500 

114 

22.8 

Bookcliils 

650 

131 

20.2 

.500 

113 

22.6 

Current  (;k. 

— 

— 

— 

.500 

120 

24.0 

Central 

Thistle 

1000 

206 

20.6 

12.50 

291 

23.3 

Tucker 

300 

37 

12.3 







Sheepcreek 

300 

48 

16.0 

250 

41 

16.4 

Wrnon 

452 

151 

.33.4 

375 

95 

253 

Stansl)ur\ 

— 

— 

— 

.300 

57 

19.0 

Soiitlieasteni 

Areawick' 

— 

— 

— 

830 

95 

11.4 

South  L'l'Mtral 

Fishlake 





_ 

:M){) 

14 

4.7 

Oak  Creek 

— 

— 

— 

195 

63 

32.3 

Southern 

Blooniinifton 

IfSOO 

41:] 

26.4 

1.500 

418 

27.9 

Total 

7040 

1413 

20.1 

8750 

1S7S 

21.5 

c()ni[)uter  wa.s  ii.stxl.  For  (question  number  IS. 
1990  siinex.  tlic  \;ilues  giwn  were  objectixeK 
placed  into  1 1  nionetan  cla.s.ses  foranaKsis.  The 
sipiificance  le\el  for  all  data  interactions  was  set 
atP  <  .05.  Forcjnestion  nunil)er2(),  199()sune\', 
because  the  (juestion  w  as  written  witli  innumer- 
able potential  responses  and  man\'  (juestion- 
uaires  contained  more  than  one  respon.se,  data 
w  ere  not  statisticalK  anaK-zed  but  are  reported 
ninnericalK.  The  responses  were  snbjectiveK 
'j;i()nped  into  71  cate<2;ories.  Data  from  tlie  1987 
and  I9SS  snr\('\s  are  added  and  compared 
w  here  applic'able. 

Results 

Return  l^ates 

Checking  stations  used  for  distribution  b\ 
Utah  Dixision  of  Wildlife  R(\sonrces  re<i;ions, 
tlie  nnmbc^r  ol  (|uestiomiaires  tlistributed.  and 
return  rates  are  sliown  in  Table  1.  .Althontfh 
rates  \  aried  considerably  b\  region  and  location, 
total  return  rates  were  20. 1  %  in  1989  and  2 1 .5% 
in  1990,  and  consistent  (Austin  and  Jordan  1989, 
Austin  et  al.  1990)  with  those  reported  for  1987 


(25.87f )  and  1988  (20.17r ).  ExjK^cted  statewide 
return  rates  using  this  method  are  thus  about 
20-257f. 

Hunter  Demographics.  Success, 
and  Satishiction 

Most  resident  himters  are  mak^  (  >90'%  ),  age 
25—14  {529f ),  and  haxe  more  than  10  \ears  of 
Utah  dv(^v  hunting  experience  (>6()'^).  During 
1989  and  1990.  hunters  had  le.ss  than  50%  part)' 
success  lor  bucks  on  ojx'uing  weekend  and  rel- 
ati\el\-  low  hunter  satisfaction  (Table  2 ).  I  lunter 
partx'  success  noticeabK  declined  betw een  1 987 
and '1988  and  again  betAwen  1 988  and  1 989.  but 
remained  about  the  same  between  1989  and 
1990. 

The  percentage  of  hunters  (<20%)  in  the 
\()un«iest  age  class  (14—24  \ears)  is  lower  than 
expected.  Participation  b\-  hunters  in  this  age 
class  should  be  highest  because  few  people 
begin  hunting  after  about  age  25.  These  figures, 
consistent  oxer  four  \ears,  alone  suggest  possi- 
])le  (nture  declines  in  the  number  of  Utah  deer 
hunters.  However,  the  sharp  drop  in  hunter 
participation  between  the  third  and  fourth  age 


366 


Great  Basin  Naturalist 


[Volume  52 


TaBLK  2.  Demographics,  paitv  success  (%),  and  hunter  satisfaction  of  Utah  resident  deer  hunters  sampled,  19S7-1990 
(sample  sizes  in  parentheses). 


Sex 

Age  class" 

Year 

Male 

Female 

.V 

1 

2 

3 

4 

5 

6 

N 

1987 

90.4 

9.6 

(863) 

19.0" 

31.8 

23.0 

14.2 

8,1 

4.0 

(869) 

198S 

89.6 

10.4 

(444) 

19.7 

33.0 

23.4 

13.8 

8.3 

2.0 

(458) 

1 989 

92.8 

7.2 

(925) 

18.6 

28.1 

25.9 

13.9 

9.0 

4.6 

(936) 

1990 

92.7 

7.3 

(1429) 

22.0 

26.6 

26.2 

12.9 

8.2 

4.1 

(1429) 

Mi'ans 

91.4 

8.6 

19.8 

29.9 

24.6 

13.7 

8.4 

3.7 

E-xperience  cl 

ass' 

Success*^ 

Satisf 

action 

Year 

1 

2 

3 

4 

N 

1987 

18.7 

21.3 

27.2 

32.8 

(867) 

69.3* 

(411) 

5.3 

(871) 

1988 

21.7 

18.4 

28.4 

31.5 

(461) 

55.3 

(459) 

4.3 

(456) 

1989 

22.7 

15.2 

25.2 

36.8 

(932) 

48.0 

(904) 

4.1 

(934) 

1990 

25.1 

14.2 

26.0 

34.6 

(1418) 

47.9 

(1406) 

4.6 

(1413) 

Means 

22.1 

17.3 

26.7 

33.9 

55.1 

4.6 

'.\ge  c-l^usses:  1  =  1-4-24.  2  =  2.5-04.  3  =  35-44. 4  =  4.5-54,  5  =  5.5-fi4.  6  =  65+  wars 
''Experience  c-lasse.s:  1  =  1-5,  2  =  6-10.  3  =  1 1-20.  4  =  21+  yeans 
'  I  IiLiitiiig  paitv  success  for  one  or  more  bucks  on  opening  weekend 

'  Hunter  satisfaction  of  current  year's  hunt  in  comparison  to  all  previous  deer  hunts.  A  score  of  5.0  would  be  expected  for  the  average  hunt. 
'.\ge  class  16-24.  Hunters  aged  14  and  15  years  were  ineligible  tor  big  game  licenses. 
Hunting  party  success  for  bucks  and  antlerless  deer  on  this  himt.  For  the  19S7  season  62.516  bucks  and  1168  antlerless  deer  were  harve.sted. 


classes  (35-44  and  45-54)  is  also  of  concern 
because  in  these  age  groups  mam  hunters"  chil- 
dren are  beginning  to  hunt,  and  parent  partici- 
pation is  a  key  factor  in  long-term  sustained 
interest  of  new  hunters  (Decker  and  ConnelK' 
1989).  Mean  age  of  all  hunters  was  36.3,  35.4, 
37.0,  and  36.0  \'ears  for  1987-90,  respecti\'elv. 
In  a  completeh-  randomized  survey  of  Utah 
hunters,  Krannich  et  al.  ( 1991 )  reported  a  mean 
age  of  37  years  and  similar  hunter  age  and  sex 
characteristics. 

One  probable  explanation  for  the  shaip  drop 
in  hunters  in  the  45-54  age  class  is  the  signifi- 
cant  interaction  between  age  and  hunter  expe- 
rience with  hunter  satisfaction  (P  <  .04). 
Hunters  with  20+  years  of  experience,  who  gen- 
erally hunted  deer  before  the  1970s  when  the 
number  of  hunters  was  lower  (Fig.  1)  and 
hunter  success  rate  was  higher  (Jense  and 
Shields  1990),  show  lower  satisfaction  scores 
than  younger,  less-e.xperienced  hunters.  Mean 
satisfaction  scores  of  experience  classes  1-3 
versus  4  (Table  2)  for  both  years  combined  were 
4.5  and  3.9,  respecti\eK'.  Similarly,  mean  satis- 
faction .scores  of  age  classes  1-3  versus  4  were 
4.5  and  3.7,  respectively.  Consequently,  hunting 
moti\  ation  for  hunters  with  20+  years  of  experi- 
ence has  likely  decreased  because  of  perceived 
lower-qualit)'  hunting. 


Another  concern  for  hunter  participation  is 
noted  b\  comparing  the  trend  of  hunter  partic- 
ipation by  experience  classes  between  survey 
years  (Table  2).  No  trends  in  hunter  participa- 
tion were  evident  for  hunters  v\ith  1 1  or  more 
years  of  experience.  However,  hunters  v\ith  6- 
10  years  of  experience  decreased  7.1%  between 
1987  and  1990,  while  hunters  with  1-5  years  of 
experience  increased  6.4%. 

Comparison  Between  Hunt  T\pes 

Utah  has  had  four  basic  t)pes  of  hunts  since 
1951,  v\ith  each  hunt  tvpe  having  a  variable 
number  of  antlerless  control  permits.  Either-sex 
hunts  dominated  from  1951  to  1973.  with  buck- 
only  hunts  dominating  from  1974  to  1990,  as 
well  as  before  1951.  From  1985  to  1990  hunter- 
number- restrictive  (limited-entn"  and  high- 
countn )  hunts,  and  from  1984  to  1989 
antler-restrictive  (three-point-and-better)  hunts 
were  established  on  some  units. 

Buck-only  hunts. — Total  buck  hanest 
averaged  63,250  per  year  v\  ith  8633  antlerless 
hanest  and  181,235  hunters  afield  (Fig.  1).  The 
number  of  unretrieved  deer  reported  per  100 
buck-only  hunters  in  these  surxevs  for  1987-90 
was  19.9,  21.7.  15.9.  and  16.0,  respectively. 
Using  the  weighted  mean  of  17.9,  total 
unretrieved  deer  for  this  period  was  32,441  per 


1992] 


Hunter  Opinions  ()\  Dkkh  Man\(;i:mknt 


36- 


250000    T 


200000    -- 


150000    -■ 


100000 


50000    i 


.A 


^^ 


♦     ♦ 


■  -  -  . 


*v/"'' 


i-O^^ 


.'■~^V 


/    ■     \ 


A-'\ 


\y 


/     --■' 


I    I    I    I    I    I    I 


-+-  I    I    I    I    I    I    I    I    I    I    I    I    I    I 


I    I    I    I 


Fig.  1.  Total  Iianrst  ui  liuck  ami  aiitlfrless  derr  and  coinhiiird  liiiiitrrs  alifld  iroiii  all  hutk  liiiuts  in  L'tali,  1951-90. 


\"ear,  and  mean  total  annual  limiting  niortalit\- 
was  104.324.  Mean  hnnter  sati.sfaction  (19S7- 
90),  with  0  representing  the  worst  hiuit  and  10 
the  best  hnnt,  was  4.4.  Hunting  pait\"  success 
was  45.8%. 

ElTHER-SEX  HUNTS. — During  23  \ears  of 
either-sex  hunting,  the  statewide  total  buck  har- 
xest  axeraged  66,992,  and  the  antlerless  hanest 
was  39,228.  Using  the  estimated  mean  for 
unretrieved  deer  (Robinette  et  al.  1977.  Staplex 
1970^  of8.0  deer  per  100  hunters  and  the  mean 
number  of  rifle  liunters  afield  ( 153,666),  a  cal- 
culat(^d  \earl\  loss  of  12,293  unretrieved  deer  is 
obtained,  bringing  the  mean  total  annual  hunt- 
ing mortalit\' to  118,513.  Hunter  j)referenc(^  for 
buck-onl\'  \ersus  either-sex  hiniting  has  not 
been  addressed. 

ANTLER-RESTKKTIXE  hunts. —  Three- 
point-and-better,  antler- rest ricti\e  hunts  were 
a\ailable  on  some  units  during  1984-89,  and 
then  discontinued.  In  coiiiparison  with  biuk- 
onl\  hunts.  three-point-and-better  limits 
showed  a  riHluction  in  hunters  afi(4d.  buck  har- 
\est,  and  hunter  success  (Jense  1990).  Howexcr. 
these  hunts  also  showed  a  small  increase  in  the 
post-season  total  buck  to  doe  ratios,  but  a  large 
decrease  in  the  number  of  post-.season,  mature 
bucks  counted.  These  areas  also  showed  a  larjie 


decrease  in  the  small  buck  (hvo-point-and-less) 
to  doe  ratio  between  preseason  and  post-season 
classification  counts  (Jense  1990). 

Our  anaKsis  confirmed  the  adxerse  impacts 
of  three-point-and-bett(M-  hunts  reported  b\ 
fense  (1990),  with  the  highest  mimber  of 
unretriexed  deer  at  39.6  per  100  hunters, 
including  21.7  bucks.  This  number  of  bucks. 
luostK  two-point-and-less.  is  compared  to  4.6 
bucks  per  l()()huut(MS  on  buck-ouK  areas.  How- 
e\er,  hunters  from  antler-restrictixe  areas  were 
mod(>ratel\  satisfied,  with  a  mean  index  of  4.8, 
and  mean  hunting  part\  success  was  55.6%. 
During  1989,  the  last  \ear  of  three-point-and- 
better  hunts,  40.0%  [n  =  931)  of  Utah  resident 
hunters  had  hunted  at  least  once  on  three-point- 
and-better  areas,  but  onl\  26.7%  (n  =  906)  pre- 
ferred to  continue  this  t\pe  of  hunt.  Indeed,  less 
than  lialf  i47.7%  )  of  hunters  who  chose  to  hunt 
these  units  in  1989  preferred  to  continue  them. 

Facii  though  antler-restrictixe  hunts  were 
not  successful  o\er  entire  deer  management 
units,  selection  of  conscientious  hunters  to 
a\()id  high  iuu-etrie\ed  deer  losses  nia\-  lead  to 
successful  antler- restrictixe  management.  For 
example,  at  the  Ea.st  Canyon  Resort  (10,000 
acres  ^  in  northern  Utah,  protecting  onk  2X2 
point    bucks    (1988-90)   increased   the    mean 


368 


Great  Basin  Nathkalist 


[N'olunie  52 


number  of  total  antler  tines  of  hai-vested  bucks 
from  4.5  (1985-87)  to  6.1  (1988-90).  The  per- 
cent of  hanested  l)uc-ks  2X2  or  smaller 
decreased  from  60  to  35%,  while  the  numl)er  of 
trophy  bucks  larger  than  4X4  increased  from 
0  to  8  (unpublished  data.  East  Canyon  Resort). 

HUNTKR-NUMBER-RESTRICTION  HUNTS. — 
Limited-entn'  hunts  have  been  used  on  some 
units  since  1985.  In  comparison  with  buck-onlv 
hunts,  the\  proxide  higher  hunter  success 
{F  <  .01)  and  satisfaction  (F  <  .001),  with  an 
index  of  6.3,  but  no  difference  in  the  total 
munber  of  unretrieved  deer  ( 1 7.7  total  deer  per 
100  hunters  wdth  9.1  bucks  and  8.6  antlerless). 
Hunting  partv  success  (1987-90)  was  high  at 
68.8%.  hi  1989,  22.8%  of  resident  hunters  (n  = 
935)  had  hunted  deer  on  limited-entn'  areas, 
and  most  (65.6%)  indicated  die  fee  of  $22.00 
was  fair.  While  most  himters  {)i  =  908)  fa\'ored 
the  same  (37.8%)  or  increased  (38.9%)  number 
of  limited-entn'  units,  hunter  preferences  for 
\  arious  permit  drawing  and  landowner  hunting 
options  were  unclear. 

A  second  t\pe  of  hunter-number-restrictive 
hunt  is  the  high-countn'  hunt.  This  uncrowded, 
high-qualitv  himt — but  one  that  han-ests  bucks 
not  then  available  during  the  Octobei"  rifle 
hunt — received  positi\'e  support  from  most 
(59.6%)  Utah  hunters. 

Vehicle  Access  to  Public  Lands 

A  strong  majorit)'  of  hunters  (76.2%)  indi- 
cated that  at  least  some  lands  should  be  closed 
to  vehicle  access  during  the  deer  hunt  to 
increase  the  qualitv  of  the  hunting  experience. 
However,  the  percentage  of  hunters  indicating 
at  least  half  of  all  public  lands  should  be  open  to 
\ehicles  was  74.5%.  Overall,  hunters  indicated 
that  a  UK^m  of  37.5%)  of  public  lands  should  be 
closed  to  \ehicle  access,  vaiying  by  location 
from  28.9  to  45.4%  The  percentage  of  hunters 
who  hunted  on  areas  with  \  (4iicle  restrictions 
was  33.8%,  while  tlie  pc^rcentage  of  hunters  who 
indicated  preference  to  hunt  on  areas  with  vehi- 
cle restrictions  was  45.2%-.  Using  the  logical 
assumption  that  the  percentage  of  areas 
restricted  to  x'ehicles  should  be  clo.selv  propor- 
tional to  the  percentage"  of  hunters  preferring 
lliem,  our  data  .suggest  the  current  amount  of 
area  with  restricted  \ehicle  access  is  ck)se  to 
hunter  pn^erence,  but  that  an  additional  3.77f 
(37..5-33.8)  to  1 1.4%  (45.2-33.8)  of  public  lauds 
should  be  r(\stricted.  More  information  is 
needed    on    hunter   preferences    for   vehicle- 


restricted  areas  in  terms  of  size,  locations,  and 
number  of  areas. 

License  Fees 

With  the  current  cost  of  a  big  game  hunting 
license  set  at  $15.00,  hunters  were  asked  what 
they  believed  to  be  the  fair  value.  Althoush 
Schreyer  et  al.  (1989)  reported  increased 
license  fees  were  opposed  bv  most  hunters,  a 
mean  value  of  $15.90  was  determined  (/;  = 
1391 )  in  our  studv.  Mo.st  hunters  (58.8%)  indi- 
cated $  15.00  was  the  fair \alue.  Sixt) -eight  hunt- 
ers (4.9%)  indicated  the  fair \alue  was  $30.00 or 
more,  while  58  hunters  (4.1%)  indicated  the 
\alue  was  less  tlian  $10.00.  It  was  interesting  to 
note  that  costs  were  not  related  to  hunter  suc- 
cess, satisfaction,  hunter  choice  of  hunt  tvpe,  or 
whether  private  or  public  kuuls  were  hunted. 

Although  license  fees  are  strongh'  and 
broadK'  approved  In-  Utah  hunters,  few 
improvements  in  the  cjualitv  of  the  deer  hunt 
can  be  made  without  the  economic  trade-off  of 
increased  hunter  fees.  Himter  preferences  for 
balancing  potential  increased  fees  with 
increased  hunt  cjualits  need  to  be  defined. 

Hunter  Concerns 

Twent\-fi\'e  categorical  responses  were 
given  bv'  1%  or  more  hunters  as  I'eason  to  quit 
deer  hunting  (Table  3).  Although  the  list  con- 
tains several  areas  of  low  management  influ- 
ence, such  as  old  atje,  hiijh  associated  costs  of 
hunting,  and  personal  attitude,  most  areas  of 
responses  are  influenced  bv  management  deci- 
sions. The  most  connnon  reasons,  directlv  influ- 
enced bv  management  decisions,  included  too 
main  hunters,  too  few  deer,  bucks,  and  big 
bucks,  private  laud  problems,  and  poor  game 
management. 

Discussion 

Reasons  to  Quit  Deer  Hunting 

The  proportion  of  mature  bucks  in  the  har- 
V  (>st  is  an  area  of  management  control.  It  is  clear 
most  hunters  prefer  hane.sting  large  bucks 
infre([ut^ntlv  as  opposed  to  hanesting  smaller 
bucks  frecjuenth  (Austin  et  al.  1990),  as  well  as 
reducing  some  hunting  opportunitv  to  increase 
the  proportion  ol  mature  bucks  in  the  hanest 
(Austin  and  Jordan  1989,  Toweill  and  Allen 
1990).  Furthermore,  with  tlie  liunting  media 
emphasis  on  tropin  bucks,  the  pott^ntial  hanest 
of  mature  bucks  adds  consitlerably  to  hunter 


19921 


llrx  1 KK  Opinions  on  D\:\:h  M an ackment 


369 


TaI51.I':  3.  I'tali  resident  deer  liuiiters'  responses  to  the  qnestion:  li  \ou  were  to  (|nit  deer  Innitiiiij  in  Utah,  wliat  reason 
wonid  \on  list?* 


Nnniher  of  (jiiestionnaires  returned: 
Nnrnher  ot  (jnestionnaires  w  ith  no  response: 
Nnnil)erof  (|uestionnaires  with  "would  not  (juit,  none 
Nuiuher  of  questionnaires  with  responses: 
Nnnilierof  totd  responses: 


14.30 

8S 

4fi 

129f) 

2()S7 


Response  categories 


Nunil)er  of 
responses 


%  hunters 


Too  nian\  hunters 

Too  few  deer 

Private  land  prol)lenis 

Too  few  hiii  Ijueks 

Old  age  or  phwsieal  inipairnient 

I  ligh  associated  costs  of  hunting 

No  iU'eas  to  hunt  or  access  to  pnlilic  lands 

Too  few  bucks 

Poor  game  management 

L'nethical  hunters 

Low  success  or  no  limit  on  statewide  lici'use  sales 

("hildren  aged  14  and  15  \eais  can  hunt 

Deer  are  too  small 

Too  much  \T\   use  or  too  man\  road  hunters 

Safet\ 

High  costs  ot  licenses 

Personiil  attitude 

Too  few  \ehicle  access  roads 

Too  manv  nonresident  hunters 

Poor  hunt  (jualih 

Proclamation  too  long  or  complicati'il 

No  either-sex  or  antler-restriction  hunts 

Too  manv  limited-enti"\'  areas 

Too  few  limited-entrs  areas 

Too  nian\'  does 

46  otlier  categories 


479 
199 
164 
122 
lOS 
83 
SI 
79 
75 
72 
63 
4S 
44 
41 
39 


31 

30 
29 
27 
19 
17 
16 
14 
139 


37.0 
15.4 
12.7 
9.4 
8.3 
6.4 
6.3 
6.1 
5.8 
5.6 
4.9 
3.7 
3.4 
3.2 
3.0 
2.7 
2.5 
2.4 
2.3 
2.2 
2.1 
1.5 
1.3 
1.2 
1.1 
10.7 


motixation,  and  Kraniiicli  ci  al.  { 1991 )  reported 
that  about  t\vo-third.s  of  Ininters  (66.3%)  were 
di.ssatisiied  with  the  si/e  ol  bucks. 

Compared  with  either-s(^\  huutinu;,  a<i;e 
.structure  of  the  male  population  declines  und(M' 
buck-onI\  huutin<j;(Mc(.'ullou(^h  1979).  In  Utah 
(Austin  1991),  the  percenta<i;e  of  mature  bucks, 
age  3  1/2  vears  and  older,  hancsted  decrea.sed 
from  about  44%  durinti;  the  pre-1951  buck-ouK 
hunts  to  about  30%  dmnng  the  period  of  either- 
sex  hunting  (1951-73).  Th(>  percentage  of 
mature  bucks  hanested  sliarpK  decreased  and 
has  rcMuaiued  at  about  1 0%  dunng  the  ])eriod  ol 
reestablished  buck-onl\-  hunting  (1974-90).  On 
limited-entn  hunts,  the  percentage  of  mature 
bucks  in  the  hanest  has  exceeded  30%  on  most 
units.  Not  onl\-  lias  size  of  hanested  bucks 
decreased  due  to  decreasing  mean  age,  but  age- 
.specific  .size  has  also  declined  (.\ustin  v[  al. 
1989). 


The  aulhois  beliexc  a  n^isonabk  liigh  j)er- 
centage  (20-40%)  of  mature  bucks  in  the  har- 
\(^st  is  critical  to  successhil  deer  management 
and  hunter  motixation.  It  is  clear  to  us  that 
(k'creased  hunting  pressure  on  the  buck  [)oi)u- 
lation  is  necessaiA.  The  data  strongK-  suggest  a 
need  to  establish  statew  ide  minimum  standards 
(or  (1)  age  structure  of  the  buck  hanest,  (2) 
post-season  buck:doe  ratios,  and  (3)  hunter  suc- 
cess for  Inicks. 

Problems  associated  with  pri\ate  lands  are 
important  to  hunters.  These  problems  inchuk" 
poorK  marked  lands,  trespass,  pn\  ate  lands  cur- 
tailing access  to  public  lands,  and  depredation. 
Pri\ate  lands  provide  deer  hunting  for  14.8% 
(1990  snnex)  of  Utah  resident  hunters,  and 
14.7%  of  hunters  reported  owning  10  or  more 
acres  u.sed  l)\  wildlife  ( 1989  sunex).  One  possi- 
ble', partial  solution  may  be  to  give  landowniers 
more   fle.xibilitA    in   management  b)'  allowing 


370 


Great  Basin  Naturalist 


[X'olunie  52 


either-sex  hunting  on  prixate  lands.  AcKantages 
include  increased  landowner  control  over  deer 
niunbers  on  their  lands,  decreased  unretrieved 
deer  kill  (Austin  et  al.  1990),  reduced  depreda- 
tion complaints,  and  improved  opportunity  for 
lianest.  Furthermore,  liberal  hunts  on  private 
lands  mav  increase  incentives  tor  landowiiers  to 
niaik  their  boundaries  and  allow  additional 
hunting  opportunit\'. 

The  categories  of  unethical  liimtei's,  safet\', 
and  minimum  age  for  hunters  are  closelv  related 
to  hunter  education  courses.  Since  the  begin- 
ning of  the  hunter  education  program  (1958) 
and  the  recjuired  wearing  of  hunter-orange 
clf)tliing  (1973),  the  mean  number  ot  total  Utah 
hunting  accidents  and  fatalities  per  \ear  has 
averaged  11.1  and  3.4,  respectixelv,  with  about 
three  accidents  and  one  fatalitv  occurring 
during  the  rifle  hunt.  Before  about  195S  when 
neither  hunter  education  nor  hunter  orange  was 
required,  o\er  100  accidents  and  about  20  fatal- 
ities occurred  yearK'  from  all  hunts  combined. 
Hunter  preference  to  allow  persons  aged  14  and 
15  vears  to  hunt  big  game  has  not  been 
addressed. 

The  length  and  complexity  of  the  proclama- 
tion is  a  concern  of  hunters.  Before  1979,  the 
one-page  Utah  deer  proclamation  measured 
17.5  X  22.5  inches  and  was  printed  on  high- 
qualit)'  paper,  with  the  rules  and  regulations  on 
one  side  and  a  multicolored  map  of  Utah's  deer 
units  on  the  reverse.  In  1990,  the  newsprint 
proclamation  sheets  were  close  to  the  same  size 
(14.5  X  23.0  inches),  but  contained  six  pages. 

The  qualit)-  of  the  hunt  in  terms  of  the  ratio 
of  deer  or  bucks  haivested  per  hunter  is  con- 
trolled by  management.  Although  management 
can  alter  the  buckidoe  ratio,  the  total  number  of 
deer  is  limited  by  habitat,  and,  conxersely,  hunt- 
ers have  not  been  numericalK-  limited.  The  Utah 
buck  harvest  has  remained  rather  constant, 
mostly 50,000-80,000,  since  1951  (Fig.  1),  while 
the  antlerless  har\-esl  lias  shaq:)ly  decreased 
since  1974  with  the  resumption  of  buck-onlv 
hunting.  Total  buck  liimters  afield  from  all  com- 
bined hunts  increased  steadily  between  1951 
and  1964,  decreased  for  three  years  (1964-67), 
slowly  increased  during  1967-69,  but  abiuptK 
increased  between  1969  and  1973.  After  a 
second  three-year  period  of  decreasing  hunters 
afield  (1973-76),  hunter  numbers  hax'e  fluctu- 
ated but  remained  high  throughout  the  1970s 
and  198()s.  CJonsetjuentlv,  the  himter  responses 
of  poor  game  management,  poor  hunt  (|ualit\-. 


tlie  lack  of  either-sex  himts,  and  too  man\-  does, 
especiallv  since  changes  to  buck-onlv  manage- 
ment were  made  beginning  in  1974,  have  merit. 

Hunter  crowding  before  about  1969  when 
license  sales  were  less  than  180,000  (Fig.  1)  was 
probably  a  much  smaller  problem  (Biu'eau  of 
Government  and  Opinion  Research  1971). 
Howe\er,  the  crowding  problem  of  increased 
human  population  and  finite  resources  (Leo- 
pold 1930)  has  been  exacerbated  because  of  the 
long-term  (Leopold  1919)  and  more  recent 
increasing  urbanization,  closures  of  private 
lands  to  public  himting,  and  increased  vehicle 
access  on  both  prixate  and  public  lands  (\hmn 
1977,  Reed  1981). 

Our  findings  indicate  tlie  majoritx'  of  hunters 
prefer  reduced  hunting  opportunity'  for  higher 
qualih'.  When  himters  were  asked  to  indicate 
the  effect  of  crowding  on  their  hunt  quality', 
using  an  1 1  -point  scale  where  0  means  crowding 
greath'  decreased  the  quality  and  10  means 
ci'owding  had  no  negative  effect,  onlv  27.8%  of 
hunters  (scale:  8,9,10)  indicated  crowding  had 
little  effect  compared  to  60.2%  of  hunters 
(scale:  0-5)  who  indicated  a  large  effect  (.v  = 
4.92).  Krannich  et  al.  (1991)  reported  71%  of 
hunters  belie\ed  there  were  too  man\'  hunters 
in  their  areas.  Crowding  effects  were  not  signif- 
icantlv  related  to  hunter  age,  sex,  years  ot  expe- 
rience, unretrieved  deer  reported,  or  whether 
hunters  were  on  private  or  public  lands.  Suipris- 
ingl\,  the  means  for  hunters  from  successful 
(5.04)  and  unsuccessful  parties  (4.96)  were  not 
different.  These  data  indicate  the  effects  of 
crowding  are  felt  b\  almost  all  groups  ecjuallw 
Howexer,  hunters  from  limited-entn  areas 
(F  <  .002),  xvhere  hunter  numbers  are  limited, 
lated  the  effect  ot  croxxcling  less  negatixely  (.t  = 
6.16),  xx'hile  hunters  preferring  to  hunt  in  areas 
restricted  from  xehicles  xvere  more  (F  <  .001) 
negatixelx-  affected  (.v  =  4.61)  than  hunters  pre- 
terrino;  no  restrictions  (.v  =  5.50). 

Management  Options  to  Reduce 
Hunter  Croxxding 

Sex  era!  options  are  axailable  to  reduce 
hunter  crowding.  Split  deer  hunting  seasons 
\\  ere  opposed  bx'  Utah  hunters  in  recent  studies 
(Krannich  and  Cundv  1989,  Austin  et  al.  1990. 
Krannich  et  al.  1991 ).  This  option  xx'ould  likelx' 
increase  Inmting  pressure  on  bucks  bx' 
increased  hunter  tlaxs,  longer  seasons,  and 
huntinti  duriuii  the   more  xailnerable   nitting 


19921 


Hunter  Opinions  on  Deeh  Manacement 


371 


period;  it  would  therein  lurtlier  decrease  mean 
age  and  size  ot  harxested  hueks. 

A  second  option  is  to  require  hunters  to 
choose  either  a  buck  or  doe  tag.  Our  suncx 
indicated  78.4%  of  resident  hunters  would 
choose  a  buck  tag,  which  would  reduce  buck 
hunting  pressure  b\  about  21.6%. 

A  third  option  is  to  recjuire  hunters  to  choose 
and  hunt  onh^  one  season.  Since  mean  hunters 
afield  for  198.S-89  combined  were  archen'  = 
26,613,  rifle  =  180,298,  and  muzzleloader  = 
8832,  this  option  would  reduce  crowding  during 
the  rifle  liunt  up  to  approximately  20%  assum- 
ing hunter  proportions  remained  about  the 
same.  Hunters  taxor  this  option:  in  our  1 989  and 
1990  suneys,  63.8  and 64.0%,  respecti\"el\-.  In  a 
1990  completely  randomized  telephone  sui'vev' 
of  14,305  deer  hunters,  58.0%  of  Utah  hunters 
indicated  preference  for  this  option.  Krannich 
et  al.  (1991)  reported  a  similar  le\el  ot  support 
(mean  score  =  6.19)  using  a  scale  of  0-10. 

ProbabK  the  most  effectixe  option  to  perma- 
nentK  reduce  hunter  crowding,  while  at  the 
same  time  establishing  a  minimum  standard  for 
(}ualit\-  in  terms  of  hunter  pressure  on  bucks,  is 
tc;  limit  license  sales  of  buck  tags.  Hunters  con- 
sistently favor  this  option.  In  our  1990  suney 
60.6%  of  resident  hunters  preferred  to  limit 
buck  license  sales  to  150,000,  with  up  to  35,000 
antlerless  tags  available  to  unsuccessful  buck  tag 
applicants;  39.4%  favored  unlimited  license 
sales.  Since  hunters  who  fa\ored  limiting  license 
sales  also  faxored  haxing  to  choose  sex  of  tag 
(F  <  .004),  most  hunters  would  favor  having  to 
choose  sex  of  tag.  Krannich  et  al.  (1991)  deter- 
mined most  hunters  (61.7%)  supported  choos- 
ing the  sex  of  tag;  and  havino;  vearK  lianest 
restricted  to  one  deer  per  hunter 

In  the  1989  sunew  onl\"  36.6%  of  hunters 
indicated  preference  to  hunt  e\en\'ear  regard- 
less of  future  growth  in  hvmter  numbers,  while 
the  majoritv  (63.4%)  selected  some  lexel  of 
hunter  number  limitation  (Austin  et  al.  1990). 
Of  hunters  preferring  the  limitation,  38.2% 
selected  the  limit  at  160,000  and  25.2%  selected 
the  200,000  limit.  Prexiously  in  1987,  55.8%  of 
hunters  showed  preference  to  limit  hunters  to 
less  than  200,000  (Austin  and  Jordan  1989). 

It  is  apparent  to  the  authors  that  some 
restrictions  are  needed.  We  beliexc  the 
increased  buck  hunting  pressure  beginning  in 
1970  (Fig.  1)  has  had  negative  effects  on  hunter 
success,  satisfaction,  motivation,  and harxe.sted- 
buck  size.  These  negative  effects  appear  to  out- 


weigh (lie  \alues  of  increased  wildlife  manage- 
ment income  and  hunting  recreation  opportu- 
nit\.  hulccd.  hunter  responses  from  these 
suiA ('\  s  continii  our  \i(n\'  that  hunting  pressure 
on  bucks  should  be  i-cduccd  (o  the  pre-1970 
lewl. 


Lite  RATI' RE  r:iTED 


Al  sTi\  D.  D.  Ujyi.  Age  spixit'if  antler  tine  counts  anil 
c;ircass  weiglits  t)f  huntcr-lianested  mule  deer  from 
Utali  cliecking  stations,  19.32-1988.  Division  of  Wild- 
life Resources,  Salt  Lake  Citv,  Utah.  .3.5  pp. 

./Vl  STiN,  D.  D.,  S.  D.  Bunnell.  .AND  P.  J.  Uhnkss  1990. 
Responses  of  deer  lumters  to  a  checking  station  ipies- 
tif)nnaire  in  UtiJi.  We.itern  Association  of  Game  and 
Fish  Connnissioners  Proceedings  69:208-229. 

.\rsTiN,  D.  D.,  .\Nl)  L.  JoRD.AN  1989.  Responses  of  Utal) 
deer  jiunters  to  a  checking  station  (]uestionnaire.  Great 
Basin  Naturalist  49:1.59-166. 

Austin,  D.  D.,  R.  A.  Rioos,  R  J.  Uhnkss.  D.  L.  Tuhnf.h. 
.AND  J.  F.  KiMB.ALL  19S9.  Changes  in  nnile  deer  .size  in 
Utali.  Great  Basin  Naturalist  49:31-35. 

Bunnell,  S.  D.,.and  D.  D.  .Austin  1990.  Unutcropinions 
and  questionnaires.  Pages  l(V-47,  87  in  Deer  manage- 
ment workshop,  June,  .\ugust  1990.  Utah  Division  of 
Wildlife  Resources,  Salt  Lake  Citv.  94  pp. 

Buhe.auofGovern.ment.and  Opinion  Resk.muii  1971. 
Opinion  sune\'  of  the  people  of  Utali  lor  wildlile 
resources  and  outdoor  recreation,  Utah  State  Uni\er- 
sit\,  Logan.  73  pp. 

Decker.  D.  J.,  .and  N.  A.  Connolly  1989.  Motivations 
for  deer  hunting:  implications  for  antlerless  deer  har- 
vest as  a  management  tool.  Wildlife  S(K'iet\'  Bulletin 
17:4.55^63. 

Fi.vrHEH,  C.  II..  AM)  1",  W  iloKKSTH.A,  1989.  An  iuiaiv.sis 
of  the  wildlife  anil  fish  situation  in  the  United  States: 
1989-2040.  USDA  Rock-A  Mountain  Forest  and  Riuige 
E.Nperiment  Station  General  Teclinical  Report  R.M- 
17S.  147  pp. 

Jense,  G.  1990,  Three-point  strateg\  in  Utah,  Pages  78-88 
in  Deer  management  workshop.  |nne.  .\ugust  1990, 
Utiili  Division  of Wikllife  Resources.  Siilt  L;ike  Cjt\.  94  pp. 

|ense,  G.  K.,,and  W.  Shields  1990.  Utdi  big  game  annual 
report.  Utah  Division  of  Wildlife  Resources  Publica- 
tion No.  90-7. 

Kk  vnnicii.  R.  S.,  and  D.  T,  Ci  ndy  1989.  Perceptions  of 
crowding  and  attitudes  about  a  split  deer  hunting 
.season  among  resident  and  nonresident  Utah  deer 
hunters.  Institute  for  Social  Science  Research  on  Nat- 
ural Resources,  Utah  State  Universit).  Logan.  29  pp. 

Khannk  II  R.  S..J.S.  Keitiland  V.  A.  Riiev  1991.  Utidi 
deer  hunters"  opinions  about  deer  hunting  and  iilterna- 
tive  season  formats.  Project  Summarv  Report.  Institute 
for  S(Kial  Science  Research  on  Natural  Resources, 
Utal)  State  Universitv.  Logan.  75  pp. 

Leopold,  A,  1919.  Wildlifers  vs.  game  farmers:  a  plea  for 
demcx-racv  in  sport.  \C?\  Bulletin.  Pages  .54-fiO  iu 
D.  E.  Brown  and  N.  B.  Carmony  (1990).  .\ldo 
Leopold's  wilderness.  Stackpole  Books.  Harrisburg, 
Pennsvlvania.  249  pp. 

.  1930.  CJame  m;uiagenient  in  the  national  forests. 

.\nierican  Forester  (July):  412—114. 


372 


Great  Basin  Naturalist 


[Volume  52 


Mann,  D.  K.  1977.  Land  use  and  acquisition  stud\  on 
wildlife  ranges.  Utah  Division  of  Wildlife  Resources 
Job  Completion  Report  \V-65-R-D-25,  Job  A-S.  270  pp. 

McCi:llouc;ii,  D.  R.  1979.  The  George  Reserx'e  deer 
herd.  Universit\  of  Michigan  Press,  Ann  Arbor.  271  pp. 

Reed,  D.  F.  19S1.  Conflicts  with  ci\ili/.ation.  Pages  509-536 
ill  O.  C.  W'allmo,  ed..  Mule  and  black-tiiiled  deer  of 
North  America.  Uni\'ersit\- of  Nebraska  Press,  Lincoln, 

605  pp. 
Robinetpe,  W.  L.,  N.  V.  H.\n(xx:k,  .and  D.  A.  Jones.  1977. 
The  Oak  Creek  mule  deer  herd  in  Utah.  Utah  Di\ision 
of  Wildlife  Resources  Publication  No.  77-15.  14Spp. 


SciiREYEK.  R.,  R.  S.  Kh.wnich.  .\nd  D.  T.  Cindy  1989. 

Public  support  for  wildlife  resources  and  programs  in 

Uttili.  Wildlife  Society  Bulletin  17:532-538. 
St.\pley,  H.  D.  1970.  Deer  illegal  kill  and  wounding  loss. 

Final  Report.  Utah  Di\ision  of  Wildlife  Resotn-ces 

Publication  W-65-R-1-A-8.  7  pp. 
TowElLL.  D.  E.,  and  S.  T  Allen    1990.  Results  from  the 

15-vear  policy  plan  questionnaire.  Idaho  Department 

of  Fish  and  Game,  Boise.  14  pp. 

Received  17  December  1991 
Accepted  3  November  1992 


Cw'eat  Basin  Natui;Ji.st  52i4).  pp.  373— 37S 

LIST  OF  OREGON  SCOL\TIDAE  (COLEOPTERA) 
AND  NOTES  ON  NEW  RECORDS 

Malcolm  M.  Funiiss  .  )anies  B.  Jolnison  .  Hicliaixl  L.  \\estc()tt".  and  Torolf  1^.  T<)r<^ersen' 

AliSTlUCr. — Listed  arc  121  species  of  Scolvtidae  Iroin  Oregon.  Ten  species  are  reported  from  Oregon  for  tiie  first  time: 
HijUistcs  tenuis  Eiclilioff,  Phlocosinits scoptiloniin  scopiilontiii  Swaine,  Plilocosinus  hofeii  Blacknuui.  Tn/jHxIctulnm  hctulac 
Swtiine, Xylehonis xyl(><ir(ij)Iiit.s  ( Sa\),  Tn/pophlncti.s strUitulus  { Mannerheini ),  T  thatclwri  Wood,  Procnjphalus  iiuicronatus 
(LeConte),  Piti/oplitlionts  scalptor  Blackman,  and  Moiuiiilinnti  dciitit^cnim  (LeConte).  Tlie  second  Oregon  sj^x?cimen  of 
an  exotic  species.  Xi/lchDnis  cdlifonuciis  Wood,  is  reported  also. 


Ki'i/  uords:  Sc('li/ti(l(ii\  liimuil  list.  Oivj^oii. 

Oregon  i.s  a  large  state  with  dixerse  \egeta- 
tion  tliat  occurs  there  chie  largeK-  to  the  widt^ 
range  of  ph\sical  and  cHniatic  emironnients. 
The  climate  residts  in  part  from  the  inteipla\' 
hetvxetMi  maritime  and  continental  air  masses 
and  the  intenening  Cascade  Moimtain  Range 
that  divides  the  state  into  distinct  western  (mai- 
itime)  and  eastern  (continental)  regions  (Frank- 
lin and  D\niess  1973).  For  example,  axerage 
annual  precipitation  \aries  from  approxiniateK' 
60-300  cm  west  of  the  Cascades  to  20-100  cn'i 
eastward. 

The  exceptionalh"  diverse  forests  of  south- 
western Oregon  ha\e an affinits" witli California, 
whereas  those  ot  northeastern  Oregon  are 
related  to  Kock\  Mountain  forest  t\pes. 
Because  ScoKtidae  are  host-specific  to  some 
degree,  their  distribution  in  Oregon  is  linked 
closeK  to  the  distribution  of  species  of  trees  and 
sliruhs. 

Oregon  scohtids  were  listed  h\  Chamberlin 
(1917),  but  that  list  is  greatly  out  of  date.  We 
herein  update  the  list  to  include  records  and 
synonvmies  publislu^d  b\  Wood  ( 19S2).  Similar 
lists  have  been  publislK^l  recentK  for  Idaho 
(Fumiss  and  Johnson  1987)  and  Montana  (Cast 
et  al.  1989).  " 

Six  species  not  pre\  ionsK  rej)orted  Irom 
Oregon  were  collected  b\-  us  on  field  trips  in 
1990,  and  four  species  were  found  among 
nuiseum  specimens.  More  species  will  sureKbe 


found  In  further  collecting.  The\  likeK  will 
inckuk'  more  Pifijophthonis,  a  genus  that  is 
relatively-  rich  in  species  in  westeni  forests  and 
elsewhere;  and  species  of  other  genera  from  the 
diverse  California  fauna  ( Bright  and  Stark  1973) 
that  infest  trees  endemic  to  both  states. 

Other  new  scoKtids  are  likeK  to  be  intro 
duced  accidentalK  b\  connnercc".  For  example, 
tlie  exotic  Xi/Iebonis  affinis  Eichhoff  was  inter- 
cepted in  1961  at  Portland  in  DiYicacna 
niassaii^caiKi  from  Puerto  Rico.  Of  the  121  spe- 
cies listed  here.  8  are  clearK'  exotics  that  have 
become  estalilished  at  unknown  times: 
Hylastiiins  ohsctinis  (Marsham).  Scoh/tus 
niffdosiis  (Miiller),  S'.  imilfistriattis  (Marsham), 
XijJchonis  dispar  (Fabricius),  A',  xijlix^raphns 
(Sav),  X  califoniicus  Wood.  Xi/lchoriinis 
saxeseni  (Ratzeburg),  and  Monaiihnnn  dciiti- 
^criiDi  (LeConte).  Of  these,  X.  califoniicus  was 
known  heretofore  in  Oregon  from  onlv  one 
specimen  (Wood  1982);  a  second  specimen  was 
caught  (b\  IB|)  in  flight  after  sunset.  6-\'III- 
1990.  (.'hampoeg  State  Park,  Marion  Co.  It 
[)r()l)abK  was  introduced  b\  connnerce  at  Port- 
land, although  its  native  range  is  still  unknown. 
We  speculate  that  it  max  infest  distressed  decid- 
uous trees  along  the  Willamette  River. 

B\  their  habits,  Oregon  Scol\ti(kie  are  char- 
acterized as  true  bark  beetles,  living  in  cambium 
(105  speciesh  ambrosia  beetles,  living  in  xvlem 
although  thev   may  feed  entirely  or  partlv  on 


Dhision  of  Entoniolu^\.  Univer.sityol  Idaho.  Moscow,  Iclalio  S-3S4:?-4iy6. 
"Oregon  Department  of  Agriculture.  Salem,  Oregon  97.'51()-01 10 
USl3,\  Forest  Sersice,  LaGrande,  Oregon  97890. 


373 


374 


Great  Basin  Naturalist 


[Vblunie  52 


fungi  tliat  they  transmit  (14  species),  living  in 
pine  cones  (Conoplithonis  pondcrosac  Hop- 
kins), or  living  in  roots  of  clox'er  {H.  obsninis). 
Conifers  are  hosts  of  98  species,  while  the  other 
23  species  occur  in  angiospernis. 

Abbreviations  of  repositories  listed  for  spec- 
imens new  to  Oregon  are:  ODAC  =  Oregon 
Department  of  Agriculture  collection,  Salem; 
PNW  =  Pacific  Northwest  Forest  and  Range 
Experiment  Station  collection.  Forest  Service, 
USDA.  Con  allis,  Oregon;  WFBM  =  W.  F  Barr 
Entomological  Museum,  Universitv'  of  Idaho, 
Moscow,  Idiilio;  and  SLW  =  Stephen  L.  Wood 
collection,  Rrigham  Yoiuig  Univ'ersit\',  Provo, 
Utali. 


Species  New  to  Oregon 

Subfamily  Hylesininae 

Ht/Iastcs  tenuis  Eichhoff 

Biology. — Monogvnous.  Infests  Finns  spp., 
presumably  the  roots. 

Distribution  and  notes,— Mexico: 
Hildago  and  Mexico  (state);  USA:  Mass.  to  Fla., 
all  southern  states  westward  to  Calif.,  and  Ida. 
Ore(;oN:  Eugene,  Lane  Co.,  22-IX-1971,  black 
light  trap,  K."j.  Goeden  (1  ODAC).  Prineville, 
Crook  Co.,  25-VII-1934  ( 1  PNW),  VIII-1935  (1 
PNW),  Finns  pondcwsa,  Hopk.  18960-83.  W.  J. 
Buckhom  (parat\pes  of  the  s)iion)-m,  H. 
ininntns  Blackman). 

Flilocosinns  scopnlonun  scopniornnt  Swaine 

BlOLO(;v. — Monog\nous.  Infests  stems  of 
Jnniperns  scopnlonini.  Galleries  parallel  to 
grain  with  a  nuptial  chamber  just  above  the 
entrance,  appearing  as  though  the  chamber 
were  halved  antl  one  side  shifted  fonvard  half 
its  diameter  (Bright  1976,  Fig.  182). 

Distribution     and     notes.— Canada: 

Alta.,  B.C.;  USA:  Wash.  OREGON:  Sisters, 
Deschutes  Co.,  H-V-nHH.  Junipcms  sp.,  R.  L. 
Penrose  (4  9,  3  d  ODAC).  Canbv.  Klackamas 
Co.,  April  15,  1965,  K.  J.  Goeden'd  6  ODA). 
North  Plains,  Wa.shington  Co.,  20-I\'-1969. 
Thuja  plicata.  K.  J.  Goeden  (2  cJ,  1  9  ODA,  2 
6  SEW).  Portland,  Multnomah  Co.,  22-X- 1971, 
Chaniaca/pahs  laicsoniana,  R.  L.  Westcott  (1 
9,  1  cJ  ODA,  1  9  SLW).  Northbend.  Coos  Co.. 
9-VI-1974,  on  cvpress,  J.  McLaughlin.  (3  9 
ODA). 


Fhloeosinus  Jiofeii  Blackman 

BI0L0(;y. — Monogynous.  Infests  branches 
of  Junipenis  deppeana,  J.  osteospcrnuL  and/. 
scopuJoninx. 

Distribution  and  notes.— Canada: 
B.C.;  USA:  all  western  states  except  Wash. 
OREGON:  about  9  km  W  Enteqirise,  Wallowa 
Co.,  9-XI-1990,  Jnniperus  scopulonim,  M.  M. 
Furniss  and  A.  Ecjuihua  (12  9,5  S  WFBM). 
Infesting  branches,  ()..5-3.0-cm  diameter,  of  a 
small  standing  tree.  Lanae  parasitized  bv  an 
abundant  braconid  wasp,  EcpJiylus  sp.,  proba- 
bly californicns  Rohwer  Host  is  restricted  in 
Oregon  to  the  vicinity  of  the  \\  allovva  River,  for 
a  distance  of  approximatelv  30  km  dovvaistream 
from  Enteiprise. 

Subfamily  Scolvtinae 

Tnjpodcndron  bcfulac  Swaine 

BI0L0(;y. — Monog)'nous.  Infests  Bctula 
spp.,  rare  in  Alnus  spp.  Tunnels  are  constiiicted 
bv  females  radiallv  into  sapwood.  Other  females 
constmct  branches  from  the  radiallv  aligned 
tunnel  at  close  intenals,  left  or  right,  in  the 
horizontal  plane.  Eggs  are  laid  in  niches  ori- 
ented above  and  below  the  galleiv.  Larvae  exca- 
vate short  cradles  in  which  thev  develop  and 
feed  on  ambrosia  fungus.  Males  are  active  in 
keeping  the  tunnels  clean  and  aerated. 

Distribution  and  notes.— Canada: 
Alta.,  B.C.,  Man.,  N.B.,  N.S.,  N.W.T.,  Ont., 
Que.;  USA:  Ida.,  Me.,  Mass.,  Minn.,  Mont., 
N.H.,  N.J.,  N.Y.,  S.D.,  Wise.  OREGON:  Mill 
Creek,  Umatilla  Co.,  8-XI-1990,  Bctula 
papt/rifcra.  M.  M.  Furniss  and  A.  Equihuad  9, 
3  (5' WFBM,  19,1c?  ODAC).  Infesting  lower 
stem  of  a  23-cm-diameter  wind-felled  tree.  Also 
present  were  Xi/lchorus  dispar  (Fabricius)  and 
Xijicborinns  saxcscni  (Ratzeburg). 

Xt/lchorns  xi/lo^rapltns  (Sav) 

Biology. — Unstudied.  In  species  of  this 
genus  that  are  studied,  haploid  males  are  pro- 
duced parthenogeneticallv.  They  are  dwarfed 
and  flightless.  Diploid  females  are  produced  by 
mating  between  siblings  or  between  a  female 
parent  and  a  male  offspring.  Infests  Qucrcns 
spp.,  i-are  in  other  hardwoods.  The  galleries  are 
made  obliquelv  into  sapwood  in  a  horizontal 
plane  to  a  depth  of  an  inch  or  more,  after  which 
thev  branch,  the  arms  follow  iii<j;  tlu^  annual  rings 
(Beal  and  Massev  1945). 


19921 


List  oi'  Ohhcon  Scoiatidae 


375 


DlSTRIBUTIOX  AND  NOTES.— CANADA: 
Out.,  Que.;  USA:  hvenh-hvo  .states  (and  D.C.) 
east  of  lOOth  iiieridiaii;  Calif.  ( 1  specimen),  Te.x. 
Ol^E(;()N:  5  km  \\\'  Xewberg,  Yamhill  Co.,  20- 
\I-1970,  black  light  trap,  K.  J.  Coeden  (1 
ODAC). 

Tn/f>(>f)]th>cu\  sthtifithis  ( Mannerheim) 

B|()L()(;Y — Monogxnous.  Infests  outer  bark 
ot  Salix  spp.,  most  connnonK'  S.  scoulcriaiia- 
also  recorded  from  Ahnis  spp.  Ma\"  reinfest  stem 
progressi\el\'  downward  for  sexeral  genera- 
tions. Ca\e  t\pe  egg  gallen;  larx^ae  mine  shal- 
low 1\  under  bark. 

Distribution  and  notes.— Canada: 
New'f..  N.S..  Que.,  Yukon;  USA:  Alas.,  Colo., 
Ida.,  Minn.,  Ut.  Orecx)N:  Hot  Springs  Camp- 
ground, Hart  Mtn.  Natl.  Antelope  Refuge,  Lake 
Co.,  14-MII-I99(),  Salix  scoulcriana,  M.  M. 
Furniss  and  J.  B.  Johnson  (34  WFBM,  5 
ODAC).  Infesting  necrotic  bark  lesions  in  a  li\e 
stem  ha\ing  a  deep  frost  crack.  Diameter  of 
infested  part:  5-10  cm.  Mature  lar\ae  present. 

Tn/)H>phIociis  thatchcri  Wood 

BlOLOCiY. — Monogx  nous.  Infests  outer  bark 
of  standing.  vmhealth\'  or  dving  Popitliis 
tirnmloides.  Ca\"e  t)pe  egggaller)';  lanal  mines 
confined  to  outer  bark. 

Distribution  and  notes.— Canada: 
B.C.;  USA:  Calif,  Ida.  OREGON:  Hot  Springs 
Campground.  Hart  Mtn.  Natl.  Antelope 
Refuge,  Lake  Co.,  14-\III-199{),  Fopulus 
trcnuiloklcs,  M.  M.  Furniss  and  J.  B.  Johnson 
(27  WFBM,  5  ODAC).  Adults  attacking  and 
walking  on  bark  of  a  d\ing,  15-cm-diameter 
tice. 

PnuTi/pluihis  mucroiuitiis  (LeConte) 

BioLOC;Y. — MonogN'uous.  Infests  Populus 
trcDiitloidcs.  Prefers  soft,  fermenting,  dead 
bark;  usualK  follows  primary'  invasion  by 
Tn/f)(>plil()('uspopuU  Hopkins  (Pett\"  1977).  The 
gallen  is  narrower  and  the  bark  o\erl\ing  the 
gallen  is  thicker  than  that  of  T.  popnli  Hopkins 
(and presumably  T.  thatclwri).  One  and  one-half 
to  two  annual  generations  (Utcili),  ovenvintering 
as  lanae  and  adults.  Eggs  appear  first  in  late 
Ma\. 

Distribution  and  notes.— Canada: 
Alta.,  B.C.;  USA:  Alas.,  Colo.,  Ida.,  Mont.,  Nev., 
N.M.,  Ut.  OREGON:  Hot  Springs  Campground. 
Hart  Mtn.  Natl,  .\ntelope  Refvige,  Lake  Co., 
U-\'m-l99(),  Popuhis  tremulokle.s,  M.  M.  Fur- 


niss and  J.  B.  John.son  (9  WFBM).  Infesting 
stem  of  a  26-cm-diameter  tree.  Jackman  Park, 
Steens  Mtn.,  Harney  Co.,  14-VIII-199(),  Pop- 
iiJiis  tronuloidt's.  M.  M.  Furniss  and  J.  B.  John- 
son (14  WFBM,  10  ()D.\C).  Attacking  lower 
stem  of  a  25-cm-dianieter  dead  tree  (foliage 
shed,  l)ark  moist  i. 

Pitijophthonts  scalptor  l^lackman 

BlOLO(;V. — Presnmal)l\  pol\g\iious.  Infests 
small  branches  of  living  pines. 

Distribution  and  notes— Canada: 
B.C.;  USA:  Calif.,  Ida.  OREGON:  15  km  \ 
Palmer-Junction,  Union  Co.,  Ifi-\1I1-1990, 
Piniis  poiulerosa,  M.  M.  Furniss  and  J.  B.  John- 
son (29,2c?  WFBM).  Infesting  1-cm-diameter 
freshlv  faded  lower  branch  on  a  li\e,  merchant- 
able tree.  Each  gallerv  contained  only  one 
female  and  one  male,  no  eggs  (jr  lanae;  thev 
appeared  destined  to  o\en\'inter  before  repro- 
ducing. 

Monatiltnuii  (lcnfi<i('nini  (LeCJonte) 

Biology. — Not  studied.  Infests  Qiicrcus 
spp.  Most  species  q(  MonaiihniDi  are  poKgN- 
nous  and  their  galleries  branch  from  a  radialK 
oriented  entrance  tunnel  in  the  x\4em.  Lanae 
of  this  genus  develop  in  niches,  apparentk'  feed- 
ing on  a  mi.xture  of  ambrosial  fungus  that  grows 
on  gallen'  walls  and  .wlem  of  the  host  tree. 

Distribution  and  notes— Me.xigo:  Baja 
California;  USA:  Ariz.,  Calif.,  Te.x.  OREGON: 
Medford,  Jack.son  Co.,  18-\1II-1968.  black  light 
trap  (1  ODAC). 

Oregon  sgolytidae 

HVI.F.SIMN.VE 

Hylastini 

Scients  aiuicctcnsLi'Conte 

Hi/lur<i(>ps  porosits  (LeGonte) 

lli/hirf^ops  reticuldtus  Wood 

Hi/htr^ops  nt^ipcniiis  ni^ipcinii.s  (Mannerheim) 

Uillnronps  subcostuhitua  sithcosUdatiis  ( Vhuinerheim) 

Ui/lastcs  'gracilis  LeGonte 

Hi/lastcs  l(>n<iic<>Uis  Swaine 

lli/lastcs  macer  LeCJonte 

Utjlastes  ni'^rimis  (Mannerheim) 

Hi/lti.stcs  n/ter  Swaine 

Ihllastcs  tenuis  Eichlioff 

H>lesinini 

Hijlastinm  ohsainis  (Mar.shani) 
Hylesinus  califonticus  (Swaine) 
Hi/lesiniis  oregomis  (Blackman) 
.\iniplui<ius aspericollis  (LeConte) 
Abiiphaf^us  hirsutus  Schedl 


376 


Great  Basin  Naturalist 


[N'olume  52 


Tomicini 

Xylcchinii.s  inoiildiiiis  Blafkinan 
PsL'udolu/li'sinits  ilispar  dispar  Blacknum 
Pseudoln/lesiuus  dispar  pullafiis  Blackmail 
P.seiul()ln/Icsiini.s  '^ninuhitits  ( LeCx)nte) 
Pseudohi/lesiiui.s  iiebtdosns  itcbulosns  (LeConte) 
Pseudolu/U'siniis  iiobilis  Swaine 
Psviidi>lu/lcsiiut.s  j)ini  Wood 
Psciidohi/lcsiiuts  scriceiis  (Mannerlieim) 
Psciidohi/lcsiiuts  sifclicnsis  Swaine 
Pscuthiln/h'siiius  tsiioac  Swaine 
Doidnxinuiis  hrcvicoinis  LeConte 
Dcndntcfonus  jeffrcyi  Hopkins 
Doulroctonus  pondewsac  Hopkins 
Dendroctomis  pscudotsugac  Hopkins 
Deudroctonus  nifij)emus  (Kirby) 
Dciidrocfoiiiis  valcns  LeConte 

Phloeotribini 

PhltHofiihiis  Iccoiitci  ScheiU 

Phloeosinini 

Pldocosinus  (iiitciiiKitiis  Swaine 
Phloco.siiiits  cuprcssi  Hopkins 
Phlocosimis  frdgois  Sw;iine 
Pidocosiims  hoferi  Blackman 
Pidoeo.sinits  punctatiis  LeConte 
Phlocosimis  scopidonim  scoptilontni  Swaine 
Phlocosimis  secpioiae  Hopkins 
Phlocosimis  scrnitiis  (LeConte) 
Phlocosimis  laiidi/kci  Swaine 

Hvpoborini 

Cluictojtldociis  hctcrodoxiis  (Casev) 

Polygraphini 

Cdijiholninis  iittcrnwdiiis  Wood 
Caii)hohonis  piccac  Wood 
Caqdiohorus  pinicolcns  Wood 
Cui-fdwbonis  pondcrosae  Swaine 
Cdqdiobonis  sansoni  Swaine 
Carfdioborus  vandi/kci  Brnck 
Poli/'gniphtis  nifijH'nnis  (Kirby) 

ScoLvriNAK 

Scolytini 

Scoli/tiis  laricis  Bkicknian 
Scoli/tits  monlicolac  Swtiine 
Scoh/liis  limit isfriattis  (Marsliam) 
Scoli/tu.s  opticus  Blackman 
Scoli/tits  orctgoni  Blackman 
Scolt/tiis  piccae  (Swaine) 
Scolytns  pracceps  LeConte 
Scolytits  niffdosns  (Miiller) 
Scolytns  siibscabcr  Le(4)nte 
Scolytns  tsiigac  (Swaine) 
Scolytns  nuispinosus  LeConte 
Scolytns  vciitralis  Le(;onte 

Micracini 

Hylocnnis  hiiicllns  (LeC^ontc) 

Cryphirgini 

Di>liir<iiis  pninilis  (MainuTlieim) 
Cnj])lui-<ins  borcidis  Swaine 

Drjocoetini 

Dn/ococtcs  rtjffrt/^('r  (Mannerlieim) 
Dryococtes  mtto'^raplms  (Ratzebnrg) 
Dnjococtcs  confnsns  Swaine 
Dn/ococtcs  scchclti  Swaine 


Ipini 

Piti/o'^ciics  caiimdcitiis  (LeCJonte) 

PittjO'^citcs  jossijroiis  ( LeConte) 

Pityoficiics  kncclitcli  Swaine 

Pityoktciiics  clegaiis  Swaine 

Pityoktcines  lasiocmyi  (Swaine) 

Pityoktciiics  inimifns  (Swaine) 

Pityoktciiics  onuitns  (Swaine) 

Oiihotoinicns  caclatns  (EichliofO 

//)s  coiiciiiuiis  (Mannerlieim) 

Ips  cilia rf^inatns  (LeConte) 

Ips  iiitc'S,cr  (Eiehhoff") 

Ips  latidcns  (LeConte) 

Ips  mcxicaims  (Hf)pkins) 

Ips  iiiontamis  (Eiclihoft") 

Ips  paracoiifnsns  Lanier 

Ips  piiii  (Sav) 

Ips  plastographus  niaritiiniis  Lanier 

Ips  plastographns  plastographiis  (LeConte) 

Ips  trich'iis  ciigclmamii  Swaine 

Ips  tridcns  tridcns  (Mannerheim) 

Xyloterini 

Tn/podcndroii  bctnlac  Swaine 
Tn/podciidroii  liiicatuin  (Olivier) 
Trypodciidron  rctiisniii  (LeConte) 
Trypodcndroii  nifitarsis  (Kirbv) 

Xyleborini 

Xylcbonis  ctdifoniiciis  Wood 
Xiltcbonis  dispar  (Fabricins) 
Xylcborns  iiitrnsns  Blandford 
Xylcbonis  xylographns  (Sa\) 
Xylclxirimis  saxcsciii  (Ratzebnre;) 

Ci-yphalini 

Trypoplilociis  sidicis  Hopkins 
Trypophlociis  striatidus  ( Mannerlieim) 
Tnjpophlocus  thatchcii  Wood 
Procniplialus  iimcronatns  (LeC'onte) 
Procn/jihaln.s  utahciisis  Hopkins 
Cn/jilidhts  j)ul)csccns  Hopkins 
Cn/phalns  nificollis  rnficollis  Hopkins 

Corthylini 

Psciuhyiityophthonis  pnbipcniiis  ( LeConte) 
Coiiophthonis  pondcrosae  Hopkins 
Pityophthorns  boi/cci  Swiiine 
Pityophtlionis  coiifciins  Swaine 
Pityojihtlionis  confiiiis  LeConte 
Pityoj)litlionis  ilif^cstiis  (LeConte) 
Pityophthorns  clcctiis  Blackman 
Pityophthorns  jeffrcyi  Blackman 
Pitijophthorns  imirrayanac  Blackman 
Pilyoplithonis  nitidnlns  (Mannerlieim) 
Pitt/ophthorns  iiitidns  Swaine 
Pityophthorns  jiscndotsiiffic  Sw;une 
Pityophlhonis  scalptor  Blackman 
Pityophthorns  foralis  Wood 
Pityophthorns  tnbcrcnlatns  Eiclilioli 
C'.iiathot helms  rctnsns  (LeConte) 
Ciiathotriclms  snlcatns  (LeConte) 
Monaiihrniii  dciitii^eniiii  (LeConte) 
Monaiihniin  scntcllare  (LeConte) 

Ac  K  N  ()  W  L  E  D  C  M  E  NTS 

Garv'  L.  Peters  provided  collet'tion  data  for 
specimens  in  tlie  ODAC.  Fred  H.  Schmidt, 


1992] 


Lisi- oi"  ()Ht:(;()N  Scoi.viii) \i 


37 


HSl^A  Forest  Senice,  LaCiramle,  Oregon,  seg- 
regated undetermined  SeoKtidae  in  the  PIV\V 
collection.  Locations  ol  Bctula  papi/rifcra.  host 
of  Tnipodcndron  hetitlac  S\\ain(\  Audjunipenis 
sci)piil(>nim.  host  of  Phloeosinus  hofcri  Black- 
nuin,  \\  ere  proxided  bv  Charles  Johnson,  USDA 
Forest  Service,  Baker,  Oregon.  The  manuscript 
was  rexiewed  in  Frank  W.  Merickel,  Uni\ersit\ 
of  Idaho,  and  Dr.  Stephen  L.  \\ood.  Brigham 
Young  Uni\ersit\,  Pro\o,  Utah,  who  also  ick'uti- 
tied  -Y.  caUfoniicus.  X.  xijlo^iyiplitis.  and  /'  .v, 
■scopiiloruni  other  than  those  collectetl  1)\  us. 
This  is  Uni\ersit\"  of  Idaho  Agriculture  Experi- 
ment Station  Research  Paper  No.  92714. 

Literature  Cited 

Bfai,  ].  A.,  .\ND  C.  L.  .M\ssEY  1945.  Bark  heetle.s  and 
ambrosia  beetles  (Coleoptera:  SeoKtidae)  witli  special 
reference  to  species  occurring  in  North  C^arolina.  Duke 
Universit\'  School  of  Forestr\  Bulletin  10.  ITS  pp. 

BiuciiT  D.  E'..  Jr.  1976.  The  bark  beetles  of  Canada  and 
.Alaska.  The  insects  and  arachnids  of  Canada,  Part  2. 
Biosystematics  Research  Institute,  Research  Board, 
Canada  Department  of  Agricultm-e  Publication  1576: 
1-241. 


Bkiciit  n.  E.,  JH  AM)  U.  W.  .Stahk  1973.  The  bark  and 
anibro.sia  beetles  of  California.  Coleoptera:  SeoKtidae 
and  Plat\podidae.  Bulletin  of  the  California  Insect 
Sur\i\.  \  ol.  16.  169  pp. 

Cll,\.\liii:iu.lN,  W.  J.  1917.  An  annotated  list  of  the  .scoKtid 
beedes  of  Oregon.  C>'anadian  Entomologi.st  49:  321- 
328,  35.3-356.  ' 

Fha\ki.i\.  J.  P.,  AND  C.  T.  Dyhness.  1973.  Natural  vegeta- 
tion of  Oregon  and  Washington.  USDA  Forest  Ser\ice 
Ccniral  Technical  Hcpoit  PNW-S.  Portland.  Oregon. 
417  pp. 

FiKMss,  M.  M.,  AND  J.  B.  Johnson.  19S7.  List  of  Idaho 
SeoKtidae  (Coleoptera)  and  notes  on  new  records. 
Creat  Basin  Naturalist  47;  37.5-382. 

Cast  S.  J..  .\I.  M.  Fi  HNiss,  J.  B.  Johnson,  and  .\I.  .\.  1\  if. 
1989.  List  of  Montana  SeoKtidae  (Ccjleoptera)  and 
notes  on  new  records.  Creat  Basin  .Naturalist  49:  3SI- 
386. 

Petty,  J.  L.  1977.  Bionomics  of  t^vo  aspen  bark  bc-edes, 
Tn/popltl(H'iis  popnli  and  Fnun/phaltis  iniicroiidtiis 
(Coleoptera:  ScoKtidac^i.  Creat  Basin  Naturalist  .37: 
10.5-127. 

Wood  S.  L.  1982.  The  bark  ;ukI  ambrosia  beetles  of  North 
and  Central  .\nierica  (Coleoptera:  SeoKtidae).  a  t;L\o- 
nomic  monograph.  Great  Basin  Naturalist  Memoirs 
No.  6.  13.59  pp. 


Received  3  April  im-I 
Accepted  1  October  1992 


Grt-at  Basin  Naturalist  52(4),  pp.  378^381 


RIFFLE  BEETLES  (COLEOPTERA:  ELMIDAE)  OF 
DEATH  VALLEY  NATIONAL  MONUMENT,  CALIFORNIA 


William  D.  Sheparcr 

Abstract. — Three  species  of  Elmidae  occur  in  Death  Valley  National  Monument:  Stenelmis  ccilicia  is  in  three  springs 
in  the  Ash  Meadows  area;  MicroajUoepiisfonnicoUleiis  is  only  in  Travertine  Springs;  and  MicroctjUocpus  -similis  is  in  several 
springs  throughout  Death  Vallev  and  Ash  Meadows.  Only  permanent  springs  support  elmids.  Considerable  morphological 
variation  occurs  in  the  disjunct  populations  of  A/,  siinilis.  The  evolution  of  elmids  in  Death  VaOey  National  Monument  is 
e(juivalent  to  that  of  the  local  pupfish  (C.ij}>rin()ilt>)i  spp.). 

Ki'i/  iconls:  Death  Vallcij.  In.sccta.  Coleoptcrii.  Ehtud(U\  (listribiitions.  (Icsci-tifiration.  ciohitioiL 


Death  Valley  National  Monument  (D\'NM) 
is  located  mostly  in  southeastern  California, 
with  two  small  extensions  into  southwestern 
Ne\ada.  D\'NM  includes  Death  VallcN'  proper, 
its  adjacent  mountain  ranges,  and  the  Ash 
Meadows  area  of  Nevada  which  surrounds 
Devil's  Hole.  Biogeographically  this  is  a  transi- 
tion area  between  the  Mojave  Desert  and  the 
Basin  and  Range  Desert.  Desert  conditions 
here  are  the  result  of  the  drier  and  warmer 
post-Pleistocene  climate  and  a  rain-shadow 
effect  from  the  Panamint  Mountains,  the  Sierra 
Nevada,  and  the  Coast  Range  mountains  to  the 
west. 

Water  sources  in  DVNM  are  une.xpectedh' 
common.  Palmer  (1980)  cites  over  100  springs 
alone.  Hunt  (1975)  has  classified  these  springs 
into  four  tvpes  based  upon  volume  of  discharge 
and  geomoiphic  origin.  The  Amargosa  Ri\er 
flows  (when  it  does!)  into  the  southern  end  of 
Death  Valley.  Two  permanent  streams.  Salt 
Creek  and  Furnace  Creek,  are  located  in  the 
central  portion  of  DVNM.  Numerous  "wells" 
(shallow,  subsurface  water  sources)  and  "seeps" 
are  to  be  found  scattered  throughout  DVNM. 
These  are  not  reliable  water  sources,  being 
more  or  less  intermittent.  Wherever  a  water 
source  does  occur,  however,  it  may  not  be  \en 
amenable  to  aquatic  organisms  because  of  lethal 
temperatures  and/or  salinities.  Discussions  of 
the  local  hvdrolog)'  can  be  found  in  Hunt  ct  al. 
(1966)  and  Soltz  and  Nai man  (1978). 


Of  the  aquatic  organisms  occurring  in 
DVNM,  onl\-  the  fishes  ha\e  been  studied 
extensively.  Soltz  and  Naiman  (1978)  re\-iewed 
the  past  work  and  presented  an  excellent  s\ni- 
thesis,  particularly  so  hrCyprinoclo)i  spp.  (pup- 
fish).  For  aquatic  insects,  studies  ha\e  been 
primarilv  descriptions  of  new  species  and  their 
t\pe  localities  [e.g..  Chandler  (1949),  Usinger 
(1956)];  onlv  one  studv  (Colbum  1980)  directly 
addressed  the  ecolog)' of  anv  species.  Howe\er, 
Deacon  ( 1967, 1968)  discussed  insects  as  part  of 
the  community  ecology  of  Saratoga  Spring. 

During  a  vacation  I  found  a  single  specimen 
of  a  riffle  beetle  in  Saratoga  Spring  at  the  south 
end  of  D\'N  M .  That  chance  discoverv  led  me  to 
embark  on  a  survey  of  the  water  sources  in 
DVNM  to  determine  if  other  elmids  (riffle  bee- 
tles) occurred  there,  and,  if  so,  in  which  sources. 

Methods 

Water  Sources 

The  water  sources  examined  were  chosen 
primariK-  because  of  their  accessibilitA".  Those 
that  rec^uired  more  than  a  da\s  tra\el  by  auto 
and/or  foot  were  not  examined.  In  all,  27  water 
sources  were  examined  in  Death  Ville)-  and  its 
enxirons,  and  in  Ash  Meadows.  Death  Valley 
water  sources  include:  Crape\ine  Spring, 
Scottv's  Castle  Spring,  Mescjuite  Spring,  Day- 
light Spring,  Hole-in-the-\Vall  Spring,  Midway 
Well,   Sto\epipe  Wells,   Salt  Creek,   Nexares 


Department  of  Entoimilog)'.  Caliloriiia  Acadcinv  of  Sciences.  Colden  CJate  Park,  San  FraTic 
Fair  Oaks,  California  9.5628. 


.  (:.ilil()inia94ns   M.iilm.' .iddress  :fiS24  l.uula  Sue  Wav. 


378 


1992] 


Death  \'ai,i.i;y  Hifki,k  Bkktlks 


379 


Springs,  Texas  Spring,  Tra\eriine  Springs,  Emi- 
grant Spring,  Naxel  Spring,  Tnle  Spring, 
Badwater,  Slioi-h"s  Well,  Eagle  Borax  Spring, 
Warm  Spring,  Ibex  Spring,  and  Saratoga  Spring. 
Ash  Meadows  water  sources  include:  Indian 
Spring,  School  Spring.  Dexil's  Hole,  Point  of 
Hocks  Spring,  Jackrahhit  Spring,  Big  Spring, 
and  an  unnamed  spring. 

The  onK'  permanent  water  sources  are  large- 
xolume  springs  on  the  east  side  of  Death  \''alle\' 
and  in  .-Xsh  Meadows,  and  Devils  Hole.  No 
water  flows  froiu  Devil's  Hole,  hut  here  the 
surface  of  the  ground  intersects  the  Indrologic 
head  of  the  groundwater  so  water  is  always 
present  in  the  bottom  of  a  large  crevice.  These 
permanent  .sources  are  all  connected  with  the 
Ash  Meadows  Groiuidwater  Basin. 

Collections 

.\11  of  the  above  water  sources  were  exam- 
ined for  the  presence  of  riffle  beetles.  Where 
possible,  collecting  was  accomplished  with  a 
standard  kick-net.  Howe\er,  manv  of  the  seeps 
and  wells  had  such  lov\'  discharge  and/or  narrow 
width  that  collection  coidd  be  done  onlv  bv 
manual  removal  of  rocks  and  sticks  for  visual 
examination.  Voucher  specimens  for  all  species 
collected  were  deposited  in  the  author's  collec- 
tion at  California  State  Universitv'-Sacramento. 


Results 

()t  the  27  water  sources  examined,  8  were 
found  to  contain  populations  of  elmids  (Table 
1  ).  Steiwhiiis  calicla  Chandler  was  still  resident 
in  Devil's  Hole,  its  t\pe  localitv.  However, 
during  this  survey  two  additional  populations 
v\ere  located  in  nearbv  Indian  Spring  and  F(jint 
of  Rocks  Spring.  La  Rivers  reports  unsuccess- 
fnllv  searching  springs  near  Devil's  Hole  in  an 
attempt  to  locate  additional  populations  (CJhan- 
dler  1949).  It  is  not  known  whether  these  ackli- 
tional  populations  were  missed  or  if  thev  are  the 
result  of  colonization  or  transplantation.  The 
spring  nm  coming  from  Indian  Spring  is  vcn 
narrow  and  de(^plv  incised  into  the  desert  floor, 
making  it  extremelv  inconspicuous. 

MicwcijUocpns  fonnicoklens  Shepard 
occurred  onlv  at  Travertine  Springs  (Shepard 
1990).  Near  the  spring  heads  (a  complex  of 
sev  eral  upvvellings)  and  for  manv  meters  below. 
M.fonnicoidetis  was  the  only  elmid  to  be  found. 
Further  downstream,  though,  it  co-occurs  with 
M.  siiuilis  (Horn).   In  the  lower  third  of  the 


Tahi.i:  1.  Tlie  occurrence  of  rii'lle  beetles  (C^oleoptera: 
Eliiiklac)  ill  water  .sources  of  Deatli\'alIe\Nation;il.VI()iiimient. 


Average 

temp. 

Elexation 

\\'ater  source 

(C) 

(m) 

Species'' 

Death  \alley 

1 .  C;rape\ine  Spring 

25-29 

S20 

2 

2.  Nexare's  Springs 

- 

300 

2 

.).  Traxertine  Springs 

32-.36 

122 

2.3 

4.  Saratoga  Spring 

26-29 

46 

2 

A.sh  Meadows 

5.  inilian  Spiing 

24-30 

705 

1,2 

6.  Devils  Hole 

7.3.5 

1 

7.  Point  of  Hocks  Spring 

705 

1.2 

S.  Big  Spring 

681 

2 

Miiniajll(ici>us    similis.    3    =    MUnirijUoqm 


spring  nm  M.  siinilis  completelv  replaces  M. 
formicoidciis.  Microcifllocpiissiniilis  also  occurs 
in  several  other  springs:  (Grapevine  Spring. 
Nevare's  Springs,  Saratoga  Spring,  Indian 
Spring,  Point  of  Rocks  Spring,  and  Big  Spring. 

All  v\'ater  soiu'ces  inhabited  bv  elmids  v\ere 
located  either  on  the  east  side  (jf  Death  X'allev 
or  in  Ash  Meadows.  \Mth  the  exception  of 
Devil's  Hole,  these  springs  all  exhibit  perma- 
nent flow  of  a  relativelv  large  volume.  Most  of 
the  water  sources  not  inhabited  bv  elmids  are 
low-volume  .seeps  (e.g.,  Davlight  Spring),  sub- 
surface sources  (e.g.,  Shortv's  Well),  or  ])()ole{l 
water  (e.g.,  Badwater). 

(Note  added  after  author  review:  Ricliard 
Zack  [Washington  State  UnivxM'sitv]  has  found 
S.  calida  in  Skruggs  Spring  and  Mexican  Spring 
inAshMea(k)ws|WDS].) 

Discussion 

The  major  factor  linking  those  sjiringsinliab- 
ited  bv  elmids  is  their  association  with  tlu>  .Ash 
Meackms Cyroiuidwater  Basin. This  large v\ater- 
shed  nn(k)ubtedlv  maintains  tlu^  c-onstant  fkm- 
recjuired  bv  elmids.  The  onlx  laige-xolnnie 
spring  not  inhabited  bv  elmids,  jackrabbit 
Spring,  v\'as  pumped  diA  during  a  local  battle 
o\(M-  water  rights. 

.Although  it  mav  at  first  seem  incongruous  to 
find  riffle  beetles  in  a  desert  area,  one  must 
icnieiuber  that  the  regional  desertification  is  a 
rather  recent  event,  geologicallv  and  ecologi- 
callv  speaking.  During  the  .several  Pleistocene 
glacial  periods,  and  perhaps  even  before,  the 
Basin  and  Ranee  Desert  was  far  cooler  and 


380 


Great  Basin  Naturalist 


1\  olunie  52 


wetter.  Tlie  IDeatli  Valley  area  is  thought  then  to 
have  had  a  climate  much  like  the  present-day 
Lake  Mono  area,  240  km  (150  mi)  to  the  north 
(Hildreth  1976).  E\idence  from  the  distribu- 
tions of  fishes  in  the  desert  of  California  and 
Nevada  and  along  the  East  Front  of  the  Sierra 
Ne\ada  suti^iests  that  many  of  the  Pleistocene 
lakes  ovei-flowed  their  basins  and  were  con- 
nected by  extensive  river  systems  (Miller  1946, 
Hubbs  and  Miller  1948,'  Soltz  and  Naiman 
1978).  Thus,  pre-Pleistocene  distributions  ol 
aquatic  organisms  would  ha\e  been  subject  to 
changes  during  the  Pleistocene.  Ultimately, 
these  distributions  were  then  subjected  to  the 
influences  of  the  warmer  and  drier,  current 
interglacial  period.  Present  distributions  are, 
therefore,  the  sum  of  pre-Pleistocene  distribu- 
tions. Pleistocene  dispersals,  and  post- 
Pleistocene  \icariant  strandings. 

Small,  isolated  populations  that  were 
stranded  in  reliable  water  sources  presented 
ideal  situations  for  rapid  evolution,  given  the 
small  gene  pools  and  lack  of  gene  flow  from 
other  populations.  These  factors  ha\'e  been 
responsible  for  the  quick  proliferation  of  pup- 
fish  taxa  in  the  Death  ValUn  area  (Solt/  and 
Naiman  1978).  This  may  also  account  for  the 
speciation  oi  M .  fonuicoicleits,  the  dexelopment 
of  subspecies  in  S.  calicla,  and  the  inter- 
populational  variation  in  M.  siiniUs. 

Stenehnis  calicla  had  been  prexiousK' 
reported  from  Ash  Meadows  in  the  form  of  its 
nominate  subspecies.  A  second  subspecies,  S.  c. 
inoapa  La  Rivers,  occurs  southeast  of  DVNM 
along  the  Muddy  River  in  southeni  Nevada. 
Each  of  the  various  populations  of  M.  siniilis 
exliibits  minor  morphologic  variations,  some 
even  in  tlie  aedeagus.  I  ha\e  vacillated  for  a  long 
tiuie  concerning  the  taxonomic  status  of  these 
disjunct  populations.  Howe\er,  since  the  genus 
needs  revision,  and  because  I  suspect  that  tlie 
variation  is  ecologically  induced,  I  have  chosen 
to  be  conservative  and  not  assign  separate  taxo- 
nomic status  to  any  of  the  populations.  Perhaps 
some  enteiprising  future  student  will  examine 
how  constant  warm  temperatures  influence 
moq3hologic  expression  in  riffle  beetles.  If  so, 
the  springs  of  DVNM  and  the  Basin  and  Range 
Desert  would  offer  an  excellent  natural  experi- 
ment, and  the  numerous  populations  ol  .^/. 
siniilis  in  tho.se  springs  and  spring  runs  would  be 
choice  stud\  material. 

The  elmids  of  DVNM  represent  an  inverte- 
brate analog  of  the  alreadv  well-documented 


evolution  of  pupfish  of  DVNM  (see  Soltz  and 
Naiman  1978).  Microciflloeptis  fonnicoideus  is 
similar  to  Ci/priiiodou  diabolis  in  being  located 
in  only  one  water  source  and  in  being  very 
distinct  from  and  smaller  than  its  congeners. 
Stenehnis  calida  is  similar  to  C.  salinus  and  C. 
milled  in  that  there  are  two  taxa  (subspecies) 
that  inhabit  two  separate  locations  along  a  once 
free-flowing  water  course  (La  Rivers  1949). 
Mierocijlloepus  siniilis  is  similar  to  C.  nevaden- 
sis  in  being  widely  distributed  but  luning  iso- 
lated, somewhat  moqihologicalK  distinct 
populations  throughout  D\'NM  and  surround- 
ing areas. 

Elmids,  like  most  acjuatic  insects,  accom- 
plish dispersal  primariK  In  living  adults.  As  the 
post- Pleistocene  desertification  proceeded, 
water  soiux-es  in  the  D\'NM  area  became 
smaller,  fewer,  and  farther  apart.  A  point  even- 
tually had  to  be  reached  at  which  aerial  dispersal 
became  hazardous.  Mutations  reducing  the 
abilit)'  to  fly  would  then  be  favored;  indeed, 
relatively  rapid  fixation  of  these  mutations  in  the 
population  would  be  expected.  It  is  not  suipris- 
ing,  then,  that  adults  of  all  three  elmids  occur- 
ring in  D\^NM  are  either  apterous  (wingless)  or 
brachvpterous  (with  incompletelv  developed 
wnigs),  and  subse(juentlv  incapable  of  flying. 

Acknowledgments 

I  thank  the  staff  of  Death  Valley  National 
Monument  for  granting  permission  to  collect, 
for  access  to  their  libraiT,  and  for  a  mvTiad  of 
helpful  comments  about  this  remarkable  area. 

LiTER.\TURE  Cited 


CllANDLKH  11.  p.  1949.  A  new  specie.s  oi'  Stviicl mis  from 
Nevada.  Pan-Pacifie  Entomologist  25(3):  133-1.36. 

CoLHUHN,  E.  A.  1980.  Factors  influencing  the  distribution 
and  abundance  of  the  caddistlw  Liiiuwphilis  assiinilis. 
in  Death  Nallev.  Unpul)lisiied  doctoral  tlissertation, 
Uni\('isit\  of  Wisconsin. 

Dk.vcox.  |.  E.  1967.  The  ecolog\-  of  Saratoga  Springs, 
Death  N'allev  National  Monument.  Pages  1-26  in  Stud- 
ies on  the  ecology-  of  Saratoga  Springs,  Death  \';illey 
National  Monument.  Final  report  of  research  accom- 
pli.shed  under  NPS  Contract  No.  14-l()-()4.34-1989. 
Nevada  Southern  Uni\'ersit\-,  Las  \egas  (Universit\ ol 
Nevada  at  Las  \egas). 

.    196S.    Ecological   studies  of  aquatic   habitats  in 

Death  \'allev  National  Monmnent  witli  .special  refer- 
ence to  Saratoga  Springs.  Final  report  of  research 
accomplished  imder  NPS  Contract  No.  14-10-0434- 
19S9.  Ne\ada  Southeni  Uniwrsib.,  LasWgas  (Univer- 
sit\^  of  Nevada  at  Las  Vegas). 


1992] 


Dkviii  \'ai,i,i;y  iiiKKU';  Bkktles 


381 


HlLDKETH.  W.  1976.  Death  N'dlev  gcoloy>.  IX'atli  \'alk'\ 
Natural  I  listorv  Association.  "2  pp. 

Ill  UBS.  C.  L.,  .^M)  H.  R.  MiLLKK  194S.  The  zoolosiical 
e\idence:  correlation  bet\veen  fish  distribution  and 
Indrologic  histors'  in  the  desert  basins  of  western 
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glacial  and  postglacial  times.  Bulletin  of  the  University 
of  Utah  38(20).  Biological  Series  10(7):  17-166. 

Ik  NT,  C.  B.  1975.  Death  \alle\:  geolog\.  ecologw  iux-heol- 
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Ill  NT,  C.  B.,  T  \V.  Robinson,  \V.  A.  Bow  i.ks.and  A.  L. 
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Suney  Professional  Pajier  494-B.  138  pp. 

L\  Ri\  KKS.  I.  1949.  A  new  subspecies  of  Stcnclinis  froiu 
Nexada.  Proc'eedings  of  the  Entomological  Societ\  of 
Washington  51(5):  218-224. 

Mii.i.KR.  R.  R.  1946.  Correlation  between  fish  distribution 
and  Pleistocene  hvdrologv  in  eastern  California  and 


southwestern  Nevatki,  with  a  map  of  the  Pleistocene 

waters.  Journal  of  Geolog\'  .54:  4.3-.53. 
Pai.MKK.  T.  S.  1980.  Place  names  of  the  Death  \alle\  region 

in  CiJifoniia  and  Nevada.  Sagebmsh  Press.  Vlorongo 

X'alley,  California.  SO  pp.  (  Reissue  of  die  1948  printing.) 
SiiK.l'AKD.  W.  D.  1990.  MicwnjUacpus formicoklcus  (Col- 

eoptera:  Elmidae),  a  new  riffle  beetle  from  Death 

N'allev  National  Monument,  California.  Entomological 

News  101(3):  147-1,53. 
Soi.i/..  D.  L..  AND  R.  J.  Nai.man    197S  The  natural  histor\- 

of  native  fi.shes  in  the  Death  \;ille\  Svstem.  Science 

Series  No.  .30.  Natinal  I  listorv  .\lu.seum  of  Los  Angeles 

Coiintv,  Los  Angeles.  76  pp. 
UsiNCKH,  R.  L.  19.56.  Aquatic  Ilemiptera.  Pages  182-228 

in  R.  L.  Usinger,  ed..  Aquatic  in.sects  of  Cdifornia  with 

keys  to  North  .Americiui  genera  and  Califoniia  sjx^cies. 

Universit)  of  Ciilifomia  Press,  Berkele\-.  .508  pp. 

Received  IGJulii  1991 
Accepted  10  September  1992 


Great  Basin  Naturalist  52(4),  pp.  382^384 

SIPHONAPTERA  (FLEAS)  COLLECTED  FROM  SMALL  MAMMALS 
IN  MONTANE  SOUTHERN  UTAH 

James  R.  Kucera    and  Glenn  E.  Haas" 


Kci/  words:  flfds.  SipluiiKiptcni.  I'tali.  ituittiuuils. 

Recent  collections  troni  \arions  small  mam- 
mals of  sontheni  Utah  have  helped  to  elncidate 
the  distribution  of  fleas  (Siphonaptera)  wathin 
the  state.  Of  special  interest  were  fleas  of  mam- 
mals found  in  forested,  high-mountain  areas  of 
the  southernmost  part  of  Utah — an  area  of  com- 
plex topography  containing  habitat  varying  from 
low  desert  to  subalpine  coniferous  forests.  In 
particular,  we  sampled  the  small  mammal  flea 
fauna  of  the  Abajo  Mountains  (San  Juan 
County),  the  La  Sal  Mountains  (Grand/San  Juan 
counties),  and  the  Pine  Valley  Mountains 
(Washington  Count)').  These  ranges  have  been 
sparsel)'  suneved  in  this  respect,  as  evidenced 
by  review  ol  the  seminal  work  of  Stark  (1959). 
After  excluding  22  records  (13  SS.Ti  9  9  )  of 
the  ubicjuitous  deer  mouse  {\e-dActhcca  iva^neri 
(Baker),  which  occurs  in  all  counties  of  Utah 
(Beck  1955),  we  present  and  discuss  the  signif- 
icance of  42  new  records  of  12  species  of  tleas. 
A  parallel  survey  of  fleas  found  in  mammal  nests 
will  be  presented  elsewhere  (Haas  and  Kucera, 
in  preparation). 

Mammal  nomenclature  is  that  of  Hall 
(1981).  However,  designations  of  long-tailed 
\  f)l(^  subspecies  should  be  considered  tentative 
becau.se  of  the  present  confused  state  of  their 
taxonomy.  Mammals  were  collected  with  Sher- 
man li\{'-traps  at  all  localities  e.xcept  Pines 
campground.  Pine  Vallev  Mountains,  and  snap- 
traps  were  used  at  all  threc^  localities  in  the  Pine 
Valley  Mountains  and  at  Oowah  Lake  camp- 
ground. La  Sal  Mountains.  An  asterisk  (") 
denotes  that  the  host  specimen  (or  at  least  one 
host  .specimen)  was  deposited  in  the  manunal 
collection  of  the  Universitv of  Utah  Museum  of 


Natural  Histon'.  Flea  specimens  are  retained  bv 
the  authors. 

Hi/sfricliopsi/Ua  clij)piei  tnincata 
Holland,  1957 

Pcroiui/sciis  nuniiciilatiis  nifiiius.  San  Juan 
Co.:  Abajo  Mts.,  Dalton  Springs  campground, 
2560  m,  8  Septeml^er  1991;  IS.  Microttis 
lou<licaii(Jiis  alficola" .  idem,  1  9  . 

HijstnchopsijUa  occidentaJis  sylvaticus 
Canipos  &  Stark,  1979 

Pcromijscus  hoi/lii  iit(ihciisis° .  Washington 
Co.:  Pine  Vallev  Mts.,  North  Juniper  Park  camp- 
ground, 2122 'm,  10  November  1991;  3  9  9. 
Peronufscu.s  manictdatus  sonoriensis.  Washing- 
ton Co.:  Pine  \'allev  Mts.,  Pines  campground, 
2079  m,  12  June  1991;  19. 

Fleas  ol  the  genus  Hi/strichopsi/lld  are  found 
on  a  \arietv  of  small  mammals  in  mesic  to  moist 
habitats.  These  are  the  first  records  ofH.  dippiei 
tnincata  from  southeastern  Utiili.  Campos  and 
Stark  (1979)  record  H.  o.  si/Ivatictis  from  San 
Juan  Co.,  but  our  records  ol  this  taxon  are  the 
first  from  southwestern  lUah. 

CorrodopsijUa  ctirvata  curvata 
(Rothschikl  1915) 

Sorcx  p<dtistri.\  naviij^dtor  (Baird)°.  Grand 
Co.:  LaSal  Mts.,  meadow  at  Oowtili  Lake,  2769 
m,  15  June  1991;  Id,  19  from  each  of  two 
shrews. 

The  onK  pre\ious  published  records  from 
Utah  of  C.  r.  ciinata  are  both  from  northern 
Utah:  Rich  Co.  (Bear  Lake;  collection  b\'  Stan- 
ford published  bv  Tipton  and  Allretl  [1951: 
107])  and  Salt  Lake  Co.  (Wasatch  Mts.;  Egoscue 
1988).   These    records   are    from    unidentified 


^4655  South  LociLst  Lane  #17,  Salt  Lake  City,  Utalj  S41 17. 
"5.57  California  Street  #7,  Boulder  Citv,  Ne\'acla  89005. 


382 


19921 


Notes 


383 


Sorcx  spp.  Records  of  C.  r.  ohtnsnta  c.\  water 
shrews  are  gi\en  from  Tooele  (^o.  In  Eo;()seiu^ 
(1966,  1988X  Published  records  of  C'.'r'.  cundia 
from  southwestern  states  are  sparse:  Haas  et  al. 
(  1973)  record  it  from  New  Mexico.  Additional 
collecting;  ma\  rexeal  its  presence  in  the  Ahajo 
Mts.,  since  water  shrews  are  knowii  to  occur 
tlierelSchafer  1991). 

Ixhddiitopst/lla  scdilis  wclilis 
i Jordan  c\- Rothschild.  1923) 

Pcro)iujscus  mtinicitlatiis  sonohcusis.  W'ash- 
ington  Co.:  Pine  \'alle\  Mts.,  North  Juniper  Park- 
campground,  21 19  m,  10  \o\(Miil)er  1991;  16. 

This  species  is  known  from  onK  one  area  of 
southern  Utah  (Garfield  (-O..  \ic.  Panguitch, 
Stark  1959),  but  not  pre\iousl\  from  southwest- 
ern Utah.  This  specimen  possesses  5  spines  in 
the  genal  comb,  as  do  other  Utah  specimens 
(Stark  1959,  and  vmpublished  data),  but  charac- 
ters of  the  genitalia  are  clearK-  referable  to  R.  s. 
scctilis  rather  than  R.  s.  goodi  (Hubbard  1941). 
Stark  ( 1959)  noted  that  the  genal  spine  number 
of  Utah  specimens  is  not  consistent  with  the 
original  description  of  R.  s.  scctilis. 

Catallagia  (iccipiciis  Rothschild,  1915 

Feromijscus  nunuciiJatiis  ntfiiuis.  San  fuan 
Co.:  Abajo  Mts.,  Dalton  Springs  campground, 
2560  m,  8  September  1991:  I  6 .  P.  m.  nifiniis. 
idem,  1  6 .  Tamias ,sp.  idem, 2  6  6 .P.m.  nifimis. 
Grand  Co.:  LaSal  Mts.,  Oowah  Lake  camp- 
ground, 2682  lu,  15  June  1991;  1  c5,  2  9  9  . 

These  are  apparentK'  the  first  specific 
records  published  for  that  part  of  Utiili  south  of 
the  Colorado  River.  Beck  ( 1955:  Table  3)  lists  it 
as  (occurring  in  San  fuan  Count\'. 

PcroinijsccpsijUd  sclciiis  (  Rothschild,  1906) 

Pcroiiii/sciis  maniculdlii.s  nijiuus.  San  |uan 
Co.:  ,\bajo  Mts.,  Dalton  Springs  campground, 
2560  m,  8  September  1991;  1  6.  Microfns 
hntgicdiidns  cdticola" .  idem,  1  6 .  M.  /.  alticola. 
idem,  1  9.  M.  1.  kittts.  Washington  Co.:  Pine 
\'allev  Mts.,  Pines  campground,  2079  m,  12  June 
1991';  2  66 A  9. 

This  species  was  not  prexionsK  known  from 
Utali  south  of  the  Colorado  Ri\er.  jolinson  and 
Traub  (1954)  gi\e  a  record  from  Iron  Conntx. 
bordering  Washington  Count\-  to  the  north. 
This  species  is  most  commouK  collected  from 
Microtiis  spp.,  but  also  from  other  small  mam- 
mals sympatric  with  the  voles. 


Pcniiiit/sci)j)si/Ila  hcspcwmijs  adclj)lui 
(Rothschild,  1915) 

Pcronit/scus  lUdnicuUilns  nijiuus.  (irand 
Co.:  LaSal  Mts.,  Oowah  Lake  campgroinid. 
2682  m.  15  June  1991;  1  c5 ,  2  9  9 . 

Johnson  and  Traub  (1954)  record  this  spe- 
cie's from  Beaxer,  Box  Elder,  Millard,  San  |uan. 
and  Washington  counties. 

Opisoddsi/s  kccui  (Baker.  1896) 

Pcwiuifscus  )U(iuicul(itus  souoricusis.  Wash- 
ington Co.:  Pine  N'allex'  Mts.,  N  |uni[)er  Park 
campground,  2119  m,  io  November  1991;  1  9 
from  each  of  three  hosts.  Peromijscus  botjlii 
rowlctji.  idem  (2122  m.),  3  9  9 .  F.  nt.  sonorien- 
sis.  Washington  Co.:  Pine  \alle\-  Mts.,  Pines 
campground,  2079  m.  1 1  June  1991:  1  9  .  P.  ///. 
souoricusis.  idem.  12  June  1991,  1  9 .  P.  //;. 
soiuuicusis.  idem.  1  6  .  Pcroiut/.scus  crinitiis  ssp. 
Washington  Co.:  below  dam  at  Baker  Dam  Res- 
enoir,  foothills  of  Pine  V'allev  Mts.,  1463  m.  12 
.Mav  1991;  1  9. 

No  known  records  of  this  s[)ecics  from  south- 
ern Utah  are  published  other  than  that  of  Hub- 
bard (1947:111,  Garfield  Co.).  Stark  (1959) 
noted  that  this  species  is  collected  onl\-  in  moun- 
tainous areas  or  moist  habitats.  We  luu'e  also 
found  this  to  be  tme  (unpublished  data).  Per- 
oiuif.scus  spp.  are  the  usual  host. 

Malaracus  tclcliiuus  (  Rothschild.  1905) 

Pcnuiu/scus  ))uinicul(itus  souoriensis.  Wash- 
ington Co.:  Pine  \al1ev  Mts.,  N  Juniper  Park 
campground,  2119  in,  10  November  1991;  2 
6  6 .  \  9  .  P.  ui.  sonoricnsis.  idem,  1  6,1  9  .  P. 
)u.  so)ioric)tsis.  idem,  2  9  9.  P.  tn.  .wnorietviis. 
idem.  1  9.  Pcwuujscus  hotflii  roulci/i.  idem,  1 
9. 


Mai. 


(iracus  siuoiuus  \  ordan. 


1925) 


Pcronu/scus  criuitus  ssp.  Washington  Co.: 
below  Baker  Dam  Re.ser\()ir,  foothills  of  Pine 
\alle\  Mts.,  1463  m,  11  Mav  1991,  1  c?,  4  9  9. 
/'.  criuitus  s.sp.  idem,  12  May  1991,  1  c?,  2  9  9. 
P.  criuitus  ssp.  idem,  1  6,4  99.  Peromijscus 
tnici  tnici.  idem.  3  9  9.  P.  t.  truei.  idem,  1  c?,  2 
9  9.   P.   hoijlii   rowlciji.   ickMU.    \    6 .   P.   hotjiii 


clciji.  idc 


9. 


B(xk  (1955:  Tiible  3)  lists  Malaracus 
tclchiuus  as  occurring  in  Washington  Count)'. 
Hubbard  (1947:  200)  gi\es  the  onl\-  specific 
record  from  .southern  Utah  (Garfield  Co.). 
Howe\er,  sexeral  records  of  M.  siuotnus  from 
desert  areas  of  southern  Utah  are  gi\en  b)^  Stark 


384 


Great  Basin  Naturalist 


[N'olume  52 


(1959).  Also,  many  specimens  of  Af.  .sinoiiuis 
from  Pewmyscus  crinitus  were  taken  by  tlu^ 
senior  author  in  Snow  Canyon  (W'asliington  Co.) 
incidental  to  the  search  for  Trauhdla  gnuiclnuiimi 
Ego.scue,  1989.  M.  telchimis  seems  to  be  like  O. 
kceiii  in  being  found  onK  in  nondesert  habitat. 

Me<iaholliris  abantis  (Rothschild,  1905) 

Microiii.s  longicoudus  alticola.  Grand  Co.: 
LaSal  Mts.,  meadow  at  Oowah  Lake,  2676  m, 
14  June  1991;  Ic?,  1  9  .  Pcroini/scus  nianiciilatiis 
nifimis.  idem,  1  9  .  M.  /.  alticola.  idem,  1  9 .  M. 
l' alticola.  idem,  15  June  1991,  M,  19.  A/.  /. 
alticola.  San  Juan  Co.,  Abajo  Mts.,  Dalton  Springs 
campground,  2560  m,  8  September  1991,  1  6 . 

Beck  (1955:  Table  3)  listed  this  species  as 
occurring  in  Bea\er,  Iron,  Sexier,  and  \\'a>ne 
counties.  This  was  apparentK'  oxerlooked  b\ 
Stark  ( 1959:  196),  who  stated,  ^This  flea  appears 
confined  to  the  northern  half  of  the  state." 
Egoscue  (1988)  reported  collecting  one  male 
specimen  from  a  pika  at  Johnson  Resenoir, 
Sevier  Count)',  in  south  central  Utah.  The  dis- 
tribution map  of  Haddow  et  al.  (1983:  Map  76) 
indicates  a  locality  record  in  that  same  region  of 
Utah.  Our  records  are  the  first  for  southeastern 
Utah.  Mcgabothris  abantis  is  usuall)'  found  on 
\arious  species  o^ Microtus. 

EuDiolpianus  cuinolpi  aiiicricanus 
(Hubbard.  1950) 

Tamias  sp.  San  Juan  Co.:  Abajo  Mts.,  Dalton 
Springs  campground,  2560  m,  8  September 
1991^2  9  9. 

These  specimens  seem  closer  to  E.  c.  aiuer- 
icaniis  than  to  E.  c.  eutnolpi  recorded  by  Beck 
(1955,  then  in  the  genus  Monopsijllus) .  Several 
of  the  t)pe  specimens  were  collected  in  San 
Juan  Count)-  (Hubbard  1950).  Johnson  (1961) 
indicates  that  intergradation  between  E.  c.  antcri- 
caiius  and  E.  c.  cuuiolpi  occurs  in  the  conutx. 

In  snmmaiy,  the  significant  findings  among 
64  collection  records  of  13  species  of  fleas  are  as 
foHows:  the  first  records  south  of  (he  (Colorado 
Rix'er  in  southeastcin  I'tali  (or  Hi/strichopsi/lla 
(lippici  tniiicata,  Corrodopsi/lla  c.  cunaia.  Pcr- 
oDiijscopsijlla  svlcnis,  and  Megaboihris  abantis; 
and  the  first  in  Washington  Count);  southwest- 
ern Utah,  for  //.  occidentalis  si/lvaticns,  Rhacl- 
inopst/lla  s.  scctilis,  P.  selenis,  luid  Opi.soda.sys  kccni. 


Acknowledgments 

We  thank  H.  Egoscue  and  C.  Pritchett  for 
their  review  of  the  manuscript;  and  E.  Rickart, 
Universitv  of  Utah  Museum  of  Natural  History, 
for  assistance  in  identifsing  some  Pcroint/scus 
specimens. 

Literature  Cited 

Bkck  D  E.  1955.  DistrihutioiiaLstiiclies  i)f' parasitic  arthro- 
pods in  Utah,  determined  as  actual  luid  potential  \ec- 
tors  of  Rock\'  Mountain  spotted  fever  and  plague,  with 
notes  on  vector-host  relationships.  Brighani  Young 
Universitx'  Science  Bulletin.  Biologic;il  Series  1  (D. 

Ca.mpos.  E.  a..  .\sd  H.  E.  Stark.  1979.  A  revaluation  [sic] 
of  the  HijstrichopsijUa  occidentalis  group,  with  descrip- 
tion of  a  new  subspecies  (Siphonaptera: 
Hvstrichop.s\llidae).  Journal  of  Medical  Entomology 
1.5:  431-444'. 

EcoscLF,,  H.  E.  1966.  New  ami  additional  host-flea  associ- 
ations and  distributional  records  of  fleas  from  Utah. 
Great  Basin  Natrn-alist  26:  71-75. 

.  19SS.  Noteworthv  flea  records  from  Utah,  Ne\ada, 

and  Oregon.  Great  Basin  Naturalist  48:  530-532. 

.    19S9.   A  new  species  of  the  Genus   Tmnhclhi 

(Siphonaptera:     Ceratoph\llidae).     Bulletin    of    the 
Southern  Ctilifornia  Academv  of  Sciences  SS:  131-134. 

R\AS,  G.  E.,  R.  P.  Mahtin.  M'.  Swickard.  and  B.  E. 
Miller  1973.  Siphonaptera-mammiJ  relationships  in 
northcentral  New  Me>dc().  [ournal  of  Medical  Ento- 
niolog\- 10:  281-289. 

Haddow.  ].,  R.  Traub,  and  M.  Rothschild  1983.  Dis- 
tribution of  ceratoph\  Hid  fleas  and  notes  on  their  hosts. 
Pages  42-163  in  R.  TraTib,  .M.  Rothschild,  ;uid  J.  F. 
Haddow,  The  Rothschild  collection  of  fleas — the 
Ceratoplnllidae;  ke\s  to  die  genera  and  host  relation- 
ships with  notes  on  their  e\olution,  zoogeograplu'  and 
medical  importance.  288  pp.  [Privateh  published.] 

Rall,  E.  R.  1981.  The  mammalsof  North  America.  2nded. 
2  \olunies.  John  W'ilev  &  Sons,  Inc..  Ne\v  York.  1181  pp. 

Hubbard,  G.  A.  1947.  Fleas  of  Western  Nordi  America. 
Iowa  State  Gollege  Press.  533  pp. 

.  1950.  A  pictorial  rexiew  of  the  North  .Americiui 

chipmunk  fleas.  Entoniologic;il  News  60:  25.3-261. 

Johnson.  P.  T.  1961.  .\  re\ision  of  the  species  oi  Mono- 
pstjlhis  Kolenati  in  North  America  (Siphonaptera. 
Geratophvllidae).  Technical  Bulletin  No.  1227,  Agri- 
cultural ReseLUX'h  Sel^ice,  U.S.  Depiutment  of  .Agiicul- 
tiue.  69  pp. 

JOHNSON  P.  T.  AND  R.  Tr.M  R  1954.  Rexision  of  die  flea 
genus  Pcroini/.scop.si/lla.  Smithsonian  Miscellaneous 
Gollections.  \'ol.  123.  No.  4.  68  pp. 

S(  HAFKR,  T.  S.  1991.  Mammdsof  die  Abajo  Mountains,  an 
isolated  nioimtain  range  in  San  Juan  Gountw  southeast- 
em  Utah.  Occasional  Papers  No.  137.  The  Museum. 
Texas  Tech  University,  Lubbock. 

.Stark.  II.  E.  19.59.  The  Siphonaptera  of  Utali.  U.S.  Dep;ut- 
ment  of  Health,  Education  and  \\'elfare.  Gommunica- 
ble  Di.sease  Genter,  Atlanta,  CJeorgia.  2.39  pp. 

Thton  \'.  J.,  AND  D.  M.  .\llrkd  1951.  New  distribution 
records  of  Utah  Siphonaptera  with  the  description  of  a 
new  species  of  Mc^ai-throfilosstis  Jordan  and  Rofli- 
.schild,  1915.  Great  Basin  Naturalist  11:  10.5-114. 

Received  22  May  1992 
.\ccej)ted  2  November  1992 


(iix'iit  Basin  Naturalist  52(4  K  pp.  385-^386 

NOTES  OX  SPIDER  (THERIDIIDAE.  SALTICIDAE)  PRED ATIOX 

OF  THE  1IAR\ESTER  AXT,  POGOXOMYRMEX  SALIXUS  OLSEX 

(HYMENOPTERA:  FORMICIDAE:  MYRMICIXAE), 

AND  A  POSSIBLE  PARASITOID  FLY  (CIILOROPIDAE) 


William  II.  Clark'  and  Paul  Iv  Bl( 


Kl'i/  uorcls:  Poiiononnnnex  saliniis.  luirvcstcrdiil'-^.  Eunopis  t( 


.\\sti(iis.  spider  predators.  Inccrtclla.  parasite. 


Spiders  are  known  predators  of  ants.  Pres- 
sure exerted  bv  consistent  spider  predation  can 
alter  the  behaxior  of  ant  colonies  (MacKax 
1982)  and  ma\'  be  a  selectixe  pressure  contrib- 
utintj;  to  the  seed-hanesting  behaxior  of 
P(><^oiii)iiii/n)wx  (MacKax'  and  MacKax*  1984). 
We  obsened  the  spider  Eim/opis  fontiosa 
Banks  (Araneae:  Tlieridiidae)  capture  and 
transport  xx'orkers  of  tlie  lianester  ant 
[Pogotiomijrmex  salinus  Olsen  [Hxiiienoptera: 
Fonnjcidae,  MxTmicinae])  in  soutlieasteni 
Idalio.  .Xdditional  obserx'ations  rexealed  a  crab 
spider  of  the  genus  Xijsticus  prexing  on  P. 
salimis  and  the  presence  of  a  clikjropid  fix' 
(IiicerteUa)  that  max-  liaxe  been  parasitizing  the 
moribund  prev  subdued  bx'  the  spider. 

Study  Sitk 

One  collection  site  is  located  along  Road 
T-2()  (Butte  Count);  T4N,  R31E,  S6)"on  the 
Idaho  National  Enxironmental  Research  Park 
(INERP)  in  the  cold  desert  of  southeastern 
Idaho.  The  second  set  of  observations  xvas  made 
on  the  INERP  (Clark  Countx;  T7N,  R31E,  S34. 
along  Highxx'ax'  28).  X'oucher  specimens  of  all 
species  have  been  deposited  at  the  Oniia  ]. 
Smith  Museum  of  Natural  Histon,  Albertson 
College  of  Idaho,  Caldxxell,  Idali()'83605  USA 
(CIDA). 

Results  .am)  Discussion 

On  3  July  1988,  1020  h.  at  the  Butt(>  ( :()uutx 
collection  .site  xx'e  collected  a  single  indixidual  of 
Eiirijopis  fontiosa  Banks  (Araneae:  TluMidiitkic) 


that  x\as  earrxing  a  worker  of  Pofj^otionu/niicx 
sdliiuis  Olsen  (Hxnienoptera:  Formicidae, 
Mxrmicinae)  across  a  large  area  of  basalt  rock. 
The  ants  xxere  actixelx'  foraging  in  the  area.  The 
air  temperature  (shaded)  xvas  31  C]  and  the  soil 
siu-face  (in  the  sun)  xvas  39.5  C.  No  other  spiders 
of  this  species  xvere  encountered.  Prev  capture 
xx'as  not  obsened. 

On  31  August  1991  at  1725  li  at  tlic  Clark 
Counts-  site  xxe  obsened  a  ciab  spider  of  the 
genus  Xi/sticiis  pre\ing  on  P.  sal  inns  about  20  cm 
from  the  ant  nest  entrance.  The  ants  xx'ere  still 
actixelx-  foraging  at  this  time.  One  spider  xxas 
riding  on  the  ant  in  the  shelter  of  an  isolated 
clump  of  Indian  ricegrass  [On/zopsis  lii/ttie- 
nuides)  at  the  edge  of  the  ant  moimd.  Tlu^  ant 
xvas  initiallv  vew  actixe.  xxalking  around  an  old 
grass  stem,  xxhile  the  spider  made  periodic 
attacks  on  the  ant.  As  time  progres.scd,  inxolim- 
tan- spasms  in  th(^  ant  increased.  The  spider  xx'as 
generallx-  oriented  toxx'ard  the  posterior  of  the 
ant,  biting  it  at  the  base  of  the  petiole.  Some- 
times the  spider  xxas  peipendicular  to  the  ant. 
holding  on  to  the  ant  xxith  onlx'  its  mandibles. 
.\fter  fixe  minutes  the  ant  fell  onto  its  side  and 
moxements  sloxxcd.  .\t  1740  h  onlx  its  antennae 
xx'ere  mox-ing  slightlx,  and  a  minute  later  the 
spider  moxed  the  ant  under  a  small  stick.  Tx\-o 
small  flies  approached  die  ant  and  one  flexx-  onto 
its  head.  Occasional  moxements  (jerks)  of  the 
ants  legs  xxere  obsened  at  1751  h.  At  this  time 
x\-e  collected  the  spider,  the  ant,  and  one  of  the 
Hies  (WHO  #9170).  The  fly  is  a  female  Incertelki 
I  Diptera:  Chloropidae)  and  may  represent  an 
undescribed  species.  Broxvn  and  Feener  (1991) 


Orma  J.  Smith  Must- uiii  of  Natural  I  liston .  Alhert.son  College  of  Ulalio,  Caldwell.  I<lalio  S.36().5  \Jii.\:  and  Departnieiit  of  Plant.  Soil,  and  Entomological 
Sciences.  Universit)'  of  Idaho.  Mo.scow.  Idaho  S.3S43  USA. 

"  Departinenl  of  Plant.  Soil,  and  Entomological  Sciences,  Universlt\-  of  Idiilio.  Moscow,  Idaho  8aS4.3  US.\. 


385 


386 


Great  Basin  Naturalist 


[\V)liuiie  52 


have  found  the  plunid  Ap(>(cf)li(ili(s  paraponerae 
selecti\eK'  parasitizing  1 1  loril  )uncl  workers  oi  Para- 
ponera  clavata.  It  may  be  that  these  InceiicUa 
flies  are  seeking  a  similar  host  and  opportunis- 
tically exploiting  the  spider  prey.  The  flies  were 
not  obsened  to  interact  with  li\'ing,  active  ants. 

At  1740  h  we  noticed  a  second  spider,  E. 
formosa,  on  tlu^  same  ant  mound.  This  spider 
oriented  uphill  on  the  side  ot  the  mound,  facing 
the  ant  nest  entrance.  At  1742  h  an  ant  walked 
over  and  slightK*  past  the  spider,  apparentlv 
failing  to  recognize  the  predators  presence.  The 
spider  remained  motionless  as  the  ant  passed, 
then  spim  around  and  mounted  the  ants  gaster. 
The  spider  released  the  ant  and  moved  to  face 
it.  The  ant  began  convxdsing  at  this  time,  while 
the  spider  sat  1  cm  away  from  the  ant  (facing 
awav  from  the  ant).  Bv  1745  h  no  motion  was 
obsened  in  the  ant  and  at  1746  h  the  spider 
climbed  onto  the  ant.  The  ant  was  on  its  side 
with  the  spider  on  top  facing  the  gaster.  A  fly 
similar  to  those  mentioned  aboxe  moved  onto 
the  head  of  the  ant.  At  1747  h  the  spider  was 
draggincr  the  ant  across  the  moimd  usingr  a  web 
sling,  as  previouslv  described  b\'  Porter  and 
Eastmond  ( 1982)  for  the  spider  E.  coki  in  south- 
eastern Idaho.  The  spider  dragged  the  ant  to  the 
edge  of  the  mound  and  into  the  grass  clump 
mentioned  earlier.  Several  other  worker  ants 
were  obsened  strung  up  in  the  grass  clumps.  At 
this  point  we  collected  the  spider  (WIIC  #9171 ). 

The  spider  genus  Eunjopis  is  known  to  pre\' 
on  ants  (Le\i  1954,  Carico  1978),  including  har- 
vester ants  of  the  genus  Pogonomijnnex  in 
North  America  (MacKay  1982,  Porter  and  East- 
mond 1982).  MacKay  (1982)  has  reported 
PJ.  ralifoniira  previngonP.  rii^osiis  in  southern 
California. 

Prey  of  E.  fonnosa  lias  not  pre\"iousl\"  been 
reported  (Levi  1954),  nor  has  the  spider  been 
reported  from  the  INERP  (Levi  1954,  Allred 
1969).  Levi  (1954)  gives  the  distribution  of  the 
species  over  most  of  kkdio  except  for  the  south- 
western comer,  so  its  presence  here  was  ex- 
pected. Allred  ( 1 969)  reported  a  related  species, 
Eiinjopissvriptipcs  Banks,  from  the  southeastern 
border  of  INERP  during  July.  Ponono»ii/n)iex 
salirms  is  tlie  dominant  .seed-luuvesting  ant  on  tlie 
INERP,  occurring  in  almost  all  of  its  plant  commu- 
nities (Blom  et  d'.  1991). 

Porter  and  Ea.stmond  ( 1982)  found  Ejinjop- 
sis  coki  Lc\i  to  be  a  comuion  preckitor  of 
Po'^oinxntjnncx  ouijhcci  {^P.  .saliniis)  in  south- 
eastern Idaho  during  July  and  August.  These 


small  gra\"  spiders  capture  ants  on  their  mounds 
and  drag  them  awa\'  bv  a  web  sling  attached  to 
the  ant  and  to  the  tip  of  the  spider's  abdomen. 
Ell rif apsis  fonnosa  is  found  from  central  Cali- 
fornia north  to  British  Columbia  and  east  to 
Wyoming  (Levi  1954).  E.  foiDiosa  mav  also  be 
an  important  predator  of  P.  salinus  at  this  site 
and  of  Pooonomijnnex  species  in  the  western 
United  States.  The  relatively  greater  precision 
and  speed  with  wliich  Eunjopsis  subdued  and 
transported  the  P.  salinus  prev  suggests  an 
established  predator-pre\'  relationship. 

ACKNOWLE  ])G  M  E  NTS 

This  work  was  conducted  under  the  IN  EL 
Radioecologx'  and  Ecology'  Programs  sponsored 
bv  the  Office  of  Health  and  Environmental 
Research,  and  the  Division  of  Waste  Products 
through  the  Fuel  Reprocessing  and  Waste  Man- 
agement Division,  United  States  Department  of 
Energ)'.  O.  D.  Markham,  T  D.  Revnolds,  and 
y.  B.  Johnson  have  provided  assistance.  ]. 
McCaffrev  and  H.  W.  Levi  provided  spider 
determinations.  C.  W.  Sabrosky  identified  the 
Inceriella  specimen  and  B.  V.  Brovvm  assisted. 
This  paper  is  published  as  Idaho  Agriculture 
Experiment  Station  Paper  No.  91767. 

Literature  Cited 

Ai.i.HED.  D.  M.  1969.  Spiders  of  the  National  Reactor  Test- 
ing Station.  Great  Basin  Naturdist  29:  10.5-108. 

Bl.OM  R  E.,  \V.  H.  ClARK  A\D  J.  B.  JOHNSON  1991. 
Golonv  densities  ot  the  seed  haivesting  ant 
P()g(>n()ini/nuex.sali)Uis  (Hviiienoptera:  Formicidae)  in 
se\en  plant  communities  on  the  Idalio  National  Engi- 
neering Laboraton'.  [onrnal  of  the  Idaho  Acadenn  of 
Science  27:  2(S-36. 

Bhown.  B.  v.,  and  D.  H.  Fekner.  Jr  1991.  Behavior  and 
host  location  cues  of  Apocctyiuilns  paraponcrac  (Dip- 
tera:  Phoridae),  a  parasitoid  of  the  giant  tropical  ant, 
Vdraponcra  clavata  ( Ihnienoptera:  Formicidae). 
Biotropica  23:  182-187. 

Carico,  J.  E.  1978.  Predaton- behavior  in  Eiin/opis  fuiicris 
(Hentz)  {Ar;mea:  Theridiidael  and  the  evohitionan 
significance  of  web  reduction.  Svmposium  of  the  Zoo- 
logical Societx  of  London  42:  .51-.58. 

I.l'A  I  II.  \V.  1954.  Spiders  of  the  genus  Eiinfopsis  from 
North  and  Central  America.  American  Museum 
Novitates  1666:  1-lS. 

MacKav,  W.  P  1982.  The  effect  of  predation  t)f  western 
widow  spiders  (Araneae:  Theiidiidae)  on  hanesterants 
( Hymenoptera:  Formicidae).  Oecologia53:  406—111. 

M  \(  Kay.  W.  R,  an  d  E.  E.  Mu:Kay  1984.  \\'h\'  do  haivester 
ants  store  .seeds  in  their  nests?  Sociobiologv  9:  31—47. 

PoRTKR,  S.  D.,  AND  D.  A.  Eastmond,  1982.  Eiinjopis  coki 
(Tlieridiidae),  a  spider  that  prevs  on  Po^oiicmi/nm'x 
ants.  Jounial  oi  .Arachnologv  10:  27.5-277. 

Received  6  Fchntan/  1992 
Accepted  10  September  1992 


THE 

GREAT   BASIN 


NATURAUST 


INDEX 


VOLUME     52     -     1992 


BRIGHAM     YOUNG     UNIVERSITY 


Great  Basin  Naturalist  52(4),  1992,  pp.  38.S-391 


INDEX 

Volume  52—1992 

Author  Index 


Acker,  Steven  A.,  284 

Agenbroad,  Larrv  D.,  59 

Allred,  Kelly  W., '41 

Anderson,  Stanley  H.,  139,  253 

Arnow,  Lois  A.,  95 

Arnovv,  Ted,  95 

Austin,  Dennis  D.,  352,  364 

Beck,  Reldon  R,  300 
Bernatas,  Susan,  335 
Blackburn,  Wilbert  H.,  237 
Block,  William  M.,  328 
Bloni,  Paul  E.,  385 
Boe,  Edward,  290 
Bonhani,  Charles  D.,  174 
Burge,  Howard  L.,  216 

Callahan,  J.  R.,  262 
Clark,  William  H.,  385 
CcMiistock,  fonathan  R,  195 
Cooke,  Lynn  A.,  288 
Cottrell,  Thomas  R..  174 
Cronquist,  Arthur,  75 
Cushing.  C.  E!.,  11 

Davis,  Russell,  262 
Deleray,  Mark  A.,  344 

Ehleringer,  James  R.,  95,  195 
Elias,  Scott  A.,59 
Evans,  R.  R,  29 

Flinders,  Jerran  T.,  25 
Furniss,  Malcolm  M.,  373 

Caines,  W  L.,  1 1 
Gill,  Ayesha,  E.,  155 

Hems,  Glenn  E.,  382 
Hcill,  Linnea  S.,  328 
Haws,  B.  A.,  160 
HoN'ingh,  Peter,  278 
Hubert,  Wayne  A.,  253 

Jehl,  Joseph  R.,  Jr.,  328 
Johansen,  Jeffrev  R.,  131 
John.son,  James  B.,  373 
Johnston,  N.  Paul,  25 

Ka\,  Charles  E.,  290 
Kaya,  C^iilvin  M.,  344 
Kitchen,  Stanlev  G.,  53 
Knapp,  Paul  A.,'  149 


Kucera,  James  R.,  382 
Kucera,  Thomas  E.,  122 
Kudo,  J..  29 

Launchbaugh,  Karen  L.,  321 
Leidy,  Robert  A.,  68 
Lindzey,  Frederick  C.  232 

Miirks,  Jeffrey  Shaw,  166 
McArthur,  E.  Durant,  1 
McNult^',  Ining  R.,  95 
Mead,  Jim  I.,  59 
Meyer,  Susan  E.,  53 
Miller,  Gary  C,  357 
Morrison,  Michael  L.,  328 
Moselev,  Robert  K.,  335 
Munay  Leigh  W.,  300 

Negus,  Norman  C,  95 
Nielsou,  M.W.,  160 
Norling,  Bradley  S.,  253 

Palmquist,  Debra  E.,  313 
Pendleton,  Rosemaiv  L.,  293 
Pickering,  Russell,  309 
Pieper,  Rex  D.,  300 

Quinnev,  Dana  L.,  269 

Rasmussen,  G.  Allen,  185 
Roberson,  Jay  A.,  25 
Roche,  Ben  F,  Jr.,  185 
Roche,  Cindv  Talbott,  185 
Rumble,  Mark  A.,  139 
Rushforth,  Samuel  R.,  131 

Saab,  \'ictoria  Aiui,  166 
Scoppettone,  G.  Gan;  216 
Sexille,  Robert  S.,  3()9 
Sheelev,  Douglas  C;.,  226 
Shepard,  \\'illiaiu  D.,  378 
Shields.  \\'es,  3(>4 
Shiozawa,  D.  K.,  29 
Simanton,  J.  Roger,  237 
Slaugh,  Bartel  f.,  25 
Smith,  Bruce  N.,  93 
Smith.  LorcMi  M.,  226 
Smith.  S,  D..  11 
Snvder,  Warren  D.,  357 
Stanton,  Nanc\' L.,  309 
Swiecki,  Steven  R..  288 


388 


19921 


Index 


389 


Taxlor.  Daniel  M.,  179 
Thomas,  Diane  M.,  309 
Torgersen,  Torolf  R.,  373 
Trost,  Chiu-lesII..  179 
Tuttle,  Peter  L..  216 
Tyser,  Robin  W'.,  189 

Uresk,  Daniel  W'.,  35 

Unless.  Philip  J..  321.  352.  364 

\an  Sickle,  Walter  D..  232 
V'icken.  Robert  K..  )r.,  145 
\'(K)rhees,  Marmierite  E..  35 


Wagstaff.  Fred  J..  293 
Wansi,  Tchouiissi,  300 
Welch,  Bruce  L.,  293 
VVeltz,  Mark  A.,  237 
Westcott,  Richard  L.,  373 
Wester,  Da\id  B.,  226 
Williams,  R.  N.,  29 
Wood.  Stephen  L.,  78,  89 
Woodward,  S.  R.,  29 

Yeiirslev,  Kurtis  H.,  131 
Yen.sen,  Eric,  155,  269 
Young,  James  A.,  245,  313 


Keyword  Index 


age-growth,  216 
alien  flora,  189 
alpine  \asculiu-  flora,  335 
aiiiilwsis 

f'actoi-.  174 

fecal,  3(X) 

dietan-,  269 

microhistological .  300 
Anas  acuta,  226 
annual  grass,  245 
Aplanusiclla,  160 
apple  trees,  352 
aquatic  habitat,  278 
arid  lands,  149 
Aristida.  41 
Artemisia 

nova,  313 

fiidcntata.  174 

tiidcntiita  tridcntata.  2S4 

tridcntata  wyoinin^ensis.  284 
arthropods,  59 
avifauna,  278 

bark  beetles,  78 
beardtongue,  53 
Beaver  Dam  Creek,  [Utali],  131 
behaxior,  25 

foraging,  293 
benthos,  11 
big  sagebrush,  284 
biochemical  differentiation,  155 
biogeography,  262 
biomtiss,  313 

leaf.  237 
birds,  278 

migrating,  179 
black  sagebrush,  313 
h()d\" 

condition,  226 

size,  216 
browse,  293 
bumblebees,  145 


bunchgrass,  284 
burrow-  stucture,  288 

California,  41,  122 

Death  \'alle\',  378 

Mono  Lake,'  328 
caribou,  321 
caves,  59 

Ccntaiirea  lirt^ata  ssp.  scjuarrosa,  185 
cerxid,  321 

checking  stations,  364 
ch  romatographv,  1 74 
chukar,  25 

reai'ing,  25 
Cicadellidae,  160 
cold  desert,  11,  195 
Coleoptera,  11,78,89,378 
colonization,  328 
colonizing  .species,  245 
Colorado,  174,  357 
Colorado  Plateau,  59,  195 
Columbian  Shaip-tailed  Grouse,  166 
competitixe  release,  68 
Cottonwood,  357 
cutthroat  trout,  29 
Cijnomys  lencunts,  288 

Death  Vallev,  [California],  378 
deer,  321 

damage  exaluation,  352 

management,  364 

mule,  122,  290,  352,  364 

white-tciiled,  290 
depredation,  352 
desert,  278 

soil  formation,  313 

streams,  131 
desertification,  378 
diatoms,  131 
dietarx  anaKsis,  269 
Diptera,  11 
dispersal,  262 
distribution(s),  160,  378 


390 


Great  Basin  Naturalist 


[y^ 


oluiiie  oz 


distuihance,  253 
DNA  sequencing.  29 
drought  response,  237 

Eiiiwria,  309 
eleetivities.  68 
electrophoresis,  155 
elk.  321 
Elniidae,  37(S 
Ephenieroi")tera,  1 1 
Ethco.stoina  /i/gn///i,  6S 
Eun/opsis fonnosa ,  385 
e\'olution,  378 

(actor  anaKsis.  174 
faunal  list,  373 
fecal  analysis,  300 
fecundit\',  216 
Felix  concolor,  232 
fish,  .344 
(leas,  382 
flora 

alien,  189 

alpine  \ascular.  335 

riUT,  335 
floristics,  41 
llowei'  colors,  145 
food  habits,  216,  269 
foraging  beha\ior,  293 
forest  management,  139 
fruit  trees,  352 
functional  groups.  1 1 

ghost  towns.  149 

Glacier  Park,  [Montana],  189 

Grand  Ganvon.  [Arizona],  59 

grassland,  95 

grayling,  344 

grcizing,  35,  245 

Great  Basin,  195,  278 

De.sei-t,  149 
green  ash,  35 
ground  squirrels,  155,  269 
Grits  ct/nadciisis.  253 

habitat 

a(|uatic.  278 

ck'scriptions,  139 

selection,  139,  253 

summer  h.  characteristics,  166 

use,  179,216 
hanester  ants.  385 
hibemaculum,  288 
hosts,  160 
hummingbirds,  145 
hunter  opinions,  364 

Idaho,  166,  269,  335 

Kane  Lake  Girque,  335 
Pioneer  Mountains,  335 

Idaho  ground  scjuirrel,  155 

imprinting,  25 

hurHrlla.  385 


induced  dormancN,  53 
Insecta,  378 
Intermountain  West,  95 
interspecific  h\'bridi/.ation,  290 
inyentoiy,  357 
iiTigation  resenoirs,  179 
islands,  328 

Kane  Lake  Girque,  [Idaho],  335 

lake.  344 

land  bridge,  328 

leaf  area  index,  237 

leaf  biomass,  237 

leaflioppers.  160 

Lcpiis  califonticiis,  300 

life  histon',  216 

long-term  site  degradation.  284 

mammals.  382 
management,  166 

deer,  364 

forest,  139 
medusahead,  245 
Mclca^ris  oallopaio,  139 
MeiTiam's  Wild  Turke\s,  139 
microhabitat  use,  68 
microhistological  anaKsis,  300 
Microius.  262 

iiioiitdiiiis.  328 
migrating  birds,  179 
migration,  122,  344 
Miiiiiiliis.  145 
MiXipa  coiiarcd.  216 
Moapa  dace,  216 
Mono  Lake,  [Galifornia],  328 
Montana 

Glacier  Park,  189 
moiphological  specializations,  68 
Muddy  Ri\er,  Ne\ada.  216 
mudflats,  179 

mule  deer,  122,  290,  352,  364 
mushroom,  321 
nncophagw  321 

Nebraska 

Platte  Ri\er,  253 
nest,  288 
Ne\ada,  216 

Muddy  River,  216 
new  genus,  160 
new  species,  160 
niche  breailth,  68 
nomenclature,  75.  78.  89 
Northern  Pintails.  226 
nutrients.  293 
nutrition.  321 

oak-majile.  95 
Ocloccilciis 

hcinioiius.  122.290,  293 

viriiiiiidnii.s.  290 
Odonata.  1  1 


19921 


Index 


391 


OiicDrJii/iirJiits.  29 
orchards.  352 
Oregon.  2<S4.  373 

l^iliiier  pen.stenion,  53 

parasite.  3S5 

])ai1  ridge.  25 

peaeli  trees.  352 

FcnstcDioii  pdlmcri.  53 

PcrcicJac.  6H 

I'croiiii/.sciis  iiuiiiiciil(/tiis,  328 

plienologx,  195 

Pioneer  Mountains,  [Idaho],  335 

plant  adaptation,  95,  195 

Platte  Rixer,  [Nebraska],  253 

Plat>podidae.  78.  89 

Pleeoptera.  1 1 

Pleistocene,  262 

Pooonomijnnex  saliniis,  385 

|)ollinator  preferences,  145 

poK merase  chain  reaction,  29 

poKsporocystic  coccidia.  309 

Populus  spp.,  357 

predictive  models,  226 

piexalence,  309 

pr()dncti\it\".  11 

propa<j;ation.  25 

ri-iiiiiis  I  in^iiiidiKi,  35 

(,)uaternaiA.  59 
{)n(rcii.s  '^(iinhclii  293 
(|iiestionnaires.  364 

radio-telenietn,  122 

rangelaiid  weeds,  185 

rare  ilora,  335 

Red  Butte  Canyon,  [Utah],  95 

reproduction  l)iolog\;  216 

Research  Natural  Area.  95 

riparian,  357 

eco](jg\',  95 
rixcr  roosts,  253 
liosidae.  75 
runiinanl.  321 

sagebrush.  174 
'  big.  284 

black.  313 
sahnit\'.  195 
sahnonids,  344 
sandbars,  253 
Sandhill  Crane,  253 
Scolytidae,  78,  89,  313 
seed 

bank.  53 

germination.  53 
sequencing,  29 
se.\  differences,  122 
sheep,  185 
sliorebirds.  179 
shrub{s).  35 

succession,  31 


Siphonaptera.  382 
soil.  174 

compaction.  149 

nitrate.  313 

organic  carbon.  284 

organic  matter.  284 

recoN'cn.  149 
speciation.  145 
Spennopliilii.s,  155 

hnaiiu'tts,  155 

towiiscndii,  269 

tridcrcinliiwatns,  309 
spider  predators,  385 
spring-streams,  1 1 
squarrose  knapweed,  185 
state  records,  335 
stream,  344 

summer  habitat  characteristics.  166 
suni\a].  25 
Sipnpliorictirpos  occidcitlalis,  35 

Tdoiidthcniiit  cdput-mcdiisdc.  245 

taxonoiuN.  75.  78.  155 

Texas.  226 

Thi/Didlhis  drcticiis,  344 

track  siuAey,  232 

Trichoptera,  11 

ti'ophic  levels,  1 1 

Tipiipatun-luis  pliasidnclhis  (■ohiinhidiiiis.  166 

Utah,  131,  232,  382 

Beaver  Dam  Creek.  131 

Red  Butte  Camon,  95 
utilization.  293 

\  icariance.  262 
\ole.  262 

water 

depth.  253 

drawdown  1,  179 

stress,  195 
waterfall,  344 
waterfowl,  226 
weather,  122 
weed  dispersal,  185 
wetlands,  278 
white-tailed  deer,  290 
wildfire(s),  245,  284 
wildlife 

habitat,  357 

methods,  364 

technicjues.  364 
winter,  293 

browsing,  352 
Wisconsin  glacial.  59 
wooded  draws.  35 
wool.  185 
\\\()ming.  290 

Xtjsticus,  385 


392  Great  Basin  Naturalist  [Volume  52 

Table  of  Contents 
Volume  52 

No.  1— March  1992 
Articles 

In  memoriam— A.  Perry  Plummer  (1911-1991):  teacher,  naturalist,  range  scientist 


E.  Durant  McArthur  1 

Secondary  production  estimates  of  benthic  insects  in  three  cold  desert  streams 

W.  L.  Gaines,  C.  E.  Gushing,  and  S.  D.  Smith  11 

Effect  of  rearing  method  on  chukar  survival Bartel  T.  Slaugh,  Jerran  T.  Flinders, 

Jay  A.  Roberson,  and  N.  Paul  Johnston  25 

DNA  extraction  from  preserved  trout  tissues D.  K.  Shiozawa,  J.  Kudo,  R.  P.  Evans, 

S.  R.  Woodward,  and  R.  N.  Williams  29 

Relating  soil  chemistry  and  plant  relationships  in  wooded  draws  of  the  northern  Great  Plains  .  .  . 

Marguerite  E.  Voorhees  and  Daniel  W.  Uresk  35 

The  genus  A  ristida  (Gramineae)  in  California Kelly  W.  Allred  41 

Temperature-mediated  changes  in  seed  dormancy  and  light  requirement  for  Penstemon  palmeri 

(Scrophulariaceae) Stanley  G.  Kitchen  and  Susan  E.  Meyer  53 

Late  Quaternary  arthropods  from  the  Colorado  Plateau,  Arizona  and  Utah 

Scott  A.  Elias,  Jim  I.  Mead,  and  Larry  D.  Agenbroad  59 

Microhabitat  selection  by  the  johnny  darter,  Etheostoma  nignim  Rafinesque,  in  a  Wyoming 

stream Robert  A.  Leidy  68 

Nomenclatural  innovations  in  Intermountain  Rosidae Arthur  Cronquist  75 

Nomenclatural  changes  and  new  species  in  Platypodidae  and  Scolytidae  (Coleoptera),  part  II.  .  . 

Stephen  L.  Wood  78 

Nomenclatural  changes  in  Scolytidae  and  Platypodidae  (Coleoptera) Stephen  L.  Wood  89 

Book  Review 

Plant  biology  of  the  Basin  and  Range  C.  B.  Osmond,  L.  F.  Pitclka,  and  G.  M.  Hidij 


Bruce  N.  Smith  93 

No.  2— June  1992 

Articles 

Red  Butte  Canyon  Research  Natural  Area:  history,  flora,  geology,  climate,  and  ecolog>- 

James  R.  Ehleringer,  Lois  A.  Arnow,  Ted  Arnow,  Irving  R.  McNulty, 

and  Norman  C.  Negus  95 

Influences  of  sex  and  weather  on  migration  of  mule  deer  in  California Thomas  E.  Kucera         122 

Diatom  flora  of  Beaver  Dam  Creek,  Washington  County,  Utah,  USA 

Kurtis  H.  Yearsley,  Samuel  R.  Rushforth,  and  Jeffrey  R.  Johansen         131 

Stratification  of  habitats  for  identifying  habitat  selection  by  Merriams  Turkeys 

Mark  A.  Rumble  and  Stanley  H.  Anderson         139 

Pollinator  preferences  for  yellow,  orange,  and  red  flowers  of  Mituuhis  vcrbenaccus  and  M. 

cardinalis Robert  K.  Vickery,  Jr.         145 

Soil  loosening  process  following  the  abandonment  of  two  arid  western  Nevada  townsites 

Paul  A.  Knapp        149 

Biochemical  differentiation  in  the  Idaho  ground  squirrel,  Spermophilus  bntuneus  (Rodentia: 

Scuridae) Ayesha  E.  Gill  and  Eric  Yensen         155 

New  genus,  Aplanusiella,  and  two  new  species  of  leattioppers  from  southwestern  United  States 
(Homoptera:  Cicadellidae:  Deltocephalinae) M.  W.  Nielson  and  B.  A.  Haws 


160 


1992]  Index  393 

Summer  hal)itat  use  hy  Columbian  Sharp-tailed  Crouso  in  western  Idaho 

\ictoria  Ann  Saab  and  Jeffrey  Shaw  Marks         166 

Notes 

Characteristics  of"  sites  occupied  by  subspecies  o(  Artr7nisia  triclrntata  in  the  Piceance  Basin, 

Colorado Thomas  K.  Cottrell  and  Charles  D.  Bonham         174 

Use  of  lakes  and  reservoirs  b\  miuratini;  shorebirds  in  Idaho 

Daniel  M.  Taylor  and  Charles  H.  Trost         179 

Dispersal  ot  scjuarrose  knapweed  (Centaurea  vir<:,ata  ssp.   squarrosa)  capitula  by  sheep  on 

rangeland  in  Juab  County,  Utah Cindy  Talbott  Roche,  Ben  F.  Roche,  Jr., 

and  G.  Allen  Rasmussen         185 

Vegetation  associated  with  two  alien  plant  species  in  a  fescue  grassland  in  Glacier  National  Park, 

Montana Robin  W.  Tyser         189 

No.  3 — September  1992 

Articles 

Plant  adaptation  in  the  Great  Basin  and  Colorado  Plateau 

Jonathan  P.  Comstock  and  James  R.  Ehleringer         195 

Life  history,  abundance,  and  distribution  of  Moapa  dace  [Moapa  coriacea) 

G.  Gar\-  Scoppettone,  Howard  L.  Burge,  and  Peter  L.  Tuttle         216 

Condition  models  for  wintering  Northern  Pintails  in  the  Southern  High  Plains 

Loren  M.  Smith,  Douglas  G.  Sheeley,  and  David  B.  Wester        226 

Evaluation  of  road  track  surveys  for  cougars  (Felis  concolor) 

Walter  D.  Van  Sickle  and  Frederick  G.  Lindzey         232 

Leaf  area  ratios  for  selected  rangeland  plant  species 

Mark  A.  Weltz,  Wilbert  H.  Blackburn,  and  J.  Roger  Simanton         237 

Ecology  and  management  of  medusahead  {Taeniathcrum  caput -medusae  ssp.  asperum  [Simk.] 

Melderis) James  A.  Young        245 

Roost  sites  used  by  Sandhill  Crane  staging  along  the  Platte  River,  Nebraska 

Bradley  S.  Norling,  Stanley  H.  Anderson,  and  Wayne  A.  Hubert         253 

Post-Pleistocene  dispersal  in  the  Mexican  vole  (Microtus  tnexicanus):  an  example  of  an  apparent 

trend  in  the  distribution  of  southwestern  mammals Russell  Davis  and  J.  R.  Callahan         262 

Can  Townsend's  ground  squirrels  survive  on  a  diet  of  exotic  annuals? 

Eric  Yensen  and  Dana  L.  Quinney         269 

Notes 

Avifauna  of  central  Tule  Valley,  western  Bonneville  Basin Peter  Hovingh         278 

Wildfire  and  soil  organic  carbon  in  sagebrush-bunchgrass  vegetation Steven  A.  Acker         284 

Structure  of  a  white-tailed  prairie  dog  burrow 

Lynn  A.  Cooke  and  Steven  R.  Swiecki         288 

Hybrids  of  white-tailed  and  mule  deer  in  western  Wyoming 

Charles  E.  Ka\-  and  Edward  Boe         290 

No.  4 — December  1992 
Articles 

Winter  nutrient  content  and  deer  use  of  gaml)el  oak  twigs  in  north  central  Utah 

Rosemary  L.  Pendleton,  Fred  J.  Wagstaff,  and  Bruce  L.  Welch         293 

Botanical  content  of  black-tailed  jackrabbit  diets  on  semidesert  rangeland 

Tchouassi  Wansi,  Rex  D.  Pieper,  Reldon  F.  Beck,  and  Leigh  W.  Murray         300 

Species  oiEimeria  from  the  thirteen-lined  ground  squirrel,  Spennophihis  thdecemlineatus ,  from 

Wyoming  .  .  Robert  S.  Seville,  Diane  M.  Thomas,  Russell  Pickering,  and  Nancy  L.  Stanton         309 


394 


Great  Basin  Naturalist  [Volume  52 


Plant  age/size  distributions  in  black  sagebrush  {Aiicinisia  nova):  etlt-cts  on  communit\  structure 

James  A.  Young  and  Debra  E.  PalnKjuist         313 

Mushroom  consumption  (mycophagy)  by  North  American  cervids 

Karen  L.  Launchbaugh  and  Philip  J.  Urness         321 

Terrestrial  vertebrates  of  the  Mono  Lake  islands,  California 

Michael  L.  Morrison,  William  M.  Block,  Joseph  R.  Jehl,  Jr.,  and  Linnea  S.  Hall         328 

Vascular  flora  of  Kane  Lake  Ciniue,  Pioneer  Mountains,  Idaho 

Robert  K.  Moseley  and  Susan  Bernatas         335 

Lakeward  and  downstream  movements  of  age-0  arctic  grayling  (rhymallus  arcticus)  originating 

between  a  lake  and  a  waterfall Mark  A.  Deleray  and  Calvin  M.  Kaya         344 

Effects  of  browsing  by  mule  deer  on  tree  growth  and  fruit  production  in  juvenile  orchards 

Dennis  D.  Austin  and  Philip  J.  Urness         352 

Changes  in  riparian  vegetation  along  the  Colorado  River  and  Rio  Crande,  Colorado 

Warren  D.  Snyder  and  Gary  C.  Miller         357 

Resident  Utah  deer  hunters'  preferences  for  management  options 

Dennis  D.  Austin,  Philip  J.  Urness,  and  Wes  Shields         364 

List  of  Oregon  Scolytidae  (Coleoptera)  and  notes  on  new  records 

Malcolm  M.  Furniss,  James  B.  Johnson,  Richard  L.  Westcott,  andTorolfR.  Torgersen         373 

Rittle  beetles  (Coleoptera:  Elmidae)  of  Death  Vallev  National  Monument,  California 

William  D.  Shepard         378 

Notes 

Siphonaptera  (fleas)  collected  from  small  mammals  in  montane  southern  Utah 

James  R.  Kucera  and  Clenn  E.  Haas         382 

Notes  on  spider  (Theridiidae,  Salticidae)  predation  of  the  harvester  ant,  Po^onomynncx  salinus 
Olsen  (Hvmenoptera:  Formicidae:  Mvrmicinae),  and  a  possible  parasitoid  fly  (Chloropidae) 

' William  H.  Clark  and  Paul  E.  Blom         385 

Index  to  Volume  52 387 


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(ISSN  0017-3614) 

GREAT  BASIN  NATURALIST  Vol  52  no  4  December  1992 

CONTENTS 

Articles 

Winter  nutrient  content  and  deer  use  of  gambel  oak  twigs  in  north  central 

Utah.  .  .  .   Rosemary  L.  Pendleton,  Fred  J.  Wagstaff,  and  Bruce  L.  Welch      293 

Botanical  content  of  black-tailed  jackrabbit  diets  on  semidesert  rangeland  .  .  . 

Tchouassi  Wansi,  Rex  D.  Pieper,  Reldon  F.  Beck,  and  Leigh  W  Murray      300 

Species  of  Eimeria  from  the  thirteen-lined  ground  squirrel,  Spermophilus  tri- 

decemlineatus,  from  Wyoming Robert  S.  Seville, 

Diane  M.  Thomas,  Russell  Pickering,  and  Nancy  L.  Stanton      309 

Plant  age/size  distributions  in  black  sagebrush  {Artemisia  nova):  effects  on 

community  structure James  A.  Young  and  Debra  E.  Palmquist      313 

Mushroom  consumption  (mycophag)')  by  North  American  cervids 

Karen  L.  Launchbaugh  and  Philip  J.  Urness      321 

Terrestrial  vertebrates  of  the  Mono  Lake  islands,  California 

Michael  L.  Morrison,  William  M.  Block,  Joseph  R.  Jehl,  Jr., 

and  Linnea  S.  Hall      328 

Vascular  flora  of  Kane  Lake  Cirque,  Pioneer  Mountains,  Idaho 

Robert  K.  Moseley  and  Susan  Bernatas      335 

Lakeward  and  downstream  movements  of  age-0  arctic  grayling  {Thymallus  arc- 

ticus)  originating  between  a  lake  and  a  waterfall 

Mark  A.  Deleray  and  Calvin  M.  Kaya      344 

Effects  of  browsing  by  mule  deer  on  tree  growth  and  fruit  production  in  juve- 
nile orchards Dennis  D.  Austin  and  Philip  J.  Urness      352 

Changes  in  riparian  vegetation  along  the  Colorado  River  and  Rio  Grande, 

Colorado Warren  D.  Snyder  and  Gary  C.  Miller     357 

Resident  Utah  deer  hunters'  preferences  for  management  options 

Dennis  D.  Austin,  Philip  J.  Urness,  and  Wes  Shields      364 

List  of  Oregon  Scolytidae  (Coleoptera)  and  notes  on  new  records 

Malcolm  M.  Furniss,  James  B.  Johnson,  Richard  L.  Westcott, 

and  Torolf  R.  Torgersen      373 

Riffle  beetles  (Coleoptera:  Elmidae)  of  Death  Valley  National  Monument,  Cali- 
fornia  William  D  Shepard      378 

Notes 

Siphonaptera  (fleas)  collected  from  small  mammals  in  montane  southern  Utah 

James  R.  Kucera  and  Glenn  E.  Haas      382 

Notes   on   spider   (Theridiidae,    Salticidae)   predation   of  the   harvester  ant, 
Pogonomyrmex  salinus  Olsen  (Hymenoptera:   Formicidae:   Myrmicinae), 

and  a  possible  parasitoid  fly  (Chloropidae) 

William  H.  Clark  and  Paul  E.  Blom      385 

Index  to  Volume  52 387