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HARVARD    UNIVERSITY 

Library  of  the 

Museum  of 

Comparative  Zoology 


Tlie  Great  Basin  Naturalist 


VOLUME  40,  1980 


Editor:  Stephen  L.  Wood 


Published  at  Brigham  Young  University,  by 
Brigham  Young  University 


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TABLE  OF  CONTENTS 

Volume  40 
Number  1  -  March  31,  1980 

Some  aspects  of  succession  in  the  spnice-fir  forest  zone  of  northern  Utah.   David  J. 

Schimpf,  Jan  A.  Henderson,  and  James  A.  MacMahon  1 

Utah  flora:  Malvaceae.  Stanley  L.  Welsh  27 

I'tah  flora:  miscellaneous  families.  Stanley  L.  Welsh 38 

The  taxononiic  status  of  the  rosy  boa  LicJianura  roseofiisca  (Serpentes:  Boidae).  John  R. 

Ottley,  Robert  W.  Murphy,  and  Geoffrey  V.  Smith  59 

Hesperoperla  hoguei,  a  new  species  of  stonefly  from  California  (Plecoptera:  Perlidae). 

Richard  W.  Baumann  and  Bill  P.  Stark  ...! 63 

Reproduction  in  three  sympatric  lizard  species  from  west-central  Utah.  John  B.  Andre 

and  James  A.  MacMahon  68 

Haplopappus  aJpinus  (Asteraceae):  a  new  species  from  Nevada.  Loran  C.  Anderson  73 

Miscellaneous  plant  novelties  from  Alaska,  Nevada,  and  Utah.  Stanley  L.  Welsh  and 

Sherel  Goodrich  78 

New  genera  and  new  generic  synonymy  in  Scolytidae  (Coleoptera).  Stephen  L.  Wood  ....  89 
The  bacterium  Thioploca  ingrica  on  wet  walls  in  Zion  National  Park,  Utah.  Samuel  R. 

Rushforth,  Sheril  D.  Burton,  Jeffrey  R.  Johansen,  and  Judith  A.  Grimes  98 


Number  2  -  June  30,  1980 

Feeding  ecology  of  Gilu  l)or(ixo})itis  (Osteichthyes:  Cyprinidae)  endemic  to  a  thermal 

lake  in  southeastern  Oregon.  Jack  E.  Williams  and  Cynthia  D.  Williams 101 

First  record  of  the  pallid  bat  {Antwzuiis  pallidus)  from  Montana.  Jeff  Shrver  and  Dennis 

L.  Flath  ' 115 

A  CJuracanthiimi  spider  bite.  Dorald  M.  Allred  116 

Identity  of  narrow-leaved  Chnjsothamnus  viscidiflorus  (Asteraceae).  Loran  C.  Anderson  117 
Ribulose  diphosphate  carboxylase  activities  in  cold-resistant  common  mallow,  Malva  ne- 

glecta  Wallr.  and  a  cold-sensitive  tomato,  Lycopersicon  esculentum  L.,  Ace  55  var. 

William  R.  Andersen  and  Jack  D.  Brotherson  121 

Recovery  of  Gambel  oak  after  fire  in  central  Utah.  L.  M.  Kunzler  and  K.  T.  Harper 127 

Relationships  among  total  dissolved  solids,  conductivity,  and  osmosity  for  five  ArtemUi 

habitats  (Anostraca:  Artemiidae).  Nicholas  C.  Collins  and  Gray  Stirling  131 

Spawning  of  the  least  chub  {lotichtlufs  pJilegethontis).  Thomas  M.  Baugh  139 

Transferrin  polymorphism  in  bighorn  sheep,  Ovi.s  canadensis,  in  Colorado.  Patrick  W. 

Roberts,  Donald  J.  Nash,  and  Robert  E.  Keiss  141 

The  genus  Eriogonum  Michx.  (Polygonaceae)  and  Michel  Gandoger.  James  L.  Reveal 143 

Parasites  from  two  species  of  suckers  (Catostomidae)  from  southern  Utah.  J.  Craig  Brein- 

holt  and  Richard  A.  Heckmann 149 

Soil  water  withdrawal  and  root  distribution  under  gnibbed,  sprayed,  and  undisturbed 

big  sagebnish  vegetation.  David  L.  Sturges 157 

Swarming  of  the  western  harvester  ant,  Pogononiywiex  occidentalis.  Dorald  M.  Allred  ...  165 
Relationship  between  environmental  and  vegetational  parameters  for  understory  and 

open-area  communities.  William  E.  Evenson,  Jack  D.  Brotherson,  and  Richard  B. 

Wilcox  167 


Seasonal  activity  pattern  ot  Columbian  ground  squirrels  in  the  Idaho  primitive  area. 

Charles  L.' Elliott  and  Jerran  T.  Flinders  I'^S 

Habitat  and  plant  distributions  in  hanging  gardens  of  the  Narrows,  Zion  National  Park, 

Utah.  Ceorge  P.  Malanson  1^^ 

Short-term  effects  of  logging  on  red-backed  voles  and  deer  mice.  Thomas  M.  Campbell 

III  and  Tim  W.  Clark 183 

Terminal  bud  formation  in  limber  pine.  Ronald  M.  Lanner  and  James  A.  Bryan  190 

Stinger  utilization  and  predation  in  the  scorpion  Paniwctonus  horeus.  Bmce  S.  Gushing 

and  Anne  Matherne  1^'^ 


Number  3  -  September  30,  1980 

Spatiotemporal  variation  in  phenology  and  abundance  of  floral  resources  on  shortgrass 

prairie.  V.  J.  Tepedino  and  N.  L.  Stanton 197 

Doe  owners  and  hvdatid  disease  in  Sanpete  County,  Utah.  Peter  M.  Schantz  and  Ferron 

L.  Andersen  -i" 

New  grass  distribution  records  for  Arizona,  New  Mexico,  and  Texas.  Stephan  L.  Hatch  ..  221 
A  comparison  of  epiphytic  diatom  assemblages  on  living  and  dead  stems  of  the  common 

grass  Phr(i<fniites  austmUs.  Judith  A.  Grimes,  Larry  L.  St.  Glair,  and  Samuel  R. 

Rushforth ' 223 

Poisonous  plants  of  Utah.  Jack  D.  Brotherson,  Lee  A.  Szyska,  and  William  E.  Evenson  ...     229 

The  successional  status  of  Cuprcssus  arizonica.  Albert  J.  Parker  254 

A  self-pollination  experiment  in  Pimis  echilis.  Ronald  M.  Lanner 265 

Comparative  floral  biology  of  Penstemon  eatonii  and  Penstenion  cyananthus  in  central 

Utah:  a  preliminary  study.  Lucinda  Bateman  268 

Differential  habitat  utilization  by  the  sexes  of  mule  deer.   Michael  M.  King  and  H. 

Duane  Smith 273 

Temporal  activity  patterns  of  a  Dipodomys  ordii  population.  Clive  D.  Jorgensen,  H. 

Duane  Smith,  and  James  R.  Garcia 282 

New  records  of  western  Trichoptera  with  notes  on  their  biology.  Bernard  G.  Swegman 

and  Leonard  C.  Ferrington,  Jr 287 

Observations  on  seasonal  variation  in  desert  arthropods  in  central  Nevada.  Robert  D. 

Pietruszka  292 


Number  4  -  December  31,  1980 

Impact  of  the  1975  Wallsburg  fire  on  antelope  bitterbrush  [Piiishid  tiidcntata).  Fred  J. 

Wagstaff .'. 299 

Terrestrial  vertebrate  fauna  of  the  Kaiparowits  Basin.  N.  Duane  Atwood,  Clvde  L.  Prit- 

chett,  Richard  D.  Porter,  and  Benjamin  W.  Wood 303 

A  new  species  of  fossil  (^hn/sothamnti.s  (Asteraceae)  from  New  Mexico.  Loran  C.  Ander- 
son       351 

New  American  bark  beetles  (Coleoptera:  Scolytidae),  with  two  recently  introduced  spe- 
cies. Stephen  L.  Wood  353 

Field  observations  on  the  response  of  the  Railroad  Valley  springfish  {Crenichtliy.s  nc- 

vadae)  to  temperature.  Thomas  M.  Baugh  and  Bruce  G.  Brown  359 

Woodrat  nest  flea  Anomiopsylhis  aniphihohis  in  southeastern  Oregon.  Harold  J.  Ego- 

scue  361 

Postemergence  development   and   interyear  residence   of  juvenile   Columbian  ground 

.squirrels  in  the  Idaho  primitive  area.  Charles  L.  Elliott  and  Jerran  T.  Flinders  362 


Flood  frequency  and  the  assenil)latj;e  of  dispersal  types  in  hanti;in<j;  (gardens  of  the  Nar- 
rows, Zion  National  Park,  Utah.  Geor<j;e  P.  Malanson  and  Jeanne  Kay  365 

Zonation  patterns  in  the  potholes  of  Kalsow  Prairie,  Iowa.  Jack  D.  Brotherson  372 

Plants  of  Angel  Island,  Marin  County,  California.  J.  D.  Kipley 385 

Additions  to  the  vascular  flora  of  Teton  (bounty,  Wyoming.   Ronald  L.  Ilartinan  and 

Robert  W.  Lichvar '. '. \ 408 

Index  to  X'oluine  40  414 


L^  f<Ll^      ^/oc 


rHE  GREAT  BASIN  NATURALIST 


Volume  40  No.  1 


March  31,  1980 


Brigham  Young  University 

-VUS,  COMP.   ZOOL 
LIBRARY 


GREAT  BASIN  NATURALIST 

Editor.  Stephen  L.  Wood,  Department  of  Zoology,  Brigham  Young  University,  Provo,  Utah 

84602. 
Editorial  Board.   Kimball  T.  Harper,  Botany;  Wilmer  W.  Tanner,  Life   Science  Museum; 

Stanley  L.  Welsh,  Botany;  Clayton  M.  White,  Zoology. 
Ex  Officio  Editorial  Board  Members.  A.  Lester  Allen,  Dean,  College  of  Biological  and  Agricul- 
tural Sciences;  Ernest  L.  Olson,  Director,  Brigham  Young  University  Press,  University 
Editor. 

The  Great  Basin  Naturalist  was  founded  in  1939  by  Vasco  M.  Tanner.  It  has  been  published 
from  one  to  four  times  a  year  since  then  by  Brigham  Young  University,  Provo,  Utah.  In  gener- 
al, only  previously  unpublished  manuscripts  of  less  than  100  printed  pages  in  length  and  per- 
taining to  the  biological  and  natural  history  of  western  North  America  are  accepted.  The 
Great  Basin  Naturalist  Memoirs  was  established  in  1976  for  scholarly  works  in  biological  natu- 
ral history  longer  than  can  be  accommodated  in  the  parent  publication.  The  Memoirs  appears 
irregularly  and  bears  no  geographical  restriction  in  subject  matter.  Manuscripts  are  subject  to 
the  approval  of  the  editor. 

Subscriptions.  The  annual  subscription  to  the  Great  Basin  Naturalist  is  $12  (outside  the 
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are  available  for  sale.  All  matters  pertaining  to  the  purchase  of  subscriptions  and  back  num- 
bers should  be  directed  to  Brigham  Young  University,  Life  Science  Museum,  Provo,  Utah 
84602.  The  Great  Basin  Naturalist  Memoirs  may  be  purchased  from  the  same  office  at  the 
rate  indicated  on  the  inside  of  the  back  cover  of  either  journal. 

Scholarly  Exchanges.  Libraries  or  other  organizations  interested  in  obtaining  either  journal 
through  a  continuing  exchange  of  scholarly  publications  should  contact  the  Brigham  Young 
University  Exchange  Librarian,  Harold  B.  Lee  Library,  Provo,  Utah  84602. 

Manuscripts.  All  manuscripts  and  other  copy  for  either  the  Great  Basin  Naturalist  or  the 
Great  Basin  Naturalist  Memoirs  should  be  addressed  to  the  editor  as  instructed  on  the  back 
cover. 


5-80  650  45971 


ISSN  0017-3614 


The  Great  Basin  Naturalist 


Published  at  Pkovo,  Utah,  by 
Brigham  Young  University 

ISSN  0017-3614 


Volume  40 


March  31,  1980 


\o.  1 


SOME  ASPECTS  OF  SUCCESSION 
IN  THE  SPRUCE-FIR  FOREST  ZONE  OF  NORTHERN  UTAH 

David  J.  Sthiiiipf'-,  Jan  A.  ilendersou,'^  and  James  A.  MacMahon' 

.\bstract.—  a  site  in  the  Rocky  Mountain  siibalpine  forest  zone  with  which  a  series  of  hypotheses  concerning 
ecosystem  succession  was  tested  is  characterized.  Succession  from  herh-dominated  meadows  to  climax  forests  of  Eng- 
ehnann  spruce  and  suhalpine  fir  can  follow  at  least  four  identified  pathwa\'s.  After  fire,  spruce  and  fir  may  reinvade 
a  site  directly,  follow  invasion  by  aspen,  or  follow  invasion  by  lodgepole  pine,  the  pathway  depending  on  a  com- 
bination of  physical  and  biotic  factors.  In  other  cases,  succession  begins  with  long-established  meadows  which  do  not 
owe  their  existence  to  fire.  In  this  latter  pathway,  aspen  invades  meadows  by  suckering  and  changes  the  environ- 
ment near  the  soil  surface  so  as  to  facilitate  establishment  of  the  climax  tree  species.  The  biota  and  soils  of  four 
characteristic  serai  stages  (meadow,  aspen,  tir,  spruce-tiri  in  this  latter  pathway  are  described. 


Succession  may  be  identified  as  the  change 
in  ecosystem  properties  of  a  specified  area 
over  a  time  interval  of  the  same  magnitude  as 
the  generation  time  of  the  conspicuous  or- 
ganisms in  the  ecosvstem.  Odum  (1969)  pro- 
po,sed  24  trends  in  ecosystem  properties  as 
successional  changes  take  place.  From  1976 
through  1978  we  and  our  colleagues  .studied 
many  of  these  properties  in  a  successional  se- 
quence of  ecosystems  in  the  subalpine  zone 
of  the  Wasatch  Mountains  of  northern  Utah. 
In  this  paper  we  review  patterns  of  succes- 
sion i-  the  subalpine  central  Rockv  Moun- 
tains, describe  our  intensively  studied  se- 
quence, and  characterize  the  environment  of 
the  study  area.  Subsequent  papers  will  report 
results  of  tests  for  trends  in  specific  ecosys- 
tem properties  along  the  successional 
gradient  in  the  context  of  Odum's  hypoth- 
eses. 


Materials  and  Methods 
Rocky  Mountain  Subalpine  Succession 

The  subalpine  zone  of  the  Rocky  Moun- 
tains is  the  uppermost  forested  part  of  the 
Cordillera,  characterized  by  climatic  climax 
ecosystems  dominated  by  Engelmann  spruce 
{Picea  engebnannii  Parry)  and  subalpine  fir 
{Abies  kisiocarpa  [Hook.]  Nutt.)  (Daubenmire 
1943,  1978,  Oosting  and  Reed  1952),  here- 
after referred  to  as  spruce  and  fir.  Some 
workers  term  this  the  upper  montane  zone 
(Love  1970).  Although  long-term  climatolog- 
ical  data  from  tliis  zone  are  comparatively 
scant,  the  occurrence  of  these  subalpine  for- 
ests is  apparently  more  closely  correlated 
with  summer  air  temperatures  than  with  pre- 
cipitation patterns  (Daubenmire  1956). 

A  variety  of  types  of  ecosystems  may  oc- 


'Department  of  Biology  and  Ecology  Center,  Utah  State  University,  Logan,  Utah  84322. 
'Present  address:  Department  of  Biology,  University  of  Minnesota,  Duliith,  Minnesota  5.5812. 

'Department  of  Forestry  and  Outdoor  Recreation  and  Erology  Center.  Utah  State  University.  Logan.  Utah  84,322.  Present  address:  U.S.  Forest  Service, 
Federal  Building,  P.O.  Box  2288.  Olympia,  Washington  98507. 


Great  Basin  Naturalist 


Vol.  40,  No.  1 


cupv  a  subalpine  site  from  the  time  it  is  her- 
baceous in  character  until  its  full  devel- 
opment as  a  spruce-fir  forest.  In  the  lower 
range  of  the  subalpine  zone,  preclimax  forests 
are  sometimes  dominated  by  tree  species 
which  form  climatic  climax  stands  in  lower 
zones;  the  particular  tree  species  involved 
depend  on  the  geographic  location  in  the 
Cordillera  (Daubenmire  1943).  Throughout 
the  subalpine  zone  the  spruce-fir  climax  is  of- 
ten preceded  by  tree  species  which  form  cli- 
max stands  only  under  restricted  topograph- 
ic/edaphic  conditions.  The  predominant 
species  of  this  type  are  quaking  aspen  {Popii- 
lus  treiniiloides  Michx.)  and  lodgepole  pine 
{Pinus  contorta  Dougl.  var.  hitifolia  Engelm.), 
hereafter  referred  to  as  aspen  and  lodgepole. 
While  lodgepole  is  usually  an  invader  of 
burned  areas,  it  also  may  invade  unforested 
sites  which  have  not  been  binned  recently 
(Patten  1969). 

Few  detailed  studies  of  successional  path- 
ways in  the  subalpine  of  the  Rocky  Moun- 
tains have  been  reported.  In  Colorado,  the  lo- 
cal presence  of  lodgepole  seeds  versus  aspen 
roots  tends  to  determine  the  composition  of 
the  preclimax  forest  which  follows  fire 
(Stahelin  1943),  though  exposure  and  soils 
also  play  a  role  (Langenheim  1962,  Feet 
1978). 

Direct  invasion  of  unforested  areas  by 
spmce  and  fir  can  be  extremely  slow,  even 
when  high  numbers  of  their  seeds  reach  the 
site  (Noble  and  Ronco  1978).  Competition 
with  vigorous  herbs  or  shrubs  in  the  open 
stands  is  undoubtedly  a  factor  (Alexander 
1974,  Dunwiddie  1977),  but  the  direct  expo- 
sure of  spruce  and  fir  seedlings  to  the  sun  and 
night  sky  may  be  more  detrimental.  Deadfall 
remaining  after  logging  or  blowdown  in- 
creases the  rate  of  reestablishment  of  spruce 
and  fir  via  shading  (Alexander  1974,  Noble 
and  Alexander  1977).  Even  greater  recruit- 
ment of  spruce  and  fir  populations  occurs  in 
the  .shade  of  lodgepole  or  aspen  if  understory 
vegetation  is  not  too  dense. 

Several  mechanisms  for  the  shading  ben- 
efaction of  Engelmann  spruce  establishment 
have  been  identified.  Shade  obviously  reduc- 
es evaporative  and  heat  .stress  in  the  seed- 
lings' environment.  Even  under  favorable  wa- 
ter balance,  spruce  seedlings  may  be 
intolerant   of   full    intensity   sunlight   (Ronco 


1970),  though  this  was  concluded  from  obser- 
vation of  seedlings  planted  at  higher  eleva- 
tions than  those  of  their  parents.  Shade-cast- 
ing objects  also  lessen  the  nocturnal  radiative 
cooling  of  seedlings;  spruce  seedling  growth 
Is  enhanced  by  higher  night  temperatures 
(Hellmers  et  al.  1970).  Under  high  soil  water 
potential,  spruce  seedlings  emerge  somewhat 
faster  than  lodgepole  at  16  C,  but  distinctly 
slower  than  lodgepole  at  35  C  (Kaufmann 
and  Eckard  1977).  The  mechanisms  for 
shade-enhanced  subalpine  fir  establishment 
are  less  well  understood. 

Subalpine  succession  involving  lodgepole 
or  aspen  preclimax  ecosystems  seems  to  qual- 
ify as  a  "facilitation"  (Council  and  Slatyer 
1977)  type  of  serai  sequence,  at  least  with  re- 
gard to  trees.  Under  certain  circumstances 
the  preclimax  .species  invade  the  site  more 
readily  than  do  the  climax  species,  and  great- 
ly facilitate  the  later  invasion  of  climax  trees. 
Aspen  and  lodgepole  are  unable  to  persist  as 
more  than  isolated  individuals  following 
spruce-fir  canopy  development,  possibly  due 
in  part  to  their  shoot  geometry  (Horn  1971) 
and  high  light  requirements  in  the  face  of 
dense  shade  cast  by  spruce  and  fir. 

Adaptive  features  conferring  this  greater 
colonizing  ability  on  lodgepole  and  aspen  ap- 
pear to  center  on  the  establi.shment  phase  of 
the  life  cycle.  Aspen  bypasses  the  seedling  es- 
tablishment barrier  of  harsh  environments  by 
vegetative  reproduction,  producing  sucker 
shoots  from  the  roots.  Establishment  from 
seed  is  thought  to  be  extremely  uncommon  in 
the  central  Rocky  Mountain  portion  of  the 
range  of  this  species  (Cottam  1954).  Thus  as- 
pen commonly  invades  only  those  sites  imme- 
diately adjacent  to  existing  clones.  Lodgepole 
establishes  only  from  seed;  in  some  portions 
of  its  range  the  seed-bearing  cones  open 
strictly  in  response  to  heat,  usually  that  from 
fire  (serotiny).  Lodgepole  seedlings  do  not 
root  deeper  than  spruce,  but  do  develop 
much  more  extensive  root  systems  (Noble 
1979),  which  should  facilitate  .seedling  survi- 
val on  drier,  exposed  sites.  Additionally, 
lodgepole  seedlings  are  not  damaged  or  in- 
hibited by  the  high  light  intensities  (such  as 
in  imforested  sites)  to  which  spruce  and  fir 
seedlings  arc  less  tolerant  (Ronco  1970). 

The  relative  proportions  of  spruce  and  fir 
in  a  stand  vary  geographicallv,  altitudinally. 


March  1980 


SCHIMPF    ET  AL.:  SpRl'CE-FiR  SUCCESSION 


and  locally  (Daubcnmire  1943.  Peet  1978). 
As  a  species,  fir  tolerates  a  wider  ranye  ot 
site  conditions  than  spruce,  and  seems  ca- 
pable of  becoming  established  on  a  greater 
variety  of  substrates  (Fowells  1965).  Fir  seed- 
lings are  more  shade  tolerant  than  spruce 
(Fowells  1965).  The  composition  of  a  voung 
spruce-fir  stand  depends  heavily  upon  which 
species  seeded  abundantly  just  prior  to  a  pe- 
riod favorable  for  establishment.  In  the 
northern  Rocky  Mountains  good  fir  seed 
crops  are  more  frequent  than  good  spruce 
crops  (LeBarron  and  Jemison  1953).  In  the 
Wasatch  Mountains  the  converse  is  true, 
though  the  cohort  of  young  fir  is  often  larger 
because  of  better  fir  survival  (T.  W.  Daniel, 
pers.  comm.). 

Both  spruce  and  fir  grow  slowly  by  com- 
pari.son  to  lodgepole  or  aspen.  Both  have  nar- 
row pyramidal  crowns,  fir  the  more  so,  and 
may  have  few  live  branches  in  the  lowest  5  m 
of  mature  closed  stands.  Individuals  of  either 
species  seldom  exceed  40  to  45  m  in  height. 
Fir  suffers  heavy  mortality  at  125  to  175 
years  because  of  its  greater  susceptibility  to 
root  rot,  but  spruce  often  lives  300  to  500 
years.  Both  species  are  highly  susceptible  to 
fire,  due  to  their  low  crowns  when  young  and 
their  thin  bark. 

The  combination  of  irregular  seed  produc- 
tion and  irregular  occurrence  of  suitable  con- 
ditions for  tree  establishment  leads  to  an  in- 
frequent incidence  in  spruce-fir  stands  of  the 
inverse  J-shaped  population  age  structure 
classically  associated  with  climax  commu- 
nities (Alexander  1974,  LeBarron  and  Jemi- 
son 1953,  Whipple  and  Dix  1979).  Stands 
may  be  even-aged  or  multi-aged,  or  may  in- 
clude suppressed  individuals  of  considerable 
age  that  show  released  growth  only  when  a 
gap  opens  in  the  canopy. 

Injurious  organisms  may  strongly  affect  the 
dynamics  of  spruce-fir  stands.  Damage  from 
the  bark  beetle  (Dryocoetes  confii.siis  Swaine) 
and  associated  fungi  often  results  in  a  reduc- 
tion in  the  proportion  of  fir  at  a  stand  age  of 
125  to  175  years  (Schmid  and  Hinds  1974). 
Spruce  becomes  more  susceptible  to  spruce 
bark  beetle  {Dendroctonus  rufijH'imi.s  Kirbv) 
(Schmid  and  Frye  1977)  damage  as  it  be- 
comes older  and  larger  in  diameter,  at  about 
250  years.  Infestations  erupt  above  endemic 
levels  when  blowdowns  of  live  spruce  are  fol- 


lowed In  temperatures  and  moisture  levels 
conducive  to  survival  of  the  larvae;  repeated 
episodes  lead  to  a  stand  of  spruce  so  few  and 
large  that  they  are  extremely  susceptible  to 
attack  and  blowdown  (Dixon  1935,  Miller 
1970,  Schmid  and  Hinds  1974).  Stand  deva.s- 
tation  results  and  the  pace  of  succession  rises 
again. 

Both  spruce  and  fir  seeds  suffer  higher 
pregermination  mortality  in  litter  of  their 
own  species  than  in  litter  from  the  other  spe- 
cies (Daniel  and  Schmidt  1972).  Originally 
described  as  Ceniculodendwn  pyrifonnc  Salt, 
the  infectious  fungu.'^  now  appears  to  be  the 
imperfect  state  of  Caloscypha  fiil^em  (Pers.) 
Boudier  (Paden  et  al.  1978,  Wicklow-Howard 
and  Skujins  1979).  Such  self-inhil)ition  may 
account  in  part  for  the  relative  rarity  of  ma- 
ture monospecific  stands  of  either  spruce  or 
fir  in  the  Rocky  Mountains. 


A  Subalpine  Site  in  the 
Northern  ^Vasatch  Moimtains 

The  Wasatch  xMountains  of  northern  Utah 
and  southeastern  Idaho  are  a  major  com- 
ponent of  the  central  Rocky  Mountains.  A 
site  for  intensive  study  of  ecosystem  proper- 
ties in  relation  to  succession  was  selected  in 
the  Utah  State  University  School  Forest,  in 
Cache  and  Rich  counties,  about  15  km  south 
of  the  Utah-Idaho  border  (Fig.  1).  Stands  in- 
vestigated ranged  between  about  2550  m  and 
2650  m  in  elevation.  The  site  is  atop  an  un- 
dissected  plateaulike  ridge  of  gentle  topogra- 
phy. Adjacent  areas  are  lower  in  elevation 
and  are  not  a  source  of  cold  air  drainage  onto 
the  study  site.  Soils  throughout  the  site  are 
derived  from  the  Knight  formation  of  the 
Wasatch  group,  a  Tertiary  red  conglomerate 
of  quartzite,  sandstone,  and  shale  (Veatch 
1907).  Tliis  parent  material  occurs  exten- 
sively south  of  the  study  site  (Stokes  1963). 
The  site  contains  no  lakes  or  permanent 
streams.  Photoperiods  range  from  15  h  2  min 
on  1  June  to  13  h  9  min  on  1  September, 
with  a  15  h  15  min  maximum  at  the  solstice. 
Daily  solar  radiation  totals  (horizontal  sur- 
face) as  high  as  768  cal  cm  -  have  been  mea- 
sured (Eaton  1971). 

The  vegetation  consists  predominantly  of 
spmce-fir  forests  of  moderate  age,  containing 


Great  Basin  Naturalist 


Vol.  40,  No.  1 


lll*>W 


4a2N 


4j<^JV 

ii  1 1 1 1 1 1 1 1 1 1 1 1 1  rti 


Fig.  L   Map  of  northern  Utah,  locating  th.  .School  Forest  (SF«).  Garden  Citv  Su„n„il  .CC  ^  '    and  the  Inuus  ot- 
unshacled  lexeept  Hear  l.akei  and  honndaries  at  I.5()(),  2  KM),  and  2700  m. 


March  1980 


SCHIMPF    ET  AL.:  SpRl  CE-FiR  SUCCESSION 


onlv  a  few  widely  spaced  spruce  older  than 
275  vears.  The  oldest  known  trees  are  367 
vears  (spruce)  and  278  years  (fir)  (T.  W.  Dan- 
iel, pers.  conmi.).  Scattered  individuals  or 
small  groups  of  lodgepole  or  aspen  occur  in 
the  spmce-fir  stands.  Within  this  broad  for- 
ested area  are  small  meadows  fringed  with 
aspen  clones  on  various  fractions  of  their  per- 
imeters (Fig.  2).  Young  spruce  and  fir  are  fre- 
quently observed  in  these  aspen  stands  but 
seldom  in  the  meadows,  except  on  the  north 
margins  of  conifer  stands,  including  the  occa- 
sional small  clumps  of  mature  spruce  or  fir 
within  the  meadows.  No  significant  logging 
has  taken  place  on  the  site.  Cattle  and  sheep 
have  grazed  the  site  since  around  1900.  The 
zonal  climax  vegetation  of  the  studv  area  be- 
longs to  the  Abies  lasiocdipa / Pcdicularis 
mcemosa  climax  comniunit\  tvpe  and  habitat 
type  (Henderson  et  al.  1976). 

The  life-form  composition  of  the  vegeta- 
tion is  rather  simple,  consisting  mainlv  of 
herbs  and,  except  in  the  meadows,  trees.  Indi- 
viduals of  shrubbv  species  are  uncommon; 
shrub  biomass  is  less  than  1  percent  that  of 
herbaceous  biomass  in  meadows  and  under 
aspen,  and  less  than  5  percent  under  spruce- 
fir.  Small  tree  species  and  lianas  are  absent. 
These  features  distinguish  the  sere  under 
study  from  those  on  most  other  forest  succes- 
sion research  sites,  especially  those  in  the  de- 
ciduous forest  biome.  Cryptogams  occur,  but 
are  not  a  conspicuous  element  of  anv  stratimi 
of  the  vegetation,  except  for  periodic  emer- 
gence of  basidiocarps  in  the  forests.  The  gen- 
eral aspect  of  the  several  categories  of  vege- 
tation is  illustrated  in  Figure  3.  For  a 
discussion  of  the  compositional  and  environ- 
mental relationships  of  this  forest  to  other  lo- 
cal forests,  see  Henderson  et  al.  (1976)  and 
Lawton  (1979). 

A  preliminary  stud)  of  the  structure  of 
over  100  stands  on  environmentally  equiva- 
lent sites  in  the  School  Forest  (Sperger  and 
Henderson,  unpubl.)  indicated  that  there  are 
probably  four  major  pathways  of  succession 
leading  to  spruce-fir  forests  (Fig.  4).  Pathway 
"1"  represents  succession  following  the  de- 
struction by  fire  of  a  forest  containing  signifi- 
cant aspen  root  biomass.  The  aspens  sucker 
within  a  short  period  of  time  to  produce  an 
aspen-dominated  stand.  Spruce  and  fir  sub- 


secjuently  invade,  and  eventuallv  outlive  and 
replace  the  aspen. 

Pathway  "2"  occurs  in  the  lower  elevation- 
al  range  of  the  habitat  type  around  the  edge 
ot  the  School  Forest  area.  If  aspen  is  not  lo- 
cally abundant,  then  lodgepole  is  the  postfire 
pioneer,  provided  that  a  local  .seed  source  ex- 
ists. Lodgepole  cones  are  not  serotinous  in 
this  area,  so  seeding  is  from  adjacent  stands. 
Spruce  and  fir  establish  and  grow  more  slow- 
ly than  lodgepole;  thus,  a  pine-dominated 
ecosystem  exists  for  some  time  prior  to 
spmce-fir  stand  recovery.  Spruce  and  fir  may 
establish  soon  after  fire,  without  site  amelio- 
ration by  aspen  or  lodgepole,  if  a  significant 
quantity  of  unburned  woody  material  re- 
mains as  protection.  This  is  pathwav  "3," 
where  the  climax  species  establish  without 
preclimax  tree  species.  The  climax  stand 
structure  typically  takes  less  time  to  develop 
than  through  pathways  "1"  and  "2." 

Pathway  "4,"  the  most  common,  and  that 
which  we  studied  intensively,  begins  with 
long-persistent  meadows,  probably  not  of  fire 
origin.  These  are  eventually  invaded  by  a.s- 
pen  suckers  followed  by  spruce  and  fir.  Typi- 
cally the  fringe  of  aspen  clones  is  dis- 
continuous around  the  meadow,  yielding 
iiregular  patterns  of  meadow  invasion  (Fig. 
2).  Once  aspen  is  present,  the  time  frame  of 
events  resembles  that  of  the  other  three  path- 
ways. However,  the  sere  as  a  whole  mav  take 
hundreds  of  years  longer,  because  of  the 
long-persistent  meadow  stage. 

Pathway  "4"  was  studied  by  sinmltaneous 
investigation  of  plots  characterized  as  mead- 
ow, aspen,  fir,  and  spruce-fir.  The  fir-domi- 
nated plots  represent  a  stage,  containing 
some  spruce,  which  sometimes  occurs  be- 
tween occupancy  by  aspen  and  the  t\pical 
spruce-fir  mixture.  The  inference  of  these 
plots  as  a  chronosequence  seems  reasonable, 
based  on  the  minimal  relief  of  the  studv  site 
and  its  uniform  soil  parent  material.  Because 
the  time  since  the  meadows  were  last  forest- 
ed could  not  be  determined,  the  stands  can- 
not be  positioned  on  an  absolute  time  axis. 

Common  herbaceous  species  in  each  stage 
are  listed  in  Table  I.  These  represent  samples 
from  one  series  of  stands  at  Big  Meadow  and 
should  not  be  interpreted  as  ranks  for  the 
study  site  in  general.  Eriogonum  is  semi- 
shnibbv   rather   than   wholh'   herbaceous.   A 


Great  Basin  Naturalist 


Vol.  40,  No.  1 


Fii;.  2.   Aoiial  pliolo  iiicliicliii<4  Ihc  Ikul'^i-r  station  in  the  Stliool  Forest.  Lower  dianrani  inclieates  type  ol  forest 
various  sections  of  tlie  photo.  .Xrrow  jxjints  to  tlie  weather  station. 


March  1980 


SCHIMPF    ET  AL.:  SpRUCE-FiR  SUCCESSION 


substantial  change  in  species  composition  oc- 
curs along  the  siiccessional  secjuence,  with  no 
species  being  abundant  in  more  than  two 
stages.  The  niunber  of  species  is  lowest  in  the 
meadow  and  highest  in  the  preclimax  aspen 
and  fir  stages,  based  on  equal  sampling  effort. 
The  proportion  of  short-lived  species  in  the 
species  list  declines  onlv  slightly  along  the 
sere.    The    semishrub    Vacciniuin    scopdiiuin 


(Leiberg),  perhaps  the  most  abundant  under- 
storv  species  in  the  Rocky  Mountain  sub- 
alpine  /.one  (Daubenmire  1978),  was  not 
found  on  the  site. 

Mannnal  species  of  the  four  stages  are  list- 
ed in  Table  2.  The  Northern  Pocket  Gopher 
{Tliomoniy.s  falpoidcs)  is  abundant  in  the 
meadows,  producing  considerable  distur- 
bance of  the  upper  portions  of  the  soil  hori- 


Fig.  3.   Photographs  showing  general  physiognomy  of  four  stages  of  succession:  meadow,  aspen,  fir,  spruce-fir.  Pho- 
tos courtesy  of  D.  .\ndersen. 


8 


Great  Basin  Naturalist 


Vol.  40,  No.  1 


zons.  The  amount  of  bare  ground  between 
meadow  plants  is  substantial  and  largely  due 
to  gopher  activity.  Gophers  are  less  abundant 
under  aspen  and  rare  imder  conifers.  Their 
effect  on  conifer  invasion  rate  is  not  well  un- 
derstood, for  though  they  can  directly  de- 
stroy young  spruce  or  fir,  their  activities  may 
also  reduce  herbaceous  species  competition 
with  tree  seedlings.  Gophers  may  also  create 
more  favorable  .seedbeds  for  conifers  by  ex- 
posing mineral  soil. 

The  dominant  mammals  in  the  coniferous 
stages  are  the  Snowshoe  Hare  {Lepus  ameri- 
canus)  and  the  Red  Squirrel  (Tamiasciiirus 
hudsonicus).  Fall  and  winter  browsing  by 
hares  may  retard  conifer  stand  development 
(Baker  et  al.  1921);  an  individual  may  con- 
sume some  300  g  (fresh  weight)  of  woody 
stems  4  mm  or  less  in  diameter  daily  in  win- 
ter (Pea.se  et  al.  1979).  Hare  population  den- 
sities in  this  area  fluctuate  little  by  com- 
pari.son  with  their  boreal  counterparts 
(Dolbeer  and  Clark  1975). 

The  Red  Scjuirrel  harvests  spruce  and  fir 


DISTURBANCE  (fire) 


GRASS -FORB  MEADOW 


ASPEN 


LONG 

PERSISTING 

MEADOW 


LODGE POLE 
PINE 


ASPEN 


FIRS  SPRUCE 


SPRUCES  FIR 


SPRUCE  /  FIR 


SPRUCE  DOMINATED 
CLIMAX  WITH  FIR 
AS  A  SUBORDINATE. 

SPARSE  UNDERSTORY 
OF  Pedicularis  racemosa 


Fi^.   4.   Summarv  ol   types  of  succfssioiial   patli\\a\> 
tli()in;lil  to  \w  operative  on  liie  Scliool  Forest. 


cones  before  the  seeds  are  shed,  transporting 
them  to  large  caches  at  shady  bases  of  trees. 
Finlev  (1969)  concluded  that  Red  Squirrels 
harvest  almost  all  the  cones  produced  in 
vears  of  poor  to  average  seed  output,  and 
that  only  in  high  seed  production  years  are 
enough  .seeds  dispersed  that  significant  tree 
recruitment  is  possible.  Further  information 
on  the  mammalian  component  of  the  ecosys- 
tems is  in  Andersen  et  al.  (1980). 

Bird  .species  of  the  four  stages  are  listed  in 
Table  3.  Only  the  spruce-fir  data  are  from 
the  site;  preceding  stages  were  inventoried  in 
more  extensive  stands  nearby.  Meadows  fea- 
tiue  low  numbers  of  both  species  and  individ- 
uals. Conspicuously  absent  from  the  co- 
niferous stands  is  the  Gray  (or  Canada)  Jay 
(Perisoreus  canadensis),  a  common  per- 
manent resident  throughout  most  of  the 
Rocky  Mountain  .subalpine  zone.  Similarity 
in  composition  of  the  vertebrate  species 
among  stages  (Jaccard  coefficient  of  commu- 
nitv)  and  its  variation  between  years  in  the 
case  of  the  avifauna  are  shown  in  Figure  5. 
Further  avian  information  is  available  in 
Smith  and  MacMahon  (submitted). 

The  comparisons  of  species  similarities  for 
all  vertebrates  (Fig.  5)  across  the  .sere  show 
that  the  most  mature  stage,  spiTice,  is  most 
different  from  the  least  mature  stage,  mead- 
ow. The  most  similar  stages  are  the  two  con- 
ifers, spruce  and  fir.  All  pair-wise  serai  stage 
comparisons  of  mammals,  though  showing 
these  patterns,  are  more  similar  than  those 
for  birds.  The  implication  is  that  the  birds,  as 
we  would  expect,  respond  more  dramatically 
to  the  physiognomic  changes  attendant  to  de- 
veloping from  meadow  to  a  deciduous  forest 
and  finally  to  a  coniferous  forest.  Mammals, 
on  the  other  hand,  respond  to  the  presence  or 
absence  of  trees,  but  do  not  vary  as  much 
with  tree  species  or  leaf  habit.  The  verte- 
brate species  count  of  13-16  mammal  .species 
and  20-30  bird  species  for  each  of  oiu'  serai 
stages  correspond  to  mammal  and  bird  spe- 
cies counts  from  similar  communities  through 
North  America  and  also  northern  Europe 
(e.g.,  Erskine  1977.  Han.sson  1974,  Jiirvinen 
and  Viiisanen  1976,  Sabo  and  Whittaker 
1979). 

The  most  abundant  insects  in  the  meadows 
are  .species  of  aphidids  and  thripids.  Follow- 
ing  a   winter   drought,   the   aphids   declined 


March  1980 


Sc:himpf  et  al.:  Spruce-Fir  Succession 


9 


precipitously  in  1977,  when  two  species  of 
cicadelhds  were  the  most  abundant.  A  cica- 
delHd  is  the  most  abundant  in  the  aspen  un- 
derstorv,  followed  bv  three  species  of  thrip- 
ids.  In  the  aspen  canopv  the  most  common 
species  is  a  serpentine  leaf  miner  (Gracila- 
riidae),  followed  bv  an  aphidid,  a  cicadellid, 
and  a  blotch  mining  gracilariid.  In  the  con- 
ifer understory  the  most  common  species  is 
an  aphidid,  followed  by  two  thripids  and  two 
more  aphidids.  During  1977  the  aphids  were 
scarce  and  a  cicadellid  was  the  most  abun- 
dant. In  the  conifer  canopy,  the  most  abun- 
dant species  are  an  encyrtid,  a  mirid,  a  thrip- 
id,  and  an  eriosomatid.  The  insects  on  spruce 
were  observed  to  be  verv  similar  to  those  on 
fir. 


Among  soil  and  litter  metazoan  in- 
vertebrates, numbers  increase  markedly 
along  the  serai  sequence.  The  relative  abun- 
dance of  Collembola  and  the  plant-feeding 
nematodes  (Tylenchida,  Dorylaimida)  does 
not  change  much.  The  forested  stages  harbor 
increased  proportions  of  bacterivorous  nema- 
todes (Rhabditida),  detritivorous  mites  (Ori- 
batei),  and  predaceous  mites  (Mesostigmata). 
Oligochaetes  are  essentially  absent,  a  charac- 
teristic of  the  region  (Gates  1967),  and  gas- 
tropods are  also  rare  on  the  site.  Populations 
of  protozoans  were  not  estimated.  Soil  micro- 
organisms also  exhibit  large  absolute  increas- 
es in  number  through  successional  time  and 
are  generally  highest  in  density  in  the  conifer 
litter.    Estimates    of   abundance    for    several 


Table  1.  Herbaceous  species  comprising  5  percent  or  more  of  the  mean  daily  herbaceous  bioinass  in  samples  at 
the  Big  Meadow  succession  stages.  1977.  Numbers  following  a  taxon  correspond  to  its  biomass  rank,  with  percent  of 
the  herbaceous  biomass  indicated  for  top-ranked  species.  Number  of  various  categories  of  species  samples  and  her- 
baceous biomass  in  relation  to  tree  leaf  biomass  are  also  included.  Only  herbaceous  tissue  of  the  semishrub  Eriogo- 
nititi  is  included. 


Meadow 


.^spen 


Fir 


Spruce-Fir 


Achillea  miUefoUiim  L.  ssp.  kinulosa  (Nutt.)  Piper 
Agropyron  trachijcaiilum  (Link)  Malte  var.  lati^himc 

(Scribn.  &  .Smith)  A.  \.  Beetle 
Agropijron  tracJiijcauhnn  (Link)  Malte  var.  ghiiicuiu 

(Pease  &  Moore)  Malte 
Aster  engelmannii  (Eat.)  Grav 
Bwmiis  cciriudttts  Hook  &  .\rn. 
Dcscurainia  nchdidsonii  (Sweet)  Schidz  var.  sonnet 

(Robins.)  C.  L.  Hitchc. 
Erigeron  speciosus  (Lindl.)  D.  C.  var.  niacninthiis  (Nutt.) 

Croncj. 
Eriogonum  herarleoidcs  Nutt. 
Cilia  aggregata  (Pursh)  Spreng. 
Ligustiriim  filicinitni  Wats. 
Lnpinits  argenteus  Pursh  var.  ruhricanlis  (Greene) 

Welsh 
Pediciilahs  raceinosa  Dugl.  var.  alba  (Pennell)  Cronq. 
Poa  nervosa  (Hook.)  Vasey  var.  wheeleri  (Vasev)  C.  L. 

Hitchc. 
Potentilld  argiita  Pursh  var.  convallaria  (Hvdb.)  Tli.  Wolf 
Riidheekia  ocridenlidis  Nutt.  var.  ocridentcdis 
'^eneeio  crasstiliis  (Jrav 
Seneeio  serra  Hook. 
Stellaria  jamesiana  Torr. 
Trisetum  spicatum  (L.)  Richter 

Total  number  of  species  sampled 
Total  number  of  annual  species  sampled 
Total  number  biennial/short-lived  perennial  species 
sampled 

Herbaceous  standing  crop 
H.S.C.  -(-  tree  leaf  standing  crop 


1  (29%) 


25%) 


Rank 
3 


1  (25%) 


1  (26%) 

7 

3 

6 

2 
6 

41 

62 

58 

49 

8 

10 

7 

8 

2 

4 

3 

2 

100% 

2% 

0.1% 

0.06% 

10 


Great  Basin  Naturalist 


Vol.  40,  No.  1 


phylogenetic  and  functional  groups  are  sum- 
marized in  Table  4. 


Methods  of  Site  Analysis 

Field  studies  were  conducted  during  1976, 
1977,  and  1978.  From  1970  through  1976  air 
temperatures  and  precipitation  had  been  re- 
corded at  a  station  (hereafter  the  Badger  sta- 
tion) near  the  edge  of  a  small  meadow  on  the 
study  site  at  2650  m  elevation  (Fig.  2).  Mean 
monthly  values  recorded  in  Lomas  (1977) 
were  used  as  dependent  variables  in  multiple 
linear  regression  models  that  employed  as  in- 
dependent variables  the  values  for  the  same 
months  at  nearby  lower  elevation  stations,  for 
which  long-term  means  are  available.  For 
both  precipitation  and  temperature  models 
these  stations  are  Logan,  USU  (elevation 
1458  m,  27  km  SW  of  the  site)  and  Laketown 
(1825  m,  16  km  E-SE);  data  from  Richmond 
(1426  m,  25  km  W-NW)  were  also  used  in 
the  precipitation  models  (see  Fig.  1).  These 
data  are  published  in  the  corresponding  years 
of  Climatological  Data,  Utah  (U.S.  Weather 
Bureau,  U.S.  Department  of  Commerce, 
Washington,  D.C.).  The  published  mean  val- 
ues for  these  stations  for  the  1941-1970  peri- 
od were  entered  into  the  appropriate  regres- 
sion models  to  produce  an  estimated  monthly 
mean  temperature  or  precipitation  total  for 


MAMMALS 
1976-77 


BIRDS 
1976 


BIRDS 
1977 


Fig.  5.  Diagrams  showing  degree  of  affinity  among 
vertebrate  assemblages  in  the  four  serai  stages.  Number 
between  stages  is  the  Jaccard  coefficient  of  community. 
M  =  meadow,  A  =  aspen,  F  =  fir.  S  =  spruce-fir. 


T.\BLE  2.     .Mammal  species  observed  or  trapped  in  various  examples  of  the  serai  stages  at  the  School  Forest  from 
1976  through  1978. 


Meadow 


.\spen 


Fir 


Spruce-Fir 


Cervus  canadensis  (Elk) 

Cletliriotwinys  ^apperi  (Boreal  Redback  Vole) 

Erctliizon  dorsatuin  (Porcupine) 

Eutamki.s  niiniiutt.s  (Least  Chipmunk) 

Eiitaiiiias  tind)rinus  (Uinta  Chipmunk) 

Glaucomijs  sabrinus  (Northern  Flying  Scjuirrel) 

Lagurus  curtatus  (Sagebrush  Vole) 

Lepus  umericanus  (Snowshoe  Hare) 

Mu.stela  crminea  (Shorttail  Weasel) 

Mustela  frenata  (Longtail  Weasel) 

Neotoma  cinereu  (Bushytaii  Woodrat) 

Odocoileus  heiniontts  (Mule  Deer) 

Pcromijsnts  manicttlatus  (Deer  Mouse) 

.Sorc.v  sp.  (Shrew) 

Spcnnophilus  latemlis  (Colden-maiitled  Stjuirrei) 

Tainiascittrus  hudsoninis  (Red  Sfjuirrel) 

Ttiomomys  talpoidcs  (Northern  Pocket  Copher) 

Zapus  princeps  (Western  Jumping  .Mouse) 

Total 


X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

■p 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

13 


16 


14 


1.3 


March  1980 


SCHIMPF    ET  AL.:  SpRUCE-FiR  SUCCESSION 


11 


the  Badger  station  for  the  same  30-vear  peri- 
od. 

Air  temperature,  relative  humidity,  and 
precipitation  were  recorded  and  pan  evapo- 
ration measured  during  the  summers  of  1976, 
1977,  and  1978  at  two  other  stations  (eleva- 
tion 2560  m)  1.5  km  \-NE  of  the  Badger  sta- 
tion. One  station  was  located  in  a  meadow 
about  20  m  from  the  edge  of  an  aspen  stand; 
an  identical  one  was  situated  about  80  m  into 
the  forest  from  the  first,  under  a  well-devel- 
oped spnice-fir  canopy.  These  are  referred  to 
as  Big  Meadow  (meadow)  and  Big  Meadow 
(conifer),  respectively.  Instruments  used  were 
tipping  bucket  recording  precipitation 
gauges  (unshielded),  recording  hvgrothermo- 
graphs  in  standard  shelters,  and  U.S.  Depart- 
ment of  Agriculture  Class  A  evaporation 
pans.  Pan  evaporation  was  measured  at  ir- 
regular intervals  which  averaged  about  7 
days.  The  instruments  were  mounted  on  a 
platform  about  2.5  m  above  ground,  higher 
than  normal  because  of  the  deep  snows 
which  cover  the  area.  Snowpack  dynamics 
were  inferred  from  periodic  observations  at 
the  site,  in  conjunction  with  measurements 
made  by  the  U.S.  Department  of  Agriculture, 
Soil  Con.servation  Service,  at  Garden  City 
Summit,  8  km  north  of  the  site  (elevation 
2400  m)  (Fig.  1). 

Stem  xylem  water  potential  (predawn)  of 
conifers  along  a  10  X  50  m  transect  from 
meadow  through  aspen  to  conifer  was  mea- 
sured with  a  FMS  pressure  bomb  (Waring 
and  Cleary  1967)  on  3  August  1977.  At  least 
three  different  branches  of  each  spruce  or  fir 
were  measured;  these  trees  were  all  about  1 
m  tall.  Tree  species  population  structure 
along  this  same  transect  was  assessed  by 
aging  stems  and  estimating  their  heights. 
Ages  were  determined  by  counting  terminal 
bud  scale  scars  or  annual  growth  rings  in  xy- 
lem cores.  In  the  latter  case,  the  age  at  the 
coring  height  (1.37  m)  was  corrected  to  total 
age  by  adding  5  years  for  aspen  and  20  years 
for  fir  or  spruce,  values  based  on  average 
number  of  bud  scale  scars  at  that  height. 

Soil  pits  were  dug  to  1  m  depth  in  two  ex- 
amples of  each  stage  of  the  sequence  in  the 
same  stands  being  sampled  for  other  ecosys- 
tem attributes.  Each  soil  horizon  was  identi- 
fied, measured,  and  sampled  prior  to  phvsical 
and  chemical  analvsis  in  the  laboratory.  Per- 


cent coarse  fragments  was  estimated  bv  eye 
in  the  field.  Composition  of  the  fine  particle 
fraction  was  determined  by  the  hydrometer 
method  and  with  sieves.  The  following  chem- 
ical parameters  were  measured:  pH  (saturat- 
ed paste),  organic  carbon  (Walkley-Black 
procedure),  cation  exchange  capacity  (sodium 
acetate  method),  total  cations  (ammonium 
acetate),  potassium  and  phosphorous  (sodium 
bicarbonate),  iron  (DTPA  extraction),  and  to- 
tal nitrogen  (Kjeldahl).  Each  of  the  eight  pe- 
dons  studied  was  classified  to  the  Great 
Group  level  using  the  system  of  USDA  Soil 
Conservation  Service  (Anonymous  1975). 

Soil  moisture  was  monitored  in  1977  and 
1978  at  the  Badger  station  as  a  continuation 
of  previous  work  (Lomas  1977).  Volume  per- 
cent moisture  was  measured  with  a  Troxler 
neutron  probe  in  five  tubes  in  a  meadow  and 
ten  tubes  in  a  spruce-fir  stand.  Values  were 
recorded  for  each  30.5  cm  increment  of  a  122 
cm  deep  profile. 

Results 

Estimated  1941-1970  monthly  mean  tem- 
peratures and  precipitation  totals  for  the 
Badger  station  are  reported  in  Table  5,  along 
with  the  multiple  coefficient  of  determina- 
tion for  the  regression  model  by  which  each 
estimate  was  derived.  These  estimates  are 
plotted  against  the  monthly  means 
(1941-1970)  for  three  weather  stations  in  the 
Rocky  Mountain  subalpine  zone  with  long- 
term  records  (Figs.  6,  7).  Most  of  the  precipi- 
tation at  Badger  is  received  as  snow,  with 
nearly  30  percent  of  the  annual  precipitation 
falling  in  December  and  January.  Rainfall  de- 
creases to  very  low  amounts  in  Julv,  rising 
somewhat  in  August.  Long-term  snowpack 
records  for  Garden  City  Summit  are  present- 
ed in  Figure  8.  Meadow  peak  snowpack 
depths  at  the  study  site  averaged  about  40-50 
cm  greater  than  those  at  Garden  City  Sum- 
mit during  the  corresponding  winters. 

Selected  temperature,  humidity,  and  pre- 
cipitation data  from  the  Big  Meadow  station 
for  the  smnmers  of  intensive  studv  are  pre- 
sented in  Table  6.  Pan  evaporation  for  these 
summers  is  plotted  in  Figure  9  against  days 
after  snowmelt  in  the  meadows;  these  can  be 
converted  to  calendar  dates  from  the  snow- 
melt   dates   in  Table   6.   The   evaporation 


12 


Great  Basin  Naturalist 


Vol.  40,  No.  1 


curves  were  integrated  planimetrically  over 
various  time  intervals  (Table  7).  Over  the  in- 
terval from  43  to  73  days  after  snowmelt, 
evaporation  in  the  conifer  forest  averaged  38 
percent  (1976),  46  percent  (1977),  and  46 
percent  (1978)  as  great  as  that  in  the  mead- 
ow. Soil  moisture  trends  at  the  Badger  station 
during  the  three  summers  of  intensive  study 
are  represented  in  Figure  10. 

From  these  data  we  can  generally  charac- 
terize the  three  summers  during  which  eco- 
system properties  were  analyzed.  During 
1976  perhaps  the  most  favorable  conditions 
for  plant  growth  and  development  occurred, 
with  above  average  July  rain,  low  evapo- 
ration, and  a  moderately  long  frost-free  peri- 
od. The  summer  of  1977  followed  an  extraor- 
dinarily dry  winter,  resulting  in  low  soil 


moisture  content.  The  growing  season  began 
very  early  and  was  much  longer  than  for  the 
other  two  summers.  June  and  August  were 
warmer  than  in  1976  and  1978,  but  not  July. 
Pan  evaporation  was  very  high,  perhaps  in 
part  because  it  was  measured  during  the 
longer  days  of  early  summer,  since  the  satura- 
tion deficits  were  not  especially  great.  Ex- 
tremely high  August  rainfall  resulted  from  a 
rare  deep  continental  intrusion  of  a  tropical 
storm.  The  1978  frost-free  season  began  late 
and  ended  early.  Temperatures  and  evapo- 
ration rates  were  intermediate,  and  satura- 
tion deficits  were  relatively  low  in  June  and 
July  but  high  in  August.  Less  rain  fell  than 
during  the  other  two  summers. 

Tree  water  potential  and  forest  height 
structure    along    the    meadow-aspen-conifer 


Table  3.     Status  of  avian  species  observed  on  study  areas  in  1976  and  1977.  P  =  permanent  resident;  B  =  sum- 
mer breeder;  F  =  feeder  in  serai  stage,  but  not  breeder;  V  =  mitirating  or  wandering  visitor;  W  =  winter  resident. 


Species 


Meadow 


Aspen 


Fir 


Spruce-Fir 


Turkey  Vulture 

Goshawk 

Cooper's  Hawk 

Sharp-shinned  Hawk 

Marsh  Hawk 

Red-tailed  Hawk 

Golden  Eagle 

Sparrow  Hawk 

Blue  Grouse 

Ruffed  Grouse 

Mourning  Dove 

CJreat  Horned  Owl 

Pygmy  Owl 

Common  Nighthawk 

Poor-will 

Broad-tailed  Hununingl)ird 

Rufous  Hiuiimingbird 

Red-shafted  Flicker 

Yellow-bellied  Sapsucker 

WilHamson's  Sapsucker 

Hairy  Woodpecker 

Downv  Woodpecker 

Northern  Three-toed  Woodpecker 

Lewis"  Woodpecker 

Dusky  Flycatcher 

Western  Wood  Pewee 

Olive-sided  Flycatcher 

Violet-green  Swallow 

Tree  Swallow 

Steller's  Jay 

Black-billed  Magpie 

Clark's  Nutcracker 

Common  Raven 

Black-capped  C^hickadee 

Monntairi  (Chickadee 


rare  permanent 

resident 

rare  permanent 

resident 

rare  permanent 

resident 

V 

V 

V 

V 

B 

V 

rare  permanent 

resident 

V 

V 

W 

W 

W 

P 

W 

W 

F 

V 

V 

V 

p 
p 

p 

V 

V 

V 
V 

F 

B 

V 

V 

F 

F 

V 

V 

F 

B 
B 

B 

V 

B 

B 

P 

P 

P 

B 

V 

P 

V 

V 

B 

B 

B? 

B 

B 
B 

B 

V 

F 

B 

P 

P 

V 

V 

V 

V 

V 

V 

V 

V 

V 

V 

B 

P 

P 

March  1980 


SCHIMPF    ET  AL.:  SpRUCE-FiR  SUCCESSION 


13 


transect  is  plotted  in  Figure  11,  and  age 
structure  in  Figure  12.  For  tabulated  age 
data,  see  Daniel  et  al.  (1979:286).  It  can  be 
seen  that  the  water  stress  of  conifers  one  to 
two  meters  tall  increases  as  the  aspen  is  re- 
placed bv  niatiue  spruce  and  fir. 

The  eight  pedons  sampled  are  character- 
ized in  Table  8.  Their  taxonomic  assignments 
may  be  interpreted  as  follows:  Cryochrepts 
are  soils  with  cold  mean  annual  temperatures 
showing  little  development.  Crvoborolls  have 
dark  surfaces,  cold  mean  annual  temper- 
atures, and  little  development  of  the  profile. 
Paleboralfs  are  similarlv  cold,  but  have  an  ar- 
gillic  horizon  (accumulation  of  clav)  deep  in 
the  profile.  This  extensive  development  is  in- 
ferred to  be  the  product  of  a  different  cli- 
mate in  the  past. 

Cone  crops  on  the  School  Forest  have  been 
estimated  annually  since  1947  (T.  W.  Daniel, 

Table  3  coutiuuccl. 


pers.  comm.).  In  1976  fir  and  spruce  had  very 
low  cone  abundance,  but  in  1977  both  bore 
heavy  crops.  In  1978  the  spruce  crop  was 
very  low,  and  fir  bore  a  moderate  crop. 

Discussion 

The  greater  similarity  of  summer  air  tem- 
peratures than  of  precipitation  patterns 
among  the  geographically  disparate  spruce- 
fir  sites  in  Figures  6  and  7  lends  support  to 
the  assertion  by  Daubenmire  (1956)  that 
lower  growing  season  temperatures  dis- 
tinguish this  zone  from  other  Rocky  Moun- 
tain forests.  A  variety  of  precipitation  re- 
gimes permits  the  existence  of  spruce-fir 
forests,  from  the  summer-dominated  precipi- 
tation south  and  east  in  the  Cordillera  to  the 
winter-dominated  patterns  north  and  west. 

Few  other  data  on  humidity  or  pan  evapo- 


Species 


Meadow 


.\spen 


Fir 


Spruce-Fir 


White-breasted  Nuthatch 
Red-breasted  Nuthatch 
Brown  Creeper 
House  Wren 
.\merican  Robin 
Townsend's  Sohtaire 
Hermit  Thrush 
Mountain  Bluebird 
Colden-crowned  Kinglet 
Ruby-crowned  Kinglet 
Northern  Shrike 
Warbling  \'ireo 
Orauge-Crowued  Warbler 
Audubon's  Warbler 
MacGillivray's  Warbler 
Wilson's  Warbler 
Townsend's  Warbler 
Western  Meadowlark 
Brewer's  Blackbird 
Western  Tanager 
Black-headed  Grosbeak 
lazuli  Bunting 
Cassin's  Finch 
.\merican  Goldfinch 
Pine  Grosbeak 
Gray-crowned  Rosv  Finch 
Pine  Siskin 
Red  Crossbill 
White-winged  Crossbill 
Green-tailed  Towhee 
\  esper  Sparrow 
IDark-eyed  Junco  (3  races) 
Chipping  Sparrow 
Brewer's  Sparrow 
White-crowned  Sparrow 
Lincoln's  Sparrow 


B 

B 

P 

P 

B 

P 

B 

B 

B 

B 
V 

B 

B 

B 

W 

P 

V 

B 
V 

B 

B 

B 

B 

V 

V 

B 

B 
V 

B 

V 

V 

V 

B 

V 

B 

B 

B 

V 

B 

B 

V 

w 

P 

P 

V 

B 

B 

B 

B 

B 

V 

B 

V 

V 

B 

F 

B 

B 

B 

F 

B 

B 

B 

B 

V 

B 

B 
V 

V 

14 


Great  Basin  Naturalist 


Vol.  40,  No.  1 


ration  in  this  zone  have  been  pubHshed.  The 
three-year  monthly  averages  for  saturation 
deficit  at  Big  Meadow  (Table  6)  are  9  per- 
cent greater  (June),  53  percent  greater  (July), 
and  28  percent  greater  (August)  than  those 
we  computed  from  one  year  of  temperature 
and  humidity  records  for  a  spnice-fir  site  in 


the  Front  Range  in  Colorado  (Marr  1967). 
Pan  evaporation  rates  in  1918  and  1919  in 
the  subalpine  of  Arizona  (Pearson  1931)  were 
far  less  than  ours  (Table  7)  in  both  forested 
and  exposed  sites.  The  Arizona  site  receives 
large  amounts  of  summer  rain,  and  may  expe- 
rience lower  saturation  deficits.  Summer  davs 


o 

o 
UJ 

a: 

UJ 
Q. 

u 


—  BRIGHTON 
--B--  BADGER 
LEADVILLE 

—  -SUMMIT 


Fig. 
zones: 
mates; 


JFMAMJJASOND 


6.  Mean  niontlilv  temperatures  during  1941-1970  period  for  four  sites  in  the  Rocky  .Mountain  spruce-fir 
Badger  station,  Utah;  Brighton,  Utah;  Leadville,  Colorado;  Summit,  Montana.  Means  for  Badger  are  esti- 
those  for  the  other  sites  are  measured  parameters. 


Table  4.  Relative  abundance  of  phylogenetic  and  fimctional  groups  of  soil  microorganisms.  Total  number  of  or- 
ganisms in  the  phylogenetic  and  functional  group  coimts  do  not  agree  because  different  methods  were  used;  func- 
tional groups  are  also  not  nuitually  exclusive.  Soil  samples  are  from  topmost  5  cm. 


l(P  organisms 
g"^  dry  substrate 

Percent  of  total 

Aerobic  bacteria 

Streptonivcetes 

Fungi 

Anaerobic  bacteria 

Sample  type 

Summer/Fall 

Summer/Fall 

Summer/Fall 

Summer/Fall 

Summer/Fall 

Meadow  soil 

2.9/5.6 

66/64 

32/34 

2/1 

1/1 

-\spen  soil 

4.9/9..3 

68/68 

29/29 

3/2 

0.2/0.2 

Fir  soil 

7.7/14.7 

77/78 

20/20 

3/2 

0.2/ <  0.1 

Spruce-fir  soil 

7.5/10.7 

77/70 

20/26 

3/4 

<0.1/<0.1 

Fir  litter 

.39.6/28.0 

86/84 

13/14 

1/2 

<0.1/<0.1 

Spnice-fir  litter 

15.0/22.0 

77/84 

21/13 

2/2 

<0.1/<0.1 

March  1980 


SCHIMPF    ET  AL.:  SpRUCE-FiR  SUCCESSION 


15 


15 


1 


BRIGHTON 


o     lOH 


I- 
< 


^      5- 

a: 
a. 


0- 


LEADVILLE 


F 


M 


A 


M 


A      S      0      N      D 


Fig 
zone 


^.7.   Mean  monthly  precipitation  totals  during  1941-1970  period  for  four  site.s  in  the  Rockv  Mountain  spruce-fi 
Badger  station,  Utah;  Brighton.  Utah;  Leadville,  Colorado;  Summit,  Montana.  Means  for  Badger  are  estimates 
those  tor  the  other  sites  are  measured  parameters. 


Table  4  continued. 


Proteolytic 
Summer/Fall 


l(fi  organisms  g-^  dry  substrate 


Hemicellulolytic 
Summer/Fall 


Chitinolytic 
Summer/Fall 


Lipolytic 
Summer/Fall 


Cellulolytic 
Summer/Fall 


0.8/1.3 
0.8/.3.0 
1.5/1.8 
1.2/1.4 
3.6/3.7 
1.8/2.0 

0.1/0.02 

0.3/0.08 

0.9/0.4 

1.0/0.4 

2.6/1.3 

1.2/1.4 

0.3/0.6 
0..3/0.7 
0.4/0.3 
0.4/0.3 
0.6/0.5 
0.6/0.4 

0.08-0.04 
0.2/0.07 
0.3/0.1 
0.3/0.1 
0.4/0.8 
0.4/0.2 

0.1/0.01 

().()6/().()3 

0.07/0.09 

0.09/0.06 

0.2/0.2 

0.3/0.2 

16 


Great  Basin  Naturalist 


Vol.  40,  No.  1 


80 

^^ 

E 

e 

60 

5  40- 
o 

Q. 
< 

u   20 


< 
a. 


40 


—J— 
50 


60 


70 


80 


90 


00 


DAYS   AFTER  SNOWMELT 


Fig.  8.  Pail  evaporation  at  the  Big  Meadow  conifer  (C)  and  meadow  (M)  stations  during  summers  1976,  1977, 
1978.  Time  is  expressed  as  days  after  snowmelt  in  the  meadow.  Evaporation  is  plotted  as  mm  wlr*  at  the  midpoint 
of  the  measuring  period,  with  curves  fitted  bv  eve.  Ciuves  not  labeled  bv  vear  are  three-vear  averages. 


a. 

LU 
Q 

O 

<: 

Q_ 
O 
C/) 


200  1 


1  50  - 


100  - 


50 


DEC 


JAN 


FEB 


MAR 


APR 


MAY 


Fig.  9.  Time  course  of  snowpack  depths  at  ('.arden  City  Summit.  Broken  lines  indicate  records  from  the  196()s, 
and  solid  lines  indicate  records  from  the  197()s.  Years  labeled  are  those  in  which  the  particular  snowpack  melted. 


March  1980 


SCHIMPF    ET  AL.:  SpRUCE-FiR  SUCCESSION 


17 


would  also  be  shorter  there.  These  com- 
parisons, coupled  with  the  paucity  of  summer 
rain  at  our  site  (Fig.  7),  lead  us  to  venture 
that  ours  is  a  comparatively  dry  example  of 
spruce-fir  forest. 

Our  pan  evaporation  statistics  indicate  that 
animals  and  plant  shoots  in  the  lower  synusia 
of  the  forest  should  have  lower  rates  of  water 
loss  than  those  in  the  meadows  (Table  7,  Fig. 
8).  Because  saturation  deficits  were  nearly 
the  same  at  both  Big  Meadow  stations,  the 
differences  nnist  be  largely  due  to  differences 
in  net  radiation  or  mean  wind  velocity.  The 
evaporation  rates  measiued  in  the  meadow 
compare  favorably  with  those  estimated  for 
tlie  Badger  station  in  1970  bv  use  of  Pen- 
mans  combination  method  (Eaton  1971). 

Conifers  of  comparable  small  size  exliibit 
greater  water  stress  in  conifer-dominated 
stands  than  in  aspen  stands  (Fig.  11).  We  in- 
terpret this  to  mean  that  competition  for  soil 
water  from  large  conifers  is  more  severe  than 
from  aspen  or  small  conifers.  From  the  pan 
evaporation  data  (Fig.  8)  we  would  expect 
greater  evaporative  stress  under  aspen  rather 
than  conifers,  so  the  low  xylem  potentials  un- 
der conifers  must  be  due  to  inability  to  ob- 
tain sufficient  soil  water.  This  is  probably  an 
important  factor  limiting  recruitment  in  ma- 
tiue  spruce-fir  stands. 

The  tree  population  age  structures  along 
the  transect  (Fig.  12)  indicate  that  aspen  in- 
vades the  meadows  at  an  average  of  about  19 
cm  yr',  and  is  followed  in  about  20  years  by 
successful  conifer  establishment.  The  results 


o 


o 
> 


50 
25 


0-305  cm 


0 
50-1 

-   25- 


50 


25 


A.      ^^M77 


^C  __M.C76 


30.5-61.0  cm 


^^ M77 


^^    C76 


"I r 


:)U- 

^   M78       61.0-91.5  cm 

'^?-^^^^^=^'^^^-i'»S3E_,^^^J^J^ 

M76 

25- 

^077 

C76 

0^ 

T r    1 1 

1 

JUN        JUL       AUG       SEP       OCT 

Fig.  10.  Soil  moisture  in  the  meadow  and  conifer 
stages  at  Badger  station  during  the  summers  of  1976, 
1977,  and  1978.  Volume  percent  water  content  is  ex- 
pressed for  four  .successive  30.5  cm  horizons  of  the  soil 
profile. 


T.\BLE  5  Estimated  mean  monthly  temperatures  and  monthly  precipitation  totals  for  the  Badger  station  during 
the  period  1941-1970.  R^  is  the  coefficient  of  determination  for  the  multiple  regression  model  bv  which  each  value 
was  estimated. 


Month 


Precipitation,  cm 

R2 

13..5 

0.74 

8.9 

0.03 

12.7 

0.73 

10.4 

0.39 

7.1 

0.81 

5.2 

0.72 

1.2 

0.61 

3.0 

0.68 

3.4 

0.95 

6.2 

0.68 

9.2 

0.90 

14.3 

0.73 

Temperature,  °C 


R2 


[anuarv 

Fehniarv 

March 

April 

May 

June 

Julv 

.\ugust 

September 

October 

.November 

December 

Annual  total  or  mean 


95.1 


-10.2 
-9.0 

-5.3 

-0.2 

2.9 

8.7 

14.5 

13.4 

8.6 

3.9 

-5.0 

-9.7 

1.1 


0.64 
0.55 
0..39 
0..38 
0.69 
0.79 
0.45 
0.96 
0.61 
0.53 
0.62 
0.79 


18 


Great  Basin  Naturalist 


Vol.  40,  No.  1 


presented  are  in  general  agreement  with  oth- 
er transects  on  the  study  site.  Altered  slopes 
in  the  aspen  age  vs.  transect  distance  rela- 
tionship suggest  that  advance  of  the  clones 
was  slowed  during  the  1930s  and  accelerated 
in  the  past  decade.  The  1930s  were  generally 
warmer  and  drier  than  the  succeeding  period, 
and  the  precipitation  since  1970  (Lomas 
1977)  generally  exceeded  the  1941-1970  esti- 
mated means. 


Aspen  dominates  a  plot  on  the  School  For- 
est for  100  to  150  years  before  spruce  or  fir 
assumes  dominance  (Fig.  12)  and  hastens  the 
demise  of  aspen  through  shading.  Persistence 
of  occasional  aspen  stems  in  more  open  por- 
tions of  the  spruce-fir  canopy  is  important 
from  the  standpoint  of  regeneration  poten- 
tial. The  transition  from  aspen  to  spruce-fir 
occurs  over  a  relatively  short  time  and  dis- 
tance (Fig.   12),  explaining  why  we  were 


i2  -20n 


<  -15- 


LU 
O 


_J 

X 


-10- 


LU 

^    -5i 


0 


spruce 


Spruce  y 


oFlrV 


r30 


h20    E 

X 
LlI 

hlO    ^ 


10 


To 


30^ 


— I — 
40 


— I — 
50 


DISTANCE  FROM  MEADOW     (m) 

Fit;.  11.  Average  tree  height  of  aspen,  fir,  and  spruce  populations,  and  water  potential  of  short  conifers  along  a 
meadow-aspen-conifer  transect.  Water  potential  (^)  of  fir  and  spruce  of  about  one  meter  height  was  measured  .3 
August  1977.  Values  are  plotted  against  distance  from  the  edge  of  a  meadow. 


Table  6.  Climatological  data  from  Big  Meadow  weather  stations.  Precipitation  is  average  of  meadow  and  conifer 
station  totals.  Snowmelt  is  first  date  when  meadow  snow  cover  v.as  estimated  to  be  less  than  10  percent.  Saturation 
deficit  is  computed  from  weekly  mean  temperature  and  mean  relative  humidity  for  the  meadow  station. 


.\ir  tempe 

ature 

,  °c 

Snowmelt 

Mean 

High 

Stage/year 

June 

July 

August 

Meadow 

1976 

7.8 

13.7 

9.9 

24.7 

June  4 

1977 

10.9 

12.7 

11.7 

23.3 

Mav  15 

1978 

- 

12.6 

10.9 

25.6 

June  16 

Conifer 

1976 

8.7 

14.6 

10.6 

27.8 

1977 

12.0 

13.9 

12.4 

26.7 

1978 

8.8 

13.9 

12.0 

26.1 

March  1980 


SCHIMPF    ET  AL.:  SpRUCE-FiR  SUCCESSION 


19 


unable  to  find  mixed  aspen-conifer  stands 
sizeable  enough  to  study.  Tabulation  of  the 
ages  of  trees  by  species  indicates  that  the  in- 
vasion of  spruce  and  fir  occurs  with  less  tem- 
poral regularity  than  aspen  invasion  of  mead- 
ows (Daniel  et'al.  1979:286). 

The  great  similarity  in  soil  physical  and 
chemical  properties  among  the  four  stand 
types  (Table  8)  strengthens  our  view  that  the 


stands  differ  mainly  as  a  consequence  of  suc- 
cession and  not  because  of  differences  in  site 
potential.  Most  of  the  edaphic  differences  ex- 
isting among  stages  are  manifested  near  the 
soil  .surface,  reflecting  the  influences  of  the 
current  resident  biota.  The  absence  of  char- 
coal in  the  upper  few  cm  leads  us  to  con- 
clude that  fire  has  not  been  a  major  force  in 
the  meadow  ecosystems  during  the  past  few 
centuries. 


10  20  30  40 

DISTANCE    FROM  MEADOW    (m) 


Fig.  12.   .\ge  stnicture  of  aspen,  fir,  and  spruce  populations  along  a  nieadow-aspen-tonifer  transect.  Mean  ages  are 
plotted  against  distance  from  the  edge  of  a  meadow. 


Table  6 

continued. 

Last 

First 
freeze 

Precipitation,  cm 

Saturation  deficit, 

g  nr3 

freeze 

July                    August 

June 

July 

August 

June  26 
May  .30 
June  26 

June  27 
May  31 
Jime  26 

Aug.  26 
Aug.  28 
Aug.  16 

Aug.  26 
Aug.  27 
Aug.  15 

4.0                        2.3 
2.2                       12.7 
0.5                       3.5 

5.10 
4.42 
3.54 

7.49 
5.65 
5.21 

4.35 
5.13 
5.49 

20 


Great  Basin  Naturalist 


Vol.  40,  No.  1 


We  must,  then,  invoke  reasons  other  than 
fire  for  the  existence  of  the  meadows  we 
studied.  We  .suggest  that  we  are  .seeing  the 
disappearance  of  subalpine  meadows  that 
were  chmatic  chmax  during  a  colder  period. 
If  this  is  true,  then  the  sequence  we  describe 
is  botii  allogenic  (warmer  temperatures  al- 
lowing forest  expansion)  and  autogenic  (cli- 
max forest  establishment  facilitated  by  aspen 
invasion)  in  character.  It  is  important  to  de- 
termine whether  these  meadows  are  old  or 
yoimg.  Even  though  phvsiognomy  and  ener- 
getics may  be  similar  in  pioneer  meadows 
and  mature  meadows,  the  life  history  charac- 
teristics of  the  organisms  and  the  structure  of 
the  communitv  mav  well  differ  due  to  the 
length  of  time  a  stand  has  been  in  existence 
(MacArthur  and  Wilson  1967). 

There  is  evidence  from  around  the  globe 
for  cooling  during  this  earlier  period  (Brav 
1971),  though  we  know  of  no  published  evi- 
dence for  colder  conditions  in  the  immediate 
vicinity  of  our  site.  A  dendrochronological 
study  of  a  site  375  km  SW  of  ours  indicates 
that  temperatures  were  cooler  than  at  pres- 
ent from  1541  to  1780  (LaMarche  and  Stock- 
ton 1974).  This  corresponds  to  the  dearth  of 
trees  older  than  275  years  on  our  site,  though 
their  absence  could  be  interpreted  as  the  re- 
.sult  of  insect  devastation  (Miller  1970). 
Whipple  and  Dix  (1979)  also  found  fewer 
.spmce  between  ages  of  300  and  400  years  in 
Colorado  than  they  expected. 

Whether  this  cooling  was  sufficient  to 
change  the  general  aspect  of  our  studv  site 
we  can  only  gue.ss.  There  is  general  agree- 
ment that  the  cooling  was  greatest  at  high 
latitudes  and  altitudes  (Bray  1971).  The  limit- 
ing summer  air  temperatures  for  normal 
growth  of  spruce  or  fir  are  not  well  under- 
stood; Wardle  (1968)  presented  evidence  that 
the  limit  lies  between  11  and  12  C  (July)  for 
spruce  in  Colorado.  A  July  decline  of  2.5  C 
(Table  5)  might  have  been  sufficient,  in  com- 
bination with  the  level  topography  of  our 
site,  to  produce  an  open  subalpine  parkland 
of  herbaceous  meadows  containing  scattered 
trees  (Billings  1969).  The  wide  dispersion  of 
old  conifer  individuals  on  the  School  Poorest  is 
reminiscent  of  the  tree  patterns  in  such  park- 
lands.  Forest  expansion  since  this  colder  peri- 
od may  have  been  anomalously  rapid;  if  our 
site  reflects  climatic  trends  in   llie   Rocky 


Mountains  as  a  whole,  then  the  years  1870  to 
1945  may  have  been  the  most  warm-moist 
75-year  period  since  1130  (Bradley  1976). 

In  conclusion,  we  would  like  to  reempha- 
size  the  integral  role  played  by  preclimax 
trees  in  the  structuring  of  the  climax  forest. 
The  successional  change  we  describe  is  not 
merely  a  consequence  of  the  passage  of  time 
and  the  differential  growth  rates  of  preclimax 
and  climax  species  (Dmry  and  Nisbet  1973). 
Species  characteristic  of  the  climax  are  not 
"present  but  inconspicuous"  early  in  succes- 
sion; they  establish  extremely  slowly  and  only 
on  the  north  margins  of  their  stands  in  the 
absence  of  aspen  or  lodgepole. 

This  is  illustrated  in  two  discrete  subalpine 
basins  we  have  observed.  Birch  Creek  and 
Summit  Creek  South  Fork,  17  km  W  of  our 
study  site.  Lodgepole  is  absent  throughout 
this  western  portion  of  the  mountain  range  at 
this  latitude.  These  two  basins  are  unusual  in 
that  they  are  also  devoid  of  aspen  in  the  sub- 
alpine zone.  Aspen  may  have  been  locally  ex- 
tirpated during  the  Pleistocene,  when  both 
basins  held  small  alpine  glaciers  (De  Graff 
1976).  Now  aspen  is  unable  to  invade  from 
adjacent  topographic  units  because  its  in- 
ability to  reproduce  by  seed.  Aspen  is  present 
at  lower  elevations  in  these  drainages,  but 
perhaps  only  as  the  late-leafing  form  which  is 
not  known  to  occur  at  higher  elevations  (Cot- 
tam  1954).  The  spruce-fir  stands  of  these  two 
topographic  units,  at  the  same  elevations  as 
our  study  site,  are  small  discrete  groves  in  a 


Table  7.  Estimated  pan  evaporation  rates  at  the  Big 
Meadow  stations,  expressed  as  mm  wk"'.  Means  for  vari- 
ous intervals  were  obtained  by  planimetric  integration 
ot  the  turves  in  Fisrure  8. 


Stage 

Period 

Rate 

Meadow 

1976,  4.3-73  days  after  melt 

40 

1977,  4.3-73  davs  after  melt 

.56 

1978.  4.3-73  days  after  melt 

43 

1976,  entire  curve 

39 

1977,  entire  ciuve 

50 

Three-year  mean,  entire  cur\e 

42 

(>oniter 

1976.  43-73  da>s  after  melt 

15 

1977,  43-73  davs  after  melt 

26 

1978,  43-73  days  after  melt 

20 

1976,  entire  curve 

16 

1977,  entire  curve 

23 

Three-year  mean,  entire  curve 

17 

March  1980 


SCHIMPF    ET  AL.:  SpRUCE-FlR  SliCC.ESSIOX 


21 


matrix  oi  herbaceous  veij;etation.  We  h\  poth- 
esize  that  this  pattern  represents  exactly  the 
result  we  would  expect  for  the  allo<Tenic  ex- 
pansion of  spruce-fir  forest  from  subalpine 
parkland  in  the  absence  of  aspen  or  lodge- 
pole. 

Acknowledgments 

The  following  contributed  data  or  other  in- 
formation used  in  this  paper:  D.  Andersen,  L. 
Bennett,  T.  Daniel.  D.  Factor,  J.  Habeck,  G. 
Hart,  P.  Lawton,  R.  Lawton,  G.  Reese,  M. 
Schwartz,  J.  Skujins,  K.  Smith,  R.  Sperger,  N. 
West,  M.  Wolfe,  D.  Zeeman.  Many  of  them 
provided  constructive  comments  on  an  ear- 
lier draft  of  this  paper  as  did  R.  Bayn  and  G. 
Waagen.  We  particulary  want  to  thank  T. 
W.  "Doc"  Daniel  for  sharing  his  wealth  of 
field  experience  in  the  Utah  State  University 
School  Forest.  R.  Bavn,  L.  Finchum,  and  B. 
Peitersen  were  instrimiental  in  the  prepara- 
tion of  the  manuscript.  This  research  was 
supported  by  Grant  DEB  78-05328  from  the 
National  Science  Foundation  to  James  A. 
MacMahon. 


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22 


Great  Basin  Naturalist 


Vol.  40,  No.  1 


Table  8.    Physical  and 

chemical  characteristics,  and 

classifications 

of  two 

pedons  in  each  of  four 

stages  of  succes- 

sion  in  the  School  Forest. 

Roots:  f  =  few 

,  c  =  common,  m   =   many. 

vf  = 

very  fine,  f  =   fine,  m 

=  mediiun,  c  = 

coarse. 

Cation 

Percent 

Percent 

Percent 

exchange 

sand, 

coarse 

organic 

capacity 

Location,  depth,  cm 

silt,  clay 

fragments 

Texture 

pH 

carbon 

meq/lOOg 

Big  Meadow  (Meadow  \) 

0-9 

.30,53,17 

30  gravel 
10  stones 

gravelly 

.silt 

loam 

63 

1.9 

14.2 

9-38 

30,53,17 

30  gravel 
10  stones 

gravelly 

silt 

loam 

6.0 

1.5 

15.7 

38-ft3 

33,47,20 

30  gravel 
10  stones 

gravelly 
loam 

5.8 

1.1 

13.8 

63-98 

45,.37,18 

40  gravel 
5  cobbles 

gravelly 
loam 

5.5 

0.3 

8.5 

98-100 

4633,21 

40  grave! 
5  cobbles 

gravelly 
loam 

5.4 

0.2 

9.5 

Doc's  Meadow  (Meadow  B) 

0-9 

35,45,20 

less  than  10 

loam 

6.3 

1.7 

14.8 

9-26 

.35,43,22 

less  than  10 

loam 

.5.8 

1.4 

14.2 

26-40 

.35,42,23 

10  gravel 

loam 

5.5 

0.7 

11.5 

40-67 

45,.33,22 

30  gravel 

gravelly 
loam 

5.2 

0.3 

10.1 

67-100 

58,26,16 

25  gravel 

gravelly 
fine 
sandy  loam 

5.3 

0.2 

62 

Big  Meadow  Fir  (Fir  A) 

0-20 

.33,56,11 

20  gravel 

gravelly 

.silt 

loam 

5.8 

1.7 

18.7 

20-62 

47,.38,15 

30  gravel 
10  cobbles 

gravelly 
loam 

5.7 

0.7 

9.1 

62-88 

.3.3,.5611 

30  gravel 
5  cobbles 

gravellv 

silt 

loam 

5.2 

0.5 

12.2 

88-100 

52,31,17 

40  gravel 

gravelly 
very  fine 
sandy  loam 

5.3 

0.3 

7.9 

Harts  Fir  (Fir  B) 

0-10 

43,4.3,14 

25  gravel 

gravelly 
loam 

5.7 

2.5 

9.9 

10-.35 

40,44,16 

30  gravel 

gravellv 
loam 

.5.8 

1.0 

8.8 

.35-62 

40,4.3,17 

.30  gravel 

gravelly 
loam 

.5.6 

0.8 

8.4 

62-100 

.37,.32,31 

30  gravel 

gravellv 
clay 

.5.4 

0.3 

19.4 

Big  Meadow  Aspen  (.\spen  A) 

4-0  litter 

0-6 


6-27 


organic 


.34,44,22 


organic 


,5.6 


30  gravel  gravelly  5.6 

20  stones  loam 


25.3 
1.7 


29.4 
19.4 


March  1980 


SCHIMPF    ET  AL.:  SPRUCE-FiR  SUCCESSION 


23 


Table  8  continued. 


64 


Percent 

base 

saturation         K,  ppm 


210 


P,  ppm 


16.3 


Fe,  ppm 


Roots 


Charcoal 


24 


nnf 
nif 


July  percent 

total 

nitrogen 

&  depth,  cm    Great  group 


0.11  top  3 


Crvohoroll 


54 


146 


52  69 

72  33 

72  44 


6.5 

3.9 
3.5 
3.1 


31 

40 
26 
24 


cvf 
cf 

cvf 

cf 

ff 

ff 


0. 1 1  25 


0.05  50 


0.03  100 


64 

128 

16.0 

.34 

cf 

0.(W  top  3 

54 

69 

7.7 

45 

ff 

+ 

0.06  25 

65 

51 

3.9 

55 

ff 

4- 

- 

42 

3.3 

47 

fvf 

+ 

0.05  50 

71 

22 

3.2 

37 

0.03  100 

Cr\oc]irept 


Paleboralf 


68 


73 


82 


87 


122 


50 


42 


31 


5.4 


5.0 


2.6 


34 


44 


20 


nivf 

inf.fin 

fc 

cvf 

cf.fm 

fc 

cvf 

cf 

ff 


1..35  litter 
0.19  top  3 

0.11  25 
0.07  50 


0.05  100 


Paleboralf 

- 

182 

42.4 

- 

cfjni 
fc 

+ 

1.. 39  litter 
0.19  top  3 

64 

200 

.37.7 

56 

cf.fm 
fc 

+ 

0.06  25 

63 

148 

32.8 

51 

ff.fm 
fc 

+ 

0.04  .50 

87 

161 

21.0 

32 

fm 

+ 

0.04  1(K) 

Crvoboroll 

62 


400  + 


192 


.30.9 


4.4 


34 


29 


invt 

mf 

mill 

nnf 

lilt. mill 

mc 


0. 1 1  top  3 


0.17  25 


24 


Great  Basin  Naturalist 


Vol.  40,  No.  1 


Kaufmann,  M.  R.,  and  a.  N.  Eckard.  1977.  Water  po- 
tential and  temperature  effects  on  germination  of 
Engelmann  spruce  and  lodgepole  pine  seeds.  For. 
Sci.  2.3:  27-33. 

La.Marche,  V.  C,  Jr.,  and  C.  W.  Stockton.  1974. 
Chronologies  from  temperature-sensitive  bristle- 
cone  pines  at  upper  treeline  in  western  United 
States.  Tree-ring  Bull.  34:  21-45. 

Langenheim,  J.  H.  1962.  Vegetation  and  environmental 
patterns  in  the  Crested  Butte  area,  Gunnison 
County,  Colorado.  Ecol.  Monogr.  32:  249-285. 

Table  8  continued. 


Lawton,  p.  M.  1979.  An  investigation  of  the  environ- 
mental relationships  of  selected  forest  habitat 
types  in  northern  Utah.  Unpublished  thesis,  Utah 
State  Univ.,  Logan. 

LeBarron,  R.  K.,  and  G.  M.  Jemison.  1953.  Ecology  and 
silviculture  of  the  Engelmann  spnice-alpine  fir 
type.  J.  For.  51:  349-355. 

Lo.viAS,  D.  A.  1977.  Soil  water  depletion  following  clear- 
cutting  small  plots  in  a  spnice-fir  forest  in  north- 
ern Utah.  Unpublished  thesis.  Utah  State  Univ., 
Logan. 


Cation 

Percent 

Percent 

Percent 

exchange 

sand, 

coarse 

organic 

capacity 

Location,  depth,  cm 

silt,  clay 

fragments 

Texture 

pH 

carbon 

meq/lOOg 

24-44 

46,39,15 

40  gravel 
10  cobbles 

gravelly 
loam 

5.6 

0.5 

10.1 

44-100 

54,29,17 

40  gravel 
5  stones 
5  cobbles 

gravelly 
fine 
sandy  loam 

5.5 

0.4 

10.1 

Hart's  Aspen  (.\spen  B) 

0-9 

43,43,14 

20  gravel 

gravelly 
loam 

.5.7 

1.5 

11.2 

9-27 

44,42,14 

20  gravel 

gravelly 
loam 

5.8 

1.3 

9.5 

27-49 

43,43,14 

.30  gravel 

gravelly 
loam 

.5.9 

0.7 

7.6 

49-75 

44,42,14 

30  gravel 
10  cobbles 

gravelly 
loam 

,5.6 

0.7 

7.3 

7,5-100 

42,4,5,13 

10  gravel 

loam 

5.3 

0.6 

12.8 

Big  Meadow  Spruce-fir  (Spruce-fir  A) 
3-0  litter 

0-19  ,39,48,13 


■30  gravel 


gravelly 
loam 


,5.8 


1.7 


12.9 


19-47 


47-72 


43,43,14 


62,18,20 


72-100  70,8,22 


Sinks  Road  Spruce-fir  (Spruce-fir  B) 
5-0  litter 

0-11  29,57,14 


30  gravel 
10  cobbles 
20  stones 
less  than  10 


gravelly 
loam 

coarse 
sandv 
clay  loam 
less  than  10  coarse 
sandy 
clay  loam 


15  gravel 


silt 
loam 


.5.5 


6.5 


5.0 


1.0 


0.4 


0.5 


1.3 


10.4 


7.1 


6.2 


II -,35 
,35-60 
60-85 
85-l(K) 


29,.57.14 
40,46,14 
43,44,13 
.54.,32,14 


20  gravel 

,30  gravel 
•5  cobbles 

,30  gravel 
5  cobbles 

40  gravel 
5  cobbles 


gravel  I V 
silt  loam 
gravelly 
loam 
gravelly 
loam 
gravelly 
very  fine 
sandv  loam 


,5.4 
5.4 
.5.4 


0.7 
0.6 
0.5 
0.3 


8.3 
7.7 
6.4 
5.1 


March  1980 


SCHIMPF    ET  AL.:  SpRUCE-FiR  SUCCESSION 


25 


LfivE,  D.  1970.  Subarctic  and  subaljMiie:  W'Iumc  and 
what?  Arc.  Alp.  Res.  2;  63-73. 

M.\c..\rthur,  R.  H.,  .a..\d  E.  O.  Wilson.  1967.  The  theo- 
ry' of  island  biogeography.  Princeton  Univ.  Press, 
Princeton,  New  Jersey. 

Marr.  ].  \\'.  1967.  Ecosystems  of  the  east  slope  of  the 
Front  Range  in  Colorado.  Univ.  Colorado  .Stu., 
Ser.  in  Biol.  \o.  8,  Bonlder,  Colorado. 

Miller,  P.  C.  1970.  .\ge  distributions  of  .spruce  and  fir 
in    beetle-killed    forests    on    the    White    River 

Table  8  continued. 


Plateau,  Colorado,  .^iner.  .Midi.  Natur.  83- 
206-212. 

Noble,  D.  L.  1979.  Roots  of  lodgepole  pine  seedlings 
reach  depth  of  only  3  to  4  inches  their  first  sea- 
son. U..S.  Dept.  .\gric.  For.  Serv.  Res.  Note  RM- 
.363. 

Noble,  D.  L.,  anp  R.  R.  Alexa.nder.  1977.  Environmen- 
tal factors  affecting  natural  regeneration  of  En- 
gelniann  spruce  in  the  central  Rockv  Mountains. 
For.  Sci.  23;  420-429. 


Percent 

l)ase 

saturation         K,  ppm 


P,  ppm 


Fe,  ppm 


Roots 


Charcoal 


July  percent 

total 

nitrogen 

&  depth,  cm    Great  group 


64 


54 


51 


4.8 


3.2 


36 


33 


cvf 
cf.fm 
fc 
ff 


0.09  50 
0.06  100 


81 
86 
75 
80 

.56 

51 

68 

61 


286 
220 
179 
172 
132 

142 

90 

48 

31 


35.0 
29.0 
25.6 
24.3 
5.6 

10.7 

9.0 

5.5 

4.2 


27 
34 
34 
42 
19 

54 

51 


28 


mf.nini 

-f- 

cf,cm 

+ 

cf,cm 

+ 

cf.cm 

+ 

ff,fm 

+ 

mvf 

mf,cm 

cc 

mvf 

mf,cm 

cc 

cvf 

cf.ftn 

fc 

ff 

Crvoboroll 
0.17  top  3 

0.06  25 

0.03  50 

0.03  1(K) 

1.29  litter         Cryochrept 
0.19  top  3 

0.07  25 

0.07  .50 

0.03  1(X) 


Cryochrept 


1.25  litter 

38 

93 

8.8 

69 

mvf 

mf,cm 

fc 

0.15  top  3 

.36 

64 

2.1 

40 

cvf 
cf.fc 

+ 

0.08  25 

39 

71 

1.4 

34 

cf.cm 

+ 

0.04  50 

47 

33 

1.1 

29 

cf.cm 

86 

.30 

1.2 

23 

cf,cm 

0,03  U)() 

26 


Great  Basin  Naturalist 


Vol.  40,  No.  1 


Noble,  D.  L.,  and  F.  Ronco,  Jb.  1978.  Seedfall  and  es- 
tablishment of  Engelmann  spruce  and  subalpine 
fir  in  clearcut  openings  in  Colorado.  U.S.  Dept. 
Agric.  For.  Serv.  Res.  Pap.  RM-200. 

Odum,  E.  p.  1969.  The  strategy  of  ecosystem  devel- 
opment. Science  164:  262-270. 

OosTiNG,  H.  J.,  AND  J.  F.  Reed.  1952.  Virgin  spruce-fir 
forests  in  the  Medicine  Bow  Mountains,  Wyom- 
ing. Ecol.  Monogr.  22:  69-91. 

Paden,  J.  W.,  J.  R.  Sutherland,  and  T.  A.  D.  Woods. 
1978.  Caloscypha  fulgens  (Ascomyce- 
tidae:Pezizales):  The  perfect  state  of  the  conifer 
seed  pathogen  Geniculodendron  pijrifomie  (Deu- 
teromycotina:  Hyphomycetes).  Can.  J.  Hot.  56: 
2.375-2.379. 

Patten,  D.  T.  1969.  Succession  from  sagebrush  to  mixed 
conifer  forest  in  the  northern  Rocky  Mountains. 
Amer.  Midi.  Natur.  82:  229-240. 

Pearson,  G.  A.  1931.  Forest  types  in  the  southwest  as 
determined  bv  climate  and  soil.  U.S.  Dept.  Agric. 
Tech.  Bull.  247. 

Pease,  J.  L.,  R.  H.  Vowles,  and  L.  B.  Keith.  1979.  In- 
teraction of  snowshoe  hares  and  woody  vegeta- 
tion. J.  Wildl.  Manage.  43:  4.3-60. 

Peet,  R.  K.  1978.  Latitudinal  variation  in  southern 
Rocky  Mormtain  forests.  J.  Biogeogr.  5:  275-289. 

Ronco,  F.  1970.  Influence  of  high  light  intensity  on  sur- 
vival of  planted  Engelmann  spruce.  For.  Sci.  16: 
.331-339. 

Sabo,  S.  R.,  and  R.  H.  Whittaker.  1979.  Bird  niches  in 
a  subalpine  forest:  .\n  indirect  ordination.  Proc. 
Natl.  .\cad.  Sci.  USA  76:1.3.38-1.342. 


ScHMiD,  J.  M.,  AND  R.  H.  Frye.  1977.  Spruce  beetle  in 
the  Rockies.  U.S.  Dept.  .\gric.  For.  Serv.  Gen. 
Tech.  Rep.  RM-49. 

ScHMiD,  J.  M.,  AND  T.  E.  Hinds.  1974.  Development  of 
spruce-fir  stands  following  spruce  beetle  out- 
break. U.S.  Dept.  Agric.  For.  Serv.  Res.  Pap.  RM- 
131. 

Smith,  K.  G.,  and  J.  A.  MacMahon.  Bird  communities 
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Stahelin,  R.  1943.  Factors  influencing  the  natural  re- 
stocking of  high  altitude  burns  by  coniferous 
trees  in  the  central  Rocky  Mountains.  Ecology 
24:  19-.30. 

Stokes,  W.  L.  1963.  Geological  map  of  northwestern 
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Veatch,  A.  C.  1907.  Geography  and  geology  of  a  por- 
tion of  southwestern  Wyoming.  U.S.  Geol.  Survey 
Prof.  Paper  56. 

Wardle,  P.  1968.  Engelmann  spruce  {Picea  engehnannii 
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Waring,  R.  H.,  and  B.  D.  Cleary.  1967.  Plant  moisture 
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1248-1254. 

Whipple,  S.  A.,  and  R.  Dix.  1979.  Age  structure  and 
successional  dynamics  of  a  Colorado  subalpine 
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Wicklow-Howard,  M.  C,  and  J.  Skujins.  1979.  In- 
fection of  Engelmann  spruce  seeds  by  Gen- 
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ca. Mycologia  (in  press). 


UTAH  FLORA:  MALVACEAE 

Stanley  L.  Welsh' 

.■\bstract.—  This  paper  is  the  third  in  a  series  dealing  with  a  revision  of  the  flora  of  Utah.  Treated  herein  are  9 
genera  and  23  species,  including  both  coniinonly  cultivated,  escaped,  and  indigenous  representatives.  Proposed  new 
taxa  include  Sphuemkcdnwssiilariifolia  (H.  and  \.)  Rydh.  var.  inoorei  Welsh,  Sphaeralcea  leptophijUa  (Gray)  Rydb. 
var.  jancac  Welsh,  and  Sphacnilcca  f)sorak>idcs  Welsh. 

This  third  paper  in  the  .series  leading  to  a  of  specimens  examined  by  me  is  indicated 
revision  of  the  flora  of  Utah  deals  with  the  following  the  discussion  of  each  species.  The 
small  but  significant  and  taxonomically  diffi-  number  in  parenthesis  is  the  number  collect- 
cult  Mallow  family.  Especially  complex  are  ed  by  me. 
members  of  the  genus  Sphaeralcea,  ably  mon- 
ographed by  T.  H.  Kearney  (1935),  and  re-  Malvaceae  Juss. 
viewed  for  Utah  by  J.  A.  M.  Jefferies  (1972).  Mallow  Familv 

As  with  previous  treatments,  the  work  con- 
siders not  only  indigenous  species  and  weeds  Herbs  or,  less  commonly,  shrubs,  usually 

or  established  escaped  species,  but  those  in-  pubescent  with  branched  or  stellate  hairs,  an- 

troduced  species  which  are  commonly  grown  nual,    biennial,    or    perennial,    with    mucila- 

as  ornamentals   or  for  other  uses.   Casually  ginous  juice;  leaves  alternate,  simple,  mostly 

grown  species,  such  as  the  okra.  Hibiscus  es-  palmately  veined,  stipulate;  flowers  perfect 

culentus  L.,  have  been  excluded.  Althaea,  Hi-  (or  imperfect),  regular,  solitary  or  in  thyrsoid 

hiscus,   and  Malta  are  included  entirely  on  cymes,  or  more  or  less  racemose  or  pan- 

the  basis  of  cultivated  ornamentals  and  weeds  iculate,  sometimes  with  an  involucel  of  sepal- 

which  have  become  established  in  the  state.  like  bractlets;  sepals  5,  more  or  less  per- 

Malva  neglecta  is  a  pest  of  cultivated  areas.  sistent;   petals  5,  separate,  adnate  to  the 

Iliamna,  Malvastrum,  Sida,  Sidalcea,  and  staminal  sheath;  .stamens  numerous,  united  by 

Spliaeralcea   are  represented  entirely  by  in-  the  filaments  (monadelphous);  ovary  superior, 

digenous  species.  Abutilon  has  one  species  in-  3-  to  many-loculed;  fruit  a  capsule  or  a  schiz- 

troduced  and  the  other  native.  The  number  ocarp. 

1.  Involucel  lacking  2 

—  Involucel  of  1   or  more  bractlets,  or  if  lacking  (as  in  some  Spliaeralcea 
specimens),  then  the  flowers  orange  (grenadine)  3 

2(1).       Petals  white,  pink,  or  lavender;  plants  of  moist  sites,  usually  at  middle  and 

higher  elevations Sidalcea 

—  Petals  yellow  or  pink  to  red;  plants  of  cultivated  lands  or  of  arid  sites,  u.sually 

at  lower  elevations  Abutilon 

3(1).        Petals   orange   or   rarely   purplish   pink;   indigenous   perennial  herbs   of  arid 

habitats  at  middle  and  lower  elevations  Spliaeralcea 

—  Petals  variously  colored,  but  not  orange;  indigenous  or  adventive  perennial, 
biennial,  or  annual  plants  or  various  distribution  4 

4(3).        Flowers  ro.se  pink  or  rarely  white;  plants  indigenous,  7-15  dm  tall,  perennial, 

of  middle  and  higher  elevations  Iliamna 


'Life  Science  Museum  and  Department  of  Botany  and  Range  Science,  Brigham  Young  University.  Provo.  Utah  84602. 


27 


28  Great  Basin  Naturalist  Vol.  40,  No.  1 

_  Flowers  white,  pink,  rose,  yellow,  or  other  hues;  plants  differing  in  one  or 

more  ways  from  above '^ 

5(4)        Flowers  mostly  6-10  cm  broad,  opening  flat;  plants  tall  adventive  or  cultivated 

biennials Althaea 

—  Flowers  less  than  6  cm  broad  or,  if  broader  then  the  plants  shrubby  6 

6(5).       Style  branches  5,  elongate;  fruit  a  capsule;  plants  low  annuals  or  shrubs 

Hibiscus 

—  Style  branches  more  than  5,  short;  fruit  a  schizocarp;  plants  annual  or  biennial 

7(6).       Style  branches  filiform,  with  elongate  stigmatic  lines;  plants  annual  or  biennial 

Malta 

—  Style  branches  with  capitate  or  truncate  stigmas  8 

8(7).       Petals  yellow,  or  orange  to  pink  or  red;  plants  annual  with  awned  carpels  or 

subshrubs  with  unawned  carpels Abutilon 

—  Petals  yellowish  white  to  lavender  or  whitish;  carpels  few  to  many,  not  awned; 
plants  spreading  annuals  or  herbaceous  perennials 9 

9(8).       Petals  yellow  white;  leaves  reniform-orbicular,  merely  crenate-serrate  Sida 

—  Petals  lavender  or  whitish;  leaves  palmately  cleft,  with  rounded  lobes 

Malvastrum 

.\butilon  Mill. 

leafy    panicles;    involucel    lacking;    calyx   5- 

Plants    herbaceous,    annual    or    perennial,  cleft;  corolla  yellow  to  orange  pink  or  red; 

with  stellate  or  simple  hairs;  leaves  alternate,  fruit    truncate-cylindric    or    subglobose,    the 

petioled,  cordate  at  base,  not  or  only  obscu-  carpels  smooth  sided,  dehiscent  nearly  to  the 

rely  lobed;  flowers  solitary  and  axillary  or  in  base;  ovules  2  or  more  per  carpel. 


Plants  perennial,  with  slender  spreading  or  trailing  branchlets;  carpels  5,  lacking 

awn-beaks;  plants  rare,  known  only  from  Washington  County  A.  parvidum 

Plants  annual,  with  robust  erect  stems;  carpels  usually  more  than  10,  each  with 

a  long  divergent  awn;  plants  uncommon,  in  agricultural  regions A.  theophrasti 


Abutilon  parvulum  Gray.  Perennial,  the 
stems  slender  and  spreading  or  trailing,  the 
caudex  woody,  grayish  tomentose  with  min- 
ute stellate  hairs,  the  branchlets  pilose;  leaves 
0.5-5  cm  long,  ovate,  cordate  basally,  den- 
tate and  sometimes  obscurely  3-lobed;  pe- 
duncles slender,  axillary,  1 -flowered,  longer 
than  the  leaves;  calyx  lobes  ovate-acuminate, 
reflexed  in  fruit;  petals  orange  pink  to  red  or 
sometimes  yellowish,  4-6  mm  long;  carpels  5, 
somewhat  tomentose,  to  8  mm  long.  Known 
in  Utah  only  from  Veyo,  Washington  County 
(Meyer  4111),  Colorado  to  California,  and 
south  to  Texas  and  Mexico,  1(0). 

Abutilon  theophrasti  Medic.  Velvet  leaf. 
Annual,  the  stems  robast,  erect,  velvety  and 
cinereous  with  short,  soft  hairs;  leaves  3-10 


cm  long  (from  sinus  to  apex)  and  as  broad  or 
broader,  orbicular-ovate,  cordate  at  the  base, 
abruptly  acuminate  at  the  apex,  velvety  pub- 
escent; peduncles  shorter  than  the  leaves,  one 
to  few  flowered;  calyx  lobes  broadly  ovate- 
acuminate;  petals  yellow,  to  about  6  mm 
long;  carpels  10  or  more,  each  with  a  long  di- 
vergent awn.  Adventive  weedy  species  of  dis- 
turbed or  cultivated  areas,  occasional  in  Utah 
and  Washington  counties  (to  be  expected 
elsewhere);  widespread  in  North  America; 
native  to  Europe;  3(0). 

Althaea  L. 

Plants   herbaceous,    biennial,    with    coarse 
stellate  hairs;   leaves  alternate,   petiolate. 


March  1980 


Welsh:  Utah  Flora,  Malvaceae 


29 


cordate  at  the  base,  lobed;  flowers  sohtarv  or 
in  racemes;  invohicel  of  6-9  bractlets,  con- 
nate at  the  base;  calyx  5-cleft;  corolla  of  vari- 
ous colors;  fruit  flattened  wheellike,  invested 
by  the  calyx,  the  numerous  carpels  separating 
at  maturity. 

Althaea  rosea  Cav.  Hollyhock.  Coarse 
biennials  to  20  dm  tall  or  more,  the  stems 
erect,  stellate-hairy;  leaves  (3-)  5-  to  7-lobed, 
mostly  3-15  cm  long  (from  sinus  to  apex)  and 
often  much  broader;  flowers  shortly  pedicel- 
late, 6-12  cm  wide  or  more,  variously  col- 
ored, often  rose  to  pink  or  lavender,  or  some- 
times white,  usuallv  with  a  dark  center;  calyx 
lobes  triangular,  investing  the  fruit  at  matu- 
rity, the  involucel  calyxlike;  carpels  numer- 
ous, stellate  along  the  margins,  and  reticulate 


on  the  sides,  5-7  mm  long.  Cultivated  orna- 
mental, persisting  and  escaping,  to  be  ex- 
pected in  all  counties  in  Utah;  widespread  in 
North    America;    introduced   from    China; 

15(0). 


Hibiscus  L. 

Plants  herbaceous  or  woody,  annual  or 
perennial,  with  stellate  or  simple  hairs;  leaves 
alternate,  petiolate,  obtuse  to  truncate  or 
cordate  basally,  lobed  to  incised;  flowers  ax- 
illary, solitary;  involucel  of  5-10  distinct 
bractlets;  calyx  5-cleft,  more  or  less  accres- 
cent in  fruit;  fruit  a  loculicidal  capsule,  the 
carpels  5;  seeds  several  in  each  locule. 


1.  Plants  annual;  calyx  strongly  veined;  petals  cream  colored,  with  a  purple  center 

H.  trionum 

Plants  shrubs;  calyx  herbaceous,  not  distinctly  veined;  petals  variously  colored, 

but  usually  rose  pink  to  lavender H.  si/riacus 


Hibiscus  syriacus  L.  Althaea;  Rose-of-Sha- 
ron.  Shrubs,  20-40  dm  tall  or  more,  glabrous 
or  softly  stellate-hairv;  leaves  2.5-8  cm  long, 
1.5-6  cm  wide,  triangular-ovate  to  rhombic, 
strongly  3-ribbed,  commonly  3-lobed;  flowers 
axillary,  4-7.5  cm  wide;  bractlets  usually  5, 
linear,  about  as  long  as  the  calyx,  glabrous  to 
obscurely  hairy;  corolla  variously  colored  and 
often  double;  fruit  oblong-ovoid,  to  25  mm 
long.  Cultivated  ornamental,  rarely  per- 
sisting; widely  cultivated  in  North  America; 
introduced  from  eastern  Asia;  3(i). 

Hibiscus  trionum  L.  Flower-of-an-Hour. 
Annual,  commonly  1.5-5  dm  tall,  the  lower 
branches  often  prostrate,  coarsely  hispid-stel- 
late to  glabrate;  leaves  3-lobed  or  more  com- 
monly 3-  to  5-parted,  the  main  lobes  cuneate 
basally,  the  middle  lobe  the  largest;  flowers 
solitary,  axillary,  mostly  3-6  cm  wide;  bract- 
lets usually  10,  linear,  often  coarsely  hispid, 
much  shorter  than  the  fruiting  calyx;  corolla 
cream  colored  to  yellowish,  with  a  purple 
Ltnter,  closing  in  shade.  Weedy  species  of 
Lultivated  land  at  lower  elevations;  wide- 
spread in  North  America;  adventive  from 
:entral  Africa;  8(i). 


Iliamna  Greene 

Plants  herbaceous,  perennial,  sparingly  and 
minutely  stellate-hairy;  leaves  alternate,  pe- 
tiolate, cordate  to  truncate  basally,  the  mar- 
gin lobed;  flowers  in  thyrsoid  panicles;  in- 
volucel of  3  narrow,  persistent  bractlets; 
calyx  5-cleft;  fruit  a  loculicidal  capsule,  the 
carpels  many;  seeds  usually  3  in  each  locule. 

Wiggins,  I.  L.  1936.  A  resurrection  and  revi- 
sion of  the  genus  lUamna  Greene.  Contr. 
Dudley  Herb.  1:  213-229. 

Iliamna  rivularis  (Dougl.)  Greene.  Wild 
Hollyhock.  {Malta  rivularis  Dougl.  ex. 
Hook.;  Sphaeralcea  rivularis  (Dougl.)  Torr. 
ex.  Gray;  Phymosia  rivularis  (Dougl.)  Rydb.). 
Perennial,  the  stems  few  to  many  from  a 
woody  caudex,  mostly  7-15  dm  tall,  minutely 
stellate-puberulent,  green;  leaves  3-  to  7- 
lobed,  cordate  to  truncate  basally,  2.5-15  cm 
long  (from  petiole  apex  to  tip),  2-16  cm 
broad,  the  lobes  triangular,  crenate-serrate, 
finely  stellate;  pedicels  mostly  less  than  1  cm 
long;  bractlets  linear-lanceolate,  shorter  than 
the  calyx;  calyx  lobes  3-5  mm  long  (to  8  mm 


30 


Great  Basin  Naturalist 


Vol.  40,  No.  1 


long  in  fruit);  petals  rose  pink  (rarely  white), 
20-37  mm  long;  carpels  6-10  mm  long  in 
fniit,  hispid  and  stellate.  Along  streams,  on 
foothills,  in  mountain  bnish,  ponderosa  pine, 
aspen,  and  spruce-fir  communities, 
1440-2900  m  elevation,  in  Daggett,  Davis, 
Duchesne,  Iron,  Juab,  Piute,  Salt  Lake,  San- 
pete, Sevier,  Summit,  Tooele,  Utah,  Wasatch, 
and  Weber  counties;  Colorado,  Idaho,  Ne- 
vada, and  Washington;  40(vi). 


Malva  L. 

Plants  herbaceous,  annual,  biennial  or  per- 
ennial, from  taproots,  the  pubescence  simple 
to  branched  or  stellate;  leaves  alternate,  pe- 
tiolate,  usually  more  or  less  cordate  basally, 
commonly  lobed;  flowers  in  axillary  clusters 
(sometimes  solitary)  or  in  subterminal  pan- 
icles; involucel  of  3  narrow  to  broad  per- 
sistent bractlets;  calyx  5-cleft;  fruit  a  schiz- 
ocarp,  the  carpels  mostly  10-15. 


1.           Petals  commonly  1.5-2  cm  long;  bractlets  of  involucel  ovate  to  oblong  .  M.  sylvestris 
Petals  asually  less  than  1  cm  long;  bractlets  of  involucel  linear  to  narrowly 
lanceolate  2 

2(1).       Stems  prostrate  spreading  from  the  caudex;  leaves  obscurely  lobed;  plant  a 

common  weedy  species M.  neglecta 

Stems  erect;  leaves  definitely  lobed;  plant  cultivated,  rarely  escaping 

M.  verticillata 


Malva  neglecta  Wallr.  Cheeses;  Mallow. 
Annual  or  biennial,  the  stems  prostrate- 
spreading,  commonly  1-6  dm  long,  stellate- 
hairy;  leaf  blades  reniform-orbicular,  0.6-3 
cm  long  (from  sinus  to  apex)  or  more,  and 
much  broader,  crenate  and  not  at  all  to  only 
shallowly  5-  to  7-lobed,  the  petioles  to  20  cm 
long  or  more;  flowers  clustered  (or  solitary) 
in  the  axils;  bractlets  linear;  calyx  (3)  4-6  mm 
long  at  anthesis,  the  lobes  acuminate;  petals 
white  to  pink  or  lilac,  about  twice  as  long  as 
the  sepals;  carpels  hairy,  roimded  on  the 
back.  Weeds  of  disturbed  sites  and  cultivated 
land,  in  much  of  Utah  (specimens  known 
from  Cache,  Iron,  Kane,  Salt  Lake,  San  Juan, 
Summit,  Utah,  and  Washington  counties); 
widespread  in  North  America;  adventive 
from  Eurasia;  22(ii).  Note:  Two  other  species, 
M.  parviflora  L.  and  M.  rotundifolia  L., 
might  be  present  in  Utah.  They  are  similar  to 
M.  neglecta  but  have  petals  subequal  to  the 
sepals.  Malva  parviflora  has  glabrous  petal 
claws,  whereas  in  M.  rotundifolia  the  claws 
are  bearded. 

Malva  sylvestris  L.  High  Mallow.  Biennial, 
the  stems  ascending,  mostly  3-10  dm  tall, 
rough  hairy  to  glabrate;  leaf  blades  3-8  cm 
long  or  more  and  often  broader,  orbicular  to 
cordate  or  reniform,  crenate  and  with  5-7 
lobes,  the  petioles  to  10  cm  long  or  more; 
flowers  clustered  in  the  leaf  axils;  bractlets 


ovate  to  elliptic;  calyx  5-7  mm  long  at  an- 
thesis, the  lobes  short  and  broad;  petals 
15-20  mm  long,  rose  purple;  carpels  glabrous 
or  nearly  so,  sharp  edged.  Cultivated  orna- 
mental, rarely  escaping  (Utah  Co.,  Larsen 
7152  BRY);  widespread  in  North  America; 
adventive  from  Europe;  1(0). 

Malva  verticillata  L.  Curled  Mallow.  An- 
nual, the  stems  erect,  mostly  to  10  dm  tall  or 
more,  sparingly  stellate-hairy;  leaf  blades 
mostly  1.5-7  cm  long  and  as  broad  or  broad- 
er, orbicular  to  reniform,  undulate-crisped 
and  distinctly  5-  to  7-lobed,  long-petioled; 
flowers  solitary  or  clustered,  subsessile  or 
some  pediceled;  bractlets  linear  to  narrowly 
lanceolate;  calyx  3.5-5  mm  long,  the  lobes 
acuminate;  petals  white,  only  somewhat  sur- 
passing the  sepals;  carpels  glabrous,  the  edges 
rounded.  Cultivated  ornamental,  rarely  es- 
caping (Washington  Co.,  Galway  in  1934 
BRY);  widely  scattered  in  the  United  States; 
adventive  from  the  Old  World;  1(0).  Our  ma- 
terial belongs  to  var.  crispa  L. 

Malvastrum  Gray 

Plants  herbaceous,  annual,  stellate-hairy; 
leaves  alternate,  petiolate,  the  blades  sub- 
cordate  to  tmncate  basally,  palmately  lobed; 
flowers  solitary  in  the  axils  or  in  terminal 
bracted  clusters;  involucel  of  usually  3  slen- 


March  1980 


Welsh:  Utah  Flora,  Malvaceae 


31 


der  bractlets;  calyx  5-cleft,  the  lobes  long- 
acuminate;  carpels  10-15;  fruit  a  schizocarp. 
Malvastrum  exile  Gray.  {Malveopsis  exile 
(Gray  Kuntze;  Eremalche  exile  (Gray) 
Greene;  Sphaeralcea  exile  (Gray)  Jepson).  An- 
nual, the  stems  spreading-decumbent  to  pro- 
strate, branching  from  near  the  base,  0.3-4 
dni  long,  rather  sparingly  stellate-hairy;  leaf 
blades  suborbicular,  0.8-3.2  cm  wide,  palma- 
tely  3-  to  5-cleft,  with  rounded  or  cuspidate 
teeth;  petioles  1-5  cm  long;  bractlets  narrow- 
ly lanceolate  to  sublinear;  calyx  3-5  mm 
long;  petals  whitish  to  pinkish  or  lavender, 
only  somewhat  surpassing  the  sepals;  carpels 
transversely  wrinkled.  Open  sites  in  black- 
brush  and  creosote  brush  communities, 
850-1200  m  elevation,  in  Garfield  (report 
probably  erroneous)  and  Washington  coun- 
ties; Arizona  and  southern  California;  6(0). 

SiDA  L. 

Plants  herbaceous,  perennial,  from  spread- 
ing rhizomes,  densely  stellate-canescent; 
leaves  alternate,  petiolate,  crenate-serrate, 
not  or  obscurely  linear,  deciduous  bractlets; 
calyx  5-lobed;  carpels  5-10,  1 -seeded;  fruit  a 
schizocarp. 

Sida  hederacea  (Dougl.)  Torr.  Alkali-Mal- 
low. {Malva  hederacea  Dougl.;  A4.  californica 
Presl.;  Disella  hederacea  (Dougl.)  Greene). 
Perennial,  the  stems  from  elongate  rhizomes, 


decumbent  to  prostrate,  the  surface  obscured 
by  overlapping  stellate  hairs,  1-4  din  long; 
leaf  blades  reniform  to  orbicular,  often 
oblique,  dentate,  obscurely  if  at  all  lobed,  the 
petioles  0.3-2.5  (3)  cm  long;  bractlets  sub- 
linear;  calyx  5-7  mm  long;  petals  yellowish 
(fading  orange),  10-12  mm  long;  carpels  reti- 
culate on  the  sides.  Saline  meadows  and 
seeps,  at  lower  elevations  in  Emery,  Salt 
Lake,  Tooele,  Uintah,  and  Utah  counties  (and 
probably  elsewhere);  Washington  south  to 
California,  Texas,  and  Mexico;  6(i). 

SiDALCEA  Gray 

Plants  herbaceous,  perennial,  from  tap- 
roots or  short  rhizomes,  usually  stellate  and 
somewhat  hirsute;  leaves  alternate,  petiolate 
often  dimorphic,  the  lowermost  merely  pal- 
mately  lobed,  the  upper  ones  commonly  cleft 
and  with  linear  lobes;  flowers  borne  in  semi- 
spicate  racemes,  of  two  types,  those  of  plants 
with  perfect  flowers  the  largest;  involucel 
lacking;  calyx  5-cleft;  carpels  5-10,  1-seeded, 
tardily  separating. 

Hitchcock,  C.  L.  1957.  A  study  of  the  per- 
ennial species  of  Sidalcea.  Univ.  Wash. 
Publ.  Biol.  18:  1-79. 

Roush,  E.  M.  F.  1931.  A  monograph  of  the 
genus  Sidalcea.  Ann.  Mo.  Bot.  Card.  18: 
117-244. 


1.  Petals  white  or  merely  pinkish-tinged,  often  drying  yellow;  anthers  bluish  pink; 

plants  rhizomatous;  stems  hirsute  below S.  Candida 

Petals  pink  to  lavender;  anthers  usually  yellow  to  white;  plants  rhizomatous  or 
not;  stems  hirsute  to  glabrous  or  tomentose  below 2 

2(1).       Plants   from   rather   fleshy   taproots;   stems   commonly   hirsute  below;   calyx 

hirsute  with  pustulose  hairs  (at  least  in  part)  S.  neomexicana 

-  Plants  often  rhizomatous;  stems  stellate  to  glabrous  below;  calyx  seldom  with 

pustulose  hairs  S.  oregana 


Sidalcea  Candida  Gray.  Plants  from  slen- 
der rhizomes,  the  stems  4-10  dm  tall, 
glabrous  to  hirsute  with  simple  hairs  below, 
more  or  less  stellate  above;  leaf  blades  6-20 
cm  wide,  the  basal  ones  shallowly  5-  to  7- 
lobed  and  coarsely  crenate,  the  upper  ones 
divided  into  3-5  entire  segments;  calyx  7-10 
mm  long,  variously  stellate-hairy  and  glandu- 
lar puberulent;  petals  white  to  pinkish,  often 
drying  yellow,  12-20  mm  long;  carpels  about 


3  mm  long.  Stream  banks,  lake  shores,  and 
seeps,  1410-2750  m,  in  Beaver,  Garfield, 
Grand,  Iron,  Millard,  Piute,  Salt  Lake,  San 
Juan,  Sevier,  Summit,  Uintah,  Utah,  and 
Wasatch  counties;  Wyoming  and  Colorado 
west  to  Nevada  and  south  to  New  Mexico. 
Our  materials  have  been  treated  as  belonging 
to  two  more  or  less  and  at  least  partially  sym- 
patric  varieties;  25(vi). 


32 
1. 


Great  Basin  Naturalist 


Vol.  40,  No.  1 


Calyx  rather  uniformly  hairy  from  base  to  apex  of  the  lobes;  plants  of  wide 

distribution S.  Candida  var.  Candida 

Calyx  more  hairy  at  the  base  than  on  the  lobes,  the  lobes  often  subglabrous; 

plants  mostly  from  mountainous  portions  of  middle  Utah 

S.  Candida  var.  glahrata 


Var.  Candida.  (S.  Candida  var.  tincta  Cock- 
erell).  Known  from  Beaver,  Grand,  Iron,  Salt 
Lake,  San  Juan,  Summit,  and  Wasatch  coun- 
ties; Colorado,  New  Mexico. 

Var.  glahrata  C.  L.  Hitchc.  Known  from 
Iron,  Millard,  Piute,  Salt  Lake,  Sevier,  Sum- 
mit, and  Uintah  counties;  Wyoming,  Colo- 
rado, and  Nevada. 

Sidalcea  neomexicana  Gray.  Plants  from 
enlarged  taproots  or  fascicled  roots,  the  stems 
2-9  (10)  dm  tall,  hirsute  below  (or  rarely 
glabrous)  with  simple  or  bifurcate  hairs;  leaf 


blades  L5-11  cm  wide,  the  basal  ones  cre- 
nate  to  shallowly  5-  to  7-lobed,  the  cauline 
ones  divided  usually  into  5  laciniate  to  entire 
segments;  calyx  5-10  mm  long,  usually  with 
some  simple  pustulose  hairs  interspersed  with 
stellate  ones;  petals  rose  pink  (fading  blue- 
purple),  11-19  mm  long;  carpels  2-3  mm 
long.  Wet  Meadows,  stream  banks,  and  seeps, 
at  1370  to  2150  m  in  Box  Elder,  Garfield, 
Juab,  Piute,  Salt  Lake,  Sanpete,  Sevier,  Sum- 
mit, Utah,  and  Wasatch  counties;  Oregon, 
Idaho,  and  Wyoming  south  to  California, 
Arizona,  and  Mexico. 


Hairs  of  lower  stem  nearly  all  simple;  calyx  coarsely  and  rather  densely  hirsute 
to  coarsely  hairy,  lacking  appressed  stellate  hairs;  upper  stems  usually 

glabrous  S.  neomexicana  var.  neomexicana 

Hairs  of  lower  stem  often  forked;  calyx  often  with  fine  appressed  stellate  hairs 

in  addition  to  the  coarse  ones;  upper  stems  often  stellate  hairy  

S.  neomexicana  var.  crenulata 


Var.  crenulata  (A.  Nels.)  C.  L.  Hitchc.  (S. 
crenulata  A.  Nels.,  type  from  Juab,  Utah;  S. 
neomexicana  ssp.  crenulata  (A.  Nels.)  C.  L. 
Hitchc).  Known  from  Box  Elder,  Juab,  Salt 
Lake,  Sanpete,  Sevier,  Summit,  Utah,  and 
Wasatch  counties;  Oregon,  Idaho,  and  Ne- 
vada; lO(ii). 

Var.  neomexicana.  Known  from  Box  Elder, 
Garfield,  Piute,  San  Juan,  Sevier,  Utah,  and 
Wasatch  counties;  Wyoming,  Colorado,  Ari- 
zona, and  Mew  Mexico;  Mexico;  12(ii). 

Sidalcea  oregana  (Nutt.)  Gray.  {Sida  ore- 
gana  Nutt.  ex  T.  &  G.;  S.  nervata  A.  Nels.). 
Plants  from  a  taproot,  lacking  or  rarely  with 
rhizomes,  the  stems  3-11  dm  tall  or  more, 
glabrous  or  usually  appressed-stellate  hairy 
below,  appressed-stellate  above;  leaf  blades 
2.5-17  cm  wide,  the  basal  ones  shallowly  5-to 
7-lobed  and  coarsely  crenate,  the  cauline 
ones  deeply  lobed,  with  3-7  coarsely  toothed 
to  entire  lobes;  calyx  3.5-9  mm  long,  various- 
ly stellate-hairy  and  sometimes  bristly;  petals 
7-23  mm  long,  pale  pink  to  rose  pink  (fading 


blue  purple);  carpels  2.5-3  mm  long.  Mead- 
ows, stream  banks,  and  open  woods,  at  1680 
to  2750  m  in  Cache,  Juab,  Salt  Lake,  San- 
pete, Summit,  Utah,  Wasatch,  and  Weber 
counties;  Washington  and  Idaho  south  to 
California,  Nevada,  and  Utah.  Our  materials 
belong  to  var.  oregana;  32  (ii). 


Sphaeralcea  St.  Hil. 

Plants  herbaceous,  perennial,  from  tap- 
roots or  rhizomes,  glabrescent  to  canescent 
with  stellate  hairs;  leaves  alternate,  petiolate, 
sometimes  dimorphic,  the  lowermost  merely 
toothed  or  palmately  lobed  (rarely  entire), 
the  upper  ones  cleft  to  entire;  flowers  borne 
in  racemose  to  thyrsoid  cymes;  involucel  of  3 
or  fewer  filiform  bractlets;  calyx  5-cleft;  car- 
pels 8-20,  the  seeds  1  or  2  per  carpel;  fruit  a 
schizocarp,  the  mature  fruit  segments  divided 
into  a  basal  indehiscent,  reticulate  portion 
and  an  apical  dehiscent  portion. 


March  1980  Welsh:  Utah  Flora,  Malvaceae  33 

Jefferies,  J.   A.   M.    1972.   A   revision  of  the  Kearney,  T.  H.  1935.  North  American  species 

genus  Sphaeralcea  (Malvaceae)  for  the  of  Sphaemlcea,   Subgenus  Eusphaeralcea. 

state  of  Utah.  UnpubHshed  thesis.  Brigham  Univ.  Calif.  Fubl.  Bot.  19(1):  1-102. 
Young  University.  92  pp. 

1.  Inflorescence  racemose,  rarely  with  more  than  one  flower  per  node  or,  if  more, 

as  in  S.  caespitosa,  then  the  plants  restricted  to  Millard  County 2 

—  Inflorescence  thyrsoid  to  thyrsoid-glomerate,  with  usually  more  than  one 
flower  per  node;  distribution  various 5 

2(1).  Leaf  blades  only  slightly,  if  at  all,  3-  to  5-lobed,  the  margins  irregularly  cre- 
nate-dentate;  hairs  with  rays  radiating  in  more  than  a  single  plane;  plants  sel- 
dom more  than  1.5  dm  tall,  known  only  from  western  Beaver  and  Millard 
counties  S.  caespitosa 

—  Leaf  blades  distinctly  3-  to  5-lobed,  -parted,  or  -divided;  hairs  of  rays  radiating 
in  a  single  plane  (except  in  S.  coccinea);  plants  often  1.5  dm  tall  or  more,  of 
different  distribution  3 

3(2).        Leaves  trifoliolate,  the  leaflets  linear  to  narrowly  oblanceolate  and  entire,  or 

the  upper  ones  simple  and  entire;  plants  of  southeastern  Utah S.  leptophylla 

—  Leaves  various,  but  if  trifoliolate  then  the  leaflets  oblanceolate  and  entire  to 
toothed,  if  the  uppermost  simple  then  toothed  or  lobed;  distribution  various 4 

4(3).  Lowermost  leaves  simple  and  entire  or  trifoliolate,  or  some  broadlv  toothed  or 
lobed;  involucel  present;  rays  of  hairs  radiating  in  one  plane;  plants  of  eastern 
Wayne  County  S.  psoraloides 

—  Lowermost  leaves  usually  3-  to  5-lobed,  the  lobes  usually  toothed  or  again 
lobed;  involucel  present  or  lacking  (caducous);  rays  of  hairs  radiating  in  several 
planes;  plants  of  broad  distribution  S.  coccinea 

5(1).        Plants  only  sparingly  pubescent,  the  herbage  bright  green 6 

—  Plants  moderately  to  densely  pubescent,  the  herbage  yellowish,  whitish,  or 
grayish 8 

6(4).  Leaves  3-  to  5-parted  or  -divided,  the  lobes  with  narrow,  regularly  pinnatifid 
margins,  the  teeth  at  nearly  right  angles  to  the  vein;  carpels  often  with 
transparent  lacunae,  4-6  mm  high;  plants  rare,  of  southern  Utah  only  S.  rusbiji 

—  Leaves  variously  lobed,  divided,  or  parted,  the  lobes  with  broader  margins  ir- 
regularly toothed  or  lobed,  but  not  as  above;  carpels  with  opaque  lacunae, 
3-4.5  mm  high  7 

7(6).  Leaves  slightly  lobed,  the  margins  unevenly  toothed  or,  in  .some,  deeply  parted 
to  divided  with  the  margin  coarsely  and  irregularly  lobed,  the  base  subcordate 
to  cuneate;  plants  of  northern  Utah  S.  munroana 

—  Leaves  3-  to  5-parted  or  -divided,  the  margins  regularly  cleft,  lobed,  or 
toothed,  the  base  subcordate  to  deeply  cordate;  plants  mostly  of  southern  Utah 

S.  grossulariifolia 

8(5).  Inflorescence  loosely  thyrsoid  (appearing  paniculate),  leafy;  flowers  not  numer- 
ous at  each  node;  peduncles  generally  elongate;  calyx  surpassing  the  fruit;  car- 
pels with  reticulae  extending  onto  back  of  carpel;  plants  of  southwestern  Utah  . 
S.  ambigua 

—  Inflorescence  contracted  thyrsoid-glomerate;  flowers  often  numerous  at  each 
node,  not  especially  leafy;  calyx  often  shorter  than  the  fruit;  carpels  with 
reticulae  confined  to  lateral  face  of  carpel;  plants  of  various  distribution 9 


34 


Great  Basin  Naturalist 


Vol.  40,  No.  1 


9(8).       Leaves  3-  to  5-cleft,  -parted,  or  -divided;  carpels  with  well-defined  reticiilae  on 
less  than  half  of  carpel  face;  plants  of  all  but  the  northeastern  one-fourth  of 

\JlJ^y^  S.  grossiilariifolia 

Leaves  shallowly  3-  to  5-lobed;  carpels  with  well-defined  to  nearly  obscure  re- 
ticulae  on  the  lower  one-third  of  the  carpel;  plants  mainly  of  eastern  and 
southern  Utah,  scattered  elsewhere  S.  parvifolia 


Sphaeralcea  ambigua  Gray.  Stems  arising 
from  a  woodv  caudex,  several  to  numerous, 
.3-10  dm  tall,  whitish  to  yellowish  canescent; 
leaf  blades  1-6  cm  long  (from  sinus  to  apex), 
0.8-5  cm  wide,  thickish,  usually  rugose,  with 
veins  prominent  beneath,  ovoid,  deltoid,  or 
nearlv  orbicular,  the  base  cordate  to  deeply 
cordate,  obscurely  to  definitely  3-  to  5-lobed, 
the  lobes  crenate;  inflorescence  an  open  pan- 
icle, sometimes  narrowly  thyrsoid;  pedicels 
usually  shorter  than  the  calyx;  calyx  uni- 
formly pubescent  to  glabrate,  6-20  mm  long 
at  anthesis,  the  lobes  lanceolate  to  acumi- 
nate; petals  15-22  mm  long,  orange  to  or- 
ange pink  (fading  pinkish);  carpels  12-16  mm 
high,  the  indehiscent  portion  comprising 
about  one-third  of  the  carpel,  prominently 
reticulate.  Creosote  bush-blackbrush  and 
mixed  warm  desert  shrub  communities, 
670-1070  m,  in  Washington  Co.;  Nevada, 
Arizona,  and  California;  and  Mexico.  Our 
material  belongs  to  var.  ambigua;  10(i). 

Sphaeralcea  caespitosa  M.  E.  Jones.  Jones 
Globemallow.  Stems  solitary  or  more  com- 
monly few  to  several  from  the  summit  of  a 
branching  woody  caudex,  0.2-2.5  dm  tall, 
whitish  to  grayish  canescent;  leaf  blades 
1.2-5.5  cm  long,  1.2-6  cm  wide,  thickish,  not 
nigose,  veins  apparent  but  not  especially 
prominent,  ovate  to  deltoid  or  orbicular,  the 
base  tnmcate  to  obtuse,  obscurely  if  at  all 
lobed,  crenate  to  crenate-dentate;  in- 
florescence thyrsoid,  the  flowers  tightly  clus- 
tered or  solitary;  pedicels  shorter  than  the  ca- 
lyx; calyx  uniformly  stellate,  the  rays  of  hairs 
not  radiating  in  a  single  plane,  the  lobes 
lance-acuminate;  petals  15-21  mm  long,  or- 
ange; carpels  12-14,  4-6  mm  high,  the  in- 
dehiscent  portion  forming  slightly  more  than 
one-third  of  the  carpel,  reticulate  on  the 
sides.  Mixed  desert  shrub  communities  (shad- 
scale,  rabbitbmsh,  winterfat),  mainly  on  Sevy 
Dolomite  formation,  at  1370-1750  m,  in  Mil- 
lard and  Beaver  counties;  endemic;  20(iii). 


Sphaeralcea  coccinea  (Nutt.)  Rydb.  Com- 
mon Globemallow.  {Malva  coccinea  Nutt.; 
Cristaria  coccinea  (Nutt.)  Pursh;  Sida  coc- 
cinea (Nutt.)  DC;  Malvastrum  coccineum 
(Nutt.)  Gray.;  Sida  dissecta  Nutt.;  M.  c.  var. 
dissectum  (Nutt.)  Gray;  M.  dissectum  (Nutt.) 
Cockerell;  S.  dissecta  (Nutt.)  Rydb.;  S.  coc- 
cinea ssp.  dissecta  (Nutt.)  Kearney;  S.  coc- 
cinea var.  dissecta  (Nutt.)  Kearney;  M.  c.  var. 
elatum  Baker;  M.  elatiim  (Baker)  A.  Nels.;  S. 
elata  (Baker)  Rydb.;  S.  c.  ssp.  elata  (Baker) 
Kearney;  S.  c.  var.  elata  (Baker)  Kearney;  M. 
cockerellii  A.  Nels.;  M.  micranthiim  W.  &  S.). 
Stems  solitary  or  few  to  many  from  the  apex 
to  a  woody  caudex,  or  less  commonly  from 
creeping  rhizomes,  0.6-4.2  dm  tall,  white  to 
yellowish  canescent;  leaf  blades  1.1-3.7  cm 
long,  1.2-5.2  cm  wide,  usually  wider  than 
long,  ovate  to  cordate-ovate  in  outline,  the 
base  often  cordate,  usually  3-to  5-lobed,  with 
main  divisions  cleft  almost  or  quite  to  the 
base,  the  lobes  usually  again  toothed  or 
lobed;  inflorescence  racemose,  sometimes 
paniculate,  rarely  thyrsoid;  pedicels  shorter 
than  the  calyx;  calyx  uniformly  stellate,  the 
rays  or  hairs  not  radiating  in  a  single  plane, 
the  lobes  lance-acuminate;  petals  8-15  mm 
long,  orange;  carpels  8-14,  2-3  mm  high,  the 
indehiscent  part  forming  two-thirds  or  more 
of  the  carpel,  reticulate  on  the  sides  and  on 
the  back.  Blackbrush,  shadscale-greasewood, 
sagebRish,  juniper-pinyon,  mountain  brush, 
and  ponderosa  pine  communities,  920-2750 
m,  in  all  counties  (except  Morgan  and 
Wasatch?);  Saskatchewan  and  Alberta  south 
to  Arizona,  New  Mexico,  and  Texas.  Our  ma- 
terials have  been  recognized  as  belonging  to 
vars.  dissecta  and  elata,  but  the  segregation 
of  these  entities  appears  to  have  been  wholly 
arbitrary,  with  intermediates  more  numerous 
than  the  supposed  taxa;  152(xviii). 

Sphaeralcea  grossulariifolia  (H.  &  A.) 
Rydb.  Gooseberry-Leaved  Globemallow. 
{Sicki  grossulariifolia   H.  &  A.;  Malvastrum 


March  1980 


Welsh:  Utah  Flora,  Malvaceae 


35 


grossulariifoUwn  (H.  &  A.)  Gray;  S.  pedata 
Torr.,  in  Gray;  S.  g.  ssp.  pedata  (Torr.) 
Kearney;  S.  g.  var.  pedata  (Torr.)  Kearney). 
Stems  few  to  many  from  a  woody  caiidex, 
1-10  dm  tall  or  more,  whitish  to  yellowish 
canescent  to  subglabrous  and  green;  leaf 
blades  1.3-5  cm  long,  1.3-5  cm  wide,  usually 
longer  than  wide,  ovate  to  cordate-ovate  in 
outline,  the  base  cordate  to  truncate  or  ob- 
tuse, usually  3-  to  5-lobed,  the  main  division 
asually  cleft  or  parted  to  irregularly  toothed; 
inflorescence  thyrsoid,  with  usually  more 
tlian  one  flower  per  node;  pedicels  shorter 
than  to  much  longer  than  the  calyx;  calyx 
luiiformly  stellate,  the  rays  of  hairs  not  radi- 
ating in  a  single  plane,  the  lobes  ovate  to 
lance- acuminate;  petals  8-20  mm  long,  or- 
ange or  rarely  rose  pink;  carpels  10-14, 
2.5-4.5  mm  high,  the  indehiscent  portion 
forming  from  two-fifths  to  three-fifths  of  the 


carpel,  reticulate  on  the  sides.  Blackbrush, 
shadscale,  rabbitbrush,  sagebrush,  juniper- 
pinyon,  and  less  commonly  mountain  brush 
communities,  920-2450  m,  in  Beaver,  Box  El- 
der, Cache,  Emery,  Garfield,  Grand,  Iron, 
Kane,  Juab,  Millard,  Morgan,  Piute,  Salt 
Lake,  San  Juan,  Sanpete,  Sevier,  Tooele, 
Utah,  Wasatch,  Washington,  and  Wayne 
counties;  Washington,  Oregon,  Nevada,  Cali- 
fornia, and  Arizona.  Two  infraspecific  taxa 
have  been  segregated,  largely  on  the  basis  of 
form  of  the  leaf  blades.  Intergradation  of  the 
phases  seems  to  be  complete.  Further,  S. 
grossulariifolia  appears  to  form  intermediates 
with  S.  coccinea,  S.  parvifolia,  and  the  more 
northern  S.  munroana.  A  phase  with  green 
herbage  and  thin  leaves  occurs  along  Glen 
Canyon.  It  seems  to  represent  a  taxonomic 
imit  worthy  of  recognition. 


1.  Herbage  bright  green;  leaves  thin-textured;  plants  of  eastern  Kane  and 

Garfield,  and  western  San  Juan  counties S.  grossulariifolia  var.  moorei 

Herbage  gray  green  to  whitish  canescent;  leaves  thick-textured;  plants 
widespread S.  grossulariifolia  var.  grossulariifolia 


Var.  grossulariifolia.  This  is  the  common 
and  widely  distributed  phase  of  the  species  in 
Utah.  The  report  by  Kearney  (I.e.,  p.  90)  of  S. 
digitata  (Greene)  Rydb.  apparently  belongs 
here;  115(xii). 

Var.  moorei  Welsh  var.  nov.  Plantae  siniilis 
var.  grossulariifoliae  sed  differt  in  folii  et 
caules  virides  et  folii  tenues.  Holotype:  Kane 
County,  Utah,  east  side  of  Last  Chance  Bay, 
Lake  Powell,  Entrada  Sandstone,  S.  L.  Welsh 
and  N.  D.  Atwood  11597,  2  May  1972  (BRY). 
Additional  specimens:  Kane  County,  mouth 
of  Escalante  River,  Lake  Powell,  S.  L.  Welsh 
and  G.  Moore  11810,  5  June  1972;  do.  Wil- 
low Tank,  D.  A.  White  111,  4  May  1962;  do, 
Escalante  Canyon,  S.  L.  Welsh  and  G.  Moore 
11827,  5  June  1972;  do,  N.  D.  Atwood  and  R. 
Allen  3211,  24  August  1971;  do,  Hole-in-the- 
Rock,  B.  F.  Harrison  12112,  14  May  1953; 
San  Juan  Co.,  1  mi.  E  of  Hole-in-the-Rock,  S. 
L.  Welsh  and  C.  A.  Toft  11869,  16  June 
1972;  do.  Three  Garden,  Lake  Powell,  ca  1 
mi.  N  of  confluence  with  San  Juan  Arm,  S.  L. 
Welsh  12420,  5  May  1974;  do.  Comb  Wash, 
S.  L.  Welsh  and  N.'D.  Atwood  9972,  6  June 
1970  (all  at  BRY).  This  variety  is  named  to 


honor  Glen  Moore,  botanist,  teacher,  collabo- 
rator, and  collector. 

Sphaeralcea   leptophylla    (Gray)    Rydb, 

{Malvastrum  leptophyllum  Gray).  Stems  few 
to  many  from  a  woody  caudes,  2.0-5.5  dm 
tall,  grayish  canescent  to  yellow  green 
throughout;  leaf  blades  1.0-3.2  cm  long,  di- 
gitately  3-lobed,  the  lobes  entire,  linear  to 
oblanceolate,  1-4  mm  wide,  or  the  upper 
leaves  simple  and  linear;  inflorescence  race- 
mose, elongate,  usually  with  one  flower  per 
node;  pedicels  from  much  shorter  to  longer 
than  the  calyx;  calyx  uniformly  stellate,  the 
rays  of  hairs  radiating  in  a  single  plane,  the 
lobes  lance-attenuate;  petals  8-12  mm  long, 
orange;  carpels  7-9,  3-3.5  mm  high,  the  in- 
dehiscent  portion  forming  two-thirds-three- 
fourths  of  the  carpel,  coarsely  reticulate, 
ridged,  or  tuberculate  on  the  back.  Black- 
brush  and  mixed  semidesert  shrub  commu- 
nities, 1200-1520  m,  in  Garfield,  Grand,  and 
San  Juan  counties;  New  Mexico,  Arizona, 
Texas,  and  Mexico.  Two  distinctive  phases 
are  recognizable  among  our  materials;  they 
can  be  distinguished  as  follows: 


36 

1. 


Great  Basin  Naturalist 


Vol.  40,  No.  1 


Plants  grayish  canescent,  the  hairs  obscuring  the  surface  of  stems,  leaves,  and 
calvx  lobes;  leaf  lobes  narrowly  oblong  to  linear;  distribution  as  for  the  species 

S.  leptophylla  var.  leptophylla 

Plants  green  to  yellow  green,  the  hairs  widely  separated,  not  obscuring  stems, 
leaves  or  calyx  lobes;  leaf  lobes,  at  least  of  lowermost  leaves,  oblanceolate  to 
spatulate-  known  only  from  San  Juan  County,  Utah S.  leptophyUa  var.  janeae 


Var.  leptophylla.  This  is  the  common  form 
of  the  species.  It  is  known  from  Garfield, 
Grand,  and  San  Juan  counties,  Utah,  and 
from  New  Mexico  and  Arizona;  7(iii). 

Var.  janeae  Welsh  var.  nov.  Plantae  similis 
var.  leptophylhi  sed  differt  in  folii  caules  et 
calvces  virides  et  lobos  foliorum  inferiorum 
oblanceolatos  vel  spathulatos.  Holotype:  San 
Juan  County,  Utah,  along  White  Rim  road, 
north  of  Turks  Head,  on  sandy  slopes  in 
blackbnish  communitv,  Canyonlands  Nation- 
al Park,  S.  L.  Welsh  7064,  17  May  1968 
(BRY).  This  variety  is  named  to  honor  Jane 
.\rdis  Murray  Jefferies,  student  of  Sphae- 
ralcea  in  Utah. 

Sphaeralcea  miinroana  (Dougl.)  Spach  in 
Gray.  Munroe  Cdobemallow.  {Malta  miin- 
roana Dougl.  in  Lindl.;  Nuttallia  miinroana 
(Dougl.)  Nutt.;  Malvastnirn  munrooniim 
(Dougl.)  Gray;  S.  suhrhomhoidea  Rydb.;  S.  m. 
ssp.  suhrhomhoidea  (Rydb.)  Kearney;  S.  m. 
var.  s.  (Rydb.)  Kearney).  Stems  several  to 
many  from  a  branching  woody  caudex,  1.8-7 
dm  tall  or  more,  yellowish  green  to  some- 
what grayish  canescent,  the  foliage  usually 
bright  green;  leaf  blades  1-6  cm  long,  0.8-6 
cm  wide,  ovate  to  orbicular  or  rhombic  in 
outline,  the  base  truncate  to  obtuse  or  sub- 
cuneate,  usually  3-  to  5-lobed,  the  sinuses 
shallow  to  very  deep,  the  main  divisions 
merely  toothed  or  the  lateral  ones  incised;  in- 
florescence narrowly  thyrsoid,  usually  with 
more  than  one  flower  per  node;  pedicels  usu- 
ally much  shorter  than  the  calyx;  calyx  uni- 
formly stellate,  the  rays  of  hairs  not  radiating 
in  a  .single  plane,  the  lobes  deltoid-ovate  to 
ovate;  petals  8-15  mm  long,  orange;  carpels 
10-1.3,  2.5-3  mm  high,  the  indehiscent  por- 
tion forming  about  half  the  carpel,  reticulate 
on  the  sides.  Mixed  desert  shrub,  or  more 
commonly,  in  sagebrush  and  mountain  brush 
conununities,  1370-2450  m,  in  Box  Elder, 
Cache,  Duchesne,  Emery,  Summit,  Tooele, 
Uintah,  Utah,  and  Wasatch  counties;  Mon- 
tana, Idaho,  Washington,  Wyoming,  Nevada, 


and  California.  This  entity  is  much  like  both 
S.  parvifoUa  and  S.  grossulariifolia.  The  green 
color  of  herbage  is  diagnostic  of  S.  miinroana 
from  both,  except  for  the  var.  moorei  which  is 
not  sympatric  with  S.  miinroana;  21(ii). 

Sphaeralcea  parvifoUa  A.  Nels.  Nelson 
Globemallow.  (S.  marginata  York,  ex  Rydb.; 
S.  arizonica  Heller,  ex  Rydb.).  Stems  few  to 
many  from  a  branching  woody  caudex, 
1.5-10  (11)  dm  tall,  grayish  canescent,  the  fo- 
liage gray  green  or  only  somewhat  yellow 
green;  leaf  blades  1.0-5.5  cm  long,  1.2-5.2 
cm  wide,  ovate  to  orbicular,  reniform,  or 
cordate-ovate,  the  base  cordate  to  truncate  or 
obtuse,  usually  shallowly  3-  to  5-lobed,  the  si- 
nuses usually  shallow,  the  lobes  crenate-den- 
tate;  inflorescence  commonly  narrowly  thyr- 
soid, usually  with  more  than  one  flower  per 
node;  pedicels  usually  shorter  than  the  calyx; 
calyx  uniformly  stellate,  the  rays  of  hairs  not 
radiating  in  a  single  plane,  the  lobes  lance- 
ovate  to  deltoid;  petals  7-15  mm  long,  or- 
ange; carpels  10-12,  3-4  mm  high,  the  in- 
dehiscent  part  forming  from  one-fourth  to 
one-third  of  the  carpel,  faintly  reticulate  on 
the  sides.  Blackbrush,  salt  desert  shrub,  sage- 
liRish,  pinyon-jiuiiper,  and  mountain  brush 
communities,  at  850  to  2700  m,  in  Box  Elder, 
Cache  and  Tooele  counties,  where  probably 
of  recent  introduction,  and  in  Duchesne, 
Emery,  Garfield,  Grand,  Iron,  Kane,  Piute, 
San  Juan,  Sevier,  Tooele,  Washington,  and 
Wayne  counties,  where  likely  indigenous; 
Nevada,  Arizona,  New  Mexico,  and  Califor- 
nia. Sphaeralcea  parvifoUa  has  been  com- 
pared by  Kearney  (I.e.)  with  S.  amhigua, 
which  it  resembles.  The  relationship  of  S. 
parvifoUa  in  Utah  seems  to  lie  with  the 
largely  sympatric  S.  gro.s.sularii folia;  144(xxii). 

Sphaeralcea  psoraloides  Welsh  sp.  nov. 
Stems  few  to  many  from  a  branching  caudex, 
1.4-2.4  dm  tall  or  more,  sparsely  yellowish 
canescent,  the  foliage  yellow  green;  leaf 
blades  1.3-3.5  cm  long,  0.4-3.8  cm  wide,  ob- 
lanceolate  to  cuneate-ovate   in  outline,,  cu- 


March  1980 


Welsh:  Utah  Flora,  Malvaceae 


37 


neate  to  obtuse  or  rounded  basally,  trifoliol- 
ate  or  simple  to  3-lobed  below,  deeply  3-  to 
5-cleft  above,  the  lobes  entire  to  few  toothed 
or  lobed,  usually  more  than  5  mm  wide;  in- 
florescence racemose,  the  flowers  solitary  in 
tlie  upper  axils;  calvx  uniformlv  stellate,  the 
rays  of  hairs  radiating  in  a  single  plane,  the 
lobes  lance-acuminate;  petals  10  (8-12)  mm 
long,  orange;  carpels  10  (fruit  unknown). 
Ephedra-Gmijia  commimity  on  Entrada  silts- 
tone,  1500  m,  in  Wavne  Countv;  endemic. 

Plantae  similis  S.  leptophylla  sed  differt  in 
foliolos  oblanceolata  vel  laminas  super iores 
confluentes  et  lobatos;  e  S.  coccinea  laminis 
inferioribus  simplicibus  vel  trifoliolati  digi- 
tatis  distinguenda. 

Caules  pauci  vel  multi  e  caudicibus  raniifi- 
cantibus  1.4-2.4  dm  alti  vel  plures  flavidi-ca- 
nescentes  parce  folia  et  caules  luteo-virides; 
laminae  foliorum  1.3-3.5  cm  longae  0.4-3.8  . 
cm  latae  oblanceolata  ad  cuneati-ovatas  cu- 
neatae  ad  obtusas  vel  rotundatas  basaliter  tri- 
foliolatae  vel  simplicia  ad  trilobata  infra  3-5 
fissa  profimde  supra  lobis  intergris  ad  pauci- 
dentatis  vel  pauci-lobatis  plerumque  plus 
quam  5  mm  latis;  inflorescentiae  racemosae, 
flores  solitari  in  axilas  supras;  calyces  stellati 
uniformiter,  radius  pilos  radiantibus  in  plan- 
item  singularem,  lobus  calycis  lanci-acumi- 
natis;  petala  10  (8-12)  mm  longa,  aurantiaca; 
carpeli     10    (fructus    ignotus).    Holotvpe: 


Wavne  County,  Utah,  Salt  Wash,  ca  17  mi. 
due  WMW  of  Hanksville,  T27S,  R8E,  Sec. 
24,  at  1500  m,  on  Entrada  siltstone,  Gmi/ia- 
Ephedra  connnunitv,  S.  L.  Welsh  13348,  1 
June  1976  (BRY).  Paratvpe:  do,  S.  L.  Welsh 
13345,  1  June  1976  (BRY). 

Sphaeralcea  riisbyi  Gray.  Stems  few  to 
many  from  a  caudex,  or  rarely  sub- 
rhizomatous,  mostly  2-6.5  (8.5)  dm  tall,  yel- 
lowish green  to  somewhat  grayish  canescent; 
leaf  blades  1.3-3  cm  long,  1.2-4  cm  wide, 
ovate  to  orbicular  in  outline,  the  base  trun- 
cate-obtuse to  prominently  cordate,  parted  to 
divided  or  merely  cleft,  the  lobes  again 
toothed  (the  teeth  spreading  at  nearly  right 
angles);  inflorescence  thrysoid  to  paniculate, 
with  more  than  one  flower  per  node;  pedicels 
usually  shorter  (to  much  longer)  than  the  ca- 
lyx; bractlets  often  dark  red;  calyx  imiformly 
stellate  (more  densely  so  than  on  the  her- 
bage), the  rays  of  hairs  not  radiating  in  a 
single  plane,  the  lobes  ovate  to  lance-ovate; 
petals  9-18  mm  long,  orange;  carpels  10-12, 
4-6  mm  high,  the  indehiscent  part  forming 
from  one-fourth  to  two-fifths  of  the  carpel, 
finely  reticulate  on  the  sides.  Blackbrush, 
creosote  brush,  and  mixed  warm  desert  shrub 
communities,  820-1070  m,  in  Washington 
County;  Arizona.  S.  rusbiji  forms  apparent 
intermediates  with  phases  of  S.  grossularii- 
folia  and  S.  pa wi folia;  4(0). 


UTAH  FLORA:  MISCELLANEOUS  FAMILIES       / 

Stanley  L.  Welsh' 

Abstract.—  Considered  in  this  treatment  are  the  families  Aquifoliaceae,  Canabinaceae,  Ericaceae,  Krameriaceae, 
Magnoliaceae,  Moraceae,  Oleaceae,  Pyrolaceae,  Resedaceae,  Tamaricaceae,  and  Tiliaceae.  These  11  families  include 
61  cultivated,  escaped,  and  indigenous  species. 


The  flora  of  Utah  is  both  large  and  diverse. 
A  portion  of  the  diversity  is  due  to  the  pres- 
ence of  a  large  number  of  cultivated  species 
in  many  plant  families.  Floras  of  regions  have 
traditionally  avoided  inclusion  of  strictly  cul- 
tivated species.  Only  those  taxa  which  escape 
and  become  acclimated  have  been  treated. 
Included  are  the  cultivated  plants  and  those 
species  which  escape.  The  present  treatment 
covers  all  taxa  in  common  cultivation,  and  es- 
pecially those  which  are  represented  in  re- 
gional herbaria.  In  Table  1  a  list  is  presented 
of  the  families  treated  herein,  the  numbers  of 
genera  and  species,  and  whether  cultivated 
or  indigenous. 

The  list  heavily  favors  the  cultivated 
and/or  escaped  species,  and,  because  of  the 
status  of  cultivated  species  collections,  the 
treatment  is  likely  to  be  incomplete.  It  is 
presented  herein  for  use  by  students  of  the 
flora  who  want  to  know  the  names  of  culti- 
vated and  of  native  plant  species. 


Aquifoliaceae 

Holly  Family 

Evergreen  Shrubs  or  small  trees:  leaves  al- 
ternate, simple,  coriaceous,  armed  with  spiny 
teeth;  stipules  minute,  caducous;  flowers  usu- 
ally imperfect,  regular,  small  and  inconspic- 
uous, solitary  or  few  in  axillary  cymes;  sepals 
usually  4,  more  or  less  connate  basally;  petals 
usually  4,  distinct  or  slightly  connate  basally; 
stamens  or  staminodes  usually  4  (-9),  alter- 
nate with  the  petals;  pistil  1,  the  ovary  supe- 
rior, 3-  to  many-loculed,  the  carpels  as  many 
as  the  locules;  fruit  a  globose,  berrylike  drupe 
with  2-8  bony  1 -seeded  divisions. 

Ilex  L. 

Evergreen;  leaves  thick  and  shining;  flow- 
ers small,  mostly  in  few-flowered  axillary 
cymes;  staminodia  usually  present  in  pistil- 
late flowers,  a  rudimentary  pistil  present  in 
most  staminate  flowers;  fruit  usually  brightly 
colored.  (Note:  Members  of  this  family  are 
known  in  Utah  in  cultivation  only). 


Flowers  in  axillary  clusters  on  branches  of  the  previous  year  /.  oquifolium 

Flowers  in  solitary  cymes  on  branches  of  the  current  year  /.  opaca 


Ilex  aquifolium  L.  English  Holly.  Tall 
shnibs  to  small  trees  of  ornamental  plantings, 
rare  in  Utah;  introduced  from  the  Old 
World;  1(0). 

Ilex  opaca  Art.  American  Holly.  Low  to 
moderate  shrubs  of  ornamental  plantings,  oc- 
casional in  Utah;  introduced  from  the  eastern 
United  States;  1(0). 


Cannabinaceae 
Hemp  Family 

Plants  herbaceous,  with  watery  juice; 
leaves  alternate  or  opposite,  palmately 
veined  and  lobed  or  divided  to  essentially 
compound;  stipules  persistent;  flowers  imper- 


'Life  Science  Museum  and  Department  of  Botany  and  Range  Science,  Brigham  Young  University,  Provo,  Utah  84602. 


38 


March  1980 


Welsh:  Utah  Flora,  Miscellaneous  Families 


39 


feet,  the  plants  dioecious,  regular,  the  stami- 
nate  in  open  racemes  or  panicles,  the  pistil- 
late in  dense  clusters;  sepals  5,  connate   in 


pistillate  flowers  and  enclosing  the  ovary; 
stamens  5;  pistil  1,  2-carpelled,  the  styles  2; 
fruit  an  achene. 


Plants  strong-smelling,  stout,  erect  herbs;  leaves  palmately  5-  to  9-parted  ..  Cannabis 
Plants  rough-stemmed  clambering  vines;  leaves  coarsely  3-  to  7-lobed Humulus 


CaNxNabis  L. 

Plants  dioecious  or  rarely  some  mon- 
oecous;  leaves  palmately  lobed  to  parted  and 
apparently  compound,  alternate  or  the  lower 
opposite;  flowers  small,  inconspicuous,  the 
staminate  in  leafy  panicles  in  upper  axils; 
sepals  5,  oblong;  stamens  5;  pistillate  flowers 
in  small  clusters  on  leafy  branches  from  up- 
per axils,  each  flower  subtended  and  enclosed 
by  an  acuminate  bract,  the  calyx  barelv 
lobed,  surrounding  only  the  base  of  the 
ovary;  stigmas  2,  elongate;  fruit  a  lenticular 
achene,  enclosed  within  the  accrescent  bract. 

Small,  E.  and  A.  Cronquist.  1976.  A  practical 
and  natural  taxonomv  for  Cannabis.  Taxon 
5:  405-435. 

Cannabis  sativa  L.  Hemp,  Marijuana, 
Hashish,  Pot,  Grass.  Plants  6-20  dm  tall  or 
more,  the  stems  simple  or  much  branched; 
leaves  long  petioled,  the  blades  3-  to  7-part- 
ed,  the  segments  oblanceolate  to  elliptic,  at- 
tenuate to  accuminate  apically,  sharply  ser- 
rate, mostly  4-12  cm  long  and  0.4-1.8  cm 
wide,  scabrous  and  more  or  less  glandular 
and  pubescent;  achenes  mostly  3.5-4.5  mm 
long.  Cultivated  historically  in  Utah  for  fiber 


Table  1.      Families,  genera,  and  species  treated. 


No.  Genera 

No. 

Species 

Family 

Cultivated    Indigenous 

.Aquifoliaceae 

1 

2 

Canabinaceae 

2 

1 

1 

Ericaceae 

6 

0 

11 

Krameriaceae 

1 

0 

2 

Magnoliaceae 

2 

4 

Moraceae 

3 

4 

Oleaceae 

6 

14 

4 

Pyrolaceae 

3 

0 

7 

Resedaceae 

1 

1 

Tamaricaceae 

1 

3 

Tiliaceae 

1 

7 

TOTAL 

27 

36 

25 

produced  from  the  stems,  the  commercial 
source  of  hemp;  currently  sporadic,  or  else 
grown  illegally  for  its  intoxicant  properties. 
Utah  materials  are  sufficiently  rare  as  to  give 
only  hints  as  to  the  classification  below  the 
species  level.  It  seems  likely,  however,  that, 
at  least  historically,  two  phases  have  been 
grown  in  the  state  (for  a  complete  review  see 
Small  and  Cronquist  1.  c).  Nineteenth-cen- 
tury plantings  for  hemp  likely  belonged  to 
ssp.  sativa,  demonstrated  to  have  only  limited 
intoxicant  ability.  At  least  some  of  the  recent 
introductions  clearly  belong  to  ssp.  indica 
(Lam.)  Small  &  Cronq.  (C.  indica  Lam.), 
which  has  demonstrated  high  intoxicant  lev- 
els. 


Humulus  L. 

Plants  herbaceous,  twining,  perennial 
vines;  stems  scabrous;  leaves  opposite,  broad- 
ly 3-  to  5-lobed;  flowers  small,  inconspicuous, 
the  staminate  in  axillary  panicles;  sepals  5, 
distinct;  stamens  5;  pistillate  flowers  in  short 
spikes,  in  pairs,  with  each  pair  subtended  by 
a  foliaceous  bract;  calyx  membraneous,  un- 
lobed,  closely  covering  the  ovary;  stigmas  2, 
elongate;  fruit  an  achene  enclosed  by  the 
persistent  calyx  and  accrescent  bracts. 

Humulus  americanus  Nutt.  American 
Hop.  Plants  twining,  the  stems  to  20  dm  long 
or  longer;  leaves  ovate  to  orbicular  in  out- 
line, deeply  cordate  ba.sally,  mostly  3-15  cm 
long  and  2.8-16  cm  wide,  the  lobes  serrate  to 
doubly  so,  attenuate  to  acimiinate  apically, 
rough-hairy  above,  glandular-dotted  beneath; 
fruiting  spikes  usually  2-3.5  (4)  cm  long  at 
maturity.  Twining  over  shrubs  and  other  veg- 
etation at  lower  and  middle  elevations  in 
Duchesne,  (iarfield.  Grand,  Millard,  Piute, 
Salt  Lake,  Summit,  Uintah,  Utah,  Wasatch, 
Washington,  and  Weber  counties,  and  prob- 
ablv  throughout  Utah;  widespread  in  North 
America.  The  hop  of  commerce,  H.  hipulus 


40 


Great  Basin  Naturalist 


Vol.  40,  No.  1 


L.,  or  European  hop,  is  grown  in  the  United 
States,  where  it  has  escaped  and  persists. 
Though  not  definitely  known  for  Utah,  the 
European  hop  might  occur  here.  It  can  be 
distinguished  by  its  unlobed  leaves,  or  when 
lobed,  the  terminal  lobe  is  less  than  twice 
longer  than  broad;  27(v). 

Ericaceae 

Heath  Family 

Shrubs  or  subshnibs;  leaves  simple,  alter- 
nate, sometimes  leathery  or  persistent;  flow- 


ers perfect,  regular,  axillary,  in  terminal  clus- 
ters, or  solitary;  sepals  mostly  4  or  5,  distinct 
or  more  or  less  connate;  petals  mostly  4  or  5, 
connate  or  distinct,  the  corolla  rotate  to  fun- 
nelform  or  urn  shaped;  stamens  as  many  as 
the  corolla  lobes  and  alternate  with  them  or 
twice  as  many,  the  anthers  dehiscent  by  ter- 
minal pores  or  by  longitudinal  slits;  pistils  1, 
the  ovary  superior  or  inferior,  usually  with 
4-10  carpels  and  locules;  styles  1,  the  stigma 
capitate  or  lobed;  fruit  a  capsule  or  a  berry. 


1.  Ovary  inferior  or  apparently  so  2 

-  Ovary  superior 3 

2(1).  Plants  prostrate  shrublets,  rooting  along  the  stems;  ovary  superior  but  sur- 
rounded by  the  fleshy  calyx  when  ripe  and  apparently  inferior  

Gaultheria  (hiimifusa) 

-  Plants  erect  or  ascending,  rooting  only  at  the  base;  ovary  inferior Vciccinium 

3(1).  Flowers  borne  in  terminal  corymbs,  white,  the  segments  of  the  corolla  much 
longer  than  the  short  tube;  leaves  punctate  below  with  yellow-glandular  dots  ... 
Ledum 

-  Flowers  solitary  and  axillary  or  in  axillary  racemes,  rarely  terminal,  pink  to 
lavender,  the  segments  of  the  corolla  much  shorter  than  the  tube;  leaves 
lacking  glandular  punctae  4 

4(3).       Corolla  broadly  saucer  shaped  or  ratate,  not  constricted  at  the  apex  Kahnia 

Corolla  campanulate  to  urn  shaped,  often  more  or  less  constricted  at  the 
throat 5 

5(4).       Corolla  campanulate;  anthers  lacking  appendages;  fruit  a  capsule  embedded  in 

a  fleshy  calyx Gaultheria  {humifusa) 

Corolla  urn  shaped;  anthers  2-awned;  fruit  a  berry  Arctostaphylos 


Arctostaphylos  Adams 

Evergreen  prostrate  to  ascending  or  erect 
shnibs,  often  with  purplish  to  orange  brown, 
smooth  bark;  leaves  alternate,  simple,  entire, 
leathery-thickened;  flowers  in  terminal  pan- 
icles or  racemes,  perfect,  regular;  sepals  usu- 
ally 5;  petals  usually  5,  united  almost  to  the 
tips;  corolla  urn  shaped;  stamens  usually  10, 
included;  anthers  opening  by  falsely  terminal 


pores,  each  with  2  hornlike  appendages; 
ovary  superior,  usually  5-loculed;  fruit  fleshy, 
berry  like,  1-  to  several-seeded. 

Adams,  J.  E.  1940.  A  systematic  study  of  the 
genus  Arctostaphylos.  J.  Elisha  Mitchell 
Soc.  56:  1-62. 

Eastwood,  A.  1934.  A  revision  of  Arctosta- 
phylos with  keys  and  descriptions.  Leaft. 
West.  Bot.  1:  105-127. 


Plants  with  creeping-prostrate  stems;  leaves  obovate-spatulate,  commonly  less 
than  1.5  cm  long X.  uva-ursi 

Plants  with  stems  ascending  to  erect;   leaves  mostlv  ovate  to  lanceolate  or 
elliptic,  often  more  than  2  cm  long 2 


March  1980 


Welsh:  Utah  Flora,  Miscellaneous  Families 


41 


2(1).        Calvx  and  pedicels  puherulent  with  spreading  glandular  hairs;  twigs  and  leaves 

puberulent  throughout  with  spreading  hairs;  plants  of  Washington  County  

A.  pringlei 

-  Calyx  glabrous  or  nearly  so;  twigs  and  leaves  puberulent  or  sessile  to  sparingly 

stipitate-glandular  or  almost  or  quite  glabrous;  plants  of  various  distribution  3 

3(2).        Twigs  and  axis  of  inflorescence  white-puberulent,  not  glandular;   plants  of 

Washington  and  Kane  counties  A.  piingens 

Twigs   and   axis   of   inflorescence   glandular   to   glandular-puberulent;    plants 
widely  distributed A.  patulu 


Arctostaphylos  patula  Greene.  Green-leaf 
Manzanita.  {Uva-iirsi  patiila  (Greene) 
Abranis;  A.  piingens  var.  phityphylla  Gray; 
A.  platyphyUa  (Gray)  Kuntze;  A.  obtiisifolia 
Piper;  A.  patula  var.  incarnata  Jeps.;  A.  pi- 
nctorum  Rollins;  A.  parryana  var.  pinetorum 
(Rollins)  Weislander  &  Schreiber).  Rounded 
shrubs  with  gnarled  stems  to  15  cm  long  or 
more,  the  bark  smooth,  cinnamon  to  reddish 
brown  or  purplish  in  color;  branchlets 
glandular-puberulent  and  sometimes  with 
long-spreading  hairs  as  well;  leaf  blades  (0.8) 
1.8-4.7  cm  long,  (0.6)  1.5-4  cm  wide,  ovate 
to  elliptic,  lanceolate,  or  orbicular,  obtvise  to 
acute  apically,  rounded  to  truncate  basally, 
glabrous  or  glandular,  yellow  green;  petioles 
pubescent  like  the  twigs;  inflorescence  pan- 
iculate, the  axis  and  bracts  glandular-pub- 
enilent  and  sometimes  with  some  long  hairs; 
pedicels  glabrous;  sepals  glabrous;  corolla 
pink  to  white,  5-8  mm  long;  ovary  glabrous; 
fruit  8-11  mm  thick,  depressed-globose, 
glabrous,  white  to  brown,  with  nutlets  sepa- 
rable or  not.  Usually  associated  with  ponde- 
rosa  pine  at  1520  to  2830  m  in  Beaver,  Duch- 
esne, Garfield,  Iron,  Juab,  Kane,  Millard,  San 
Juan,  Sanpete,  Sevier,  Summit,  Tooele,  Uin- 
tah, Utah,  Wasatch,  and  Washington  coun- 
ties; Colorado,  Nevada,  Oregon,  Arizona,  and 
California.  Arctostaphylos  patula  and  A. 
platyphyUa  both  date  as  species  from  the 
same  year,  1891;  the  question  of  which  has 
priority  is  difficult  to  ascertain;  64(x). 

Arctostaphylos  pringlei  Parry.  Pink- 
bracted  Manzanita.  Rounded,  erect  shrubs  to 
20  dm  tall  or  more,  the  bark  smooth,  dull  red 
l)rown;  branchlets  densely  glandular-hairy 
with  long-spreading  hairs;  leaf  blades  (1.2) 
1.8-4.2  cm  long,  (0.4)  0.8-2  cm  wide,  elliptic 
to  lance-elliptic  or  lanceolate,  obtuse  to 
acute  apically,  truncate  to  rounded  or  obtuse 
basally,  glandular-pubescent,  gray  green;  pe- 


tioles pubescent  like  the  twigs;  inflorescence 
paniculate  or  racemose,  the  axis  and  bracts 
glandular-hairy;  corolla  pink,  6.5-8.5  mm 
long;  ovary  glandular-hairy;  ovary  glandular- 
hairy;  fruit  6-10  mm  thick,  ovoid,  glandular- 
hairy,  red,  with  nutlets  inseparable.  Oak-juni- 
per community,  1840-2750  m,  in  Washing- 
ton County;  Arizona,  California  and  Baja 
California;  4(o). 

Arctostaphylos  pungens  H.B.K.  Mexican 
Manzanita.  Erect  or  ascending,  rounded 
shrubs  to  20  dm  tall  or  more,  the  bark 
smooth,  red  brown;  branchlets  canescent 
with  a  dense  pubescence;  leaf  blades  1.6-4.7 
(6)  cm  long,  0.5-3.2  cm  wide,  ovate  to  ellip- 
tic or  oblong,  rounded  to  acute  apically, 
acute  to  rounded  basally,  puberulent  on  one 
or  both  sides,  bright  green;  petioles  pub- 
escent like  the  twigs;  inflorescence  pan- 
iculate, the  axis  and  bracts  canescent;  pedi- 
cels glabrous;  sepals  glabrous;  corolla  pink  to 
white,  5.5-8.5  mm  long;  ovary  glabrous;  fruit 
5-8  mm  thick,  depressed-globose,  glabrous, 
brownish  red,  with  nutlets  separable  or  not. 
Pinyon,  juniper,  live  oak  communities, 
920-2750  m,  in  Washington  and  Kane  (At- 
wood  .3538  BRY)  counties;  California,  Ari- 
zona, New  Mexico,  Texas;  Mexico;  19(iii). 

Arctostaphylos  uva-ursi  (L.)  Spreng.  Kin- 
nikinnick,  Bearberry,  Sandberry.  {Arbtitus 
uva-ursi  L.;  Uva-ursi  procumhens  Moench; 
Mairania  uva-ursi  (L.)  Desv.;  U.  buxifolia  S. 
F.  Gray;  A.  officinalis  Wimm.  &  Grab.;  A. 
procumhens  in  Mey.  &  Elkan;  U.  uva-ursi 
(L.)  Britt.  in  Britt.  &  Br.;  A.  media  Greene;  A. 
uva-ursi  var.  coactilis  Fern.  &  Macbr.;  A. 
uva-ursi  var.  adenotricha  Fern.  &  Macbr.). 
Prostrate  shrub  with  stoloniferous  rooting 
steins,  mat-forming,  the  branches  ascending, 
the  intemodes  usually  apparent,  puberulent 
and  sometimes  glandular,  the  bark  exfoliating 
exposing  dull  brown  imder  bark;  leaf  blades 


42 


Great  Basin  Naturalist 


Vol.  40,  No.  1 


(0.6)  1-2.7  (3)  cm  long,  0.3-1.2  wide,  oblan- 
ceolate  to  spatulate,  rounded  apically,  cu- 
neate  to  acute  basally,  glabrous  or  pub- 
enilent,  especially  on  the  margins,  green; 
inflorescence  racemose,  the  axis  and  bracts 
glandular;  pedicels  glabrous  or  sparingly 
puberulent;  sepals  glabrous;  corolla  pink  to 
white,  4-5.2  mm  long;  ovary  glabrous;  fruit 
6-11  mm  thick,  globose,  bright  red,  with  sep- 
arable nutlets.  Ground  layer  in  coniferous 
forests,  at  2140-3350  m,  in  Daggett,  Duch- 
esne, Garfield,  Salt  Lake,  Sevier,  Summit, 
Uintah,  and  Wasatch  counties;  Alaska  and 
Yukon  east  to  the  Atlantic  and  south  to  Cali- 
fornia, New  Mexico,  Illinois,  and  Georgia; 
Eurasia;  15(ii). 


Gaultheria  L. 

Prostrate  shrubs,  the  branches  rooting; 
leaves  alternate,  thin,  serrulate;  flowers  ax- 
illary, solitary,  perfect,  regular;  calyx  5- 
lobed,  united,  enlarging  and  becoming  fleshy 
at  maturity;  corolla  campanulate,  the  lobes 
shorter  than  the  tube;  stamens  usually  10,  in- 
cluded, the  filaments  flattened,  tapering  to 
the  apex;  anthers  opening  by  terminal  pores, 
not  awned;  ovary  superior,  usually  5-loculed; 
fniit  a  loculicidally  dehiscent  capsule  en- 
closed by  the  fleshy  expanded  calyx. 

Gaultheria  humifusa  (Grab.)  Rydb.  Al- 
pine Wintergreen.  {Vaccinium  humifusum 
Grab.;  G.  myrsinites  Hook.).  Prostrate, 
scarcely  woody  plants  with  creeping,  rooting 
stems  to  2  dm  long,  glabrous  or  puberulent; 
leaves  0.6-1.5  cm  long,  0.4-1.3  cm  wide,  oval 
to  ovate  or  elliptic,  rounded  to  obtuse  api- 
cally and  basally,  serrulate;  flowers  solitary, 
axillary;  calyx  glabrous;  corolla  3-4  mm  long, 
campanulate,  pink;  fruit  5-7  mm  thick,  sub- 
globose,  red.  Ground  layer  in  coniferous  for- 
ests and  margins,  2900-3350  m,  in  Duchesne 
and  Summit  counties,  and  possibly  elsewhere; 
Colorado  westward  to  California  and  north 
to  Alberta  and  British  Columbia;  5(i). 

Kalmia  L. 

Low  shrubs  with  puberulent  branches; 
leaves  opposite,  evergreen,  leathery,  decur- 
rent,  entire,  revolute,  glaucous  beneath;  flow- 
ers in  terminal  leafy-bracted  corymbs  or  soli- 


tary, perfect,  regular;  calyx  5-lobed,  the 
segments  almost  distinct;  corolla  bowl 
shaped,  the  lobes  shorter  than  the  tube,  the 
tube  with  10  pouches  in  which  the  anthers 
are  enclosed  in  bud;  stamens  usually  10,  the 
filaments  flattened,  hairy  below;  anthers 
opening  throughout,  unawned;  ovary  superi- 
or, 5-loculed;  fruit  a  septicidally  dehiscent 
capsule. 

Kalmia  microphylla  (Hook.)  Heller.  Bog 
Laurel.  {K.  glauca  var.  microphylla  Hook.;  K. 
polifolia  var.  microphylla  (Hook.)  Rhed.). 
Erect  slender  shrubs,  0.7-1.5  dm  tall;  leaves 
0.6-1.8  (3)  cm  long,  0.2-0.8  (1.2)  cm  wide, 
lance-oblong  to  elliptic,  revolute,  shining  and 
green  above,  grayish  beneath;  corymbs  most- 
ly 2-  to  6-flowered,  the  pedicels  1-3  cm  long; 
sepals  glabrous,  ciliate;  corollas  11-14  mm 
broad,  pink;  capsules  4-6  mm  broad.  Alpine 
meadows  and  lake  margins,  2900-3800  m,  in 
Daggett  (?),  Duchesne,  Summit,  and  Uintah 
counties;  Alaska  and  Yukon  south  to  Califor- 
nia and  Colorado;  ll(i). 


Ledum  L. 

Erect  or  spreading  shrubs  with  glandular- 
pubenilent  branchlets;  leaves  alternate,  ever- 
green, leathery,  entire,  revolute,  pale  below; 
flowers  in  terminal  corymbs,  perfect,  regular; 
calyx  small,  the  segments  almost  distinct;  co- 
rolla rotate,  the  5  petals  distinct  or  nearly  so; 
stamens  usually  5-10,  the  filaments  almost 
filiform,  usually  hairy  below;  anthers  opening 
by  terminal  pores,  unawned;  ovary  superior, 
5-loculed;  fruit  a  septicidally  5-valved  cap- 
sule, opening  at  the  base.  Note:  At  least  some 
species  of  this  genus  are  poisonous  to  live- 
stock. 

Ledum  glandulosum  Nutt.  Trapper's  Tea. 
Plants  mostly  5-15  dm  tall,  the  branchlets 
puberulent  and  glandular  dotted;  leaves 
1.1-3.4  (4)  cm  long,  0.4-1.4  (1.8)  cm  wide,  el- 
liptic to  oblong,  rounded  to  acute  apically 
and  basally,  green  above,  pale  to  grayish  be- 
neath, glandular,  the  margin  more  or  less 
revolute;  flowers  white,  the  segments  to  5 
mm  long  or  more;  pedicels  commonly  1-2.5 
cm  long,  puberulent  near  the  base;  capsules 
3-6  mm  long,  puberulent  and  glandular. 
Meadows,  stream  banks,  and  bogs  in  open 
forest,  2600-3050  m,  in  Duchesne^  Salt  Lake, 


March  1980 


Welsh:  Utah  Flora,  Miscellaneous  Families 


43 


Summit,  and  Uintah  counties;  British  Cohim- 
bia  east  to  Montana  and  south  to  Cahfornia, 
Nevada,  and  Wyoming;  9(o). 

Vaccinium  L. 

Decumbent-ascending  to  erect  shrubs; 
leaves  alternate,  deciduous,  or  more  or  less 
evergreen,  entire  or  serrulate,  flat,  green  or 
pale  beneath;  flowers  solitary,  axillary,  or  in 
terminal  clusters,  perfect,  regular;  calyx  4-  to 


6-lobed,  united  at  the  base;  corolla  urn 
shaped  or  campanulate,  the  4-6  lobes  shorter 
than  the  tube;  stamens  8-12,  the  fllaments 
usually  glabrous;  anthers  opening  by  pores  at 
the  ends  of  tubular  beaks,  usually  2-awned; 
ovary  inferior,  usually  4-locular;  fruit  a  sever- 
al-seeded berry. 

Camp,  W.  H.  1942.  A  survey  of  the  Ameri- 
can species  of  Vaccinum,  subgenus  Eu- 
vaccinium.  Brittonia  4:  205-247. 


1.  Branches  bright  green  and  angled;  plants  often  less  than  3  dm  tall  2 

-  Branches  neither  bright  green  nor  angled,  or  sometimes  irregularly  angled 
when  dry;  plants  often  more  than  3  dm  tall 3 

2(1).       Fruit  red;  grooves  of  branches  usually  glabrous;  leaves  often  less  than  12  mm 

long V.  scoparium 

-  Fniit  blue  black  or  black;  grooves  of  branches  usually  puberulent;  leaves  often 
over  12  mm  long  V.  myrtillus 

3(1).       Flowers  in  clusters  of  2-4,  or  solitary;  leaves  entire;  calyx  deeply  lobed,  the 

lobes  persistent  in  fruit  V.  occidentale 

Flowers  solitary  in  leaf  axils;  leaves  more  or  less  serrate;  calyx  shallowly  lobed, 

the  lobes  deciduous  in  fruit  4 

4(3).       Plants  mostly  1-3  dm  tall;  leaves  serrate  above  the  middle  and  uncon- 
spicuously  below  the  middle,  mainly  1-3  (4)  cm  long,  oblanceolate  to  obovate  . 

V.  caespitosiwi 

Plants  mostly  4-7  dm  tall  or  more;  leaves  serrate  to  the  base  or  nearly  so, 
commonly  2-6  cm  long,  elliptic  to  ovate V.  membranaceum 


Vaccinium  caespitosum  Miehx.  Dwarf 
Huckleberry.  Plants  mostly  1-3  dm  tall; 
twigs  brownish,  somewhat  angled,  pub- 
erulent or  glabrous;  leaves  0.7-4  cm  long, 
0.3-2  cm  wide,  oblanceolate  to  obovate,  ob- 
tuse or  less  commonly  acute  to  rounded  api- 
cally,  usually  cuneate  basally,  serRilate  from 
tip  to  below  the  middle;  flowers  solitary,  ax- 
illary, whitish  to  pink,  the  corollas  5-6  mm 
long,  twice  as  long  as  thick;  calyx  obscurely 
lobed,  the  lobes  deciduous  in  fruit;  berries 
blue  glaucous,  subglobose,  5-8  mm  broad, 
edible  and  good.  Streamsides,  meadows,  and 
rock  outcrops,  2,227-3,416  m  elevation,  in 
the  Uinta  Mountains  and  Boulder  Mountains, 
in  Daggett,  Duchesne,  Garfield,  Summit,  and 
Uintah  counties;  Alaska  and  Yukon  east  to 
Newfoundland  and  New  Hampshire,  and 
south  to  California  and  Colorado.  Materials 
from  Utah  have  previously  passed  under  the 
names  V.  membranaceum  Dougl.  (see  below) 


and  V.  globulare  Rydb.  The  latter  is  not 
known  for  the  state;  10(i). 

Vaccinium  membranaceum  Dougl.  Moun- 
tain Huckleberry.  Shrubs  mostly  3-7  dm  tall 
or  more;  twigs  brownish,  glabrous  or  pub- 
CRilent;  leaves  1.8-7  cm  long,  1-3.4  cm  wide, 
elliptic  or  less  commonly  ovate  or  obovate, 
acute  to  obtuse  apically,  acute  to  rounded 
basally,  serrate  almost  throughout;  flowers 
solitary,  axillary,  yellowish  pink,  the  corollas 
about  6  mm  long,  about  one-third  longer 
than  broad;  calyx  obscurely  lobed,  the  lobes 
deciduous;  berries  purple,  not  glaucous,  7-9 
mm  broad,  edible  and  good.  Slopes  in  aspen- 
conifer  and  spruce-fir  woods,  2,500  to  2,775 
in,  in  Cache,  Carbon,  Duchesne,  Salt  Lake 
(?),  and  Summit  counties;  British  Columbia 
southward  to  California,  Idaho,  and  Mon- 
tana; 6(0). 

Vaccinium  myrtillus  L.  Dwarf  Billberry. 
(V.  oreophilum  Rydb.,  in  part,  the  type  from 


44 


Great  Basin  Naturalist 


Vol.  40,  No.  1 


the  Uinta  Mountains.)  Plants  mostly  0.5-3  dm 
tall;  twigs  seldom  numerous  and  broomlike, 
green,  sharply  angled,  puberulent;  leaves 
1.1-3.9  cm  long,  0.6-1.6  cm  wide,  ovate  to 
lanceolate  or  elliptic,  acute  to  obtuse  api- 
cally,  obtuse  to  roimded  basally,  serrulate  al- 
inos't  or  quite  from  base  to  apex;  flowers  soli- 
tary, axillary,  pink,  the  corollas  4-5  mm  long; 
calyx  shallowly  lobed;  berry  usually  bluish, 
5-8  mm  broad.  Ground  layer  in  coniferous 
forests,  2750-3200  m,  in  the  Uinta,  Wasatch, 
and  LaSal  moimtains  (Daggett,  Duchesne, 
San  Juan,  Summit,  and  Uintah  counties), 
where  evidently  not  common;  British  Colum- 
bia and  .\lberta  south  to  Arizona  and  New 
Mexico;  Eurasia.  Vaccinium  myrtillus  is  a 
near  congener  of  the  very  common  V.  scopa- 
riwn  and  can  be  distinguished  by  the  larger 
size  of  its  leaves  and  flowers  and  by  the  pub- 
erulent stems;  6(i). 

Vaccinium  occidentale  Gray.  Western 
Huckleberry.  Plants  mostly  2-6  dm  tall,  the 
twigs  round,  usually  glabrous;  leaves  0.6-2.1 
cm  long,  0.4-1.2  cm  wide,  oblanceolate, 
rounded  to  obtuse  apically,  acute  basally,  en- 
tire; flowers  2-4,  or  less  commonly  solitary  in 
the  axils,  pinkish,  the  corollas  3.5-6  mm  long; 
calvx  definitely  lobed,  the  lobes  persistent  in 
fniit;  berries  blue,  glaucous,  4-6  mm  thick. 
Meadows,  streamsides,  and  forest  margins, 
2750-3100  m,  in  the  Uinta  Mountains  in 
Daggett  (?),  Duchesne,  Summit,  Uintah,  and 
Wasatch  counties;  British  Columbia  south  to 
California  and  Idaho;  12(iv). 

Vaccinium  scoparium  Leiburg.  Grouse- 
berry.  (V.  myrtillus  var.  microphijUum  Hook.; 
V.  microphijllwn  (Hook.)  Rydb.,  not  Rein.;  V. 
enjthrococcum  Rydb.).  Plants  mostly  1-2.5 
dm   tall,   the   twigs  numerous,  broomlike, 


sharply  angled,  usually  glabrous;  leaves 
0.6-1.3  cm  long,  0.3-0.7  mm  wide,  ovate,  ob- 
tuse to  acute  apically,  roiuided  to  obtuse  ba- 
sally, serrulate  throughout;  flowers  solitary, 
axillary,  pinkish,  the  corollas  2.5-3.5  mm 
long;  calyx  very  shallowly  lobed;  berry  bright 
red,  drying  red  purple,  4-6  mm  thick.  Com- 
mon component  of  ground  layer  in  con- 
iferous forests  and  forest  margins,  2450-3200 
m,  in  the  Uinta  Mountains  in  Daggett,  Duch- 
esne, Summit,  Uintah,  and  Wasatch  counties; 
British  Columbia  and  Alberta  south  to  Cali- 
fornia and  Colorado;  20(i). 

Krameriaceae 
Ratany  Family 

Shrubs,  with  divaricate  branches;  herbage 
grayish  pubescent;  leaves  alternate,  simple, 
entire  extipulate;  flowers  perfect,  irregular, 
solitary,  axillary;  pedicels  usually  with  2  op- 
posite foliacious  bracts;  sepals  4  or  5,  un- 
equal; petals  5,  the  upper  3  long  clawed,  dis- 
tinct or  partially  connate  and  often  purplish 
in  color,  the  2  others  broad,  thick,  sessile, 
usually  greenish  and  glandlike;  stamens  4, 
free  or  adnate  to  claw  of  upper  petal,  the  an- 
thers dehiscent  by  pores;  ovary  superior,  1- 
loculed;  ovules  2;  fruit  an  indehiscent  pod, 
armed  with  prickles. 

A  family  of  the  Western  Hemisphere  of  a 
single  genus  with  about  25  species  from 
South  America  to  southern  United  States. 

Krameria  L. 

A  single  genus  with  characteristics  of  the 
family. 


Branchlets  modified  as  thickened  thorns  0.8-1.2  mm  in  diameter  at  base;  spines 

of  fruit  barbed  at  apex  only K.  graiji 

Branchlets  not  modified  as  thorns  or  if  so  then  less  than  0.6  mm  in  diameter; 
spines  of  fruit  with  barbs  scattered  or,  rarely,  barbless K.  pawiflora 


Krameria  grayi  Rose  &  Painter.  White  Ra- 
tany. Shrubs  branched,  2.5-6  dm  tall  and  as 
wide;  leaves  5-7  (25)  mm  long,  1-3  mm 
wide,  lance-ovate  to  lanceolate,  elliptic  or 
oblong,  more  or  less  spinulose-tipped,  tomen- 
tose  on  both  surfaces;  pedicels  not  glandular- 


pubescent;  upper  petals  2.5-3.5  mm  long, 
0.3-0.5  mm  wide,  yellowish  with  a  purplish 
tip;  sepals  4.5-6.5  mm  long,  villous-pilose 
dorsally,  pilose  to  glabrate  within,  purplish; 
prickles  of  the  fruit  2-6  mm  long  at  maturity, 
each  with  a  whorl  of  barbs  at  the  apex;  pods 


March  1980 


Welsh:  Utah  Flora,  Miscellaneous  Families 


45 


subglobose,  6-10  mm  in  diameter,  hirsute 
over  the  surface  and  on  bases  of  prickles. 
Blackhmsh  and  creosote  bush  communities  at 
670-1170  m  in  western  Washington  Co.;  Cal- 
ifornia, Nevada,  Arizona,  New  Mexico, 
Texas,  and  Mexico;  2(i). 

Krameria  parvifolia  Benth.  Range  Ratany. 
(K.  glandulosa  Rose  and  Painter;  K.  parvi- 
folia var.  glandulosa  (Rose  and  Painter) 
Macbr.;  K.  iniparata  (Britton)  Macbr.) 
Shrubs,  intricately  branched,  2-6  dm  tall  and 
as  wide;  leaves  3-15  mm  long,  0.3-1  mm 
wide,  linear  to  oblong,  callous-  to  spinulose- 
tipped,  tomentose  on  both  surfaces;  pedicels 
glandular  or  not;  upper  petals  2.5-2.8  mm 
long,  0.7-1.2  mm  wide,  yellowish;  sepals  4-6 
mm  long,  strigulose  dorsally,  glabrous  within, 
pinkish  to  purplish;  prickles  of  fruit  2-5  mm 
long,  retrorsely  barbed  along  the  rachis;  pods 
subglobose,  5-9  mm  in  diameter,  pilose-hir- 
sute on  the  surface.  Joshua  tree,  blackbrush, 
creosote  bush,  and  bursage  communities, 
750-1600  m,  in  Washington  Co.;  California, 
Nevada,  Arizona,  New  Mexico,  Texas,  and 
Mexico.  The  materials  demonstrate  variation 


in  glandular  condition  of  pedicels,  sepals,  and 
bracts.  The  variation  seems  to  be  haphazard, 
with  little  or  no  correlation  with  other  fea- 
tures or  with  ecology.  Hence,  included  herein 
as  synonyms  are  those  names  involved  with 
recognition  of  glandular  and  nonglandular 
phases;  16(i). 


Magnoliaceae 
Magnolia  Family 

Deciduous  or  evergreen  trees  or  shrubs; 
leaves  alternate,  simple,  entire  or  lobed,  stip- 
ulate, the  stipules  enclosing  the  buds,  de- 
ciduous or  caducous,  and  leaving  a  circular 
scar;  flowers  regular,  perfect,  solitary,  termi- 
nal and  axillary,  large  and  showy,  the  floral 
parts  spirally  arranged;  sepals  often  3,  the 
petals  6  to  many;  stamens  numerous,  sepa- 
rate, hypogynous,  the  anthers  2-loculed;  pis- 
tils several  to  many,  each  1-loculed  and  1- 
carpelled;  style  1,  the  stigma  1;  fruit  a  follicle 
or  samara. 


1.  Leaves  lobed,  truncate  or  broadly  retuse  at  the  apex;  flowers  borne  after  the 

leaves Liriodendron 

Leaves  entire,  acute,  or  acuminate;  flowers  borne  before  or  after  the  leaves 

Magnolia 


Liriodendron  L. 

Trees,  the  leaves  large  and  4-lobed;  flowers 
large,  inconspicuously  colored;  sepals  3,  soon 
reflexed;  petals  6,  ascending  to  erect,  forming 
a  tuliplike  corolla;  anthers  extrorse;  pistils 
many,  en  masse  becoming  conelike,  the  indi- 
vidual samaras  eventually  deciduous. 

Liriodendron  tulipfera  L.  Tulip  Tree;  Yel- 
low Poplar.  Deciduous,  cultivated  trees  to  40 
m  tall  or  more,  the  tmnks  to  10  dm  in  diam- 
eter or  more;  leaves  long-petioled,  the  blades 
6-15  cm  long  and  almost  as  wide;  flowers 
solitary,  terminal;  sepals  green;  petals  3.7-6 


cm  long,  yellow  green,  with  a  basal  orange 
spot  within;  samaras  narrow,  3-4  cm  long. 
Occasional  shade  tree  in  more  moderate  low 
elevation  portions  of  Utah;  introduced  from 
the  eastern  United  States;  8(o). 

Magnolia  L. 

Trees  or  shrubs;  leaves  large,  entire;  flow- 
ers large,  conspicuous  or  inconspicuous;  se- 
pals 3,  colored  like  the  petals;  petals  6-12, 
erect  or  spreading;  anthers  introrse;  pistils 
many,  en  masse  becoming  conelike,  the  indi- 
vidual follicles  finally  dehiscent. 


Plants  shrubs  or  small   trees,   deciduous;   flowers  showy,   cream   to   pink   or 

suffused  with  rose  or  lavender,  borne  before  the  leaves  appear  M.  soulangeana 

Plants  moderate  to  large  trees,  deciduous  or  evergreen;  flowers  greenish  and 
inconspicuous  or,  if  showy,  then  white  in  color  and  the  trees  evergreen  2 


46 


Great  Basin  Naturalist 


Vol.  40,  No.  1 


Plants  evergreen,  the  leaves  dark  green,  leathery;  flowers  white  M.  grandiflora 

Plants  deciduous,  the  leaves  not  both  dark  green  and  leathery;  flowers  greenish 
ygllovv ^-  acuminata 


Magnolia  acuminata  L.  Cucumber-tree. 
Deciduous  trees  to  30  m  tall  or  more;  leaves 
deciduous,  short-petioled,  the  blades  8-25  (3) 
cm  long  and  4-15  cm  wide;  flowers  solitary, 
terminal;  perianth  greenish  yellow,  5-8  cm 
long.  Cultivated  shade  tree,  uncommon, 
hardy  in  the  major  cities  of  the  state;  in- 
troduced from  the  eastern  United  States;  2(o). 

Magnolia  grandiflora  L.  Bull  Bay.  Ever- 
green trees  to  30  m  tall;  leaves  evergreen, 
short-petioled,  the  blades  mostly  8-20  cm 
long  and  3-8  cm  wide;  flowers  solitary,  ter- 
minal; perianth  white,  mostly  8-12  cm  long. 
Cultivated  ornamental,  uncommon,  not 
hardv  except  in  favorable  sites  in  moderate  to 
warm  portions  in  Utah;  introduced  from 
southeastern  United  States;  l(o). 

Magnolia  soulangeana  Soul.  Showy  Mag- 
nolia. Shrubs  or  small  trees  to  about  4  m  tall; 
leaves  deciduous,  short-petioled,  the  blades 
mostly  8-14  cm  long  and  3.5-10  cm  wide; 
flowers  solitary,  terminal;  perianth  cream  to 
pink  or  suffused  with  rose  or  lavender,  6-12 
cm  long  or  more.  Cultivated  ornamental,  oc- 


casional in  more  moderate  climatic  areas  of 
Utah;  a  hybrid  of  M.  denadata  Descr.  and  M. 
lili flora  Descr.,  both  native  of  China;  3(o). 


MORACEAE 

Mulberry  Family 

Deciduous  trees  or  shrubs  with  milky  juice; 
leaves  alternate,  simple,  pinnately  or  pal- 
mately  veined,  entire,  serrate,  or  lobed,  stipu- 
late, the  stipules  small  and  distinct  or  each 
pair  forming  a  cap  over  the  bud  and  leaving 
a  scar  around  the  stem;  flowers  imperfect, 
minute,  regular,  borne  in  cymes  or  much 
modified  inflorescences;  perianth  of  usually  4 
sepals;  staminate  flowers  with  usually  4  (2  in 
Ficus)  stamens,  the  filaments  distinct;  pistil- 
late flowers  with  or  without  a  4-lobed  per- 
ianth; pistil  1,  the  ovary  superior  to  inferior, 
1-loculed,  the  styles  and  stigmas  2  (1  in  Ma- 
dura)., fruit  a  multiple  (Morus,  Madura)  or  a 
syconium  (Ficus). 


1.  Fniit  a  fleshy  hollow  receptacle  with  flowers  borne  inside  (syconium);  leaves 
palmately  veined  and  lobed;  cultivated  plants  of  Washington  County,  and  of 
greenhouses  elsewhere  Ficus 

-  Fruit  a  multiple  (formed  of  several  flowers  and  a  common  axis);  leaves  various  . 

2 

2(1).  Leaves  crenate-serrate,  palmately  veined  and  often  palmate  lobed  as  well; 
flowers,  both  sterile  and  fertile,  borne  in  catkinlike  spikes;  fruit  seldom  more 
than  1  cm  thick  Morus 

Leaves  entire,  pinnately  veined,  not  lobed;  flowers  borne  in  dissimilar  in- 
florescences, the  sterile  in  racemes,  the  fertile  in  globular  heads;  fruit  globular, 
more  than  5  cm  thick Madura 


Ficus  L. 

Trees  or  large  shrubs;  leaves  alternate, 
simple,  palmately  veined  and  lobed,  the  stip- 
ules forming  a  circular  scar  around  the  stem; 
flowers  minute,  numerous,  borne  inside  a  hol- 
low receptacle  which  ripens  to  form  a  syco- 
nium; staminate  perianth  2-  to  6-parted,  with 
1  or  2  stamens;  pistillate  perianth  reduced  or 


lacking;    receptacles    perfect    or    imperfect; 
fruits  of  individual  flowers  of  achenes. 

Ficus  carica  L.  Common  Fig.  Deciduous 
trees  to  5  m  tall,  rarely  more,  often  sprawling 
in  age;  leaves  prominently  veined,  thick,  to 
25  cm  long  or  more  and  to  20  cm  broad,  3-  to 
5-lobed,  the  lobes  undulate-serrulate;  fruits 
obovoid,  mostly  2.4-4.5  cm  long  and  2-3  cm 
thick.  Cultivated  fruit  plant  in  Washington 


March  1980 


Welsh:  Utah  Flora,  Miscellaneous  Families 


47 


(and  formerly  Garfield,  at  Hite)  County,  frost 
sensitive  elsewhere  except  under  glass;  in- 
troduced from  the  Mediterranean  region  of 
the  Old  World.  This  is  the  fig  of  commerce; 

5(iii). 

Maclura  Nutt. 

Dioecious  trees  with  hard  yellow  wood; 
leaves  entire,  the  stipules  minute,  the  scar  not 
encircling  the  stem;  staminate  flowers  nu- 
merous in  loose,  peduncled,  axillary  heads  or 
umbels,  the  calyx  4-parted  and  with  4  sta- 
mens; pistillate  flowers  coherent  in  dense, 
globose,  axillary  heads,  the  calyx  4-lobed,  the 
single  filiform  style  very  long;  fruit  a  globose 
multiple. 

Maclura  pomifera  (Raf.)  Schneid.  Osage 
Orange.  {Toxylon  pomiferum  Raf.)  Trees  to 
10  m  tall,  rarely  more;  stems  usually  armed 
with  stout  thorns  1-2  cm  long;  leaves  petio- 
late,  the  blades  5-10  cm  long  and  1.8-6.5  cm 


wide,  ovate,  entire,  rounded  to  obtuse  ba- 
sally,  attenuate  to  acuminate  apically;  clus- 
ters of  staminate  flowers  2.5-3.5  cm  across; 
heads  of  pistillate  flowers  2-2.5  cm  across; 
multiple  fruit  mostly  8-14  cm  thick.  Culti- 
vated ornamental  and  botanical  curiosity  of 
low  elevation  regions  in  Utah,  long  per- 
sisting; introduced  from  the  eastern  states. 
The  wood  of  this  tree  is  very  strong,  and  has 
served  as  a  source  of  bows  for  American  In- 
dians and  others;  5(i). 


Morus  L. 

Dioecious  trees;  leaves  palmately  veined, 
serrate  to  dentate,  sometimes  lobed;  stipules 
lanceolate,  the  scar  not  encircling  the  stem; 
flowers  monoecious  or  dioecious,  those  of 
both  sexes  borne  in  stalked,  axillary,  catkin- 
like clusters;  calyx  4-parted;  stamens  4;  styles 
2,  deeply  parted;  fruit  a  multiple. 


Leaves  glabrous  above  and  beneath  or  pubescent  beneath  only  along  main 

veins  and/or  in  vein  axils;  our  common  mulberry M.  alba 

Leaves  pubescent  over  much  of  the  lower  surface,  scabrous  above;  rarely 
cultivated  M.  nisra 


Morus  alba  L.  White  Mulberry.  Cultivated 
ornamental  and  shade  tree  to  10  m  tall  or 
more;  leaves  obliquely  ovate  and  crenate-ser- 
rate  or  irregularly  lobed,  mostly  3.5-14  cm 
long  and  2.5-10  cm  wide,  truncate  to  sub- 
cordate  basally,  acute  to  acuminate  apically, 
glabrous  above  and  below  except  along  veins 
and  in  vein  axils;  fruit  1-2  cm  long  and  0.6-1 
cm  thick,  white,  pink,  red  purple,  or  nearly 
black.  Persisting  and  occasionally  escaping  in 
most  of  Utah  at  lower  elevations;  introduced 
from  China;  widespread  in  North  America. 
This  plant  was  introduced  to  southern  Utah 
to  provide  food  for  silkworms  in  an  attempt 
to  develop  a  silk  industry.  The  fruit  is  edible, 
but  is  consinned  mainly  by  birds.  Reports 
from  Utah  of  red  mulberry,  M.  rubra  L.,  be- 
long here.  Red  mulberry  is  easily  recognized 
by  the  densely  hairy  lower  and  scabrous  up- 
per leaf  surfaces.  So-called  fruitless  phases 
are  known;  24(v). 


Morus  nigra  L.  Black  Mulberry.  Small 
trees  to  about  10  m;  leaves  cordate-ovate, 
crenate-serrate,  seldom  lobed,  5-20  cm  long, 
3-15  cm  wide,  cordate  basally,  obtuse  to 
acuminate  apically,  scabrous  above  and  hairy 
over  veins  and  at  least  some  intervein  areas 
below;  fniit  1-2.5  cm  long  and  to  1  cm  thick, 
purple  to  black.  Sparingly  cultivated  orna- 
mental, mainly  in  warm  regions  of  Washing- 
ton County;  widely  cultivated  in  temperate 
regions  of  the  earth  for  its  fruit;  introduced 
from  Asia;  5(o). 

Oleaceae 
Olive  Family 

Trees  or  shrubs;  leaves  opposite  (or  rarely 
alternate),  simple  or  pinnately  compound, 
stipulate;  flowers  perfect  or  imperfect,  borne 
in  axillary  or  terminal  racemose,  paniculate, 
or  thyrsoid  inflorescences;  calyx  commonly  4- 


48 


Great  Basin  Naturalist 


Vol.  40,  No.  1 


lobed  or  absent;  corolla  usually  of  4  united  or 
distinct  petals,  or  lacking;  stamens  2,  distinct; 
pistil  1,  the  ovary  superior,  2-carpelled  and 
2-loxuled;  style  1,  or  lacking,  the  stigmas  1  or 


2;  fruit  a  berry  (Ligustrinn),  drupe 
(Forestiera),  loculicidal  capsule  {Syringa,  For- 
sythia),  circumscissile  capsule  (Menodora),  or 
samara  {Fraxiniis). 


1.  Leaves  pinnately  compound;  fruit  a  samara Froxinus 

Leaves  simple,  or  rarely  compound;  fruit  various  2 

2(1).       Leaves  ovate  to  orbicular,  crenate-serrate;  fruit  a  samara;  plants  indigenous  

Fraxiniis 

Leaves  various,  but  seldom  ovate  to  orbicular  and  crenate-serrate;  fruit  a 
drupe,  capsule,  or  berry;  plants  cultivated  or  indigenous 3 

3(2).       Shrubs  with  yellow  flowers  appearing  before  the  leaves;  plants  cultivated 

Forsythia 

Shmbs,  subshrubs,  or  trees  with  flowers  variously  colored,  but  if  yellow  then 

not  as  above,  and  appearing  with  or  after  the  leaves  (before  in  Forestiera)  4 

4(3).       Corolla  none  or  rudimentary,  the  flowers  often  unisexual;  fruit  a  drupe;  shrubs 

of  stream  banks  in  southeastern  Utah Forestiera 

-  Corolla  well  developed,  the  flowers  perfect;  fruit  a  berry  or  a  capsule  5 

5(4).       Corolla  yellow;  fruit  a  membranous,  circumscissile  capsule;  plants  indigenous 

subshrubs  of  southern  Utah Menodora 

-  Corolla  commonly  lavender  to  red,  purple,  white,  or  cream;  fruit  a  loculicidal 
capsule  or  a  berry;  plants  cultivated  shrubs  or  trees 6 

6(5).       Flower  clusters  usually  less  than  6  cm  long;  flowers  white  to  cream;  fruit  a 

berry  Ligustrwn 

Flower  clusters  usually  6-30  cm  long  or  more;  flowers  lavender  to  red,  purple, 
lilac,  white  or  cream  Syringa 


Forestiera  Poir. 

Sprawling  indigenous  shrubs;  leaves  oppo- 
site, simple,  serrate  to  entire;  flowers  incon- 
spicuous, polygamo-dioecious,  borne  sessile 
or  in  cymes,  appearing  before  the  leaves;  ca- 
lyx minute,  unequally  5-  to  6-cleft,  or  lack- 
ing; corolla  lacking,  or  rarely  with  2  or  3  pet- 
als; stamens  2  or  4;  ovary  2-loculed,  with  2 
ovules  per  locule;  style  slender;  stigma  1; 
fniit  a  dnipe. 

Forestiera  pubescens  Nutt.  Desert  Olive. 
(F.  neo77iexicana  Gray;  Adelia  neomexicana 
(Gray)  Kuntze;  A.  parvifolia  Gov.).  Shrubs  to 
2  m  tall  or  more;  leaves  (0.8)  L5-5.5  cm 
long,  (0.3)  0.5-2  cm  wide,  oblanceolate  to  el- 
liptic, entire  to  serrulate;  staminate  flowers 
sessile;  pistillate  flowers  pedicellate;  drupe 
5-7  (8)  mm  long,  ellipsoid,  blue  black.  Sandy 
terraces  along  the  Colorado  and  San  Juan  riv- 
ers and  tributaries,   1280-1750  m  in  Grand 


and  San  Juan  counties;  California  eastward  to 
Oklahoma  and  Texas,  and  south  to 
Chihuahua.  The  fruit  is  eaten  by  fox  and  by 
coyotes,  and  tlie  purple-stained,  stone-laden 
fecal  pellets  are  to  be  found  far  from  the  riv- 
ers. Long  known  as  F.  neomexicana,  our  ma- 
terials form  a  portion  of  a  complex  whose 
definition  includes  those  portions  known  as  F. 
pubescens,  and  that  name  has  priority;  8(iii). 

Forsythia  Vahl 

Cultivated  shrubs;  leaves  opposite,  simple 
or  some  compound,  entire  to  serrate;  flowers 
perfect,  showy,  borne  in  axillary  cluster  of 
3-5,  or  solitary,  appearing  before  the  leaves; 
calyx  4-lobed;  corolla  4-lobed,  campanulate; 
stamens  2,  inserted  at  corolla  base;  ovary  2- 
loculed,  with  several  ovules  per  locule;  fruit 
a  loculicidal  capsule,  with  many  winged 
seeds  (ours  seldom  fruiting). 


March  1980  Welsh:  Utah  Flora,  Miscellaneous  Families  49 

Forsythia  suspensa   (Thunb.)  Vahl.  C.old-  Fraxinus  L. 
en-bell.  {St/rin^a  suspciisa  Thunb.).  Shrub  to 

2  m   tall   or  more;  branchlets  somewhat  4-  Deciduous,    cultivated   and/or    indigenous 

angled;  leaves  6-10  cm  long,  ovate  to  lanceo-  trees  or  .shrubs;  winter  buds  often  prominent, 

late,  acute  apically,  cuneaie  to  rounded  ba-  g^y  to  brown  or  black;  leaves  opposite,  pin- 

.sally,  usually  serrate;  flowers  to  25  mm  long,  nately    compound    (simple    in   F.    anomala); 

golden  yellow;  fruit  lance-ovoid,  to   15  mm  flowers  perfect  or  uni.sexual,   inconspicuous, 

long,    .seldom    developing.    Cultivated    orna-  borne  in  panicles;  calyx  4-lobed  or  lacking; 

mental,  common,  persisting  but  not  spread-  toroUa  lacking  or  of  2  or  more,  usually  dis- 

ing  at  lower  elevations  throughout  Utah;  tinct  petals;  stamens  commonly  2;  ovary  2-lo- 

widespread;  introduced  from  China.  Numer-  <-uled;  styles  1;  stigmas  1  or  2;  fruit  a  samara, 
ous  horticultural  varieties  are  present;  2(o). 

1.  Leaves  normally  simple,  sometimes  with  1  or  2  leaflets  below  the  terminal  one; 
indigenous  shrubs  or  .small  trees  of  eastern  and  southern  Utah F.  anomala 

—  Leaves  normally  pinnately  compound  with  5-9  or  more  leaflets;  trees,  either 
indigenous  or  cultivated  2 

2(1).  Branchlets,  petioles,  and  axis  of  panicle  commonly  spreading  hairy,  seldom 
glabrous;  leaflets  usually  5  or  fewer;  trees,  indigenous  in  southwestern  Utah, 
cultivated  elsewhere  F.  velutina 

2.  Branchlets,  petioles,  and  axis  of  panicle  variously  hairy  or  glabrous,  but  seldom 
spreading  hairy;  leaflets  usually  7  or  more;  trees,  cultivated  and  sometimes 
escaping  3 

3(2).        Flowers  appearing  after  leaves  formed;  corolla  present  F.  ornus 

—  Flowers  appearing  before  leaves  formed;  corolla  lacking  4 

4(3).       Fiiiit  with  calyx  persisting  as  a   campanulate  cap;  anthers  oblong;   leaflets 

usually  5-7 5 

—  Fruit  with  calyx  early  deciduous  or  lacking  (except  in  F.  qiiadrangulata); 
anthers  often  cordate;  leaflets  u.sually  9-11  or  more 6 

5(4).  Petiolules  of  middle  and  lower  mature  leaflets  wingless  nearly  their  entire 
length;  winter  buds  black;  leaf  scars  horseshoe  shaped;  wing  of  fruit  terminal, 
not  or  only  slightly  decurrent  F.  americana 

—  Petiolules  of  middle  and  lower  mature  leaflets  winged  nearly  to  the  base;  win- 
ter buds  brown;  leaf  scars  semicircular  or  shield  shaped;  wing  of  fruit  decur- 
rent to  below  the  middle  F.  pennsylvanica 

6(4).        Branchlets  4-sided,  4-angled;  bark  broken  into  plates;  flowers  with  a  minute, 

deciduous  calyx  F.  qtiadrangulata 

—  Branchlets  terete,  not  or  only  slightly  4-angled;  bark  smooth  or  irregularly 
roughened;  flowers  with  calyx  lacking 7 

7(6).        Leaflets  glabrous  or  somewhat  hairy  along  veins  beneath;  commonly  cultivated 

tree F.  excelsior 

—  Leaflets  definitely  pubescent  beneath,  especially  along  the  veins,  the  long  red- 
dish hairs  extended  onto  and  along  the  leaf  rachis;  unconunon  to  rarely 
cultivated  tree F.  nigra 

Fraxjnus  americana  L.  White  Ash.  Moder-  usually  7  (5-9)  6-15  cm  long,  petiolulate, 
ate  to  large  trees;  branchlets  terete,  green  to  ovate  to  lanceolate,  acuminate  apically,  cu- 
brown,  glabrous;  winter  buds  black;  leaflets      neate  to  rounded  ba.sally,  entire  to  serrate, 


50 


Great  Basin  Naturalist 


Vol.  40,  No.  1 


glaucous  beneath  and  usually  glabrous;  an- 
thers oblong,  apiculate;  calyx  persistent;  co- 
rolla lacking;  samaras  (20)  25-35  (50)  mm 
long,  4-7  mm  wide,  the  wing  not  decurrent 
along  the  terete  base.  Shade  tree  of  lower 
elevations  in  Utah;  introduced  from  eastern 
North  America;  5(o) 

Fraxinus  anomala  Torr.  ex  Wats.  Single- 
leaf  Ash.  Shrub  or  small  tree,  commonly 
2.5-4  m  tall,  usually  with  many  stems; 
branchlets  4-angled;  leaves  glabrous,  ovate, 
crenate-serrate  to  subentire,  1.5-6.5  cm  long, 
1-6  cm  wide,  acute  to  obtuse  or  subcordate 
basally,  acute  to  rounded  or  emarginate  api- 
cally,  sometimes  2-  or  3-foliolate  or  transi- 
tional to  simple;  flowers  usually  perfect;  an- 
thers oblong:  calyx  campanulate,  persistent; 
petals  lacking;  samaras  winged  almos^  to  the 
base,  12-27  mm  long,  5-11  mm  wide.  Mixed 
desert  shrub,  mainly  on  rimrock  or  along 
drainages,  and  in  pinyon-juniper  woodland, 
900-2150  m,  in  Emery,  Garfield,  Grand, 
Iron.  Kane,  San  Juan,  Uintah,  Washington, 
and  Wayne  counties;  Colorado,  New  Mexico, 
Arizona,  and  California;  80(xv). 

Fraxinus  excelsior  L.  European  Ash.  Mod- 
erate to  large  trees;  branchlets  terete, 
glabrous;  winter  buds  black;  leaflets  7-11, 
5-12  cm  long,  .sessile,  ovate  to  oblong  or  lan- 
ceolate, acuminate  apically,  cuneate  basally, 
serrate,  green  Ijeneath,  glabrous  except  along 
midrib,  the  hairs  sometimes  extending  to  the 
rachis;  flowers  polygamous;  anthers  ovoid; 
calyx  lacking;  corolla  lacking;  samaras  25-35 
(40)  mm  long,  5-11  mm  wide,  the  blade  de- 
current  almost  or  quite  to  the  base  of  the 
flattened  body.  Shade  tree  of  habitations  and 
streets  at  lower  elevations  throughout  Utah; 
introduced  from  Europe;  11(0). 

Fraxinus  nigra  Marsh.  Black  Ash.  Moder- 
ate trees;  branchlets  terete,  glabrous;  winter 
buds  black;  leaflets  7-11,  mostly  6-12  cm 
long,  sessile,  lanceolate  to  oblong,  obtuse  to 
rounded  basally,  long-acuminate  apicallv, 
serrate,  green  and  glabrous  except  reddish- 
hairy  along  veins,  the  pubescence  extending 
along  the  leaf  rachis;  flowers  dioecious;  an- 
thers oblong;  calyx  lacking;  corolla  lacking: 
siunaras  mostly  25-35  nun  long  and  6-10  mm 
broad,  the  blade  decurrent  to  the  ba.se  of  the 
flattened  body.  Sparingly  cultivated  shade 
tree  at  lower  elevations  in  at  least  the  major 


population  centers;  introduced  from  eastern 
North  America;  4  (o). 

Fraxinus  ornus  L.  Flowering  Ash.  Small  to 
moderate  trees;  branchlets  terete;  winter 
buds  gray  to  brownish;  leaflets  usually  7 
(7-11),  mostly  2.5-7  cm  long,  petiolulate, 
lance-ovate  to  obovate  (terminal  one), 
rounded  to  obtuse  basally,  acuminate  api- 
cally, crenate-serrate,  glabrous  except  along 
midrib;  flowers  perfect;  calyx  present,  per- 
sistent, with  4  triangular-acuminate,  spread- 
ing lobes;  petals  present,  linear;  samaras 
20-25  mm  long,  3-6  mm  wide,  the  blade  ter- 
minal on  the  terete  base.  Rarely  cultivated 
shade  and  ornamental  tree  of  lower  eleva- 
tions in  Utah;  introduced  from  Europe;  2(o). 

Fraxinus  pennsylvanica  Marsh.  Red  Ash. 
Moderate  trees;  branchlets  terete,  pubescent 
to  glabrous,  sometimes  glandular;  winter 
buds  olive  to  brown;  leaflets  usually  7  (5-9), 
6-15  cm  long,  petiolulate,  lanceolate  to 
lance-oblong,  acuminate  apically,  acute  to 
obtuse  or  rounded  basally,  serrate  to  entire, 
green  and  glabrous  or  hairy  (especially  along 
the  veins)  beneath;  anthers  oblong,  apiculate; 
calyx  campamulate,  persistent;  corolla  lack- 
ing; samaras  27-40  (50)  mm  long,  the  blade 
decurrent  to  the  middle  of  the  terete  body  or 
below.  Common  shade  tree  of  lower  eleva- 
tions throughout  Utah,  persisting  and  escap- 
ing, in  Box  Elder,  Cache,  Davis,  Iron,  Juab, 
Millard,  Salt  Lake,  Utah,  and  Washington 
counties;  introduced  from  eastern  North 
America.  The  escaped  plants  have  become 
established  along  streams  and  on  lake  mar- 
gins at  lower  elevations.  Much  of  our  mate- 
rial has  glabrous  branchlets  and  petioles,  and 
has  been  designated  as  F.  pennsylvanica  var. 
lanceolata  (Borkh.)  Sarg.  (F.  lanceolata 
Brokh.).  This  phase  is  known  as  green  ash;  28 
(o). 

Fraxinus  quadrangulata  Michx.  Blue  Ash. 
Small  to  moderate  trees;  branchlets  sharply 
4-angled,  glabrous;  winter  buds  black;  leaflets 
7-11,  mostly  5-12  cm  long,  petiolulate,  lan- 
ceolate to  ovate-lanceolate,  acute  to  rounded 
basally,  acute  to  acuminate  apicallv,  serrate, 
glabrous  except  along  the  midrib  or  rarely 
hairy  over  the  lower  surface;  flowers  perfect; 
calyx  minute,  caducous;  corolla  lacking;  an- 
thers cordate-oblong,  blunt;  samaras  20-40 
(50)  nmi  long,  the  blade  decurrent  to  the  base 
of  the  flattened  body.   Sparingly  cultivated 


March  1980 


Welsh:  Utah  Flora,  Miscellaneous  Families 


51 


shade  tree,  at  lower  elevations  in  Utah;  in- 
troduced from  eastern  North  America;  l(o). 

Fraxinus  vehitina  Torr.  Velvet  Ash,  Ari- 
zona Ash.  [F.  pcnnsylvcmica  Marsh,  ssp.  veln- 
tina  (Torr.)  G.  N.  Miller]  Moderate  trees; 
hranchlets  terete,  denselv  spreading  hairv  to 
merelv  sparingly  so,  or  glabrous;  winter  buds 
brown;  leaflets  3-5  (or  leaves  simple),  lan- 
ceolate to  ovate,  elliptic,  or  orbicular,  pe- 
tiolulate,  cuneate  to  acute  basally,  acuminate 
to  rounded  apicallv,  serrate,  glabrous  or  hairv 
over  the  lower  surface;  flowers  imperfect;  ca- 
lyx campanulate,  persistent;  corolla  lacking; 
anthers  oblong,  apiculate;  samaras  16-34  mm 
long,  4-6  mm  wide,  the  blade  decurrent 
about  half  way  along  the  terete  body.  In- 
digenous tree  of  stream  courses  and  flood 
plains  in  Washington  and  Iron  counties,  and 
cultivated  there  and  elsewhere  in  Utah;  Ari- 
zona and  New  Mexico.  The  phase  with  co- 
riaceous leaflets  has  been  treated  as  var.  co- 
riacea  (Wats.)  Rehd.  (F.  coriacea  Wats.),  but 
seems  not  to  be  worthy  of  taxonomic  recogni- 
tion, at  least  in  Utah;  25(ii). 

Note:  The  shrubby  Fraxinus  dipetala 
Hook.  &  Am.  is  reported  for  Utah  in 
Kearney  &  Peebles,  1961.  Flora  of  Arizona, 
Supplement  p.  1063.  The  related  F.  cuspi- 
data  Torr.  is  known  from  adjacent  Mohave 
and  Coconino  counties,  Arizona,  and  might 
occur  in  Utah.  Both  species  have  corollas 
present;  the  former  has  two  petals  and  the 
latter  has  four. 

LiGUSTRUM  L. 

Shrubs;  leaves  opposite,  simple,  entire; 
flowers  perfect,  white,  showy  through  small, 
borne  in  terminal  penicles,  appearing  after 
the  leaves;  calyx  4-toothed;  corolla  4-lobed, 
funnelform;  stamens  2,  inserted  on  the  co- 
rolla tube;  ovary  2-loculed,  1-  or  2-seeded; 
fruit  a  berry. 

Ligustrum  vulgare  L.  Common  Privet.  De- 
ciduous or  semievergreen  shrub  to  3  m  tall  or 
more,  with  puberulent  to  glabrate  branchlets; 
leaves  2-6  cm  long,  0.8-2  cm  wide,  oblong  to 


elliptic  or  ovate-lanceolate,  glabrous;  panicle 
dense,  3-6  cm  long;  corolla  tube  shorter  than 
the  lobes,  white;  anthers  exserted;  fruit  6-8 
mm  long,  black,  ovoid  to  subglobose.  Culti- 
vated hedge  plant  throughout  Utah  at  lower 
elevations,  persisting  and  escaping;  in- 
troduced from  Europe;  3(o). 

Menodora  Humb.  &  Bonpl. 

Subshrubs;  leaves  alternate  or  the  lower- 
most opposite,  simple,  sessile  or  nearly  so; 
flowers  perfect,  arranged  in  cymes;  calyx  5-to 
15-lobed;  corolla  yellow,  subrotate,  5-  to  6- 
lobed;  stamens  2,  inserted  on  the  corolla 
tube;  ovary  2-loculed,  with  2-4  ovules  per 
locule;  style  slender,  the  stigma  capitate; 
fiiiit  a  circumscissile  capsule. 

Steyermark,  J.  A.  1932.  Revision  of  the  genus 
Menodora.  Ann.  Missouri  Bot.  Card.  19: 
87-176. 

Menodora  scabra  Gray.  Plants  erect  or  as- 
cending, commonly  2-3.5  dm  tall,  woody  at 
the  base  only;  leaves  0.5-2.9  cm  long,  0.2-0.5 
cm  wide,  narrowly  elliptic  to  oblong  or  lan- 
ceolate, glabrous  or  scaberulous;  calyx  mi- 
nutelv  puberulent,  the  lobes  linear;  corolla 
bright  yellow,  subrotate,  the  lobes  5-9  mm 
long;  capsule  8-12  mm  thick,  membranous; 
seeds  4-5  mm  long.  Pinyon-juniper  commu- 
nity, known  in  Utah  only  from  Garfield,  San 
Juan,  and  Washington  counties;  California, 
Arizona,  New  Mexico,  Texas,  and  Mexico; 
3(1). 

Syringa  L. 

Shrubs  or  small  trees;  leaves  opposite, 
simple,  petiolate;  flowers  perfect,  in  terminal 
or  lateral  panicles;  calyx  campanulate,  4- 
toothed  to  nearly  truncate,  persistent;  corolla 
tubular,  the  limb  4-lobed  and  rotate  or  nearly 
so;  stamens  2,  inserted  on  the  corolla  tube; 
ovary  2-loculed,  each  locule  with  usually  2 
ovules;  style  with  a  2-lobed  stigma;  fruit  a 
loculicidal  capsule. 


Flowers  cream  to  whitish,  borne  in  large  panicles;  corolla  tube  1-2.2  mm  long, 
only  half  as  long  as  the   calyx;   fragrance   musky,   not   that   of  lilac;   plants 

flowering  in  summer,  often  treelike  2 

Flowers  lilac,  violet,  purpli.sh,  or  white;  corolla  tube  mostly  6-12  mm  long  or 
more,  several  times  longer  than  the  calyx;  fragrance  usually  of  lilac;  plants 
commonly  shrubs,  flowering  in  spring  or  .summer 3 


52  Great  Basin  Natuhalist  Vol.  40,  No.  1 

2(1).       Leaves  ovate,  rounded  or  subcordate  basally,  the  veins  prominent  on  the  lower 

surface  S.  amiirensis 

—  Leaves  lanceolate  to  elliptic  or  ovate-lanceolate,  obtuse  to  cuneate  basally,  the 
veins  not  prominent S.  pekinensis 

3(1).        Panicles  from  terminal  buds;  leaves  of  current  season  borne  on  branch  with 

panicle;  plants  flowering  in  summer S.  villosa 

—  Panicles  from  lateral  (or  terminal)  buds,  the  terminal  buds  often  lacking;  leaves 
of  current  season  not  borne  on  the  branch  with  panicle;  plants  flowering  in 
springtime 4 

4(3).       Leaves  ovate  to  cordate,  the  base  subcordate  to  obtuse;  our  most  common 

species  S.  vulgaris 

—  Leaves  lanceolate  to  elliptic  or  ovate,  obtuse  to  cuneate  basally;  common  to 
uncommon  5 

5(4).        Leaves  mostly  less  than  4  cm  long,  some  often  irregularly  lobed;  individual 

panicles  short,  mostly  7  cm  long  or  less S.  persica 

—  Leaves  often  over  4  cm  long,  entire;  individual  panicles  usually  8-12  mm  long 
S.  X  chinensis 


Sijringa  amiirensis  (Rupr.)  Rupr.  Amur  Li- 
lac. Shrubs  or  small  trees  to  5  nun  tall  or 
more;  leaf  blades  3.5-13  cm  long,  1.3-8  cm 
wide,  ovate,  rounded  to  obtuse  of  short 
acuminate  basally,  acuminate  apically,  the 
lower  surface  hairy  to  glabrous,  the  veins 
prominent;  petioles  mostly  1-2  cm  long;  pan- 
icles 10-15  cm  long,  the  clusters  of  panicles 
usually  much  longer;  flowers  cream  to  white; 
stamens  exserted.  Sparingly  cultivated  orna- 
mental of  lower  elevations  in  Utah;  in- 
troduced from  Japan;  flowering  in  summer; 
4(0). 

Sijringa  \  chinensis  Willd.  Chinese  Lilac. 
Shrub  to  4  m  tall  or  more,  with  spreading 
and  often  arching  branches;  leaves  2.5-8  cm 
long,  1.5-4  (5)  cm  wide,  ovate-lanceolate,  ob- 
tu.se  to  cuneate  basally,  acuminate  apically, 
glabrous,  the  veins  not  prominent;  petioles 
0.5-1.5  cm  long;  panicles  mostly  8-12  cm 
long,  the  clusters  of  panicles  much  longer; 
flowers  purple  lilac,  or  otherwise;  stamens  in- 
cluded. Commonly  cultivated  ornamental  al- 
most throughout  Utah;  introduced  from  the 
Old  World.  This  plant  is  evidently  of  hybrid 
origin,  having  resulted  from  a  cross  between 
S.  persica  and  S.  vulgaris,  (j.v.;  flowering  in 
springtime;  3(o). 

Syringa  pekinensis  Rupr.  Peking  Lilac. 
Shrub  or  small  tree  to  5  m  tall  or  more,  with 


spreading  branches;  leaves  5-12  cm  long,  2-4 
(6)  cm  wide,  lanceolate  to  ovate,  cuneate  ba- 
sally, acuminate  apically,  glabrous,  the  veins 
not  prominent;  petioles  1.5-3  cm  long;  pan- 
icles mostly  8-15  cm  long,  the  clusters  of 
panicles  to  30  cm  long  or  more;  flowers 
cream  to  yellow  white;  stamens  exserted.  Un- 
common, cultivated  ornamental  in  northern 
Utah,  but  to  be  expected  elsewhere;  in- 
troduced from  China;  flowering  in  early  sum- 
mer; l(o). 

Syringa  persica  L.  Persian  Lilac.  Shrub  to 
2  m  tall,  with  upright  to  arching  branches; 
leaves  1.5-6  cm  long,  0.6-3  cm  wide,  lan- 
ceolate to  elliptic,  sometimes  lobed,  cuneate 
to  obtuse  basally,  acute  to  acuminate  api- 
cally, glabrous,  the  veins  not  prominent;  pet- 
ioles 0.5-1  cm  long;  panicles  mostly  3-7  cm 
long;  flowers  usually  lilac  but  purple  phases 
are  known;  stamens  included.  Uncommonly 
cultivated  ornamental,  especially  in  northern 
Utah;  introduced  from  Asia  Minor;  flowering 
in  springtime;  5(o). 

Syringa  villosa  Vahl.  Shrub  to  3  m  tall, 
rarely  more,  with  erect  branches;  leaves  4-15 
cm  long  (or  more),  2.5-9  cm  wide,  ovate  to 
elliptic,  acute  basally,  abruptly  acuminate 
apically,  spreading  hairy  below,  especially 
along  the  prominent  veins;  petioles  0.8-2  cm 
long;  panicles  mostly  10-18  cm  long;  flowers 


March  1980 


Welsh:  Utah  Flora,  Miscellaneous  Families 


53 


pink  lilac  to  white;  stamens  included.  Spar- 
ingly but  widely  planted  ornamental,  mainly 
in  northern  Utah;  introduced  from  China; 
tlowering  in  summer;  4(o). 

Syringa  vulgaris  L.  Common  Lilac.  Shrubs 
to  4  m  tall  or  more,  the  branches  usually 
erect;  leaves  3-12  cm  long.  1.5-8  cm  wide, 
ovate  to  cordate,  cordate  to  rounded,  trun- 
cate or  obtuse  basally,  acute  to  acuminate 
apically,  glabrous;  petioles  0.8-3  cm  long; 
panicles  mostly  10-20  cm  long;  flowers  lilac 
or  white,  seldom  purple;  stamens  included. 
Abundantly  cultivated  ornamental,  long  per- 
sisting, in  most  of  Utah;  introduced  from  Eu- 
rope; flowering  in  springtime.  Many  horticul- 
tiual  forms  are  known;  7(o). 


Pyrolaceae 
VVintergreen  Family 

Suffrutescent  or  herbaceous  perennials; 
leaves  simple,  alternate,  opposite,  or  appear- 
ing whorled,  evergreen  or  much  reduced  and 
lacking  chlorophyll;  flowers  usually  perfect, 
regular,  or  irregular;  calyx  with  4  or  5  more 
or  less  distinct  sepals;  corolla  with  4  or  5 
more  or  less  distinct  petals  (united  in  Ptew- 
spora):  stamen  twice  as  many  as  the  petals, 
the  anthers  pendulous,  opening  by  appa- 
rently terminal  pores  or  by  slits,  or  the  an- 
thers erect,  awnless  or  2-awned;  pistil  1; 
ovary  superior,  4-  or  5-loculed;  style  1;  fruit  a 
capsule. 


1.  Plants  lacking  chlorophyll;  leaves  reduced  and  scalelike,  reddish,  brownish, 

purple,  or  yellowish  when  fresh,  often  drying  dark  Ptewspom 

Plants  with  chlorophyll  (rarely  without);  leaves  not  reduced  to  scales,  except 
rarely,  commonly  evergreen  2 

2(1).        Flowers  solitary,  the  petals  rotate  or  nearly  so Moneses 

Flowers  few  to  several,  the  petals  concave  3 

3(2).        Stems  leafy,  though  short,  the  leaves  apparently  whorled;  flowers  corymbose; 

staminal  filaments  dilated  near  the  base;  styles  very  short  or  lacking 

ChimaphUa 

Stems  leafy  at  base  only;  flowers  in  elongate  racemes;  filaments  not  especially 
dilated  at  the  base;  styles  in  most  species  over  2  mm  long Pyrola 


Chimaphila  Pursh 

Low  shrubs  from  creeping  rhizomes,  the 
stems  erect  or  ascending;  leaves  evergreen, 
leathery,  apparently  whorled  or  some  alter- 
nate; flowers  (1)  2-several,  borne  in  peduncu- 
late, umbellate  corymbs;  sepals  usually  5,  dis- 
tinct nearly  to  the  base,  persistent;  petals 
usually  5,  distinct,  rotate-cam panulate;  sta- 
mens usually  10,  the  filaments  dilated  and  cil- 
iate  near  the  base;  anthers  awnless,  opening 
by  falsely  terminal  pores  on  short  tubes; 
ovary  superior,  5-lobed  and  5-loculed;  fruit  a 
loculicidallv  dehiscent  capsule. 

Cimaphila  umbellata  (L.)  Bart.  Pipsis- 
sewa.  Prince's  Pine.  {Pyrola  innbellata  L.;  C. 
occklentalis  Rydb.;  C.  umbellata  ssp.  occiden- 
talis  (Rydb.)  Hulten).  Plants  (1)  1.5-2.5  (3) 
dm  tall,  the  stems  glabrous,  only  somewhat 


woody;  leaves  1.5-4.5  (6)  cm  long,  0.5-1.5  (2) 
cm  wide,  elliptic  to  oblanceolate,  cuneate  ba- 
sally, sharply  serrate,  shining  above,  pale  be- 
neath, glabrous;  peduncles  4-7  (10)  cm  long, 
glabrous  or  minutely  glandular-puberulent, 
often  suffused  with  red  purple;  pedicels 
glandular-puberulent  or  merely  puberulent; 
flowers  1-6  or  more,  umbellate-corymbose; 
sepals  erose-ciliate;  petals  5-7  mm  long, 
pink;  stamens  with  expanded  bases  ciliate; 
capsules  5-7  mm  broad.  Coniferous  forests, 
2300-2750  m,  in  Duchesne,  Summit,  Uintah, 
and  Washington  counties;  .\laska,  southward 
to  California  and  Mexico,  east  to  New 
Mexico  and  Colorado,  and  in  the  eastern 
United  States;  Eurasia.  Our  materials  are 
referable  to  var.  occidentalis  (Rvdb.)  Blake; 
4(0). 


54 


Great  Basin  Naturalist 


Vol.  40,  No.  1 


MoNESES  Salisb. 

Rhizomatous  herbs;  leaves  with  chloro- 
phyll, leathery,  persistent,  mainly  basal,  but 
sometimes  opposite  or  in  whorls;  flowers  soli- 
tary, nodding,  borne  on  a  long  peduncle;  se- 
pals usually  5,  persistent;  petals  usually  5,  dis- 
tinct, spreading;  stamens  usually  10,  the 
filaments  tapering  to  the  apex,  the  anthers 
awnless,  nodding,  opening  by  means  of  ap- 
parently terminal  pores;  ovary  superior,  5- 
loculed,  the  stigma  borne  on  an  elongate, 
glabrous  style;  fruit  a  loculicidal  capsule. 

Moneses  uniflora  L.  Single  Delight,  Wax- 
flower.  (M.  reticulata  Nutt.;  M.  uniflora  var. 
reticulata  (Nutt.)  Blake).  Plants  0.4-1.7  dm 
tall;  leaves  (including  petioles)  0.8-4  cm  long, 
0.6-2  cm  broad,  serrate  to  crenate-serrate; 
peduncles  3-15  cm  long,  usually  with  1  or  2 
bracts  along  its  length;  flowers  1.3-2.5  cm 
broad,  white  to  cream;  sepals  1.5-2.5  m  long, 
ciliate;  petals  7-11  mm  long,  spreading;  style 
2-4  mm  long;  capsule  5-8  mm  broad.  Moist 
sites  in  coniferous  forest,  2450-3050  m,  in 
Beaver,  Carbon,  Duchesne,  Emery,  Juab,  Salt 
Lake,  and  Utah  counties;  widely  distributed 
in  North  America;  Eurasia;  9(ii). 


Pterospora  Nutt. 

Plants  herbaceous  saprophytes,  devoid  of 
chlorophyll,  tall,  reddish  or  purplish  brown, 
the  stems  arising  from  a  bulbous  cluster  of 
coralloid  roots;  leaves  alternate,  simple, 
scalelike,  colored  like  the  stems;  flowers  nu- 
merous, borne  in  an  elongate  raceme,  nod- 
ding; calyx  5-lobed;  corolla  urn-shaped,  the 
tube  much  longer  than  the  lobes;  stamens  10, 


the  filaments  flattened,  tapering  to  the  apex, 
glabrous,  the  anthers  with  2  recurved  awns, 
dehiscent  almost  throughout;  ovary  superior, 
5-loculed,  the  stigma  borne  on  a  short  thick 
style;  fruit  a  loculicidal  capsule. 

Pterospora  andromeda  Nutt.  Pinedrops. 
Plants  erect,  the  stems  simple,  2-8.5  (10)  dm 
tall,  reddish  brown,  succulent,  arising  from  a 
cluster  of  roots  to  5  cm  in  diameter,  glandu- 
lar-hairy, leafy  only  near  the  base;  racemes 
3-35  cm  long  or  more;  flowers  5-8  mm  long, 
nodding,  axillary;  pedicels  5-15  mm  long,  re- 
curved; sepals  oblong,  glandular;  corolla  pale 
yellow,  depressed  urn-shaped;  capsule  8-12 
(14)  mm  broad,  5-lobed,  depressed  globose. 
Coniferous  forest,  2300-2900  m,  in  Daggett, 
Duchesne,  Garfield,  Grand,  San  Juan,  Sum- 
mit, Uintah,  and  Washington  counties,  and  to 
be  expected  at  higher  elevations  elsewhere; 
widely  distributed  in  North  America;  15(iii). 

Pyrola  L. 

Rhizomatous  herbs;  leaves  with  chloro- 
phyll, leathery,  persistent,  all  basal  or  appar- 
ently so,  or  rarely  lacking  and  the  plants  then 
partially  or  completely  saprophytic;  flowers 
regular  to  irregular,  borne  in  terminal  ra- 
cemes; sepals  5,  united  at  the  base;  petals  5, 
distinct,  usually  concave,  deciduous;  stamens 
10,  the  filaments  tapering  to  the  apex,  the  an- 
tliers  unawned,  pendulous,  opening  by  means 
of  apparently  terminal  pores;  ovary  superior, 
4-loculed,  the  stigma  borne  on  a  straight  or 
curved  style;  fruit  a  loculicidal  capsule. 

Copeland,  H.  F.  1947.  Observations  on  struc- 
ture and  classification  of  the  Pyroleae. 
Madrono  9:  65-102. 


1.  Styles  straight   or  nearly   so;   pores  of  anthers  sessile;   stigma  usuallv   much 

broader  than  the  style  2 

Styles  bent  or  curved;  pores  of  anthers  usually  borne  on  short  tubes;  stigmas 
only  slightly  broader  than  the  styles 3 

2(1).       Styles  2  mm  long  or  less,  not  (or  seldom)  exserted  from  the  flower;  flowers  not 

secund;  petals  pinkish  to  cream P.  minor 

Styles  over  2   mm   long,   exserted   from    the   flower;   flowers  secund;   petals 
greenish  white P.  secunda 

3(1).       Flowers  pink  to  purplish;  sepals  longer  than  broad  P.  asarifolia 

Flowers  pale,  greenish  yellow;  sepals  broader  than  long P.  virens 


March  1980 


Welsh:  Utah  Flora,  Miscellaneous  Families 


55 


Pyrola  asarifoUa  Michx.  Liver-leaf  Win- 
tergreen.  {P.  rotimdifolia  var.  bracteata 
(Hook.)  Gray;  P.  asarifoUa  var.  hractcata 
(Hook.)  Jeps.;  P.  rotimdifolia  var.  purpurea 
Bunge;  P.  asarifoUa  var.  purpurea  (Bunge) 
Fern.;  P.  iiiearnata  Fisch.  in  DC;  P.  asari- 
foUa var.  inearnata  (Fisch.)  Fern.;  P.  asari- 
foUa var.  ovata  Farw.;  P.  uliginosa  T.  &  G.  ex 
Torr.;  P.  rotimdifolia  var.  uliginosa  (T.  &  G.) 
Gray;  P.  asarifoUa  var.  uUginosa  (T.  &  G.) 
Farw.;  P.  ekita  Nutt.;  P.  bracteata  var.  hiUi  J. 
K.  Henry).  Plants  1.3-4  dm  tall;  leaves  basal 
or  essentially  so,  the  blades  1.3-7.5  cm  long, 
1.1-7.3  cm  wide,  oval,  rotund,  elliptic,  or 
obovate,  subcordate  to  rounded,  obtuse,  or 
acute  basally,  rounded  to  obtuse  or  emargi- 
nate  apically,  entire  to  serrulate;  petioles  1-9 
cm  long;  racemes  mostly  2-  to  12-flowered; 
pedicels  3-8  mm  long;  sepals  longer  than 
broad,  1.5-4  mm  long;  petals  pink  to  pur- 
plish, 5-7  mm  long;  anthers  pink,  the  pores 
on  short  tubes;  style  curved,  with  a  flaring 
collar  below  the  stigma.  Coniferous  and  de- 
ciduous woods,  often  along  streams,  or  less 
commonly  in  meadows,  1750-2750  m,  in 
Daggett,  Duchesne,  Emery,  Garfield,  Grand, 
Iron,  Juab,  Piute,  Rich,  Salt  Lake,  Summit, 
Uintah  and  Washington  counties  (and  likely 
elsewhere);  Alaska  east  to  Newfoundland  and 
south  to  California,  New  Mexico,  South  Da- 
kota, and  New  England;  Asia.  Varietal  status 
of  Utah  materials  is  not  clear;  .30(iv). 

Pyrola  minor  L.  Lesser  Wintergreen. 
{Amelia  tninor  (L.)  Alef.;  Erxlebenia  minor 
(L.)  Rydb.;  P.  minor  var.  conferta  C.  &  S.;  P. 
conferta  (C.  &  S.)  Fisch.  ex  Ledeb.).  Plants 
0.8-2.4  dm  tall;  leaves  basal,  the  blades  (0.4) 
1.1-3.3  cm  long,  (0.6)  0.9-2.5  cm  broad,  oval, 
elliptic,  or  ovate,  obtuse  to  rounded  or  sub- 
cordate  basally,  obtuse  to  rounded  apically, 
crenate  to  subentire;  petioles  0.2-3  cm  long; 
racemes  mostly  5-  to  13-flowered;  pedicels 
2-3  mm  long;  sepals  1-1.5  mm  long,  erose  to 
subentire;  petals  pale  pink  to  cream,  3.5-4.5 
mm  long;  anthers  with  pores  sessile;  style 
straight,  very  short,  not  exserted  from  the  co- 
rolla, with  a  more  or  less  distinctive  collar 
below  the  stigma.  Wet  stream  sides  and  other 
moist  sites,  usually  in  coniferous  forests, 
2150-2750  m,  in  Beaver,  Daggett,  Duchesne, 
Garfield,  Juab,  Salt  Lake,  Sevier,  Siunmit, 
Uintah,  and  Washington  counties;  Alaska  and 


Yukon  east  to  Greenland  and  south  to  Cali- 
fornia and  Colorado;  circumboreal;  ll(i). 

Pyrola  secimdti  L.  One-sided  Wintergreen. 

{Ramisehia  secimda  (L.)  Garke;  Actinocijclus 
secundus  (L.)  Klotzsch;  P.  secimda  var.  obtii- 
sata  Turcz.;  Orthilia  secimda  var.  obtusata 
(Turcz.)  House;  P.  secunda  var.  pumila 
Paine;  P.  secimda  f.  eiicycla  Fern.).  Plants 
0.6-1.8  (2.1)  dm  tall;  leaves  basal  or  rarely 
some  cauline,  or  sometimes  with  a  naked 
stem  below  the  leaves,  the  blades  1.3-4  (5) 
cm  long,  1-3  cm  wide,  ovate,  oval,  elliptic, 
or  orbicular,  obtuse  ro  rounded  basally,  acute 
to  obtuse  or  rounded  apically,  crenate-ser- 
rate;  petioles  0.6-2  cm  long;  racemes  mostly 
4-  to  15-flowered,  the  flowers  secund;  pedi- 
cels 2-5  mm  long;  sepals  0.5-1.5  mm  long; 
petals  greenish  white,  4-6  mm  long;  anthers 
with  pores  sessile;  style  straight,  exserted 
from  the  corolla,  lacking  a  collar.  Ground 
layer  in  usually  coniferous  forests,  2000-3350 
m,  in  Box  Elder,  Carbon,  Daggett,  Duchesne, 
Garfield,  Juab,  Kane,  Piute,  Salt  Lake,  San 
Juan,  Sanpete,  Summit,  Uintah,  Utah,  and 
Washington  counties,  broadly  distributed  in 
North  America;  Eurasia.  Segregation  of  our 
materials  into  the  various  proposed  in- 
fraspecific  categories  seems  unwarranted; 
33(vi). 

Pyrola  virens  Schweigg.  in  Schweigg.  & 
Koerte.  Greenish  Wintergreen.  {P.  chlorantha 
Sw.;  P.  chlorantha  var.  saximontana  Fern.;  P. 
virens  var.  saximontana  (Fern.)  Fern.;  P.  chlo- 
rantha var.  paucifolia  Fern.;  P.  virens  f. 
paucifolia  (Fern.)  Fern.;  P.  chlorantha  f. 
paucifolia  (Fern.)  Camp).  Plants  0.9-2.5  dm 
tall;  leaves  basal,  the  blades  0.6-3.5  cm  long, 
0.5-3  cm  broad,  elliptic,  oval,  or  obovate,  ob- 
tuse to  rounded  basally,  rounded  to  obtuse 
apically,  crenate-serrate  to  subentire;  petioles 
0.8-6  cm  long;  racemes  mostly  2-  to  9-flow- 
ered;  pedicels  3-8  mm  long;  sepals  0.5-1.5 
mm  long;  petals  greenish  yellow,  5-7  mm 
long;  anthers  yellowish,  the  pores  on  elongate 
tubes;  style  curved,  with  a  flaring  collar  be- 
low the  stigma.  Coniferous  or  deciduous 
woods,  often  in  moist  sites,  2150-2750  m,  in 
Daggett,  Duchesne,  Piute,  Salt  Lake,  Sum- 
mit, and  Uintah  counties;  widely  distributed 
in  North  America;  Eurasia;  8(i). 


56 


Great  Basin  Naturalist 


Vol.  40,  No.  1 


Resedaceae 
Mignonette  Family 

Annual  or  perennial  herbs  with  watery 
sap;  leaves  alternate,  simple,  or  pinnately  to 
siibpalmately  divided;  flowers  perfect,  ir- 
regular, borne  in  terminal  racemes;  sepals  (4) 
5-6  (8),  distinct;  petals  (4)  5-6  (8),  unequal  in 
size,  the  upper  one  the  largest,  appendaged; 
stamens  8  or  more,  borne  on  the  upper  side 
of  a  rounded  disk,  the  anthers  2-loculed;  pis- 
til 1,  the  ovary  superior,  1-loculed,  with  usu- 
ally 3  (2-6)  carpels;  style  lacking;  fruit  a  cap- 
sule, usually  open  at  the  tip  before  maturity. 

Reseda  L. 

Erect  or  ascending  annual  or  perennial 
herbs  from  a  taproot;  leaves  alternate;  flow- 
ers greenish  yellow;  sepals  subequal;  petals 
imequal;  pistils  1,  the  carpels  usually  3,  open 
toward  the  apex. 

Reseda  lutea  L.  Yellow  Mignonette.  Plants 
simple  or  much  branched,  glabrous;  leaves 
pinnatifid  or  subpalmately  divided;  flowers 
greenish  yellow,  numerous,  borne  in  elongate 
racemes;  petals  usually  6,  each  commonly 
with  3  connate  or  distinct  appendages;  ovary 
and  capsule  usually  with  3  apical  lobes.  Cul- 
tivated ornamental;  rarely  escaping  in  Utah; 

1(0). 


Tamaricaceae 
Tamarisk  Family 

Shrubs  or  small  to  moderate  trees;  leaves 
alternate,  scalelike,  exstipulate,  entire;  flow- 
ers mostly  perfect,  regular,  borne  in  spikelike 
racemes  arranged  in  panicles;  sepals  4  or  5, 
overlapping;  petals  4  or  5,  separate,  more  or 
less  overlapping,  arising  from  the  base  of  a 
nectiferous  disk;  stamens  usually  as  many  or 
twice  as  many  as  the  petals,  the  anthers  2-lo- 
culed; pistil  1,  the  ovary  superior,  unilocular, 
usually  3  or  5  carpelled,  the  placentation  bas- 
al; stigmas  2-5,  separate;  ovules  2  per  pla- 
centa; fruit  a  capsule,  the  seeds  comose. 

Baum,  B.  R.  1967.  Introduced  and  natural- 
ized tamarisks  in  the  United  States  and 
Canada  (Tamaricaceae).  Baileya  15: 
19-25. 

Tamarix  L. 

Deciduous  or  evergreen  shrubs  or  trees, 
the  branchlets  deciduous;  leaves  clasping  or 
sheathing;  flowers  small,  shortly  pedicelled; 
petals  white  to  pink  or  lavender,  inserted  be- 
low the  disk;  capsules  dehiscent  by  3-5 
valves. 


1.  Leaves  sheathing;  evergreen  trees  of  moderate  size,  restricted  to  Washington 

County T.  ophylla 

Leaves  not  .sheathing,  at  most  merely  clasping;  deciduous  trees  of  small  size  or 
merely  shrubs  of  broad  distribution 2 

2(1).  Flowers  4-merous,  or  the  stamens  sometimes  more  than  4;  stamens  emerging 
gradually  from  the  di.sk-lobes;  plants  uncommon  both  in  cultivation  and  as 
e.scapes T.  pairiflora 

Flowers  5-merous,  or  the  stamens  sometimes  more  than  5;  stamens  inserted  un- 
der disk  near  the  margin  between  the  emarginate  lobes;  plants  abundant, 
cultivated  and  otherwise  T.  ramosissima 


Tamarix  aphylla  (L.)  Karst.  Athel  Tamar- 
isk. {Thuja  aphylla  L.)  Trees  to  10  m  tall  and 
6  dm  in  diameter  or  more,  the  bark  reddish 
brown  to  gray;  branchlets  jointed;  leaves 
sheathing,  minute,  evergreen;  bracts  longer 
than  the  pedicels;  flowers  5-merous;  sepals 
entire,  the  inner  ones  slightly  larger;  petals 
elliptic-oblong  to  ovate,  2-2.2  mm  long, 
early  deciduous  or  with  1  or  2  persisting;  sta- 


minal  filaments  inserted  between  the  disk 
lobes.  Cultivated  sparingly  in  Washington 
County,  where  it  seldom  flowers;  native  to 
•Africa  and  the  Middle  East;  introduced  in 
California,  Nevada,  Arizona,  and  Texas;  2(i). 

Tamarix  parviflora  DC.  Small-flowered 
Tamarisk.  Shrubs  or  small  trees  to  5  m  tall; 
bark  brown  to  deep  purple;  branchlets  not 
jointed;  leaves  merely  sessile,  not  .sheathing. 


March  1980 


Welsh:  Utah  Flora,  Miscellaneous  Families 


57 


deciduous  with  the  branchlets;  bracts  longer 
than  the  pedicels,  more  or  less  translucent; 
flowers  4-merous;  sepals  erose-denticulate, 
the  outer  two  keeled  and  acute,  the  inner  flat 
or  slightly  keeled  and  obtuse;  petals  oblong 
to  ovate,  1.9-2.3  mm  long,  persistent;  stainin- 
al  filaments  arising  gradually  from  disk-lobes. 
Cultivated  and  naturalized  along  streams  and 
seeps,  in  Emery,  Kane,  Utah,  and  Washing- 
ton counties,  and  to  be  expected  elsewhere; 
introduced  from  southern  Europe  and  now 
widespread  in  Canada  and  the  United  States; 
7(i). 

Tamarix  ramosissima  Ledeb.  Branched 
Tamarisk;  Salt  Cedar.  (T.  gaUica  authors,  not 
L.;  T.  pentandra  authors,  not  Pall.).  Shrubs  or 
small  trees  to  6  m  tall,  or  rarely  more;  bark 
reddish  brown;  branchlets  not  jointed;  leaves 
merely  sessile,  not  sheathing,  deciduous  with 
the  branchlets;  bracts  longer  than  the  pedi- 
cels, scarious  but  scarcely  translucent;  flow- 
ers 5-merous;  sepals  erose-denticulate,  the 
outer  2  narrower  than  the  inner,  all  more  or 
less  acute;  petals  obovate,  1-1.8  mm  long, 
persistent;  filaments  inserted  under  the  disk 
near  the  margin  between  the  emarginate 
lobes.  Cultivated  and  naturalized  along  seeps, 
streams,  and  reservoirs,  almost  throughout 
Utah  (Carbon,  Davis,  Duchesne,  Emery,  Gar- 


field, Grand,  Juab,  Kane,  Millard,  San  Juan, 
Sevier,  Tooele,  Uintah,  Utah,  Wasatch, 
Washington,  Wayne,  and  Weber  counties); 
introduced  from  Eurasia,  now  widespread  in 
the  southern  United  States;  99(xix). 

Tiliaceae 
Linden  or  Bas.swood  Family 

Trees;  leaves  alternate,  simple,  serrate  to 
obscurely  lobed,  usually  oblique,  stipulate; 
flowers  regular,  perfect,  borne  in  cymes;  sep- 
als 5,  distinct  or  more  or  less  connate;  petals 
5,  alternate  with  the  sepals;  stamens  numer- 
ous, the  filaments  free  or  connate  in  bundles 
of  5-10;  ovary  superior,  5-loculed;  fruit  dru- 
paceous. 

TiLIA  L. 

Cultivated  trees;  leaves  long-petioled,  the 
blades  obliquely  cordate,  serrate  or  doubly 
so,  sometimes  obscurely  lobed;  flowers  in 
long-peduncled  cymes,  the  peduncle  adnate 
at  its  base  to  a  ligulate  bract;  sepals  5;  petals 
5;  stamens  numerous,  distinct  or  in  5  clusters, 
sometimes  bearing  petaloid  staminodia  oppo- 
site the  petals;  ovary  5-loculed,  the  stigma  5- 
lobed;  fruit  subglobose,  1-  to  3-seeded. 


1.  Branchlets  and  petioles  densely  white-hairy;  leaf  blades  white  stellate  hairy 

beneath  T.  tomentosa 

Branchlets  and  petioles  glabrous  or  nearly  so;  leaf  blades  variously  pubescent 

or  glabrous 2 

2(1).        Leaf  blades  hairy  (sometimes  thinly  so)  over  the  lower  surface  and  usually 

along  the  veins  beneath  '^ 

Leaf  blades  glabrous  beneath,  except  in  vein  axils  5 

3(2).       Leaf  blades  densely  white  or  brown  stellate  hairy  beneath T.  heterophylla 

Leaf  blades  variously  hairy  but  the  surface  not  obscured  by  hairs 4 

4(3).        Hairs  of  lower  leaf  surface  stellate,  at  least  some;  flowers  with  staminodes  

T.  neglecta 

Hairs  of  lower  leaf  surface  all  simple;  flowers  without  staminodes 

T.  platijpliyllos 

5(2).       Leaf  blades  deflnitely  glaucous  beneath,  usually  less  than  8  cm  long;  flowers 

lacking  staminodes  T.  cordata 

-  Leaf  blades  green  or  merely  pale  green  beneath,  the  largest  usually  more  than 

8  cm  long;  flowers  with  or  without  staminodes 6 


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Great  Basin  Naturalist 


Vol.  40,  No.  1 


6(5).       Flowers  with  staminodes;  leaves  serrate  to  doubly  serrate  with  long-acuminate 

teeth,  the  largest  blades  on  flowering  stems  to  10  cm  long  or  more  

T.  atnericana 

Flowers  without  staminodes;  leaves  serrate  with  short  acute  teeth,  the  largest 
blades  on  flowering  stems  usually  less  than  10  cm  long T.  europaea 


Tilia  atnericana  L.  American  Linden. 
Moderate  to  large  trees  of  streets  and  other 
ornamental  plantings,  common  in  Salt  Lake, 
Utah,  and  Weber  counties,  and  probably 
grown  elsewhere;  indigenous  to  the  eastern 
states  and  Canada;  9(o). 

Tilia  cordata  L,  Small-leaved  European 
Linden.  Small  to  large  trees  of  ornamental 
plantings;  common  in  Box  Elder,  Cache, 
Juab,  Salt  Lake,  Utah,  and  Weber  counties; 
widely  cultivated  in  North  America;  in- 
troduced from  Europe;  12(o). 

Tilia  X  europaea  L.  Common  or  European 
Linden.  Moderate  to  large  trees  of  ornamen- 
tal plantings,  uncommon  in  Utah;  indigenous 
to  Europe.  This  tree  is  reputed  to  be  a  hybrid 
derivative  of  T.  cordata  x  T.  platijphyllos; 
2(0). 


Tilia  heterophylla  Vent.  White  Basswood. 
Large  ornamental  trees,  uncommon  in  Utah; 
indigenous  to  the  eastern  United  States;  2(o). 

Tilia  neglecta  Spach.  Moderate  to  large  or- 
namental trees,  uncommon  in  Utah;  in- 
digenous to  the  eastern  United  States  and 
Canada.  This  taxon  resembles,  and  apparent- 
ly intergrades  with,  T.  atnericana,  with  which 
it  is  very  closely  allied;  2(o). 

Tilia  platyphyllos  Scop.  Large-leaved 
Linden.  Moderate  to  large  ornamental  trees, 
common  in  Salt  Lake,  Utah,  and  Weber 
counties,  and  probably  elsewhere;  indigenous 
to  Europe;  8(o). 

Tilia  tomentosa  Moench.  Silver  Linden. 
Moderate  to  large  ornamental  trees,  moder- 
ately common  in  Cache,  Juab,  Salt  Lake, 
Utah,  and  Weber  counties;  indigenous  to 
eastern  Europe  and  Asia  Minor;  5(o). 


THE  TAXONOMIC  STATUS  OF  THE  ROSY  BOA 
LICHANURA  ROSEOFUSCA  (SERPENTES:  BOIDAE) 

John  R.  Ottley',  Robert  W.  Murphy-,  and  Geoffrey  V.  Smith' 

Abstract. —  Evidence  is  presented  indicating  that  lAchanura  roseofusca  and  Lkhanum  triiirgata  are  conspecific. 
Data  include  the  report  of  an  intermediate  specimen  from  El  Arco,  Baja  California  Norte,  a  site  midway  between 
the  previously  known  peninsular  ranges  of  the  two  species;  captive  hybridization  provides  additional  support  for  the 
conclusion. 


The  close  relationship  of  the  boas  Lich- 
anurci  triiirgata  Cope  and  Lichanura  roseo- 
fusca Cope  has  long  been  recognized,  due 
principally  to  the  overlap  of  most  scale  char- 
acters and  because  the  desert  boa  L.  roseo- 
fusca gracia  Klauber  appears  to  be  an  inter- 
mediate between  the  two  species  (Klauber 
1931).  The  problem  in  establishing  their  rela- 
tionship stems  from  the  rather  broad  gaps  be- 
tween their  known  ranges  in  central  Baja 
California  and  southwestern  Arizona. 

Subsequent  to  the  description  of  L.  r. 
gracia,  Klauber  (1933)  reported  a  single  spec- 
imen from  Guaymas,  Sonora.  This  specimen 
agrees  exactly  with  L.  trivirgata  in  coloration 
but  has  scale  counts  resembling  those  of  L.r. 
gracia.  He  stated  that  the  specimen  might  be 
considered  an  intergrade  of  L.  trivirgata  and 
L.r.  gracia.  This  is  somewhat  surprising  since 
he  restricted  L.  trivirgata  to  the  cape  region 
of  Baja  California,  thus  necessitating  a  trans- 
gulfian  dispersal  of  trivirgata  to  facilitate 
hybridization.  Gorman  (1965)  reemphasized 
the  wide  variation  in  meristic  characters 
within  the  genus,  as  first  demonstrated  by 
Stejneger  (1891),  and  referred  to  Klauber  s 
(1933)  scale  counts  and  color  descriptions  as 
evidence  indicating  that  the  populations 
from  southern  Arizona,  Sonora,  and  southern 
Baja  California  are  all  one  form,  L.  trivirgata 
(all  have  three  primary  stripes  of  chocolate 
brown  on  a  light  drab  background). 

The  variation  seen  in  the  genus  led  Klau- 


ber (1931,  1933)  to  speculate  that  we  might 
be  dealing  with  a  single,  polytypic  species,  L. 
trivirgata.  He  suggested,  however,  that  before 
such  a  designation  be  considered  we  should 
await  the  collection  of  more  material  from 
regions  of  potential  hybridization. 

Gorman  (1965)  and  Bostic  (1971)  com- 
mented on  new  material  from  the  range  gaps 
and  stated  that  the  basis  was  yet  lacking  for 
uniting  the  two  species  because  of  the  great 
uniformity  of  L.  trivirgata  throughout  its 
range  and  the  absence  of  obvious  intergrades. 
The  range  gaps  were  shown  to  be  separations 
of  approximately  160  km  (100  miles)  in  both 
central  Baja  California  and  southwestern  Ari- 
zona. In  spite  of  these  appraisals,  several  au- 
thors (Miller  and  Stebbins  1964,  Lowe  1964, 
Soule  and  Sloan  1966)  have  proposed,  in  ad- 
vance of  adequate  evidence,  to  imite  the  two 
species.  The  needed  evidence  is  reported  in 
this  paper. 

During  the  summer  of  1979,  an  unusual 
specimen  of  L.  trivirgata  (Fig.  1)  was  collect- 
ed at  the  town  known  as  El  Arco,  Baja  Cali- 
fornia Norte  (28°02'N,  113°27'W).  The  spec- 
imen, taken  as  it  was  crossing  the  road  in 
front  of  the  military  base  on  17  July  at  2225 
hours  by  Kenneth  A.  Stockton,  is  unique  for 
two  reasons.  First,  its  coloration"  and  scale 
counts  are  intermediate  between  the  two  spe- 
cies. Second,  the  geographic  location  of  El 
Arco  is  midway  between  the  previously  re- 
ported limits  for  the  two  species  (Bostic 


'Life  Science  Museum,  Brighani  Young  University,  Provo,  Utah  84602. 
^Department  of  Biology.  UCLA,  Los  Angeles,  California  90024. 
'Alia  Mira  .\nimal  Clinic,  VLsta,  California  9208.3. 


Color  characters  with  numbers  refer  to  the  color-name  charts  by  Kelly  (1958). 


59 


60 


Great  Basin  Naturalist 


Vol.  40,  No.  1 


1971).  Scale  counts  are  as  follows:  224  ven- 
trals,  48  subcaudals,  41  dorsal  scale  rows, 
15-14  supralabials,  15-16  infralabials,  and 
10-11  oculars.  The  specimen  is  an  adult  male 
measuring  577  mm  total  including  the  85  mm 
tail.  The  coloration  and  color  pattern  consists 
of  three  primary  stripes  of  deep  brown  (No. 
56)  on  a  ground  of  light  gray  olive  (No.  109). 
When  one  considers  all  these  characters,  the 
El  .\rco  specimen  appears  to  be  the  obvious 
intergrade  spoken  of  by  Gorman  (1965).  Al- 
though this  report  essentially  closes  the  range 
gap  on  the  Baja  California  peninsula,  a  gap 
vet  remains  between  the  Kofa  Mountains  and 
Organ  Pipe  Cactus  National  Park  in  south- 
western Arizona.  Fowlie  (1965)  has  indicated 
in  a  range  map  that  trivirgata  and  grcicia 
overlap  in  the  region  of  the  Growler  Moun- 
tains southwest  of  Ajo.  If  two  subspecies  are 
in  fact  found  together  in  the  area,  we  would 
expect  to  see  the  effects  of  intergradation.  No 
such  evidence  has  ever  been  reported  or  are 
we  aware  of  any  specimens  that  substantiate 
such  a  claim.  We  must  therefore  (juestion  the 
validity  of  Fowlie's  range  for  gracia  in  the 
Ajo  region. 


Notes  on  Captive  Breeding 

Recent  captive  breeding  experiments  have 
produced  enlightening  results.  In  April  1975 
a  male  L.t.  roseofusca  from  San  Diego,  Cali- 
fornia, was  bred  to  a  female  L.t.  trivirgata 
from  Cabo  San  Lucas,  Baja  California  Sur. 
On  7  August  1975  three  young  were  born, 
two  of  which  died  within  a  few  days;  how- 
ever, the  third  specimen,  a  male,  is  alive  at 
the  time  of  this  writing  and  in  our  possession 
(Fig.  2).  Coloration  and  color  pattern  consist 
of  three  primary  stripes  of  medium  brown 
(No.  56)  on  a  light  olive  gray  (No.  112)  back- 
ground. The  stripes  are  moderately  serrated, 
yet  fairly  uniform.  Scale  counts  are  as  fol- 
lows: ventrals  2.31,  subcaudals  47,  dorsal  scale 
rows  41,  supralabials  14-14,  infralabials 
17-15,  and  oculars  10-10.  Another  cross,  in- 
volving a  male  L.t.  trivirgata  from  near  San 
Bartolo,  Baja  California  Sur,  and  a  female  of 
the  same  subspecies  from  the  vicinity  of  Esta- 
cion  Ortiz,  Sonora,  occurred  in  March  1976. 
Four  young  were  born  on  29  July  1976.  A  fe- 
male from  that  litter  (Fig.  3)  yet  remains  in 
our  possession.  Coloration  and  pattern  are  of 


Fig.  1.   Dorsal  view  of  a  Lkhunum  triim^ala  x  wscofusai  iiitcnnfcliato  tioin  El  Arco.  Baja  California  \ortt 


March  1980 


Ottley  et  al.:  Rosy  Boa 


61 


Fig.  2.   Dorsal  view  of  a  Lichaniini  tritirgata  x  roscufiisca  hybrid;  male  parent  is  a  L.  t.  wscofttsca  from  San  Diego, 
California,  and  the  female  parent  is  a  L.  t.  triiirgata  from  Cabo  San  Lucas,  Baja  California  Sur. 


Fig.  .3.  Dorsal  view  of  a  Lichanura  t.  triiin^dta  transgnlfian  cross;  male  parent  is  a  L.  t.  trivirgata  from  near  San 
Bartolo,  Baja  California  Sur,  and  the  female  parent  is  a  L.  t.  trivirgata  from  Estacion  Ortiz,  Sonora. 


62 


Great  Basin  Naturalist 


Vol.  40,  No.  1 


three  primary  stripes  of  chocolate  brown  on 
a  cream  ground.  The  stripes  are  uniform  with 
shghtly  serrated  edges.  Scale  counts  are  as 
follows:  ventrals  220,  subcaudals  44,  dorsal 
scale  rows  38,  supralabials  13-13,  infralabials 
15-14,  and  oculars  10-11.  The  female  L.t.  tri- 
virgata  transgulfian  cross  and  a  male  desig- 
nated as  L.t.  gracia  from  near  Punta  Prieta, 
Baja  California  Norte,  were  observed  cop- 
ulating on  16  May  1979.  On  24  October  1979 
three  young  were  bom,  all  males,  each  bear- 
ing well-delineated  medium  brown  stripes 
and  a  ground  of  color  intermediate  between 
the  parents. 

Acknowledgments 

We  thank  Wilmer  W.  Tanner  and  Kent  M. 
Van  De  Graaff  for  their  constructive  criti- 
cisms and  comments  in  reviewing  this  paper, 
Vickie  R.  Ottley  for  typing  the  manuscript, 
and  Lawrence  E.  Hunt,  Kenneth  A.  Stockton, 
and  Dale  M.  Stockton  for  their  help  and 
companionship  in  the  field.  Scientific  collec- 
ting permit  30/832/79  was  issued  by  Ignacio 
Ibarrola  Bejar,  director  general  of  the  De- 
partamento  de  la  Conservacion  de  la  Fauna 
Silvestre. 


Summary 

In  coloration  the  El  Arco  specimen  and  the 
captive  bred  trivergata  x  roseofusca  hybrid 
are  very  similar,  differing  only  in  the  latter 
having  moderately  serrated  stripes.  These 
data  support  our  consideration  of  the  El  Arco 
specimen  as  an  intermediate.  Since  El  Arco  is 
situated  in  a  geographical  region  midway  be- 
tween "pure"  trivergata  and  roseofusca,  in- 


dicating a  continuous  range,  we  find  no  alter- 
native to  considering  the  two  taxa  as  being 
conspecific.  The  binomial  Lichanura  trivir- 
gata  Cope  has  priority  over  L.  roseofusca 
Cope  by  publication  date.  Accordingly,  we 
recognize  L.  trivirgata  as  a  single,  polytypic 
species  with  four  subspecies  (L.  t.  trivirgata 
Cope,  L.  t.  roseofusca  Cope,  L.  t.  gracia 
Klauber,  and  L.  t.  bostici  Ottley). 

Literature  Cited 

BosTic,  D.  L.  1971.  Herpetofauna  of  the  Pacific  Coast  of 
north  central  Baja  CaUfomia,  Mexico,  with  a  de- 
scription of  a  new  subspecies  of  PhyUodactylus 
xanti.  Trans.  San  Diego  Soc.  Nat.  Hist. 
16(10):237-263. 

FowLiE,  J.  A.  1965.  The  snakes  of  Arizona.  Azul  Quinta 
Press,  Fallbrook,  California,  164  pp. 

Gorman,  G.  C.  1965.  The  distribution  of  Lichanura  tri- 
virgata and  the  status  of  the  species.  Herpetolo- 
gica  21(4):2a3-287. 

Kelly,  K.  L.  1958.  ISCC-NBS  color-name  charts  illus- 
trated with  centroid  colors.  National  Bureau  of 
Standards,  NBS  Circular  533. 

Klauber,  L.  M.  1931.  A  new  subspecies  of  the  Califor- 
nia Boa,  with  notes  on  the  genus  Lichanura. 
Trans.  San  Diego  Soc.  Nat.  Hist.  6(20):305-318. 

19.33.  Notes  on  Lichanura.  Copeia  (4):214-215. 

Lowe,  C.  H.  1964.  The  vertebrates  of  Arizona:  Anno- 
tated check  lists.  Tucson,  University  of  Arizona 
Press. 

Miller,  A.  H.,  and  R.  C.  Stebbins.  1964.  The  lives  of 
desert  animals  in  Joshua  Tree  National  Mon- 
ument. Berkeley  and  Los  Angeles,  University  of 
California  Press. 

Ottley,  J.  R.  1978.  A  new  subspecies  of  the  snake  Lich- 
anura trivirgata  from  Cedros  Island,  Mexico. 
Great  Basin  Nat.  .38:411-416. 

SouLE,  M.,  AND  A.  J.  Sloan.  1966.  Biogeography  and  dis- 
tribution of  the  reptiles  and  amphibians  on  is- 
lands in  the  Gulf  of  California,  Mexico.  Trans. 
San  Diego  Soc.  Nat.  History  14(11):137-156. 

Stejneger,  L.  1891.  On  the  snakes  of  the  California 
genus  Lichanura.  Proc.  U.S.  Nat.  Mus. 
14:511-515. 


HESPEROPERLA  HOGUEI,  A  NEW  SPECIES  OF  STONEFLY 
FROM  CALIFORNIA  (PLECOPTERA:  PERLIDAE) 

Richard  \V.  Baiiinaiin'  and  Bill  P.  Stark- 


.\bstr.\ct.—  a  new  species  of  Hcspcropcila  is  named  from  northern  California.  The  adult  male  and  female, 
nymph,  and  egg  are  described  and  figured.  Hespcroperla  lioouci  adds  a  second  species  to  this  previously  monotypic 
genus. 


The  genus  Hcsperoperla  Banks  (1938)  was 
not  accepted  by  other  workers  until  the  re- 
cent world  catalog  (lilies  1966).  Studies  by 
Needham  and  Claassen  (1922),  Claassen 
(1940),  and  Frison  (1942)  placed  six  species  in 
synonymy  luider  Acroneiiria  pacifica  Banks, 
including  Hesperoperia  obscura  (Banks),  the 
designated  type  species. 

Stark  and  Gaufin  (1976),  in  their  revision 
of  the  Perlidae,  confirmed  that  Hesperoperia 
was  indeed  a  separate  genus  with  one  valid 
species,  Hesperoperia  pacifica  (Banks). 

In  the  fall  of  1976,  Charles  L.  Hogue  of 
the  Natural  History  Museum,  Los  Angeles 
County,  sent  a  distinctive  female  perlid  to 
the  senior  author  for  identification.  When  it 
proved  to  belong  to  Hesperoperia,  the  help  of 
numerous  colleagues,  including  Bill  P.  Stark, 
was  enlisted. 

The  types  of  several  species  in  the  A.  pa- 
cifica synonymy  were  examined,  including: 
A.  pacifica  Banks,  A.  nigrita  Banks,  A.  pu- 
mila  Banks,  and  A.  obscura  Banks.  The  spe- 
cific name  nigrita  suggested  that  its  type 
might  be  dark  and  distinctive,  but  this  was 
not  the  case.  The  type  locality  of  A.  pumila. 
Three  Rivers,  California,  which  is  near  Vis- 
alia,  is  a  short  distance  south  of  the  known 
range  of  this  new  species,  but  the  specimen 
was  positively  H.  pacifica. 

Several  additional  specimens  were  sub- 
sequently found  that  confirmed  that  this  was 
an  undescribed  species  of  Hesperoperia  with 
a  known  range  that  included  most  of  north- 
em  California. 


Both  species  of  Hesperoperia  occur  in  the 
same  streams  with  no  intergradation;  thus 
they  are  sympatric  species. 

Hesperoperia  hoguei,  n.  sp. 

Figs.  1-9 

Male.—  Macropterous.  Length  of  fore- 
wings  (18-20  mm;  length  of  body  16-18  mm. 
Dorsum  of  head  mostly  yellow,  with  brown 
U-shaped  pattern  connecting  ocelli,  posterior 
lateral  margins  brown  behind  compound 
eyes,  sometimes  with  faint  brown  area  near 
frontal  margin.  Pronotum  with  broad  yellow 
median  stripe,  lateral  margins  dark  brown, 
rugosities  distinctive  and  slightly  lighter  than 
lateral  margins  (Fig.  4).  Legs  brown.  Wings 
light  brown,  veins  brown.  Abdomen  yellow 
with  narrow  brown  lateral  stripes.  Tergum 
10  with  median  tergite,  segment  covered 
with  medium-length  hairs.  Sternum  9  with 
large  quadrangular  hammer.  Paraprocts 
sclerotized,  broad  basally,  apex  narrow  and 
pointed  (Fig.  2).  Aedeagus  with  two  bands  of 
large  spinules,  apical  band  broad,  consisting 
of  12  or  more  closely  set  spinule  rows,  basal 
band  narrow,  interrupted  on  both  dorsomesal 
and  ventromesal  surfaces;  small  spinules  pres- 
ent in  patches  near  base  and  at  dorsal  apex 
(Figs.  5-6). 

Female.—  Macropterous.  Length  of  fore- 
wings  26-28  mm;  length  of  body  24-26  mm. 
Color  pattern  similar  to  male.  Subgenital 
plate  strongly  produced,  with  more  darkly 
sclerotized  area  along  mesoposterior  margin 


'Monte  L.  Bean  Life  Science  Museum  and  Department  of  Zoology,  Brigham  Young  University,  Provo,  Utah  84602. 
'Department  of  Biological  Sciences,  Mississippi  College,  Clinton,  Mississippi  39056. 


63 


64 


Great  Basin  Naturalist 


Vol.  40,  No.  1 


Fig.  1.  Hesperoperht  /logi/ei,  mature  nymph,  habitus. 


March  1980 


Baumann,  Stark:  California  Stonefly 


65 


2 


3 


11 


5 


Figs.  2-6.  Hesperoperla  hogtici:  (2)  male  terminalia,  dorsal;  (3)  female  terminalia,  ventral;  (4)  adult,  head  and  pro- 
notum;  (5)  aedeagiis,  dorsal  (.50X,  inset  lOOX);  (6)  aedeagiis,  lateral  (SOX,  inset  lOOX). 


66 


Great  Basin  Naturalist 


Vol.  40,  No.  1 


(Fig.  3).  Vagina,  spermathecum,  and  acces- 
sory glands  membranous. 

Egg.-  Outline  oval;  cross-section  circular. 
Collar  stalked,  margin  flanged  and  irregu- 
larly incised;  collar  end  of  egg  with  regular 
indentations  which  end  at  terminal  margin. 
Chorion  smooth.  Micropyles  arranged  cir- 
cumlinearly  in  apical  end  opposite  of  collar 
(Figs.  7-9). 

Nymph.—  General  color  dark  brown,  pat- 
terned with  yellow  markings.  Occiput  with 
an  irregularly  spaced  row  of  spinules.  Post- 
ocular  fringe  present.  Head  mostly  dark,  with 
distinctive  inverted  yellow  W-shaped  pattern 
located  anterior  to  compound  eyes,  frontal 
margin  entirely  yellow.  Pronotum  with  later- 
al setae  sparse  or  absent.  Abdominal  terga 
with  numerous  small  intercalary  spinules. 
Cerci  with  fringe  of  spines  at  segmental 
joints,  never  longer  than  segments;  few  tiny 
intersegmental  spinules  present.  Proventricu- 
lus  with  teeth  in  12  longitudinal  bands;  acces- 
sory bands  and  structures  absent.  Thoracic 
and  anal  gills  present  (Fig.  1). 

Diagnosis.—  Hespemperla  fwguei  can  be 
easily  separated  from  H.  pacifica  in  the  nym- 
phal  stage  by  the  difference  in  head  pattern 
and  the  abdominal  spinulation.  Hesperoperla 
pacifica  exhibits  a  large,  inverted,  mushroom- 
shaped  pattern  anterior  to  the  compound 
eyes  that  terminates  in  an  enlarged  base  on 
the  mesoanterior  margin,  and  H.  hoguei  bears 
an  inverted  W-shaped  pattern  and  a  separate 
broad  yellow  band  running  the  full  width  of 


the  anterior  margin.  Intercalary  spinules  are 
entirely  absent  from  the  abdominal  terga  of 
mature  H.  pacifica  nymphs  but  are  numerous 
in  H.  hoguei  specimens.  Claassenia  sabulosa 
nymphs  also  have  an  inverted  W-shaped  pat- 
tern on  the  head,  but  they  possess  a  complete 
occipital  ridge. 

Adults  of  H.  pacifica  have  a  plain  yellow 
brown  pattern  on  their  head  and  pronotum. 
Those  of  H.  hoguei  have  a  broad  yellow  me- 
dian area  set  off  by  distinctive  dark  lateral 
margins.  The  external  genitalia  are  quite  sim- 
ilar, but  the  basal  spinule  band  on  the  ae- 
deagus  of  the  males  is  different.  In  H.  pacifi- 
ca the  band  is  only  broken  ventrally,  but  in 
H.  hoguei  the  band  is  broken  both  dorsally 
and  ventrally. 

Types.-  Holotype:  $  ,  Gibson  Creek,  800 
ft,  1  mile  west  of  Ukiah,  Mendocino  Co.,  Cal- 
ifornia, 6-IX-1976,  C.  L.  Hogue.  Allotype: 
Toadtown,  3000  ft,  4  miles  SW  Stirling  City, 
Butte  Co.,  California,  9-IX-1976,  C.  L. 
Hogue. 

Paratypes:  Butte  Co.:  Paradise,  25-V-1966, 
Lowe,  2  females  (CSUC);  Tehama  Co.:  Big 
Chico  Creek,  Hwy.  32,  14-IX-1979,  G.  L. 
Boles,  13  males,  7  females  (GLB)  (BYU).  Ad- 
ditional specimens:  Butte  Co.:  Big  Chico 
Creek,  580  ft,  V4  mile  below  Salt  Springs, 
Bidwell  Park,  P-VIII-1972,  M.  W.  Kainu, 
nymph  (UCD);  2  miles  SW  Stirling  City,  20- 
VI- 1979,  J.  A.  Stanger,  nymphs  (BYU). 
Plumas  Co.:  Sulphur  Creek,  Hwy.  89,  5-VII- 
1979,  B.  P.  Stark  and  K.  W.  Stewart,  nymphs 


Figs.  7-9.  Hesperoperla  hoguei:  (7)  egg,  lateral  (200X);  (8)  egg,  collar  end  {400X);  (9)  egg,  micropyles  (700X). 


March  1980 


Baumann,  Stark:  California  Stonefly 


67 


(BPS).  SJiasta  Co.:  unnamed  spring  tributary 
to  Lake  Britton,  20-IX-1978,  G.  L.  Boles, 
nvniphs  (GLB);  South  Fork  Bear  Creek,  12- 
v'lI-1979,  G.  L.  Boles,  nymphs  (GLB).  Te- 
hama Co.:  Big  Chico  Creek,  Hwv.  32,  14- V- 
1978,  G.  L.  Boles,  nymph  (BYU),'  same  data, 
2-IX-1978,  nymphs  (2  females  emerged  28- 
IX- 1978)  (GLB)  (BYU). 

Holotype  and  allotype  deposited  at  the 
Natural  History  Museum,  Los  Angeles  Coun- 

ty- 

Etymology.—  This  species  is  named  in 
honor  of  Dr.  Charles  L.  Hogue,  Senior  Cura- 
tor of  Entomology,  Natural  History  Museum, 
Los  Angeles  County.  He  has  collected  many 
interesting  stoneflies  during  his  studies  on  the 
torrenticolus  insects  of  the  New  World. 


Acknowledgments 

We  thank  the  following  individuals  for  the 
help  that  we  received  during  this  study:  Ger- 
ald L.  Boles,  California  Department  of  Water 
Resources,  Red  Bluff,  California  (GLB);  Dr. 
Charles  L.  Hogue,  Natural  History  Museum, 


Los  Angeles  County,  California  (LACM);  Dr. 
David  H.  Kistner,  California  State  University, 
Chico,  California  (CSUC);  Dr.  Robert  O. 
Schuster,  University  of  California,  Davis, 
California  (UCD);  Jean  A.  Stanger,  Brigham 
Young  University,  Provo,  Utah  (BYU);  Dr. 
Kenneth  W.  Stewart,  North  Texas  State  Uni- 
versity, Denton,  Texas. 

The  drawings  were  made  by  Connie  A.  Be- 
van  Bhagat. 

Literature  Cited 

Banks,  N.  1938.  A  new  genus  of  Perlidae.  Psyche 
45:136-1.37. 

Claassen,  p.  W.  1940.  A  catalogue  of  the  Plecoptera  of 
the  world.  Mem.  Cornell  Agr.  Exp.  Station 
232:1-235. 

Prison,  T.  H.  1942.  Descriptions,  records  and  systematic 
notes  concerning  western  North  .American  stone- 
flies  (Plecoptera).  Pan-Pac.  Entomoi.  18:61-73. 

Illies,  J.  1966.  Kataiog  der  rezenten  Plecoptera.  Das 
Tierreich,  Berlin,  82,  632  pp. 

Needham,  J.  G.,  AND  P.  W.  Claasse.n.  1922.  The  North 
American  species  of  the  genus  Acronetiriti  (Order 
Plecoptera).  Canad.  Entomoi.  54:249-255. 

Stark,  B.  P.,  and  A.  R.  Gaufin.  1976.  The  nearctic  gen- 
era of  Perlidae  (Plecoptera).  Misc.  Pubi.  Entomoi. 
Soc.  Amer.  10:1-77. 


REPRODUCTION  IN  THREE  SYMPATRIC  LIZARD  SPECIES 
FROM  WEST-CENTRAL  UTAH 


John  B.  Andre'  and  James  A.  MacMahon' 

.\bstr.-vct.-  Data  on  reproduction  by  the  lizards  Uta  stanshiiriana.  Ciotaphytus  colkiris,  and  Cnciindophorus  tigris 
are  presented  from  a  communitv  where  they  are  sympatric  in  west-central  Utah.  Data  are  compared  to  a  previous 
studv  of  these  species  at  the  same  site  and  to  data  from  other  sites  in  the  United  States. 


Lizard  reproductive  data  from  geographi- 
callv  separated  populations  are  important  to 
ecologists  attempting  to  explain  a  highly  var- 
iable species  characteristic.  Reproduction  by 
Uta  stansburiana  has  been  well  documented 
in  the  literature  (Fautin  1946,  Medica  and 
Turner  1976,  Nussbaum  and  Diller  1977, 
Tanner  1965,  Tinkle  1961,  1967,  Turner  et 
al.  1970,  1973,  1976,  Worthington  and  Ar- 
vizo  1973,  Parker  1974,  Parker  and  Pianka 
1975,  Tinkle  and  Hadley  1975,  and  Goldberg 
1977).  Reproductive  studies  of  Ciotaphytus 
coUaris  and  Cnetnidophorotis  tigris  are  scarce 
(Fautin  1946,  Turner  et  al.  1969,  Pianka 
1970,  Burkholder  and  Walker  1973,  Parker 
1973,  and  Vitt  and  Ohmart  1977).  This  paper 
presents  data  on  the  reproduction  by  three 
lizard  species  {Uta  stansburiana  stansbu- 
riana, Crotaphytus  collaris  bicinctoris,  and 
Cnemidophorus  tigris  tigris),  coexisting  in 
west-central  Utah. 

We  are  cognizant  of  the  limitations  im- 
posed by  the  short  period  covered  by  our  col- 
lections. Despite  this,  there  is  clearly  a  need 
for  carefully  collected  data  on  reproduction 
by  lizards,  or  other  taxa,  so  that  one  might 
gather  such  data  into  a  body  of  information 
used  to  address  general  evolutionary  theorv 
(Tinkle  1969a,  b.  Tinkle  et  al.  1970). 

Methods 

Lizards  were  collected  in  Tule  Valley,  Mil- 
lard   County,    Utah    (lat.    39°13'N,'  long. 


113°27'W).  Tule  Valley,  bordered  on  the  east 
and  west  by  mountain  ranges,  is  typical  of 
the  basin  and  range  topography  of  the  Great 
Basin  Desert  (MacMahon  1979).  Bajadas  (coa- 
lesced alluvial  fans)  slope  from  the  bases  of 
both  mountain  ranges  to  the  playa  that  cov- 
ers most  of  the  valley  floor. 

The  study  site  was  located  in  the  Tetra- 
dy?nia  glabrata  and  Atriplex  confertifolia 
communities  described  by  Fautin  (1946).  The 
common  plant  species  were  T.  glabrata,  A. 
confertifolia,  Chrysothamnus  viscidiflorus, 
Artemisia  spinescens.  Ephedra  nevadensis, 
Ceratoides  (Eurotia)  lanata,  and  Hilaria 
jamesii.  The  substrate  was  mostly  small  rocks 
embedded  in  packed  soil,  with  localized 
areas  containing  large  boulders,  which  were 
used  as  basking /perching  sites  by  C.  coUaris. 

Lizards  were  collected  (shot)  throughout 
the  day,  at  three-week  intervals  between  1 
April  and  29  August  1976.  Specimens  were 
preserved  in  10  percent  formalin  within  two 
hours  of  collection.  Analysis  of  reproductive 
state  (for  females)  and  measurement  of  snout- 
vent  length  (SVL)  were  made  in  the  labora- 
tory. 

The  sex  of  each  specimen  was  determined 
by  dissection.  The  reproductive  tracts  of  the 
females  were  removed  and  the  number  of 
corpora  lutea,  yolked  follicles  and/ or  oviduc- 
al  eggs  were  recorded.  Estimates  of  clutch 
size  were  based  on  the  number  of  yolked  fol- 
licles >  2.5  mm  diameter  and/ or  oviducal 
eggs  and  corpora  lutea  for  U.  stansburiana 


'This  paper  is  a  contribution  from  the  Department  of  Biology  and  the  Ecology  Center,  Utah  State  University,  Logan,  Utah  84322.  Reprint  requests  sh 
be  sent  to  James  A.  MacMahon.  Present  address  for  John  B.  Andre  is  Cape  Romain  National  Wildlife  Refuge,  R.R.  1,  Box  191,  Awendaw,  South  Car, 


should 
rolina 


68 


March  1980 


Andre,  MacMahon:  Utah  Lizards 


69 


and  volked  follicles  >  5.0  mm  diameter 
and/or  oviducal  eggs  and  corpora  liitea  for 
C.  coUam  and  C.  tigris. 

Results 

Mean  clutch  size  and  mean  SVL  of  sex- 
ually mature  females  of  each  species  are  list- 
ed in  Table  1.  The  relationship  between 
clutch  size  and  SVL  is  illustrated  for  U. 
stanshuriona  in  Figure  I.  Analyses  of  similar 
data  for  C.  coUaris  and  C.  tigris  showed  no 
significant  correlation  (F-tests).  The  line  in 
this  figure  was  determined  by  linear  regres- 
sion, the  correlation  coefficient  is  given  for 
the  data  set. 

Uta  stanshuriana  females  reach  sexual 
maturity  in  their  second  growing  season  (10 
mo.  old)  at  about  40  mm  SVL  (the  smallest 
female  having  yolked  follicles  was  37.0  mm 
SVL,  see  Table  2).  Most  U.  stansburiana 
emerged  from  hibernation  bv  the  first  week 
of  April  and  bred  shortly  after  this  time. 
Yolked  follicles  and  oviducal  eggs  were  pres- 


ent from  1  April  to  16  May;  only  oviducal 
eggs  were  found  from  6  Jiuie  to  29  June. 
From  17  July  through  29  August  no  yolked 
follicles  or  oviducal  eggs  were  found  in  the 
females  collected.  Though  yolked  follicles 
and  oviducal  eggs  were  present  in  the  fe- 
males from  April  to  the  end  of  June,  females 
contained  the  most  oviducal  eggs  between  24 
April  and  15  May.  While  we  believe  that  fe- 
males laid  one  or  two  clutches  of  eggs  in 
1976,  our  data  are  not  exten.sive  enough  on 
this  point.  Turner  et  al.  (1970)  have  warned 
of  the  problem  of  determining  clutch  fre- 
quency with  too  few  observations. 

Crotaplujtus  coUaris  females  are  sexually 
mature  at  about  85  mm  SVL.  Yolked  follicles 
were  present  in  the  single  specimen  collected 
15  May.  All  females  collected  in  the  first 
week  of  June  contained  oviducal  eggs.  At  the 
end  of  June  no  females  contained  yolked  fol- 
licles or  oviducal  eggs. 

Cnemidophorus  tigris  females  attain  sexual 
maturity  abovit  73  mm  SVL.  Specimens  col- 
lected during  the  first  and  last  weeks  of  June 


CLUTCH 
SIZE 


5 

.Ufa 

• 

4 

, 

t  • 

• 
• 

4         4 

3 

• 

10        9 

5 

f%   \    •    I   V    • 

2 

, 

t-rr 

^4    4 
V  #^#^   *•    V     • 

•        • 

1 

y  =  -6.448+0.216  X 
r  =  0.605 

36 


40 


42 


44 


46 


48 


50 


SNOUT-VENT  LENGTH     (mm) 


Fit;.  1.   Relationship  hctufi-ii  tlutcli  si/c  iiiuiiil)t'n  and  S\'I,  mini)  tor  I'.  sldiisbitrUm 


Table  1.     Clutch  size  and  SVL  of  adult  female  Vta  stansburiana,  Crotaphytus  collaris  and  Cnemidophorus  tigris. 


N 

SVL  (mm) 

Clutch  size 

Species 

X 

Range 

X                   Range 

Uta  stanshuriana 
Crotaphytus  coUaris 
Cnemidopliorus  tigris 

96 
13 
15 

44.61 
90.69 
a3.13 

.39.5-49.0 
85.5-99.0 
73.0-96.0 

2.99                  2-5 
5.38                 3-7 
3.07                  2-5 

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Great  Basin  Naturalist 


Vol.  40,  No.  1 


contained  yolkfed  follicles  and/or  oviducal 
eggs.  After  mid-July,  no  females  contained 
yolked  follicles  or  oviducal  eggs. 

The  first  hatchlings  observed  were:  U. 
stansburiana,  17  July;  C.  collaris,  9  August; 
C.  Tigris,  7  August. 


Discussion 

Mean  clutch  size  for  U.  stansburiana  was 
2.99  (range  2-5).  Fautin  (1946)  reported  a 
mean  clutch  size  of  4.1  (range  3-5)  from  the 
vicinity  of  our  study  site.  For  northern  popu- 
lations (Tooele  Coimty,  Utah)  Parker  and 
Pianka  (1975)  found  a  mean  clutch  size  for 
Vta  of  4.6;  Nussbaum  and  Diller  (1977)  in 
Oregon  found  3.3.  Both  Parker  and  Pianka 
(1975)  and  Nussbaum  and  Diller  (1977)  re- 
ported that  Vta  produces  one  or  two  clutches 
per  season.  At  our  site,  one,  and  for  some  fe- 
males perhaps  two,  clutches  of  eggs  were  laid 
by  Uta  in  1976. 

Parker  and  Pianka  (1975)  also  reported 
that  oviducal  eggs  were  present  during  a 
three-  to  four-month  period;  our  data  agree. 


The  relationship  between  SVL  and  clutch 
size  indicates  that  larger  females  produce 
larger  clutches.  An  F  test  shows  a  significant 
relationship  between  SVL  and  clutch  size 
(0.05  level).  Other  workers  report  the  same 
relationship  between  SVL  and  clutch  size  for 
Uta  stansburiana  from  other  parts  of  its 
range  (Tinkle  1961,  Turner  et  al.  1973,  Park- 
er and  Pianka  1975,  and  Goldberg  1977). 

By  mid-July  Uta  is  in  postreproductive 
condition:  the  reproductive  tracts  of  both 
males  and  females  have  decreased  in  size. 
This  size  decrease  is  accompanied  by  an  in- 
crease in  the  size  of  the  fat  bodies.  Fat  bodies 
continue  to  enlarge  as  the  growing  season 
progresses. 

Little  information  exists  on  the  reproduc- 
tion of  C  collaris.  In  southern  New  Mexico, 
Parker  (1973)  reported  a  mean  clutch  size  of 
5.3  (range  3-7).  Robison  and  Tanner  (1962) 
reported  a  mean  clutch  size  of  6.7,  but  the 
lizard  was  collected  from  many  different 
parts  of  its  range.  Our  data  show  a  mean 
clutch  size  of  5.38  (range  3-7). 

The  relationship  between  SVL  and  clutch 
size  is  illustrated  in  Figure  2.  A  loose  correla- 


CLUTCH   ^ 
SIZE      5 


88  90  92  94  96 

SNOUT-VENT    LENGTH     (mm) 

Fig.  2.  Relationship  between  clutch  size  (number)  and  SVL  (mm)  for  C.  collaris. 


98 


lOO 


Table  2.     Monthly  clutch  size  and  SVL  of  adult  female  Uta  stansburiana. 


Month 


April 

May 

June 

July 

August 


SVL  (mm) 


Clutch  size 


N 

X 

Range 

X 

Range 

21 

43.17 

37.0-47.5 

2.39 

2-4 

13 

44.38 

41.5-48.0 

3.46 

2-5 

20 

42.73 

38.0-47.5 

2.45 

2-4 

20 

42.80 

38.0-49.0 

2.47 

2-4 

22 

43.50 

40.0-48.0 

2.76 

2-4 

March  1980 


Andre,  MacMahon:  Utah  Lizards 


71 


tion  exists,  with  larger  females  producing 
larger  clutches;  an  F  test  (0.05  level)  is  not 
significant. 

Yolked  follicles  and/ or  oviducal  eggs  were 
present  from  mid-May  to  mid-June  in  C.  col- 
laris  females.  By  the  end  of  June  this  species 
is  postreproductive;  the  reproductive  tracts 
are  decreasing  in  size  and  the  fat  bodies  are 
enlarging. 

Cneinidophoms  tigris  becomes  active  near 
the  end  of  April.  Mating  occurs  shortly  after. 
Yolked  follicles  and/or  oviducal  eggs  were 
present  from  June  to  the  first  week  of  July. 
Bv  mid-July  this  species  is  postreproductive; 
they  exhibit  small  reproductive  tracts  and  en- 
larging fat  bodies. 

Mean  clutch  size  was  3.07  (range  2-5), 
whereas  Fautin  (1946)  reported  a  mean 
clutch  size  of  6.7  (range  5-9).  The  relation- 
ship between  SVL  and  clutch  size  is  similar 
to  that  of  C.  coJIaris,  with  larger  females  ten- 
ding to  produce  more  eggs  (Fig.  3),  although 
an  F  test  (0.05  level)  is  not  significant.  Pianka 
(1970)  and  Vitt  and  Ohmart  (1977)  report  a 
relationship  between  SVL  and  clutch  size 
that  is  "loosely  correlated"  for  C.  tigris. 

Acknowledgments 

This  work  was  made  possible  by  the 
US /IBP  Desert  Biome  fimded  by  the  Nation- 
al Science  Foundation  (Grant  GB32139). 
Linda  Finchum  typed  the  manuscript.  Robert 
Bayn  executed  the  figures. 


Literature  Cited 

BuRKHOLDER,  G.  L.,  AND  J.  M.  Walker.  1973.  Habitat 
and  reproduction  of  the  desert  whiptail  lizard, 
Cncmidophonis  tigris  Baird  and  Girard  in  south- 
western Idaho  at  the  northern  part  of  its  range. 
Herpetolot^ica  29:76-8.3. 

Fautin.  R.  W.  1946.  Biotic  communities  of  the  northern 
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Goldberg,  S.  R.  1977.  Reproduction  in  a  mountain  pop- 
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11:31-35. 

MacMahon,  J.  A.  1979.  North  American  deserts:  Tlieir 
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Perry  and  D.  Goodall,  eds.  Arid  land  ecosystems: 
Their  structure,  fvmctioning  and  management. 
Vol.  1.  Cambridge  University  Press,  Cambridge. 

Medica,  p.  a.,  and  F.  B.  Turner.  1976.  Reproduction 
by  Uta  stansburiana  (Reptilia,  Lacertilia,  Igua- 
nidae)  in  southern  Nevada.  J.  Herpetol. 
10:123-128. 

Nussbaum,  R.  a.,  and  L.  V.  Diller.  1977.  The  life  his- 
tory of  the  side-blotched  lizard,  Uta  stansburiana 
Baird  and  Girard,  in  north-central  Oregon. 
Northwest  Sci.  50:24.3-260. 

Parker,  W.  S.  1973.  Notes  on  the  reproduction  of  some 
lizards  from  Arizona,  New  Mexico,  Texas  and 
Utah.  Herpetologica  29:258-264. 

1974.  Home  range,  growth  and  population  den- 
sity of  Uta  stansburiana  in  Arizona.  J.  Herpetol. 
8:135-139. 

P.ARKER,  W.  S.,  AND  E.  R.  PiANKA.  1975.  Comparative 
ecology  of  populations  of  the  lizard  Uta  stansbur- 
iana. Copeia  1975:61.5-6.32. 

PiA.NiCA,  E.  R.  1970.  Comparative  autecology  of  the  liz- 
ard Cneinidophorus  tigris  in  different  parts  of  its 
geographic  range.  Ecology  51:703-719. 

RoBisoN,  W.  G.,  Jr.,  a.nd  W.  VV.  Tanner.  1962.  A  com- 
parative study  of  the  species  of  the  genus  Crota- 
phi/tus  Holbrook  (Iguanidae).  Brigham  Young 
Univ.  Sci.  Bull.,  Biol.  Ser.  2(1):1-21. 


CLUTCH 
SIZE 


Cnemidophorus 


72 


76 


80 


84 


88 


y  =  -1.423  +0.051  X 


r=  0.333 


92 


96 


SNOUT- VENT  LENGTH    (mm) 


100 


Fig.  3.  Relationship  between  clutch  size  (number)  and  SVL  (mm)  for  C.  tigris. 


72 


Great  Basin  Naturalist 


Vol.  40,  No.  1 


Tanner,  W.  W.  1965.  A  comparative  population  study 
of  small  vertebrates  in  the  uranium  areas  of  the 
Upper  Colorado  River  Basin  of  Utah.  Brigham 
Young  Univ.  Sci.  Bull.,  Biol.  Ser.  7(1):  1-31. 

Tinkle,  D.  W.  1961.  Population  structure  and  reproduc- 
tion in  the  lizard  Uta  stansburiana  stejnegeri. 
Amer.  Midi.  Natur.  66:206-234. 

1967.  The  life  and  demography  of  the  side- 
blotched  lizard  Vta  stansburiana.  Misc.  Publ. 
Mus.  Zool.  Univ.  Michigan  132:1-182. 

1969a.  The  concept  of  reproductive  effort  and  its 

relation  to  the  evolution  of  life  histories  of  liz- 
ards. Amer.  Natur.  103:501-516. 

1969b.  Evolutionary  implications  of  comparative 

population  studies  in  the  lizard  Uta  stansburiana. 
Pages  133-154  in  Systematic  biology,  Proc.  of  In- 
ternational Conf.,  National  Acad.  Sci.  Publ.  No. 
1692,  Washington,  D.C. 

TiNKJLE,  D.  W.,  .\ND  N.  F.  H.\uLEY.  1975.  Lizard  repro- 
duction effort:  caloric  estimates  and  comments 
on  its  evolution.  Ecology  56:427-4.34. 

Tinkle,  D.  W.,  H.  M.  Wilbur,  and  S.  G.  Tilley.  1970. 
Evolutionary  strategies  in  lizard  reproduction. 
Evolution  24:5.5-74. 

Turner,  F.  B.,  G.  A.  Hoddenbach,  P.  A.  Medica,  and  J. 
R.  La.n.nom.  1970.  The  demography  of  the  lizard, 


Uta  stansburiana  Baird  and  Girard,  in  southern 
Nevada.  J.  Anim.  Ecol.  .39:50,5-519. 

Turner,  F.  B.,  P.  A.  Medica,  and  B.  W.  Kowalewsky. 
1976.  Energy  utilization  by  a  desert  lizard  (Ufa 
stansburiana).  US/IBP  Desert  Biome  Monogr. 
No.  1.  Utah  State  Univ.  Press,  Logan. 

Turner,  F.  B.,  P.  A.  Medica,  J.  R.  Lannom,  and  G.  .\. 
Hoddenbach.  1969.  A  demographic  analysis  of 
fenced  populations  of  the  whiptail  lizard,  Cne- 
midophorus  tigris,  in  southern  Nevada.  South- 
west. Nat.  14:189-202. 

Turner,  F.  B.,  P.  A.  Medica,  and  D.  D.  Smith.  1973. 
Reproduction  and  survivorshp  of  the  lizard,  Uta 
stan.iburiana,  and  the  effects  of  winter  rainfall, 
density  and  predation  on  these  processes.  US/IBP 
Desert  Biome  Res.  Memo.  RM  74-26,  Utah  State 
Univ.,  Logan. 

ViTT,  L.  J.,  and  R.  D.  Ohmart.  1977.  Ecology  and  repro- 
duction of  lower  Colorado  River  lizards:  II.  Cne- 
midophorus  tigris  (Teiidae),  with  comparisons. 
Herpetologica  33:223-234. 

WoRTHiNGTON,  R.  D.,  AND  E.  R.  Arvizo.  1973.  Density, 
growth  and  home  range  of  the  lizard  Uta 
stansburiana  stejnegeri  in  southern  Dona  Ana 
County,  New  Mexico.  Great  Basin  Nat. 
.33:124-128. 


HAPLOPAPPUS  ALPINUS  (ASTERACEAE):  A  NEW  SPECIES  FROM  NEVADA 


Loran  C'.  Anderson' 


Abstr.\ct.—  The  new  species,  llaplopapptis  alpinus  of  section  Toncstus,  is  forniallv  described  and  illustrated.  It  is 
endemic  to  the  high  mountains  of  central  Nevada.  Vegetative  and  floral  morphology  of  related  species  is  detailed. 
The  new  species  appears  to  he  most  closely  related  to  H.  exirnius  but  also  demonstrates  close  affinity  to  H.  aberrans 
(all  three  are  diploids  with  n  =  9). 


In  North  America,  Haplopappus  (Aste- 
raceae)  contain.s  about  95  .species  represented 
in  17  sections.  Additional  species— including 
the  type  species,  H.  ghttinosus—Sire  in  South 
.\merica.  Chronio.somally,  two  major  group- 
ings can  be  identified  in  the  genus  (Anderson 
at  al.  1974).  One  group  is  generally  her- 
baceous and  chromosomallv  based  on  x  =  4, 
5,  or  6.  The  other,  more  woody  group,  is 
based  on  x  =  9.  These  groupings  are  poorly 
distinguished  by  growth  form.  Woodiness  in 
the  "herbaceous  group"  is  seen  in  sections 
Isoconia  and  Hazardia  (correlated  with  their 
xeromorphy?),  and  reduced  woodiness  is  seen 
in  the  "shrubby  group"  in  sections  Tonestus 
and  Stcnotus  where  their  growth  forms  are 
apparently  related  to  their  montane  or  alpine 
habitats. 

The  generic  integrity  of  this  assemblage 
has  been  challenged.  Many  feel  that  Hap- 
lopappus  is  unnatural  and  should  be  broken 
up  (Shinners  1950,  Anderson  1966,  Turner 
and  Sanderson  1971,  Clark  1977,  Urbatsch 
1978);  some  would  raise  each  section  to  ge- 
neric standing.  Others  feel  many  of  the  sec- 
tions are  interrelated;  i.e.,  the  South  Ameri- 
can taxa  and  Hazardia  (Grau  1976).  Jackson 
(1966)  has  demonstrated  genetic  relationship 
among  many  of  the  sections  of  the  "her- 
baceous group"  through  intersectional  hybr- 
idizations; he  has  recently  (pers.  comm.) 
hybridized  South  American  taxa  with  those 
of  .section  Hazardia. 

Until  more  is  known  about  the  biology  of 
Haplopappiis  (especially  the  South  American 
taxa),  Hall's  con.servative  generic  treatment 


(1928)  seems  better  at  the  moment  than  the 
alternative  of  elevating  each  section  to  ge- 
neric status.  Possibly  a  half  dozen  genera  are 
represented  in  the  North  American  material, 
but  I  can  not  envision  a  precise  treatment 
now. 

Description  of  a  new  species  in  section  To- 
nestus  presents  a  problem.  That  section  is  in 
the  x  =  9  group,  whereas  the  type  species, 
H.  ghitinosus,  is  n  =  5  (personal  count  in 
1971  from  Kew  Garden  material;  Grau  1976). 
So,  in  retaining  the  wider  generic  inter- 
pretation at  this  time,  a  species  of  Hap- 
lopappus  will  be  named  that  most  likely  will 
be  transferred  to  another  genus  when  the 
complex  is  better  known.  Deferring  descrip- 
tion of  this  species  until  a  comprehensive  ge- 
neric revision  is  available  might  possibly 
withhold  additional  data  that  would  be  sup- 
portive of  the  ultimate  revision. 

Methods  and  Materials 

Fresh  and  dried  materials  were  processed 
as  in  Anderson,  1964.  Five  heads  were  mea- 
.sured  for  involucral  and  floral  data.  Cytolo- 
gical  methods  are  those  of  Anderson,  1966. 

Plant  materials  were  collected  personallv 
in  the  field  or  .supplied  by  Sherel  Goodrich. 
Vouchers  for  various  measurements  and 
chromosome  counts  are  at  FSU. 

Taxonomy 

Haplopapfnis  alpinus  L.  C.  Anderson  &  S. 
Goodrich,  sp.  nov. 


'Department  of  Biological  Science,  Florida  State  University,  Tallahassee,  Florida  .33206. 


73 


74 


Great  Basin  Naturalist 


Vol.  40,  No.  1 


Herba  perennis  et  lignosa,  0.5-2.0  dm  alta; 
stirpes  glandulosae;  folia  in  basi  obovata  vel 
oblanceolata,  serrata  vel  dentata,  3-7  cm 
longa,  10-36  mm  lata,  folia  caulina  aliquan- 
temis  angustiora  et  serrata,  3-5.5  cm  longa, 
8-18  mm  lata;  inflorescentia  vel  mon- 
ocephala  vel  cyma  paucis  cum  capitibus;  in- 
volucra  10-12  mm  longa,  circa  7  mm  lata, 
phyllariis  21-28,  exterioribus  ovatis  et  folio 
similibus  et  glandulosis,  interioribus  bracteis 
angustioribus;  disci  florum  29-55,  flavi,  co- 
rollis  5.8-7.1  mm  longis,  lobis  circa  1.3  mm 
longis,  lineis  stigmaticis  saepissime  longitu- 
dine  paribus  styli  appendicibus;  achaenia  4-5 
mm  longa  et  pubescentia. 

Type:  Nevada,  Nye  Co.,  granitic  rocks  at 
10,600  ft  on  11,077-ft  peak  on  Toiyabe  Crest 
between  Washington  Creek  and  Aiken 
Creek,  24  air  mi  SSW  of  Austin,  21  Jun  1979, 
L.  C.  Anderson  4885  (BRY-holotype!,  FSU!, 
m^!,  UC!). 

Perennial  herb,  woody  only  at  base,  short 
rhizomatous,  (0.5)0.7-1.0(2.0)  dm  tall;  stems 
branched  only  in  the  inflorescence  or  mon- 
ocephalous,  densely  glandular-pubescent;  fo- 
liage dark  green,  glandular-pubescent,  basal 
leaves  obovate  to  oblanceolate,  petiolate,  ser- 
rate to  deeply  toothed  above  the  middle,  3-7 
cm  long,  10-36  mm  wide,  moderately  vis- 
cous, cauline  leaves  oblanceolate  to  spatu- 
late,  cuneate  or  clasping  the  stem,  saliently 
dentate,  3-5.5  cm  long,  8-18  mm  wide;  in- 
florescence usually  monocephalous  (open 
sites)  or  with  up  to  5  heads  in  an  elongate  or 
flat-topped  cyme  (deep  crevices  or  protected 
sites);  heads  campanulate  to  hemispheric, 
10-12  mm  long,  7-10  mm  wide  (pressed), 
phyllaries  21-28,  outer  ones  nearly  as  long  as 
involucre,  leaflike,  broadly  ovate,  3-nerved, 
glandular,  slightly  spreading,  obtuse  with 
small  mucro,  inner  bracts  narrower,  lanceol- 
ate-spatulate,  with  finely  ciliate  margins, 
acuminate-cuspidate;  ray  flowers  absent;  disk 
flowers  (29)35-50(55),  golden-yellow,  corollas 
(5.8)6.4-7.1(7.6)  mm  long,  lobes  (1.0)1.3(1.6) 
mm  long,  lanceolate,  slightly  spreading  to  re- 
curved; anthers  about  2.6  mm  long,  appen- 
dages 0.6  mm  long,  style  branches  slender, 
stigmatic  lines  nearly  as  long  as  style  appen- 
dages; achenes  cylindric  to  hisiform,  4-5  mm 
long,  pubescent,  pappus  dull  white,  6-7  mm 
long;  n  =  9  (Fig.  1).  Infrequent  on  boulders, 
talus,  or  rocky  summits  near  treeline  (primar- 


ily on  light-colored  granites  but  occasionally 
on  basalt,  andesite,  metamorphics,  or  lime- 
stone), 9,000-11,000  ft,  Toiyabe  and  To- 
quima  mountains  of  southern  Lander  and 
Nye  counties,  Nevada.  Mid-July-September. 

Additional  specimens  examined:  Nevada, 
Lander  Co.,  peak  between  Aiken  and  Carsely 
Creek,  S.  Goodrich  12137  (FSU,  UTC);  Nye 
Co.,  type  locality,  S.  Goodrich  12126  (FSU, 
NY,  UTC),  head  of  left  fork  San  Juan  Creek, 
S.  Goodrich  11997  (BRY,  UTC),  S.  Goodrich 
12006  (FSU,  UTC),  McLeod  Creek,  S.  Good- 
rich 13437  (BRY,  FSU),  crest  between  Tim- 


Fig.  1.   Representative,  but  somewhat  stout,  specimen 
of//,  alpinus;  drawn  largely  from  Goodrich  12233  (FSU). 


March  1980 


Anderson:  A  Nevada  Haplopappus 


75 


blin  Creek  and  Marysville  Canyon,  S.  Good- 
rich 12226  (BRY),  toiyabe  crest  at  French 
VABM,  S.  Goodrich  12233  (FSU),  right  fork 
Stewart  Creek,  S.  Goodrich  13502  (BRY),  top 
of  Shoshone  Mtn.,  Toquima  Range,  S.  Good- 
rich 6  F.  Smith  13267  (FSU).  All  collections 
but  the  last  came  from  the  Toiyabe  Range. 
Goodrich  (pers.  comm.)  also  reports  seeing  a 
population  on  Mt.  Jefferson  at  11,000  ft  at 
the  head  of  left  fork  of  Barker  (Shipley) 
Creek  in  the  Toquima  Range.  The  species  is 
found  for  about  23  miles  along  the  crest  of 
the  Toiyabe  Range  from  the  Lander-Nye 
County  line  south  to  the  head  of  Stewart 
Creek  and  is  reported  for  two  sites  in  the  To- 
quimas. 

At  the  type  locality,  H.  alpinus  occurs  on 
windswept  slopes  above  the  treeline  with  H. 
macronema,  ChrysotJiamnus  viscidiflorus, 
Erigeron  compositus,  and  Eriogonian  iim- 
bellatU7n.  Some  sites  are  at  or  just  below 
treeline,  where  the  species  occurs  on  rocks  in 
scattered  Pinus  flexilis  or  Cercocarpus  ledi- 
foUiis.  Other  alpine  endemics  from  central 
Nevada  that  have  been  found  in  the  vicinity 
of  H.  alpinus  are  Draba  arida,  Eriogonum 
ovalifolium  var.  caelestinum,  Geranium 
toquimense,  Hackelia  sp.  nov.,  Senecio  sp. 
nov.,  and  Smelowskia  holmgrenii. 

Relationships  and  Phytogeography 

This  species  belongs  to  section  Tonestus 
and  is  related  to  H.  aberrons  and  H.  eximius 
(Figs.  2-4).  Comparative  floral  features  for 


the  three  species  plus  the  more  distantly  re- 
lated H.  peirsonii  are  given  in  Table  1;  all  are 
distinctive.  The  Nevadan  endemic  is  like  H. 
abcrrans  in  its  eradiate  heads  of  similar  size; 
however,  the  latter  has  differently  shaped 
cauline  leaves  that  overtop  the  racemosely 
disposed,  turbinate  to  narrowly  campanulate 
heads.  Also,  the  bracts  are  more  numerous, 
narrower,  and  somewhat  squarrose  in  H. 
abcrrans  compared  to  H.  alpinus.  The  new 
species  is  more  similar  to  H.  eximius  in  leaf 
size  and  shape,  and,  although  the  phyllaries 
are  fairly  similar,  the  campanulate  heads 
differ  with  those  of  H.  eximius,  being  radiate 
with  shorter  involucres.  Haploppapus  alpinus 
also  differs  from  H.  eximius  in  the  following 
minor  floral  features:  H.  alpinus  has  stouter 
corolla  tubes  (as  in  Chrijsothamnus  spathu- 
latus  versus  C.  viscidiflorus;  illustrated  in  An- 
derson 1964);  its  pappus  is  shorter  than  the 
corolla  length,  whereas  pappus  equals  corolla 
length  in  H.  eximius;  and  it  has  longer  corolla 
lobes. 

Original  meiotic  chromosome  counts  for 
the  taxa  include:  H.  alpinus,  n  =  9  {Ander- 
son 4885,  the  type  collection);  H.  abcrrans,  n 
=  9  {Anderson  3660  from  Blaine  Co.,  Idaho); 
and  H.  peirsonii,  n  =  45  (Anderson  4326 
from  Inyo  Co.,  California).  The  count  for  H. 
peirsonii  agrees  with  the  earlier  count  by 
Stebbins,  who  also  reported  H.  eximius  as  a 
diploid  (Howell  1950).  The  other  counts  rep- 
resent first  reports  for  those  species.  Meiosis 
appeared  normal  with  pairing  as  bivalents  in 
all  instances.  Pollen  stainability  for  Anderson 


'■JU 


Figs.  2-4.  Flowering  heads  and  individual  outer  bracts;  heads  scaled  to  5  mm  bracket,  individual  bracts  slightly 
enlarged.  Fig.  2.  H.  aherrans  (Anderson  3660,  FSU).  Fig.  3.  //.  alpinus  (C^oodricli  12137,  FSU).  Fig.  4.  //.  eximius 
[Anderson  4320,  FSU). 


76 


Great  Basin  Naturalist 


Vol.  40,  No.  1 


4885  was  97.7  percent,  and  there  were  no  in- 
dications of  apomixis  in  H.  alpinus. 

Haplopappus  aberrans  was  originally  de- 
scribed as  an  aberrant  member  of  section 
Macronema  and  was  later  tentatively  as- 
signed to  Tonestus  (Hall  1928),  but  until  now 
it  has  been  considered  poorly  placed  in  To- 
nestus (Cronquist  1955).  With  the  addition  of 
H.  alpinus,  the  section  houses  H.  aberrans 
more  comfortably. 

Haplopappus  alpinus  is  .somewhat  inter- 
mediate between  H.  aberrans  and  //.  eximius 
morphologically  and  geographically.  Harper 
et  al.  (1978)  consider  the  Great  Basin  moun- 
tain ranges  flori.stically  as  islands  in  the  sur- 
rounding desert  possibly  populated  by  migra- 
tions from  the  "mainland"  mountain  systems 
of  the  Rockies  or  Sierras.  The  introduction  of 
H.  alpinus  from  eximiuslike  precursors  from 
the  Sierra  Madre  seems  very  plausible.  The 
possible  origin  of  H.  aberrans  in  the  Saw- 
tooth Mountains  of  Idaho  from  alpinuslike 
stock  poses  an  interesting  situation.  Hap- 
lopappus aberrans  occurs  in  the  Rocky 
Mountain  "mainland"  system  (in  which  H. 
hjallii  and  H.  pijgmaeus  of  Section  Tonestus 
occur).  Still,  its  affinities  lie  with  H.  alpinus 


and  H.  eximius.  The  species  could  represent 
the  culmination  of  a  migration  from  the 
western  Sierra  mainland  across  the  Great  Ba- 
sin deserts  to  the  Sawtooths  and  the  eastern 
mainland.  These  mountain  groups  were  less 
isolated  in  the  relatively  recent  past  (Harper 
et  al.  1978),  and  such  a  migration  is  plausible. 
Billings  (1978)  suggests  the  alpine  flora  in 
the  Great  Basin  may  have  resulted  from  "up- 
ward evolution"  of  preadapted  desert  species 
of  lower  elevations.  This  doesn't  .seem  to  ap- 
ply to  H.  alpinus  or  other  Tonestus  taxa.  Bil- 
lings further  suggests  that  due  to  reduced 
habitat  diversity  in  Great  Basin  mountains 
there  is  a  trend  toward  edaphic  endemism. 
Again,  H.  alpinus  does  not  follow  the  trend; 
it  has  been  collected  on  granite,  basalt,  meta- 
morphics,  andesite,  and  limestone  in  the  geo- 
logically diverse  Toiyabe  Mountains.  Clearly, 
these  alpine  areas  of  central  Nevada  do  need 
more  vegetational  work,  as  Billings  (1978) 
observed.  Our  ideas  of  that  region  may  be 
greatly  changed  with  further  study;  endem- 
ism apparently  is  not  as  low  as  Harper  et  al. 
(1978)  record.  Goodrich,  for  example,  has 
found  several  imdescribed  endemics  in  his 
current  survey  of  the  region. 


T.\BLE  1.—  .\veraged  floral  data  (and  ranges  entered  parenthetically)  for  .selected  taxa  of  Haplopappus,  section  To- 
nestus. 


Taxon  anc 

i  collection 

H.  aberrans 

Anderson 
3660 

H.  alpinus 

H.  eximius 

H.  peirsonii 

Feature 

Anderson 
4885 

Goodrich 
12137 

Anderson 
4320 

Anderson 
4899 

Anderson 
4326 

Involucre 

Bract  number 
Length,  mm 

42.0  (37-49) 
11.0(10-12) 

24.2  (21-25) 
10.6(10-11) 

23.8  (21-28) 
11.0(10-12) 

27.2  (25-30) 
8.5  (8-9.5) 

29.0  (24-35) 
9.0  (8.2-10) 

27.0  (23-29) 
15.2  (13-16) 

Rav  flowers 

Flower  number 
Flower  length,  mm 
Flower  width,  mm 

- 

- 

- 

13.6(11-19) 
10.5 
2.2 

15.0  (12-20) 
10.2 
2.1 

21.8(21-23) 
10.4 
4.4 

Disk  flowers 

Flower  number 
Flower  length,  mm 
Lobe  length,  mm 
Stigmatic  area— total 
style  branch,  % 

38.0  (29-48) 
7.6  (7.0-8.2) 
1.0(0.9-1.1) 

51.7  (50-54) 

41.6  (.35-49) 
6.3(5.8-7.1) 
1.3(1.0-1.6) 

50.0  (43-55) 

44.4  (29-55) 
7.1  (6.4-7.6) 
1.3(1.2-1.4) 

48.1  (45-50) 

59.2  (54-69) 
7.1  (6.6-7.4) 
1.1(1.0-1.2) 

56.3  (44-62) 

51.6(44-60) 
6.4  (6.0-7.0) 
1.0(0.7-1.2) 

53.7  (50-56) 

55.8  (45-75) 
8.4  (7.0-9.5) 
1.1(1.1-1.2) 

.38.1  (33-43) 

March  1980 


Anderson:  A  Nevada  Haplopappus 


77 


Acknowledgments 

Appreciation  is  expressed  to  Sherel  Good- 
rich for  guiding  me  to  the  alpine  collection 
sites  and  for  supplying  habitat  data  and  sev- 
eral collections.  Melanie  Darst  prepared  the 
line  drawings;  Dr.  Walter  Forehand  is 
thanked  for  assistance  with  the  Latin  diag- 
nosis. This  study  was  supported  by  National 
Science  Foundation  grant  DEB  76-10768. 


Literature  Cited 

Anderson,  L.  C.  1964.  Taxonomic  notes  on  the  Chrijso- 
thamnus  viscidiflorus  complex  (Astereae.  Com- 
positae).  Mandrono  17:222-227. 

1966.  Cytotaxonomic  studies  in  Chitj.sothdmiuis 

(Astereae,  Conipositae).  .\nier.  ].  Bot. 
53:204-211. 

.\nderson,  L.  C,  D.  W.  Kyhos,  T.  Mosquin,  A.  M. 
Powell,  .\nd  P.  H.  Raven.  1974.  Chromosome 
numbers  in  Conipositae.  IX.  Haplopappus  and 
other  .\stereae.  .\mer.  J.  Bot.  61:66.5-671. 

BiLLLNGS,  D.  W.  1978.  .\lpine  phytogeography  across  the 
Great  Basin.  Great  Basin  Nat.  Mem.  2:105-117. 


Clark,  W.  D.  1977.  Chemosystematics  of  the  genus 
HaMrdia  (Conipositae).  J.  Ariz.  .\cad.  Sci.  12:16. 

Cro.nquist,  .v.  1955.  Conipositae.  V.  In:  C.  L.  Hitch- 
cock, A.  Cronquist,  M.  Ownbey,  J.  W.  Thompson 
(eds.).  Vascular  plants  of  the  Pacific  Northwe.st, 
Univ.  of  Washington  Press,  Seattle. 

Grau,  J.  1976.  Chromosomenzahlen  von  Siidanierikanis- 
chen  Haplopappus  ,\rten.  Mitt.  Bot.  .Miinchen 
12:40.3-410. 

IL\LL,  H.  M.  1928.  The  genus  Haplopappus,  a  phyloge- 
netic  studv  in  the  Conipositae.  Carnegie  Inst. 
Publ.  .389:i-.391. 

Harper,  K.  T.,  D.  C.  Freeman,  W.  K.  Ostler,  and  L. 
G.  Klikoff.  1978.  The  flora  of  Great  Basin  moun- 
tain ranges:  diversity,  sources,  and  dispersal  ecol- 
ogy. Great  Basin  Nat.  Mem.  2:81-103. 

Howell,  J.  T.  1950.  Studies  in  California  Aplopappus. 
Leafl.  West.  Bot.  4:84-88. 

Jackson,  R.  C.  1966.  Some  intersectional  hybrids  and 
relationships  in  Haplopappus.  Univ.  Kans.  Sci. 
Bull.  46:47.5-488. 

Shlnners,  L.  H.  1950.  Notes  on  Texas  Conipositae,  IV, 
V.  Field  &  Lab.  18:25-42. 

Turner,  B.  L.,  and  S.  Sanderso.n.  1971.  Natural  hybridi- 
zation between  the  Composite  "genera"  Machae- 
ranthera  and  Haplopappus  (sec.  Blepharodon). 
Amer.  J.  Bot.  .58:467. 

Urbatsch,  L.  E.  1978.  The  Chihuahuan  Desert  species 
of  Ericaincria  (Compositae:  .\stereae).  Sida 
7:298-303. 


MISCELLANEOUS  PLANT  NOVELTIES  FROM  ALASKA,  NEVADA,  AND  UTAH 

Stanley  L.  Welsh'  and  Sherel  Goodrich' 

\BSTRACT  -  Described  as  new  to  science  are  Ahwnia  ar^ilhsa  Welsh  &  Goodrich,  from  Utah  and  Colorado;  An- 
drosace  alaskana  Gov.  &  Standi,  var  recdae  Welsh  &  Goodrich,  from  Kohlsaat  Peak,  Alaska;  Lepidium  osten  Welsh 
&  Goodrich,  from  Beaver  Gounty,  Utah;  Lygode.mia  entrada  Welsh  &  Goodrich,  from  Grand  Connty,  Utah;  Pedto- 
cactm  des-painii  Welsh  and  Goodrich,  from  Emery  County,  Utah;  and  Senecio  toiyabensis  Welsh  and  Goodrich,  from 
the  Toiyabe  Range  in  Nye  Coimty,  Nevada. 


Several  undescribed  and  unnamed  taxa 
have  accumulated  at  the  herbarium  of  Brig- 
ham  Young  University  in  recent  years. 
Mainly  they  represent  materials  which  have 
been  treated  within  other  taxa,  or  they  are 
oddities  of  exceedingly  limited  areal  extent. 
Some  have  been  known  for  long  periods  of 
time.  Others  are  only  recently  discovered.  All 
are  unique  in  one  or  more  ways  and  of  suffi- 
ciently limited  apparent  distribution  as  to  be 
candidates  for  inclusion  on  lists  of  sensitive 
species. 

Abronia   argillosa    Welsh    &   Goodrich,   sp. 
nov. 

Plantis  similis  Abronia  fragranti  Nutt.  et 
Abronia  elliptica  in  habitas  sed  differt  in  an- 
thocarpus  alarum  nuUis  rostro  nuUo  gracilior- 
ibus,  in  receptaculo  conico  et  fructibus  in- 
structis  superioris  foliis  floribus  et 
anthocarpis  glabris,  et  floribus  paucioribus. 

Plantae  perennes  e  caudicibus  ramifican- 
tibus  gracilibus  (6)  15-30  cm  altae;  caules 
glabri  ad  basim  nibellos  saepe  frondosi  om- 
nino;  folia  (5)  15-35  mm  longa,  3-35  mm  lata 
elliptica  cvato  obovata  vel  suborbiculares 
glabra;  pedunculi  1-8  cm  longi  glabri  vel 
puberulentes  raro;  bractae  7-15  mm  longae, 
6-15  mm  latae  ovales  vel  orbiculares  sca- 
riosae  glabrae  vel  ciliolatae;  flores  15-22  in 
quoque  inflorescentiam;  perianthi  tubus 
10-15  mm  longus  viridis  glaber  vel  raro  pub- 
enilentus  limbus  ca.  6  mm  latus  albidus;  re- 
ceptaculiun  breve  conicum  ferens  fnictus  in 
dimidio  superiore  anthocarpus  sine  alis  sine 


rostris  plicatus  leviter  vel  nullus  7-9  mm 
longus  3-4  mm  latus  scariosus  glabri  rugosus 
alborostratus;  semina  2.5-3  mm  longa  1  mm 
lata. 

Holotype:  Utah,  Grand  Co.,  T22S,  R24E, 
Sec  18,  ca.  6  miles  due  south  of  Cisco  at  ca. 
4300  feet  elev.,  on  Mancos  Shale  Formation, 
in  an  Atriplex  community,  S.,  E.,  and  M. 
Welsh  16689,  30  May  1979  (BRY,  8  isotypes 
to  be  distributed). 

Paratypes:  Utah,  Grand  Co.,  Fifteen  miles 
east  of  Thompson,  B.  F.  Harrison  et  al  10403, 
16  June  1941  (BRY);  first  escarpment  north  of 
Thompson,  west  of  Sego  Canyon,  S.  L.  Welsh 
6943,  1  May  1968  (BRY);  T22S,  R24E,  Sec  7, 
ca.  4  mi.  south  of  Jet.  50-6  and  U-128,  S.  L. 
Welsh  and  K.  Taylor  14637,  28  April  1977 
(BRY);  T18S,  R25E,  Sec  27,  ca.  20  mi.  NE  of 
Cisco,  S.  L.  Welsh  14916,  8  June  1977  (BRY). 
Uintah  Co.,  Ca.  2  mi.  S  of  Dragon,  S.  L. 
Welsh  5379,  13  May  1966  (BRY).  Colorado, 
Mesa  Co.,  ca.  5  mi.  W  of  Mack  along  US 
Hwy  6-50,  L.  C.  Higgins  and  S.  L.  Welsh 
1034,  14  June  1967  (BRY);  ca  13  Km  due  NW 
of  Mack,  T9S,  R104W  ca.  Sec  11,  A. 
Cronquist  11427,  25  May  1976  (BRY;NY). 

The  clay  verbena,  Abronia  argillacea 
Welsh  and  Goodrich,  is  restricted  to  the 
Grand  River  Valley,  and  less  commonly  in 
the  drainage  of  the  White  River  in  east-cen- 
tral Utah  and  west-central  Colorado,  where  it 
occurs  on  heavy  soils  derived  from  Mancos 
Shale  and  Green  River  formations.  It  seems 
probable  that  the  taxon  was  taken  much  pri- 
or to   the  specimens  cited  above,  but  was 


'Life  Science  Museum  and  Department  of  Botany  and  Range  Science,  Brigham  Young  University,  Provo,  Utah  84602. 


78 


March  1980 


Welsh,  Goodrich:  Plant  Novelties 


79 


KHT 


Fig.  L  Abronia  argillosa:  A,  habit  of  plant. 


80 


Great  Basin  Naturalist 


Vol.  40,  No.  1 


overlooked  due  to  the  superficial  resem- 
blance to  phases  of  A.  elliptica  A.  Nels.  and 
to  A.  fragrans  Nutt.  ex  Hook. 

The  plants  are  glabrous  or  essentially  so 
and  possess  orbicular  leaves  which  are 
glaucous.  The  genus  has  been  revised  by  Gal- 


loway (1975),  who  distinguishes  A.  elliptica 
(common  and  widespread  in  Utah)  from  A. 
fragrans  (cited  from  San  Juan  County  only) 
by  technical  characters  of  the  anthocarps. 
The  following  key  will  distinguish  A.  argil- 
losa  from  those  taxa. 


1.  Anthocarps  with  2  wings,  these  folded  together  to  form  a  deep  groove;  plants 

of  western  Colorado  and  westward A.  elliptica  A.  Nels. 

—  Anthocarps  wingless,  or,  if  winged,  then  the  wings  not  folded  together;  plants 

of  eastern  Utah  and  eastward 2 

2(1).  Anthocarps  beaked,  deeply  grooved  or  narrowly  winged;  leaf  blades  variable 
in  shape,  glandular-puberulent  to  glandular-pubescent;  plants  usually  of  sandy 

soils  in  San  Juan  Co.,  Utah,  and  eastward  in  Colorado  

A.  fragrans  Nutt.  ex  Hook 

—  Anthocarps  beakless,  slightly  or  not  at  all  folded;  leaf  blades  mainly  orbicular, 
glabrous;  plants  usually  of  clay  soils  in  Uintah  and  Grand  counties,  Utah  and 
Mesa  Co.,  Colorado  A.  argillosa  Welsh  &  Goodrich 


Androsace  alaskana  Gov.  &  Standi,  ex  Hul- 
ten 

var.  reedae  Welsh  &  Goodrich,  var.  nov. 

Similis  Androsace  alaskana  Gov.  &  Standi, 
in  scapis  numerosis  floribus  sessilibus  solitariis 
vel  binatum,  sed  differt  scapis  numerosioribus 
brevioribus  gracilioribus  pubescentioribus, 
foliis  integris  parvioribus  dense  villosis,  et  flo- 
ribus parvioribus. 

Herbae  annuae  vel  biennis;  scapi  25-40  in 
quoque  rosellam  10-45  mm  longi  filiformes 
0.3-0.5  mm  in  diametrum  pubentes  parce  vel 
dense  pilis  furcatis  apprime  infra  flores  juxta; 
folia  5-10  mm  longa  1-3  mm  lata  linearia  vel 
spathulata  integra  vel  denticulata  cum  1-2 
dentibus  glabra  infra  vel  pilis  ad  costam  juxta 
apicem  supra  dense  villosa  pilis  multi- 
cellulosis  furcatis  vel  simplicibus  apprime  ul- 
tra medium;  flores  solitari  vel  imparibus  ali- 
quando  bractaea  subtenda  tubus  calycis 
subaequali;  tubus  calycis  2-3  mm  longus, 
dentibus  circa  1.5  mm  longis;  corolla  alba 
tubo  calyce  subaequali  lobus  circa  2  mm 
longis;  capsulae  maturae  ignotae. 

Holotype:  Alaska,  Lat.  62°12'N.,  Long. 
152°47'W,  ca.  2  mi.  SSW  of  Kohlsaat  Peak, 
near  VABM  5048,  at  4900  feet  (1495  m) 
elev.,  on  rocky  ridge  top,  K.  Reed  5857,  29 
June  1977  (BRY,  isotype  at  Leningrad). 

Par.\type:  Yukon  Territory,  Canada, 
Mount  St.  Elias  Quad.:  Outpost  Mt.  at  south 


end  of  Kluane  Lake,  60°56'N,  138°22'W,  at 
ca.  2140  m.,  D.  F.  Murray  3014,  22  Julv  1969 
(BRY;  ALA). 

The  materials  herein  segregated  as  var. 
reedae  represent  the  slender  peduncled,  sub- 
entire-  to  entire-leaved,  smaller,  flowered 
high  elevation  phase  of  A.  alaskana  in  interi- 
or Alaska  and  southwestern  Yukon.  Tliat  the 
morphological  differences  noted  in  the  pro- 
tologue  might  be  the  result  of  ecological  re- 
sponse has  been  considered.  The  tendency  to 
entire  leaves  and  less  pronounced  ciliate  mar- 
gins, along  with  slender  peduncles  and  flow- 
ers that  seem  to  average  smaller,  indicate  a 
syndrome  of  characteristics  which  should  re- 
ceive at  least  some  taxonomic  recognition. 

The  plant  is  named  to  honor  the  collector 
of  the  holotype,  Katherine  Reed  of  Ancho- 
rage, Alaska. 

Lepidium  ostleri  Welsh  &  Goodrich,  sp.  nov. 

Habitu  Lepidio  nana  S.  Wats  sed  differt  in 
inflorescentia  longiore  floribus  numer- 
osioribus foliis  longioribusque  pubentioribus 
et  sepalis  pubentioribus. 

Herbae  perennes  pulvinatae  caespitosae 
caudice  ramoso  folium  basim  marcescentibus; 
caules  fnictifri  10-35  mm  alti  hirsuti;  folia 
4-15  mm  longa  hirsuta  linearia  et  integra  vel 
basalia  3-5  lobatis  vulgo  caulina  nulla  vel 
pauca;  racemi  circa  1  cm  longi  in  florem  et 


March  1980 


Welsh,  Goodrich:  Plant  Novelties 


81 


mm 


Fig.  2.  Androsace  alaskana:  A,  adaxial  side  of  leaf;  B,  abaxial  side  of  leaf;  C,  elose-iip  of  stem  and  inflorescence;  D, 
habit  of  plant. 


82 


Great  Basin  Naturalist 


Vol.  40,  No.  1 


:pi.^      I  mm 


mm  '•'iiLv-M  /•; 


Fig.  3.  Lepidiiim  ostleri:  A,  habit  of  plant;  B.  close-up  of  leaves  showing  variation;  C,  close-up  of  flower;  D,  close- 
up  of  fniit;  E,  close-up  of  inflorescence. 


March  1980 


Welsh,  Goodrich:  Plant  Novelties 


83 


1-2  cm  longi  in  fructem  5-25  floribus;  pedi- 
celli  in  fructem  2-3  mm  longi;  sepala  1.3-2 
mm  longa  obtusa  hyalina  aspre  pilosa  saepe 
purpurascentia;  petala  2-3  mm  longa  alba 
purpiirascentia;  fnictus  2.5-3  mm  longa, 
2.3-2.5  mm  lata  ovata  late  usque  ad  1  mm 
crassa  sinus  0.2  mm  profundus;  stylus  0.4-0.7 
mm  longus. 

Holotype:  Utah,  Beaver  Co.,  T27S, 
R13W,  Sec  23  (SW'/4),  San  Francisco  Moun- 
tains, at  Frisco,  on  rocky  ridge,  in  pinyon- 
jimiper  community,  K.  Ostler  and  D.  Ander- 
son 1258,  6  June  1978  (BRY). 

Paratypes:  Utah,  Beaver  Co.,  T27S, 
R13W,  Sec  23  (NW1/4),  near  Frisco,  6900 
feet,  rocky  slopes,  K.  Ostler  and  D.  Anderson 
1210,  1  June  1978  (BRY);  do,  T27S,  R13W, 
Sec  16  (SE^/i),  San  Francisco  Mountains,  near 
Utah  Hwy  21,  at  5800  ft,  on  west  slope,  dry 
limestone,  Atriplex  community,  K.  Ostler 
1415,  19  June  1978  (BRY);  do,  west  slope  of 
San  Francisco  Mountains,  bristlecone  pine, 
ponderosa  pine,  Douglas  fir,  white  fir  com- 
numity,  K.  Ostler  1588,  5  July  1978  (BRY). 

The  obvious  relationship  of  this  perennial 
dwarf  species  lies  with  the  Lepidium  nantim, 
an  endemic  of  Nevada.  The  longer  racemes, 
greater  flower  number,  and  dense  pub- 
escence constitute  the  most  important  diag- 
nostic features.  The  petals  of  Lepidium  ostleri 
are  white,  while  those  of  L.  nanum  are  yel- 
low to  cream  yellow.  While  probably  of  less 
importance  than  other  features,  the  color  of 
the  petals  becomes  important  when  taken 
with  the  differences  in  raceme,  flower  num- 
ber, and  pubescence  features. 

The  species  is  named  in  honor  of  its  collec- 
tor. Dr.  Kent  Ostler,  an  enthusiastic  collector 
and  botanist. 

Lygodesmia  entrada  Welsh  &  Goodrich,  sp. 
nov. 

Ab  Lygodesmia  grandiflora  Nutt.  in  caul- 
ibus  rigidioribus  ramosissimis  foliis  brevior- 
ibus  et  paucioribus  et  radiis  albis  differt. 

Herbae  perennes  caudice  subterraneo 
ramosissimo  omnino  usque  ad  45  cm  altae; 
folia  integra  linearia  vel  acicularia  5-30  mm 
longa;  pedunculi  potius  numerosi  bracteati 
elongati  12-20  cm  longi  in  capitulum  termi- 
nans;  bracteae  involucrorum  hyalinae  exte- 
riores  5-10  mm  longae  fimbrillatae  interiores 
circa  sex  16-18  mm  longae  apex  puberulus; 


radii  albi  circa  3  cm  longi;  pappus  barbel- 
latus  sordidus  setae  10-15  mm  longae;  ach- 
enia  costata  glabra. 

Holotype:  Utah,  Grand  Co.,  T24S,  R19E, 
Sec  25,  Tusher  Canyon,  ca.  15  mi.  due  WNW 
of  Moab,  4800  feet  elev.,  Entrada  Sandstone 
Formation,  juniper  community,  S.  L.  and  S. 
L.  Welsh  16725,  3  June  1978  (BRY,  four  iso- 
types  to  be  distributed). 

This  white-flowered  material  has  been 
identified  by  A.  S.  Tomb  (pers.  comm.)  as  a 
probable  triploid  assignable  to  Lygodesmia 
arizonica  Tomb.  The  triploid  hypothesis  can- 
not herein  be  questioned  because  of  lack  of 
knowledge  concerning  the  cytological  nature 
of  the  plants  in  question.  However,  despite 
the  ultimate  disposition  of  these  plants  fol- 
lowing future  determinations  of  chromosome 
numbers,  the  strikingly  different  morpholo- 
gical features  dictate  taxonomic  recognition, 
if  for  no  reason  other  than  the  fact  that  the 
plants  are  so  different  from  other  plants  of 
Lygodesmia  in  Utah. 

A  second  collection  at  BRY,  here  assigned 
to  L.  entrada,  is  J.  S.  Allen  132,  from  north  of 
Courthouse  Wash  Ridge  in  Arches  National 
Park.  Dried  flower  remnants  appear  to  be 
pink,  but  the  tall  nidularius  habit  and  defi- 
nitely ligneous  stems  and  branches  are  appar- 
ent. 

Lygodesmia  arizonica  Tomb  is  a  low  her- 
baceous plant  usually  of  more  southern  distri- 
bution in  Utah.  Even  in  late  anthesis  the 
stems  are  herbaceous  and  lack  the  character- 
istic bird's  nest  appearance  of  L.  entrada. 
Lygodesmia  entrada  differs  from  L.  grandi- 
flora in  ways  similar  to  those  discussed  for  L. 
arizonica. 

Pediocactus  despainii  Welsh  &  Goodrich,  sp. 
nov. 

Ab  Pediocacto  hradyi  L.  Benson  differt  in 
spinis  paucioribus  brevioribus  gracilioribus  et 
floribus  colorum. 

Plantae  carnosae  hemisphericae  depressae 
3-6  cm  in  diametrum  4-8  cm  longae;  tuber- 
culi  ovata  numerosa  ordinata  in  serialia  cir- 
cularia  vel  spiralia;  areolae  spinis  8-14;  spin- 
ae  2-5  mm  longae  serialia  stellatim;  pilis 
coactis  instructis  interdum;  flores  2.5-4  cm 
diametrum  fragrantes  ad  apicem  gerenti;  se- 
pala numerosa;  petala  numerosa  albida  suf- 
fusa  rosea  et  flava;  stamina  numerosa  lutea 


84 


Great  Basin  Naturalist 


Vol.  40,  No.  1 


Fig.  4.  Lygodesmia  entrada:  A,  habit  of  plant;  B,  close-up  of  achene. 


March  1980 


Welsh,  Goodrich:  Plant  Novelties 


85 


cm 


B 


<c:^ 


Fig.  5.  Pcdiocactus  despainii:  A,  habit  of  plant;  B,  habitat  of  plant. 


86 


Great  Basin  Naturalist 


Vol.  40,  No.  1 


omnino;  stigmata  numerosa  lutea  omnino; 
fmctus  8-9  mm  diametrum  10-12  mm  longus 
operculo  circumscissili  umbone  8-9  mm  dia- 
metnim  5.5-6  mm  alto;  fructus  corpi  8-9 
mm  diametrum  5.5-6  mm  longus  findens 
longirostronim  coloris  viridis  ad  porphyreus; 
seminum  3-3.5  mm  longum  2-2.5  mm  latum 
varicosis  tuberculatibus  minutibus. 

Holotype:  Utah,  Emery  Co.,  San  Rafael 
Swell,  Despain  266a,  5  May  1978  (BRY). 

Paratypes:  Utah,  Emery  Co.,  San  Rafael 
Swell,  Despain  445,  15  May  1979  (BRY);  do, 
E.  Neese  &  K.  Thome  504,  7  May  1979. 

The  Despain  pediocactus  is  a  diminutive 
cactus  of  very  local  distribution  in  the  San 
Rafael  Swell  of  Emery  County,  Utah.  Exact 
locality  is  not  given  so  as  to  provide  a  mea- 
sure of  protection  of  this  species  from  ama- 
teur and  commercial  fanciers.  The  species  is 
compared  in  the  protologue  to  P.  bradyi, 
from  which  it  differs  as  indicated.  It  is  similar 
to  the  newly  described  P.  winkleri  (Heil 
1979)  from  Wayne  County,  but  differs  inter 
alia  in  the  monocephalous  nature,  stems 
which  average  larger  (3-6  cm  in  diameter  vs. 
2-2.6  cm),  larger  flowers  (2.5-4  cm  broad  vs. 
1.7-3  cm),  and  shorter  capsules  (5.5-6  mm 
long  vs.  7-9  mm). 

The  species  is  named  in  honor  of  its  discov- 
erer, Mr.  Kim  Despain,  student  of  the  flora  of 
the  San  Rafael  Swell. 

Senecio  toiyabensis  Welsh  &  Goodrich,  sp. 
nov. 

Ab  Senecio  fremontii  T.  &  G.  foliis  integris 
angustis  et  statura  elata  differt. 

Herbae  perennes  caudicibus  ramificantibus 
ligneis;  caules  15-60  cm  altae  glabri  vel  pub- 
escentes  parce;  folia  accrescentia  sursum 
magniora  linearia  integra  vel  denticulata  2-8 
cm  longa  2-7  mm  lata;  bracteae  diminutae; 
inflorescentia  corymbosa;  bracteae  in- 
volucromm  ca  13,  6-8  mm  longae  margines 
hyalini  apices  acuti  pilosi  breves  vel  glabri 
raro  rubentes  interdum;  radii  8  vel  pauciores 
circa  1  cm  longi  lutei;  achenia  scabra. 

Holotype:  Nevada,  Nye  Co.,  Toiyabe  Na- 
tional Forest;  Toiyabe  Range,  just  under  the 
crest  of  the  range  on  the  leeward  side,  above 
Timblin  Cr.,  35  air  miles  SW  of  Austin, 
100-500  yards  N  of  French  VABM,  T13N, 
R42E,  near  center  of  Sec.  4,  10,500  feet,  in 
cracks  of  metamorphic  rocks  and  talus,  with 


Artemisia  michauxiana,  Penstemon  watsonii, 
Eriogonwn  microthecwn,  Philadelphiis  micro- 
plujllus,  and  Sphaeromeria  cana,  Goodrich 
12235,  30  Aug.  1978  (BRY;  numerous  iso- 
types  to  be  distributed). 

Paratypes:  Nevada,  Nye  Co.,  Toiyabe  Na- 
tional Forest,  Toiyabe  Range,  along  or  near 
the  crest  of  the  range  between  San  Juan- 
Tierney  Creeks  and  McLeod  Cr.,  T14N, 
R43E,  in  or  near  W1/2  of  Sec.  11,  9800-10,000 
feet,  on  metamorphic  precambrian  outcrops, 
talus  and  rocky  ground,  Goodrich  &  Schlatter- 
er  12156,  10  Aug.  1978  (BRY);  do,  north  rim 
of  Aiken  Creek,  very  near  the  Lander-Nye 
Co.  marker;  T15N,  R43E,  Sec.  17,  NE1/4  of 
SEV4,  10,800  feet,  on  rocky  slope;  growing 
with  Cymopterus  petraeus,  Haplopappus 
macronema,  Penstemon  speciosus,  Oryzopsis 
hymenoides,  and  scattered  Pinus  flexilis, 
Goodrich  12138,  5  Aug.  1978  (BRY);  Nevada, 
Lander  Co.,  east  side  of  Bunker  Hill,  16.5  mi. 
187°  from  Austin,  N39n5'25-35"  W117°7' 
10-20",  11,000  ft.,  steep,  rocky  limestone 
slopes,  Goodrich  13338,  10  July  1979  (BRY). 

The  Toiyabe  groundsel  is  a  near  congener 
of  S.  fremontii  from  which  it  differs  in  having 
linear  entire  leaves  (not  ovate  or  obovate  to 
oblanceolate  and  dentate).  The  leaves  are  2-7 
mm  wide,  compared  to  1-4  cm  wide  in  S. 
fremontii.  Stems  are  mostly  erect  and  are 
1.5-6.0  cm  tall  when  in  an  thesis. 

In  the  key  to  group  X.  Triangulares  by 
Barkley  (1978),  S.  toiyabensis  would  require 
modification  as  follows: 

2.  Plants  taprooted,  or  with  a  sub- 
rhizomatous  caudex  surmounting  a  tap- 
root. 

3.  Leaves  linear,  entire;  plants  1.5-6 
dm  tall,  restricted  to  the  Toiyabe 

Range,  Nevada  S.  toiyabensis 

3.  Leaves  ovate  or  obovate  to  ob- 
lanceolate, dentate;  plants  1-3(4) 
dm  tall,  distribution  not  as  above 

S.  fremontii 

2.  Plants  variously  fibrous  rooted  or  with  a 
persistent  caudex  but  not  taprooted. 

Literature  Cited 

Barkley,  T.  M.  1978.  Senecio.  Fl.  N.  Amer.  Series  11, 
Part  10:  50-139.  New  York  Bot.  Card.,  New  York. 

Galloway,  L.  A.  1975.  Systematics  of  the  North  Ameri- 
can desert  species  of  Ahronui  and  Thptcrocahjx 
(Nyctaginaceae).  Brittonia  27:328-347. 


March  1980 


Welsh,  Goodrich:  Plant  Novelties 


87 


Fig.  6.  Senccio  toiyabensis:  A,  close-up  of  flower;  B,  habit  of  plant. 


Great  Basin  Naturalist 


Vol.  40,  No.  1 


Heil,  K.  D.  1979.  Three  new  species  of  Cactaccae  from 

southeastern  Utah.  Cact.  &  Succulent  J.  (U.S.)  51: 

25-30. 
Hitchcock,  C.  L.  1936.  The  genus  Lepidium  in  the 

United  States.  Madroiio  3:  265-320. 
To.MB,  .\.  S.  1979.  Novelties  in  Lygodesmia  and  Stephan- 

omeria     (Compositae-Cichorieae).     Sida     3(7): 

530-5.32. 


Welsh,  S.  L.  1974.  Anderson's  flora  of  Alaska  and  adja- 
cent parts  of  Canada.  Brigham  Young  University, 
Provo,  Utah. 

Welsh,  S.  L.,  and  J.  L.  Reveal.  1977.  Utah  flora:  Brassi- 
caceae  (Cruciferae).  Great  Basin  Nat.  37: 
279-365. 


NEW  GENERA  AND  NEW  GENERIC  SYNONYMY  IN  SCOLYTIDAE  (COLEOPTERA) 

Steplieii  L.  Wood' 

Abstract.—  New  generic  synonymy  in  the  world  fauna  of  Scolytidae  includes:  Acanthotomictis  Blandford 
{  =  Isophthorus  Schcdl),  Acranttt.s  Broun  {  =  Chaetophorotis  Fuchs,  Chaetopteliiis  Fuchs),  Cosmudercs  Eichhoff 
{  =  Erioschiilia.s  Schedl),  Ernoporicus  Berger  (  =  Ernopoceius  Balachowsky),  Ernoporus  Thomson  {  =  Euptiliits  Schcdl), 
Hyliirdri'ctonii,s  Schedl  ( =  .\'(//og(>/JHiw,y  Schedl),  Ozopanon  Hagedorn  {Dnjocoetiops  Schedl),  Scolijtogenes  Eichhoff 
(  =  Cnjpliahmiorplius  Schauhiss).  Stepluinopodius  Schedl  {^Crijphdloniinictes  Browne),  and  Xijlechintis  Chapuis 
(^Sijuaina.'iinulus  Nimberg).  Genera  new  to  science  and  their  tvpe-species  include:  Anaxylchont.'i  {Tomicits  tnin- 
ccittis  Erichson),  Apoxi/lchorus  (Xt/lcborus  mancus  Blandford),  Crifphulogenes  {Cri/pluilo^enes  euphorbiae  Wood),  Er- 
nochidius  {Cn/phahis  corpulcntu.s  Sampson),  lladrodcniius  (Xijlebonis  globus  Blandford),  Leptoxt/lebonis  (Phloeo- 
trootis  sordicauda  Motschulsky),  Micropenis  (Xijlebonis  theae  Eggers),  Taphrodasus  [Xijlebonis  penorthi/liis  Schedl), 
and  Taiirodemiis  (Xijlebonis  sharpi  Blandford).  The  new  name  Hijliirdn'ctoniis  corticiiiiis  is  presented  to  replace  H. 
araiicariae  (Schedl  1972).  Dnjocoetes  coffeae  Eggers  is  transferred  to  Eiilepiops.  The  following  genera  are  treated  in  a 
revised  context:  Cn/togeniiis.  Dnjocoetes,  Eiilepiops,  Enioporiciis,  Enwponis,  Xijlebonis,  and  Xylechinus.  Cnjphalo- 
genes  euphorbiae  and  C.  exiguus  (Sri  Lanka)  are  named  as  new  to  science. 


In  a  review  of  the  genera  of  Scolytidae  in 
the  world  fauna,  several  problems  that  relate 
to  synonymy  were  encountered.  The  new 
svnonvmv  listed  in  the  above  abstract  is  re- 
ported here  in  order  that  names  might  be 
used  in  their  new  context  before  the  generic 
revision  is  completed.  In  addition,  several 
genera  are  treated  in  a  sense  somewhat  dif- 
ferent from  the  traditional.  The  basis  for 
these  departures  is  established.  The  genera 
are  treated  alphabetically  for  convenience  of 
reference.  They  include  representatives  from 
the  subfamily  Hylesininae,  tribe  Tomicini 
{Acmntiis,  Hyhirdrectonus,  Xylechinus)  and 
from  the  subfamily  Scolytinae,  the  tribes 
Ipini  {Acanthotornicus),  Dryocoetini  {Cijrto- 
genitis,  Eulepiops,  Ozopemon),  Xyleborini 
(Xyleborus),  and  Cryphalini  {Cosmoderes,  Er- 
noporictis,  Ernoporus,  Scolytogenes,  Steph- 
anopodius).  Nine  new  genera  represent  the 
tribes  Xyleborini  {Anoxyleborus,  Apoxyle- 
borus,  Hadrodcmius,  Leptoxyleborus,  Micro- 
perns,  TapJirodasus,  Taurodemus)  and 
Cryphalini  (Cryphalogenes,  Ernocladius). 
Xylechinosomus  Schedl  is  removed  from  syn- 
onymy with  Pteleobius.  The  new  name  Hy- 
hirdrectonus corticinus  is  presented  to  re- 
place the  junior  homonym  H.  araucariae 


(Schedl  1972).  The  species  Cryphalogenes  eu- 
phorbiae and  C.  exiguus  (Sri  Lanka)  are 
named  as  new  to  science. 


Acanthotornicus  Blandford 

Acanthotornicus  Blandford,  1894,  Trans.  Ent.  Soc.  Lon- 
don 1894:89  (Type-species:  Acanthotornicus  spin- 
osus  Blandford,  monobasic) 

Isophthorus  Schedl,  1938,  Archiv  Naturgesch.  7(2):  173 
(Type-species:  Isophthorus  quadrituberculatus 
Schedl,  present  designation).  .Vpir  synonymy 

In  the  original  description  of  Isophthorus 
Schedl,  two  .species  were  definitely  included 
and  a  third  species  was  doubtful,  but  a  type- 
species  was  never  designated.  Since  then, 
Schedl  has  transferred  all  three  species  else- 
where. To  anchor  the  generic  name,  Isoph- 
thorus quadritidierculatus  Schedl  is  here  des- 
ignated as  the  type-species  of  Isophthorus. 
Because  this  species  and  Myeloborus  bico- 
nicus  Schedl  have  been  transferred  to  Acan- 
thotornicus and  the  unrelated,  doubtfiil  spe- 
cies, Pityophthorus  heteae  Hagedorn,  has 
been  transferred  to  Cryptocarenus,  the  fix- 
ation of  a  type-species  requires  that  Isoph- 
fliorus  be  placed  in  .synonymy  under  Acan- 
thotornicus. 


'Life  Science  Museum  and  Department  of  Zoology.  Brigham  Young  University.  Provo.  Utah  84602.  Scolytidae  contribution  number  69. 


89 


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Great  Basin  Naturalist 


Vol.  40,  No.  1 


Acrantus  Broun 

Hfliminis  Broun,  1881,  Manual  of  New  Zealand  Coleop- 

tera  2:720  (Tvpe-species:   Homanis  inunduliis 

Broim,  monobasic).  Preoccupied 
Acrantus  Broun,   1882,  Ann.   Mag.   Nat.  Hist.  (5)9:409. 

Replacement  name 
Chaetophorus  Fuchs,  1912,  Morphologische  studien  iiber 

Borkenkafer,   II.  die  europiiischen   Hyiesinen,   p. 

46  (Tvpe-species:  Hylesiniis  vestitus  Mulsant  6c 

Re\',  monobasic).  Preoccupied 
Clwetoptelius  Fuchs,  191.3,  in  Reitter,  Wiener  Eut.  Zeit. 

.32(Beiheft):4.3.  {Replacement  name).  Xeic  synon- 

!/'"!/ 
The  names  Acrantus  Broun  and  Chaetop- 
telitis  Fuchs  have  been  treated  as  synonyms 
of  Pteleobius  Bedel  (Schedl  1963:262)  and 
Pseudohijlesinus  Swaine  (Schedl  1966:75),  re- 
spectively. However,  in  a  review  of  the  char- 
acters of  the  type-species  of  these  genera,  it 
was  demonstrated  (Wood  1978)  that  Pte- 
leobius must  be  placed  in  the  tribe  Hylesinini 
and  that  Chaetopteliiis  and  Pseudohijlesinus 
belong  in  the  tribe  Tomicini.  For  that  study, 
Schedl's  (1963:262)  placement  of  Acrantus 
was  not  challenged. 

In  a  subsequent  review  of  the  genera  of 
Tomicini,  di.ssection  demonstrated  that  Ho- 
marus  mundulus  Broun,  type-species  of  Acr- 
antus, clearly  belongs  to  the  Tomicini  and  is 
quite  unrelated  to  Pteleobius.  Furthermore, 
Pseudohijlesinus  totally  lacks  pronotal  aspe- 
rities, it  has  three  distinct  sutures  on  the  an- 
tennal  club,  and  the  male  frons  is  not  noticea- 
bly impres.sed.  Acrantus,  Chaetopteliiis,  and 
Xijlechinosomus  all  have  numerous  pronotal 
asperities,  two  or  four  poorly  marked  sutures 
on  the  antennal  club,  and  the  male  frons 
strongly  impressed  and,  thus,  form  a  group 
quite  distinct  from  Pseudohijlesinus.  Xijle- 
chinosomus, which  Schedl  (1966:75)  also 
placed  in  synonymy  with  Pteleobius,  has  the 
antennal  club  less  elongate,  less  strongly 
compressed,  and  (apparently)  with  four  ob- 
•scure  sutures  and  the  frontal  rectangle  much 
more  elongate.  Acrantus  and  Chaetopteliiis 
have  the  antennal  club  more  elongate, 
strongly  flattened,  and  marked  by  two  su- 
tures and  the  frontal  rectangle  comparatively 
broad.  Biological  differences  also  support  the 
continued  recognition  of  Xijlechino.wmus. 
However,  I  can  find  no  characters  that  sepa- 
rate Acrantus  and  Chaetoptelius.  For  this  rea- 
son, Chaetoptelius  is  placed  in  synonvmv  un- 
der Acrantus,  as  indicated  above. 


Acrantus  includes  mundulus  and  vestitus, 
cited  above,  and  most  if  not  all  of  the  species 
from  New  Zealand,  Australia,  New  Guinea, 
and  neighboring  areas  placed  by  Schedl  in 
Leperisinus  and  Xijlechinus. 

Anaxyleborus,  n.  gen. 

This  genus  is  distinguished  from  Eii- 
wallacea  Hopkins  and  allied  genera  by  the 
truncate,  concave  elytral  declivity  which  has 
a  complete,  .sharply  elevated,  circum- 
declivital  costa  from  base  to  apex.  The  discal 
interstrial  punctures  are  uniseriate;  in  the  su- 
perficially similar  Apoxijleborus  they  are  con- 
fused. 

Description.—  Antennal  club  with  one  su- 
ture visible  on  posterior  face,  anterior  face 
with  segment  1  corneous,  2  conspicuous, 
sometimes  rather  large.  Procoxae  contiguous. 
Protibia  armed  by  more  than  11  socketed 
teeth.  Declivity  and  discal  punctures  on  in- 
terstriae  as  described  in  above  diagnosis. 

Type-species:  Tomicus  truncatus  Erichson. 

Species  assigned  previously  to  the  Xijle- 
borus  truncatus  group  belong  here. 


Apoxyleborus,  n.  gen. 

This  genus  is  distinguished  from  Tauro- 
demus  by  the  presence  of  only  four  to  seven 
socketed  teeth  on  the  protibia,  by  the 
obliquely  truncate  elytral  declivity,  with  an 
abrupt  (not  acute)  circumdeclivital  costa,  and 
the  face  flat  to  weakly  concave.  It  is  distin- 
gui-shed  from  the  superficially  similar  Ana- 
xyleborus by  the  rather  widely  separated  pro- 
coxae,  by  the  strongly  confused  interstrial 
punctures  on  the  disc,  and  by  the  less  dis- 
tinctly concave  elytral  declivity. 

Description.—  Body  stouter  than  1.9  times 
as  long  as  wide.  Antennal  club  with  .segment 
1  corneous,  with  no  sutures  evident  on  poste- 
rior face,  apical  margin  of  segment  1  on  ante- 
rior face  acutely  elevated  into  a  continuous 
co-sta  forming  a  complete  circle.  Procoxae 
moderately  separated.  Protibia  armed  by  four 
to  seven  socketed  teeth.  Elytral  disc  with  in- 
terstrial punctures  strongly  confused,  declivi- 
ty as  described  in  above  diagnosis. 

Tvpe-species:  Xijleborus  mancus  Bland- 
ford. 


March  1980 


Wood:  American  Bark  Beetles 


91 


Species  assigned  previously  to  the  Xyle- 
bonis  mancus  group  belong  here. 

Cosmoderes  Eichhoff 

Cosmoderes  Eichhoff,  1878,  preprint  of  Mem.  Soc.  Roy. 
Sci.   Liege  (2)8:49.5  (Type-species:  Cosmoderes 
moiiilUcollis  Eichhoff,  monobasic) 
Erioschidias  Schedl,  1938,  Trans.  Roy.  Soc.  S.  Austraha 
62:42  (Type-species:   Crijphalus  sctistriattis   L^a, 
subsequent  designation  by  Wood,  1960,  Insects  of 
Micronesia  18(1):21).  New  sijnoiuit)nj 
The  Beeson  Collection  at  the  Forest  Re- 
search Institute,  Dehra  Dim,  India,  contains 
series  of  three  species  that  were  placed  by 
Beeson  in  Cosmoderes.  One,  from  Samsingh, 
Kalimpong,    Bengal,    is   labeled   monillicoUis 
Eichhoff;   the  other  two  bear  manuscript 
names    not    yet    validated.    Beeson 's    private 
notes,  of  which  two  volumes  treating  Scoly- 
tidae  are   in  my  possession,   contain   no   in- 
dication under  this  name   that  he  saw   the 
type  of  moniUicoUis.  However,  elsewhere  in 
his  notes  there  are  several  indications  that  he 
saw  the  Eichhoff  Collection  at  Hamburg  be- 
fore it  was  destroyed  during  World  War  II. 
Blandford  also  saw  the  Eichhoff  Collection, 
but  there  is  some  doubt  (Blandford  1894:86) 
that  he  actually  examined  the  type  of  mon- 
illicoUis. 

Both  the  Beeson  and  Blandford  specimens 
are  congeneric  with  Erioschidias  Schedl.  Bee- 
son's  .specimens  of  monillicoUis  match  the  dis- 
tinctive characters  of  Eichhoff's  description 
in  every  detail.  It  is,  therefore,  propo.sed  that 
Erioschidias  be  placed  in  synonymy  under 
Eichhoff's  name,  as  indicated  above. 


Cryplialogenes,  n.  gen. 

This  genus  is  distinguished  from  the  closely 
allied  Scolytogenes  Eichhoff  by  the  3-seg- 
mented  antennal  funicle,  by  the  antennal 
club  with  sutures  1  and  2  weakly  procurved, 
marked  by  .setae,  and  1  grooved  and  partly 
.septate,  and  by  tlie  horizontal  venter  of  the 
abdomen. 

Description.—  Frons  convex,  not  sexually 
dimorphic.  Eye  elongate-oval,  entire.  Anten- 
nal scape  elongate,  simple;  fimicle  3-seg- 
mented;  club  oval,  a  slight  constriction  and 
groove  at  suture  1,  sutures  1  and  2  moder- 
ately procurved,  1  partly  septate  at  lea.st  on 
lateral  half.  Pronotum  with  basal  and  lateral 


margins  marked  by  a  fine,  raised  line;  ante- 
rior slope  asperate,  anterior  margin  armed  by 
low,  poorly  formed  serrations.  Elytral  punc- 
tures largely  replaced  by  rows  of  rounded 
strial  and  interstrial  granules;  vestiture  of 
rows  of  -Strial  hair  and  interstrial  scales.  Pro- 
tibia  armed  by  four  socketed  teeth.  Venter  of 
abdomen  horizontal.  Sexes  subequal  in  size. 

Type-species:  Cryphalogenes  euphorbiae 
Wood. 

Cryphalogenes  euphorbiae,  n.  sp. 

This  species  is  distinguished  from  exiguus 
Wood  by  the  larger  size,  by  the  absence  of 
reticulation  of  the  pronotum  (except  in  ex- 
treme lateral  areas  of  some  specimens),  and 
by  the  comparatively  .smaller  pronotal  and 
elytral  granules. 

Male.—  Length  1.2-1.4  mm,  2..3  times  as 
long  as  wide;  color  dark  brown. 

Frons  broadly  convex,  very  feebly  so  on 
longitudinal  axis;  surface  largely  reticulate, 
minute  punctures  moderately,  uniformly 
abundant,  most  of  them  feebly  granulate.  An- 
tennal club  slightly  longer  than  scape. 

Pronotum  1.0  times  as  long  as  wide;  sides 
almost  straight  and  parallel  on  basal  third, 
rather  broadly  rounded  in  front;  anterior 
margin  armed  by  about  four  to  six  irregular, 
poorly  formed  serrations;  summit  near 
middle;  anterior  .slope  rather  coarsely  aspe- 
rate, punctured  between  asperities;  posterior 
areas  smooth,  shining  (except  some  reti- 
culation present  in  extreme  lateral  areas  of 
some  specimens),  with  close,  moderately 
large,  rounded  granules,  anterior  slope  of 
each  granule  bearing  a  puncture  (punctures 
usually  visible  only  when  light  source  ceph- 
alad).  Vestiture  of  fine,  .short,  semirecumbent 
hair. 

Elytra  1.3  times  as  long  as  wide,  1.4  times 
as  long  as  pronotum;  .sides  almost  straight  and 
parallel  on  basal  two-thirds,  rather  broadly 
rounded  behind;  striae  not  impressed,  each 
puncture  largely  replaced  by  a  large  roimded 
granule  as  wide  as  striae,  puncture  confined 
to  posterior  slope  of  each  granule;  interstriae 
as  wide  as  striae,  .smooth,  shining,  punctures 
largely  replaced  by  rounded  granules  of  same 
size  and  shape  as  those  of  striae.  Declivity 
steep,  convex;  sculpture  as  on  disc.  Vestiture 
of  rows  of  fine,  short,  strial  hair  and  rows  of 


92 


Great  Basin  Naturalist 


Vol.  40,  No.  1 


erect  interstrial  scales,  each  scale  slightly 
shorter  than  distance  between  rows,  spaced 
within  a  row  by  length  of  scale,  each  four  to 
six  times  as  long  as  wide  on  disc,  two  to  four 
times  on  declivity. 

Female  —  Similar  to  male  in  all  respects. 

Type  locality-  Thirty  km  southeast  of 
Puttalam,  Sri  Lanka  (Ceylon). 

Type  material—  The  male  holotype,  fe- 
male allotype,  and  34  paratypes  were  taken 
at  the  type  locality  on  18-VI-1975,  No.  214, 
from  Euphorbia  antiquonim,  by  me;  28  para- 
types bear  the  same  data  except  they  were 
taken  24  km  SE  Puttalam.  Additional  para- 
tvpes  were  taken  in  Sri  Lanka  as  follows:  21 
at  5  km  SE  Naula,  14-VM975;  14  at  48  km 
N  Naula,  14-VI-1975;  2  at  32  km  N  Haba- 
rana,  12-VI-1975;  1  at  8  km  SW  Kurunegala, 
13-VI-1975;  and  1  at  11  km  W  Kikirawa,  19- 
VL1975;  all  from  the  same  host  and  collec- 
tor. 

The  holotype,  allotype,  and  half  the  para- 
types are  in  the  U.S.  National  Museum.  The 
remaining  paratypes  are  in  my  collection. 


Cryphalogenes  exiguus,  n.  sp. 

This  species  is  distinguished  from  eu- 
phorhiae  Wood  by  the  smaller  size,  by  the 
strongly  reticulate  pronotum,  and  by  the 
comparatively  larger  pronotal  and  elytral 
granules. 

Male.—  Length  0.8-1.0  mm,  2.2  times  as 
long  as  wide;  color  dark  brown. 

Frons  as  in  euphorbiae  except  more  strong- 
ly convex,  granules  smaller,  less  conspicuous. 
Antennal  club  with  septum  in  suture  1  less 
apparent. 

Pronotum  as  in  euphorbiae  except  reti- 
culate, shining,  granules  in  posterior  areas 
proportionately  slightly  larger. 

Elytra  as  in  euphorbiae  except  interstrial 
scales  averaging  more  slender,  those  on  de- 
clivity not  less  than  four  times  as  long  as 
wide. 

Female.—  Similar  to  male  in  all  respects. 

Type  locality.—  Thirty  km  southeast  of 
Puttalam,  Sri  Lanka  (Ceylon). 

Type  material-  The  male  holotype,  fe- 
male allotype,  and  43  paratypes  were  taken 
at  the  type  locality  on  18-VI-1975,  No.  214, 
from  Euphorbia  antiquonim,  by  me.  Addi- 


tional paratypes  were  taken  in  Sri  Lanka  dur- 
ing 1975  from  the  same  host,  by  me,  as  fol- 
lows: 13  at  24  km  SE  Puttalam,  17-VI;  5  at  5 
km  SE  Naula,  14-VI;  4  at  48  km  N  Naula,  14- 
VI.  The  specimens  were  taken  in  indepen- 
dent galleries  in  the  same  stems  with  eu- 
pliorbiae. 

The  holotype,  allotype,  and  half  the  para- 
types are  in  the  U.S.  National  Museum.  The 
remaining  paratypes  are  in  my  collection. 


Cyrtogenius  Strohmeyer 

Ki/rtogcniits  Strohmeyer,  1910,  Ent.  Blatt.  6:127  (Type- 
species:  Kijrtogeniiis  bicolor  Strohmeyer,  mon- 
obasic) 

Ciirtooenius  Strohmeyer,  1911,  Ent.  Bliitt.  7:116.  Valid 
emendation 

Carposinus  Hopkins,  1915,  U.S.  Dept.  Agric.  Rept.  99:9, 
47  (Tvpe-species:  Carposiniis  pini  Hopkins 
=  Lepicenis  nitidus  Hagedorn,  original  designa- 
tion) 

Orosiotes  Niisima,  1917,  Coll.  Essays  Y.  Nawa,  p.  1 
(Type-species:  Orosiotes  kumatoensis  Niisima, 
monobasic) 

Metahylastes  Eggers,  1922,  Ent.  Bliitt.  18:165  (Type-spe- 
cies: Metahijlastes  africanus  Eggers,  monobasic) 

Pelicenis  Eggers,  1923,  Zool.  Meded.  Roy.  Mus.  Nat. 
Hist.  Leyden  7:216  (Type-species:  Lepicerus  ni- 
tidus Hagedorn,  original  designation) 

Taphrohorus  Nunberg,  1961,  Ann.  Mag.  Nat.  Hist. 
(13)3:617  (Type-species:  Taphrohorus  vaticae 
Nunberg,  original  designation) 

Much  confusion  exists  in  the  literature  rel- 
ative to  the  identity  of  this  tropical  genus.  It 
is  characterized  by  five  socketed  teeth  on  the 
lateral  margin  of  the  protibia,  by  the  posteri- 
or face  of  the  antennal  club  with  only  one  su- 
ture, sutures  on  anterior  face  procurved,  with 
the  pubescence  extending  to  the  base,  by  the 
narrowly  separated  procoxae,  and  by  the 
slightly  elevated  or  armed  posterolateral 
margin  of  the  elytral  declivity.  Dryocoetes 
differs  from  it  by  the  recurved  suture  1  on 
the  antennal  club,  the  pubescence  never  ex- 
tending to  the  base,  by  the  contiguous  pro- 
coxae,  and  by  the  rounded,  unarmed,  post- 
erolateral margins  of  the  elytral  declivity. 
Both  genera  are  phloeophagous  and  have 
heterosanguineous  polygenous  breeding 
habits  in  which  the  male  is  subequal  in  size  to 
the  female  and  assists  in  the  formation  of  new 
parental  galleries.  Both  genera  have  been 
confused  with  Eulepiops  (see  below). 


March  1980 


Wood:  American  Bark  Beetles 


93 


Ernocladiiis,  n.  gen. 

This  genus  is  distinguished  from  the  closely 
allied  Ernoporus  Thomson  by  the  3-seg- 
mented  antennal  funicle,  by  the  uniseriate  in- 
terstrial  setae  (interstrial  ground  vestiture  al- 
ways absent  on  disc,  a  few  setae  sometimes 
present  on  declivity),  and  by  the  weakly  pro- 
curved  (often  obscure)  sutures  of  the  antennal 
club. 

Description.—  Frons  dimorphic,  moder- 
atelv  impressed  in  male,  convex  in  female. 
Eye  elongate-oval,  entire.  x\ntennal  scape 
elongate;  funicle  3-segmented;  club  rather 
large,  sutures  weakly  to  moderately  pro- 
curved,  aseptate,  marked  by  rows  of  setae, 
grooves  present  or  not.  Pronotum  with  basal 
margin  marked  by  a  fine,  raised  line,  lateral 
margin  rounded,  without  a  raised  line;  aspe- 
rities in  concentric  rows,  their  bases  often 
contiguous  or  even  reduced  to  a  continuous 
costa.  Elytra  with  basal  margins  rounded, 
strial  punctures  in  rows,  sculpture  conserva- 
tive; vestiture  of  rows  of  minute  strial  hair 
and  rows  of  erect  interstrial  scales,  interstrial 
ground  vestiture  absent  on  disc,  a  few  short 
setae  in  ground  cover  sometimes  present  on 
declivity. 

Type-species:  Cn/pJialus  corptilentus 
Sampson. 

Several  additional  species  will  be  trans- 
ferred to  this  genus  as  soon  as  their  types  can 
be  examined.  Schedl  (1940:590)  assigned  Cry- 
phuhis  corpulentus  to  Margadillius,  apparent- 
ly without  appreciating  the  significance  of 
tlie  emarginate  eye  or  the  fine,  raised  line  on 
the  lateral  margin  of  the  pronotum  of  Marga- 
dillius species. 


Ernoporicus  Berger 

Ernoporicus  Berger,  1917,  Rev.  Russc  d'Ent.  16:242 
(Tvpe-species:  Ernoporicus  spessivtzevi  Berger, 
monobasic) 

Eocnjphalm  Kurentzov,  1941,  Acad.  Sci.  USSR,  Koma- 
rov  Sta.  Sci.,  Orient,  p.  230  (Type-species:  Eocnj- 
pluilus  seinenovi  Kurentzov,  monobasic) 

Ernopocerus  Balachowsky,  1949,  Fauna  de  France 
5():211  (Type-species:  Ernoporus  caitcasicus  Lind- 
emann,  subsequent  designation  by  Wood,  19.54, 
Univ.  Kansas  Sci.  Bull.  36:986).  Xinc  sijnonyiny 

The  complex  of  genera  allied  to  Ernoporus 


Thom.son  have  been  poorly  known  and  erro- 
neously classified,  largely  due  to  the  paucity 
of  material  for  study.  Following  an  exam- 
ination of  the  type-species  of  Ernoporicus, 
Eocryphalus,  and  Ernopocerus,  it  was  con- 
cluded that  these  three  congeneric  species 
have  the  ba.sal  and  lateral  margins  of  the  pro- 
nottun  rounded  (without  a  fine,  raised  line), 
the  procoxae  narrowly  separated,  the  eye 
short  and  entire,  the  antennal  funicle  4-seg- 
mented,  and  the  antennal  club  with  the  su- 
tures procurved  and  marked  only  by  setae  or 
obsolete  (never  septate).  Ernoporus  kanawhae 
Hopkins  of  North  American  and  E.  fagi  (Fab- 
ricius)  and  a  few  species  from  A.sia  also  be- 
long here.  The  genus  Ernoporus  is  quite  dif- 
ferent, as  indicated  below. 


Ernoporus  Thomson 

Ernoporus  Thomson,  1859,  Skandinaviens  Coleoptera 
Svnoptiskt  Bearbitade,  p.  147  (Type-species:  Bos- 
trichus  tilkic  Panzer,  original  designation) 

Cryphalops  Reitter,  1889,  Wiener  Ent.  Zeit.  8:94  (Type- 
species;  Cn/phalus  lederi  Reitter  =Bostrichus 
tiliae  Panzer,  monobasic) 

Sfcphanorhopalus  Hopkins,  1915,  U.S.  Dept.  .\gric. 
Rept.  99:35  (Type-species:  Stephanorhopalus 
nulodori  Hopkins,  amended  to  melodori  by 
Schedl,  1966,  Ent.  Abh.  Mus.  Dresden  .35:19, 
original  designation) 

Euptilius  Schedl,  1940,  Mitt.  .Miincher  Ent.  Ges.  .30:.590 
(Type-species:  Ernoporus  concentralis  Eggers, 
original  designation).  New  synonymy 

Ernoporus  Thomson  has  the  basal  and  lat- 
eral margins  of  the  pronotum  marked  by  a 
fine,  rai.sed  line,  the  procoxae  contiguous, 
most  pronotal  asperities  arranged  in  con- 
centric rows,  the  antennal  funicle  4-seg- 
mented,  the  antennal  club  .sutures  strongly 
procurved  to  obsolete,  and  the  elytral  vesti- 
ture abundant  and  conhused.  Most  of  the  .spe- 
cies occur  in  tropical  .\sia  except  for  tiliae, 
the  type-species.  In  a  review  of  the  genera 
belonging  to  this  complex,  it  was  found  that 
Ernoporus  concentralis  Eggers  falls  well  with- 
in the  range  of  variability  for  Ernoporus.  Be- 
cause concentralis  is  the  type-species  of  Eu- 
ptUius  Schedl,  it  is,  therefore,  necessary  to 
place  Schedl's  genus  in  synonymy  as  in- 
dicated above.  The  structure  of  the  pronotum 
indicates  that  this  genus  is  quite  di.stinct  from 
Ernoporicus,  as  noted  above. 


94 


Great  Basin  Naturalist 


Vol.  40,  No.  1 


Eulepiops  Schedl 

Eulepiops  Schedl,  19.39,  J.  Fed.  Malay  St.  Mus.  18{,3):344 
(Tvpe-species:  EiiUpiops  glaber  Schedl,  mon- 
oba.sic) 
This  genus  has  been  confused  with  Cyrto- 
genius  Strohnieyer  and  Dnjocoetes  Eichhoff. 
It  differs  by  the  protibia  bearing  only  three 
socketed  teeth  on  the  lateral  margin,  by  the 
posterior  face  of  the  antennal  club  with  two 
sutures  indicated,  the  anterior  face  with  su- 
ture 1  straight  to  recurved  and  always  on  the 
basal  fourth.  The  male  is  either  unknown  or 
dwarfed,  deformed,  flightless,  and  does  not 
participate  in  the  formation  of  new  parental 
galleries.  Reproduction  is  either  by  con- 
sanguineous polygyny  or  possibly  by  some 
form  of  parthenogenesis.  The  habit  is  myelo- 
phagy  for  the  only  species  observed.  Dnjo- 
coetes coffeae  Eggers  and  its  allies  belong  to 
this  genus. 

Hadrodemius,  n.  gen. 

This  genus  is  distinguished  from  Eccoptop- 
tems  Eichhoff  by  the  tibiae  being  of  normal 
size  and  all  bearing  socketed  teeth,  by  the 
normal  metatarsi  (not  compressed),  by  the 
declivity  being  restricted  to  the  posterior  half 
of  the  elytra,  and  by  the  convex  to  moder- 
ately impressed,  unarmed  elytral  declivity. 

Description.—  Body  very  stout,  less  than 
1.8  times  as  long  as  wide,  usually  black.  An- 
tennal club  with  posterior  face  unmarked  by 
sutures,  on  anterior  face  costa  marking  apical 
margin  of  corneous  area  usually  forming  a 
complete  ring.  Scutellum  visible  only  on  an- 
terior declivous  slope  of  elytral  margins.  De- 
clivity and  tibiae  as  described  in  above  diag- 
nosis. 

Type-species:  Xylebarus  globus  Blandford. 

Members  of  the  Xyleborus  globus  species 
group  should  be  referred  here. 

Hylurdrectonus  Schedl 

Hylurdrectomis  Schedl,  1938,  Trans.  Roy.  Soc.  S.  .'Vustra- 
lia  62;4()  (Type-species:  Hylurdrectonus  piniarius 
Schedl,  monobasic) 

Xylogopinus  Schedl,  1972,  Papua  New  Guinea  .\5;ric.  J. 
23:64  (Type-species:  Xylogopinus  araucariae 
Schedl  =  Hylurdrectonus  corticinus  Wood,  mon- 
obasic). New  synonymy 

A  review  of  long  series  of  Hylurdrectonus 
piniarius    Schedl,    H.    araucariae    Schedl 


(1964a:213),  and  Xylogopinus  araucariae 
Schedl  indicates  the  absence  of  characters 
that  will  separate  these  two  genera.  Con- 
sequently, it  is  necessary  to  place  Xylogo- 
pinus in  synonymy  under  the  older  name  as 
indicated  above.  This  act  creates  homonymy 
as  indicated  below. 

Hylurdrectonus  corticinus,  new  name 

Xillogopinus  araucariae  Schedl,  1972,  Papua  New  Guin- 
ea Agric.  J.  23:64  (Bulolo,  Morobe  Distr.,  New 
Guinea) 

A  long  series  of  this  species  was  collected 
near  Bulolo  and  compared  to  the  holotype 
and  paratypes  in  the  Forest  Research  Labora- 
tory collection  at  Bulolo.  As  indicated  above, 
this  species  must  be  transferred  to  Hylurdrec- 
tonus. The  transfer  makes  this  species  a  jun- 
ior homonym  of  H.  araucariae  Schedl,  1964. 
Tlie  new  name  Hylurdrectonus  corticinus  is 
proposed  to  replace  H.  araucariae  (Schedl 
1972). 

Leptoxyleborus,  n.  gen. 

This  genus  is  distinguished  from  the  allied 
Theobortis  Hopkins  and  Coptoborus  Hopkins 
by  the  declivity  commencing  anterior  to  the 
middle  of  the  elytra,  its  lower  half  broadly 
impressed  and  either  flat  or  shallowly  con- 
cave. If  the  discal  interstrial  punctures  are 
uniseriate,  then  the  declivital  surface  is  den- 
sely covered  by  small,  confused  scales;  if  the 
declivital  setae  are  hairlike,  then  the  discal 
interstrial  punctures  are  confused. 

Description.—  Antennal  club  with  two  su- 
tures indicated  on  posterior  face,  anterior 
face  with  segment  2  comparatively  large, 
sclerotized,  convex,  apical  portion  beyond 
segment  2  flat  to  concave.  Protibiae  and 
metatibiae  each  armed  by  six  or  seven  sock- 
eted teeth.  Anterior  coxae  contiguous.  Scutel- 
lum visible.  Declivity  as  described  in  above 
diagnosis. 

Type-species:  Phloeotrogus  sordicauda 
Motschulsky. 

Other  species  placed  previously  in  the 
Xyleborus  sordicauda  group  also  belong  here. 

Micropenis,  n.  gen. 

This  genus  is  distinguished  from  Taphro- 
dastis  Wood  by  the  convex  elvtral  declivity 
tliat  lacks  a  circumdeclivital  costa,  by  the  ab- 


March  1980 


Wood:  American  Bark  Beetles 


95 


sence  of  declivital  scales,  and  by  the  strial 
pimctures  that  are  arranged  in  definite  rows. 

Description.—  Body  slender,  at  least  two 
times  as  long  as  wide,  color  yellowish  or  red- 
dish brown.  Posterior  face  of  antennal  club 
with  at  least  one  suture  visible,  apical  margin 
of  corneous  area  never  costate.  Scutellum  not 
visible.  Strial  punctures  usually  in  rows.  De- 
clivity convex,  variously  sculptured,  without 
a  costa. 

Type-species:  Xyleborus  theae  Eggers. 

Members  of  the  Xijleborus  theae  species 
group  should  be  referred  here.  The  name  Mi- 
cropenis was  originally  coined  by  F.  G. 
Browne  for  this  group  for  use  in  an  unpub- 
lished manuscript  a  decade  ago. 

Ozopeinon  Hagedorn 

Ozopcmon  Hagedorn,  1908,  Deutsche  Ent.  Zeitschr. 
1908:382  (Type-species:  Ozopemon  regius  Hage- 
dorn, monobasic) 

Dryocoetiops  Schedl,  1957,  Ann.  Mus.  Roy.  Congo  Beige, 
Tervairen,  Ser.  8,  Sci.  Zool.  .56:13  (Type-species; 
Ozopemon  laevis  Strohmeyer,  monobasic).  New 
sijnonymy 

A  series  of  Ozopemon  laevis  Strohmeyer 
was  compared  to  Eggers's  series  of  this  spe- 
cies and  to  representatives  of  eight  species  of 
Ozopemon.  Although  the  sculpturing  of  the 
pronotum  is  somewhat  unique  for  the  genus, 
this  species  appears  to  fall  well  within  the 
limits  of  variability  for  Ozopemon.  For  this 
reason,  Dryocoetiops  is  placed  in  synonymy 
as  indicated  above. 


Scolytogenes  Eichhoff 

Scolytogenes  Eichhoff.  1878,  preprint  of  Mem  Soc.  Roy. 
Sci.  Liege  (2)8:475,  497  (Type-species:  Scolyto- 
genes danvinii  Eichhoff.  monobasic) 

Cryphcilomorphus  Schaufnss,  1890  (1891),  Tijdschr.  Ent. 
34:12  (preprint  1890  by  Martinus  N'ijhoff,  Hagg) 
(Type-species:  Cryphahnurpbiis  conimiinif:  Schau- 
fnss, monobasic).  \eiv  synonymy 

Eggers  (1929:53)  examined  the  type-speci- 
mens of  the  type-species  of  Scolytogenes  and 
Lepicerus  and  compared  them  to  the  type- 
specimens  of  Negritus  major  Eggers  and  A'. 
minor  Eggers.  He  concluded  that  N.  major 
and  N.  minor  were  congeneric  with  Scoly- 
todes  danvinii  Eichhoff.  The  holotype  of  S. 
darwinii  apparently  was  lost  when  the  Stettin 
Museum  was  damaged  during  World  War  II. 
In  the  absence  of  that  type,  direct  com- 


parisons are  not  now  possible;  however,  if  it 
is  assumed  that  Eggers  was  correct  in  his  ob- 
servations, then  N.  major  and  S.  danvinii  are 
congeneric.  My  examination  of  the  lectotype 
of  A',  major  and  .syntypes  of  N.  ater  (type-spe- 
cies of  Negritus)  demonstrates  that  these  spe- 
cies are  congeneric;  consequently,  Negritus 
must  be  a  jimior  synonym  of  Scolytogenes. 
Because  N.  major  and  A\  ater  are  also  consid- 
ered congeneric  with  Cryplmlomorphus  com- 
munis Schaufuss  (type-species  of  Cryphalo- 
morplius)  (Schedl  1957:152),  it  must  also  be 
concluded  that  Cryphalomorphus  is  a  junior 
synonym  of  Scolytogenes. 

(Note  added  in  press:  The  list  of  types  in 
the  Schedl  Collection  at  the  Vienna  Museum, 
just  received,  includes  the  type  of  S.  darwinii. 
It  will  be  examined  as  soon  as  arrangements 
can  be  completed.) 


Stephanopodius  Schedl 

Stephanopodius  Schedl,  1941,  Rev.  Zool.  Bot.  Afr.  .34:.396 
(Type-species:  Stephanoderes  dispar  Eggers,  sub- 
sequent designation  by  Schedl,  1961,  Rev.  Ent. 
Mozambique  4:6.33) 

Cryplwlomimus  Browne,  1962,  West  .\frican  Timber  Bo- 
rer Research  Unit  Rept.  5:75  (Type-species:  Hy- 
pocryphahts  ghanaensis  Schedl.  original  designa- 
tion) 

Cryphcdmomimetes  Browne,  1963,  Ann.  Mag.  Nat.  Hist. 
(1.3)6:242  (Replacement  name).  Xcii  synonymy 

When  Schedl  named  Htjpocryphalus  gha- 
naensis and  then  later  (Schedl  1964b:305) 
transferred  this  species  from  Cryphalomi- 
metes  back  to  Hypocnjphalus,  he  overlooked 
some  very  important  characters.  In  this  spe- 
cies and  in  Stephanopodius,  the  basal  margin 
of  the  pronotum  bears  a  fine,  raised  line,  but 
the  lateral  margin  is  rounded  and  lacks  the 
fine,  raised  line  of  Hypocryphalus.  In  addi- 
tion, the  antennal  club  is  quite  different  from 
Hypocryphalus.  The  species  ghanaensis  is 
congeneric  with  Stephanopodius  dispar  (Egg- 
ers) and,  as  indicated  above,  should  be  trans- 
ferred to  that  genus.  Cryphalomimetes  is, 
therefore,  a  .synonym  of  Stephanopodius 
Schedl  and  not  of  Hypocryphalus  Hopkins. 

Taphrodasus,  n.  gen. 

This  genus  is  distinguished  from  Micro- 
penis Wood  by  the  confused  interstrial  pimc- 
tures, by  the  presence  of  scales  on  the  elytral 


96 


Great  Basin  Naturalist 


Vol.  40,  No.  1 


declivity,  and  by  the  strongly  concave  decliv- 
ity that  commences  on  the  basal  half  of  the 
elytra  and  is  marked  on  its  lateral  margins  in 
such  a  way  as  to  form  a  blunt,  elongate,  cir- 
cumdeclivital  costa. 

Description.—  Body  slender,  at  least  2.0 
times  as  long  as  wide,  color  reddish  brown. 
Posterior  face  of  antennal  club  with  one  su- 
ture visible,  apical  margin  of  corneous  area 
never  costate.  Scutellum  not  visible.  Strial 
punctures  on  disc  confused.  Declivity  as  de- 
scribed in  above  diagnosis. 

Type-species:  Xylehorus  percorthylus 
Schedl. 

Taurodemus,  n.  gen. 

This  genus  is  distinguished  from  Xijleborus 
Eichhoff  by  the  moderately  to  rather  widely 
separated  procoxae,  by  the  rather  stout  body, 
by  the  presence  of  9  to  12  socketed  protibial 
teeth,  and  by  the  distinctive  sculpture  of  the 
sulcate  elytral  declivity. 

Description.—  Body  stout,  less  than  1.9 
times  as  long  as  wide.  Antennal  club  with 
segment  1  corneous,  without  any  sutures  evi- 
dent on  posterior  face,  apical  margin  of  seg- 
ment 1  on  anterior  face  acutely  elevated  into 
a  continuous  costa  forming  a  complete  circle. 
Procoxae  moderately  to  rather  widely  sepa- 
rated. Protibia  armed  by  9  to  12  socketed 
teeth.  Elytral  declivity  moderately  to  very 
strongly  sulcate  on  at  least  basal  half,  lateral 
margins  armed  by  at  least  one  major  spine 
and  several  smaller  tubercles. 

Type-species:  Xijleborus  sharpi  Blandford. 

The  following  species  are  transferred  from 
Xylehorus  to  Taurodemus:  bicornutus  Wood, 
ebenus  Wood,  (Bostrichus)  flavipes  Fabricius, 
godmani  Blandford,  pandulus  Wood,  {Am- 
phic(iranus)  perebeae  Ferrari,  salvini  Bland- 
ford, sanguinicollis  Blandford,  sharpi  Bland- 
ford, splendidus  Schaufuss,  (Bostrichus) 
varians  Fabricius,  and  varus  Wood. 


Xyleborus  Eichhoff 

Xijleborus  Eichhoff,   1864,  Berliner  Ent.  Zeitschr.  8:37 
(Type-species:  Bostrichus  monographus  Fabricius, 
subsequent  designation  bv  Hopkins.    1914,  Proc. 
U.S.  Nat.  Mus.  48:131) 
The   genus   Xyleborus    Eichhoff,   as   inter- 
preted in  recent  years  by  Schedl,  contains 


more  than  1400  nominate  species,  that  is,  vir- 
tually all  the  species  in  the  tribe  Xyleborini. 
However,  the  diversity  of  characters  and 
habits  within  this  group  suggests  the  exist- 
ence of  several  distinct  clusters  of  species  and 
species  groups  that  could  and  should  be  char- 
acterized as  genera.  The  difficulty  in  frag- 
menting the  group  piecemeal,  as  has  been  at- 
tempted by  some  workers,  is  that  when  one 
group  is  removed  and  elevated  to  generic 
rank,  the  remainder  becomes  unclassifiable 
on  a  logical,  phylogenetic  basis.  In  order  to 
remedy  this  situation,  a  classification  is  being 
composed,  based  on  such  constant  features  as 
the  location  of  mycetangia,  structure  of  the 
antennal  club,  form  and  armature  of  the 
tibiae,  characters  of  the  scutellum,  and  many 
other  features.  A  deliberate  effort  is  being 
made  to  avoid  use  of  adaptive  characters 
such  as  the  surface  sculpturing  of  the  pro- 
notum  and  elytra. 

Tentatively,  27  groups  are  being  given  ge- 
neric status  within  the  Xyleborini.  Those  de- 
scribed previously  include:  Ambrosiodmus 
{  =  Browneia,  Phloeotrogus),  Arixyleborus 
{=Xyleboricus),  Cnestus  {  =  Tosaxyleborus), 
Coptoborus  {  =  Streptocranus),  Cryptoxyle- 
borus,  Dryocoetoides,  Eccoptoptcrus  {  =  Eu- 
rydactylus,  Platydactylus),  Euwallacea,  Ka- 
lantanius,  Mesoscolytus,  Notoxyleborus, 
Premnobius  {  =  Prernnophilus),  Pseudoxyle- 
borus,  Sampsonius,  Schedlia,  Theoborus, 
Webbia  {  =  Prowebbia,  Pseudowebbia,  Xely- 
borus),  Xyleborinus,  Xyleborus  {=Anaeritus, 
Anisandrus,  Boroxylon,  Bufonus,  Coptodryas, 
Cyclorhipidion,  Heteroborips,  Phloeotrogus, 
Progenius,  Terminalinus,  Xyleborips),  and 
Xylosandrus.  The  above  is  mentioned  to  es- 
tablish a  context  into  which  the  seven  genera 
in  this  tribe,  described  in  this  article,  can  fit. 
The  seven  include:  Anaxyleborus,  Apoxylc- 
borus,  Hadrodemius,  Leptoxyleborus,  Micro- 
penis, Taphrodasus,  and  Taurodemus. 

Xylechinus  Chapuis 

Xylechinus  Chapuis,  1869,  Synopsis  des  Scolvtides,  p.  36 
(Type-species:  Dcndroctontis  pilosiis  Knoch) 

Squamasinuhis  Nunberg,  1964,  Ann.  Hist. -Nat.  Mus. 
Nat.  Hungarici,  Pars  Zool.  56:431  (Type-species: 
S(iu(im(isinulus  chiliensis  Nunberg,  original  desig- 
nation). New  synonymy 

When  the  holotype  of  Squamasinuhis  chi- 
liensis   Nunberg    and    several    allied    species 


March  1980 


Wood:  American  Bark  Beetles 


97 


from  South  American  were  examined,  no 
characters  could  be  found  that  distinguish 
this  genus  from  Xylechinus  Chapuis.  As 
nearly  as  can  be  determined  at  the  present 
time,  the  genus  Xylechinus  consists  of  14 
Central  and  South  American,  2  North  Ameri- 
can, 5  Asian,  and  1  European  species.  Schedl 
has  referred  four  New  Guinean  and  Austra- 
lian species  to  this  genus,  all  of  which  appar- 
ently should  be  transferred  to  Acrantus.  One 
African  species  placed  in  Xijlechinus  by 
Schedl  apparently  belongs  elsewhere. 

Literature  Cited 

Blandford,  W.  F.  H.  1894.  The  Rhynochophorous  Co- 
leoptera  of  Japan.  Part  III.  Scolytidae.  Trans. 


Ent.  Soc.  London,  pp.  53-14L 

Eggers,  H.  1929.  Zvir  Synonymic  der  Borkeiikafer 
(Ipidae,  Col.).  Wiener  Ent.  Zeit.  46:41-55. 

Sc;hedl,  K.  E.  1940.  Zur  Einteilung  und  Synonvmie  der 
Cryphalinae  (Col.  Scoivt.).  Mitt.  Miichner  Ent. 
Ges.  .30:583-591. 

1957.  Bark-  and  timber-beetles  from  South  .Af- 
rica. Ann.  Mag.  Nat.  Hi.st.  (12)10:149-159. 

1963.  Zur  Synonymie  der  Borkenkafer  IX.  Ent. 

Abh.  Mus.  Tierk.  Dresden  28:257-268. 

1964a.  On  some  Coleoptera  of  economic  impor- 
tance from  New  Guinea  and  Australia.  Pacific  In- 
sects 6:211-214. 

1964b.  Zur  Synonymie  der  Borkenkafer  XV.  Rei- 

chenbachia  3:303-317. 

1966.  Neotropi.sche  Scolytoidea  VIII.  Ent.  Arb. 

Mus.  Frey  17:74-128. 

Wood,  S.  L.  1978.  A  reclassification  of  the  subfamihes 
and  tribes  of  Scolytidae  (Coleoptera).  Ann.  Soc. 
Ent.  France  (N.S.).'  14:95-122. 


THE  BACTERIUM  THIOPLOCA  INGRICA  ON  WET  WALLS 
IN  ZION  NATIONAL  PARK,  UTAH 

Samuel  R.  Riishforth',  Sheril  D.  Burton",  Jeffrey  R.  Johansen',  and  Judith  A.  Grimes' 

.\bstract.—  Hanging  gardens  and  wet  wall  habitats  have  been  studied  for  the  past  several  years  in  many  arid 
regions  of  the  Intermountain  West.  One  unusual  large  wet  wall  habitat  in  Zion  National  Park  was  found  to  be  cov- 
ered with  a  mucilaginous  red-colored  growth  of  the  filamentous  gliding  bacterium  Thioploca  ingrica  Visloukh.  This  is 
the  only  habitat  we  have  examined  where  the  predominant  matrix-forming  organism  was  a  bacterium  rather  than  an 
alga. 


Hanging  gardens  are  unusual  habitats 
found  in  several  areas  around  the  world,  par- 
ticularly in  western  North  America.  Such 
habitats  form  when  water  percolates  vertical- 
ly through  permeable  rock  facies  (generally 
sandstone)  imtil  it  reaches  an  impervious  lay- 
er. The  water  then  moves  laterally  imtil  it 
exits  the  rock  formation  often  on  a  vertical 
wall  or  cliff.  Such  exit  springs  often  occur 
along  fairly  long  horizontal  lines  to  form  lin- 
ear seep  walls  or  wet  walls.  Such  habitats 
rapidly  become  colonized  by  a  variety  of 
mesic  plant  species,  some  of  which  are  en- 
demic to  such  gardens.  Seep  walls  become 
weakened  through  time,  particularly  in  the 
massive  mesozoic  sandstone  formations  typi- 
cal of  areas  of  southern  Utah  and  northern 
Arizona.  When  this  occurs,  large  portions  of 
the  wall  slough  away  to  form  grottos  that  are 
.shaded  from  the  sim  and  are  cooler  and  more 
humid  than  .surrounding  areas. 

Such  hanging  garden  habitats  have  been 
imder  study  for  several  years  (Rushforth  et  al. 
1976,  Clark  and  Rushforth,  in  press,  Welsh 
and  Moore  1965a,  1965b).  We  have  been 
particularly  interested  in  the  algal  species 
that  colonize  hanging  garden  walls.  The  algal 
floras  of  such  habitats  are  variable  depending 
upon  several  factors,  primarily  the  amount  of 
water  available.  The  moister  walls  are  almost 
always  covered  with  heavy  growth  of  green 
or  blue  green  algae  that  secrete  copious 
mucilage.  Such  algal  mats  are  in  turn  colo- 


nized by  dozens  of  other  algal  species,  in- 
cluding blue  green  algae,  green  algae,  eu- 
glenophytes,  golden  algae,  and  diatoms.  Algal 
diversity  is  really  quite  high  in  many  of  these 
gardens,  and  the  species  are  often  imusual  in 
morphology  and/or  distribution. 

We  have  found  that  wet  wall  faces  that  are 
exposed  to  the  sun  are  often  colonized  by  al- 
gae with  dominant  nonchlorophylous  pig- 
ments. Thus,  on  certain  walls,  the  green  alga 
Palmella  miniata  Leiblein  is  common.  This 
organism  is  often  a  deep  red  color  due  to 
hematochrome  pigments,  and  in  turn  the  gar- 
den walls  colonized  by  these  organisms  are 
often  a  beautiful  brick-red  color.  Likewise, 
Scytonema  myochrous  (Dillw.)  C.  A.  Agardh 
and  Scytoneina  alatiim  (Carm.)  Borzi  are 
prevalent  on  some  walls.  These  organisms  are 
large,  prominently  ensheathed  blue  green  al- 
gae. The  sheath  and  the  cell  wall  of  these  al- 
gae become  colored  as  the  organisms  mature 
to  form  yellow  or  yellow  brown  filaments 
which  also  color  their  walls  of  colonization. 

Recently,  we  have  been  studying  algae  and 
lichens  of  Zion  National  Park.  One  promi- 
nent wet  wall  in  the  park  is  Weeping  Rock 
(Fig.  1),  a  famous  tourist  attraction.  This  wall 
is  very  large  and  is  usually  very  moist.  It  is 
heavily  colonized  by  many  algal  species.  In 
several  places  on  the  wall,  bright  orange  red 
patches  and  streaks  are  evident.  We  collected 
specimens  from  these  areas  expecting  to  find 
Palmella  miniata.  Upon  returning  to  the  lab- 


'Department  of  Botany  and  Range  Science,  Brigham  Young  University,  Provo,  Utah  84602. 
■Department  of  Microbiology,  Brigham  Young  University,  Provo,  Utah  84602. 


98 


March  1980 


RUSHFORTH    ET  AL.:  HANGING  GaRDENS  BaCTERIUM 


99 


Fig.  1-4.  1.  Habit  of  Tliioplora  and  algal  communities  of  Weeping  Rock,  Zion  National  Park;  2,  "Braid"  of  Thio- 
ploca  im^rica  filaments  in  a  common  sheath;  .3,  Thioplocd  with  Chamae.siphon  species  attached;  4,  Low  magnification 
photograph  showing  heavy  epiphytic  growth  of  Chamaesiphon  species. 


100 


Great  Basin  Naturalist 


Vol.  40,  No.  1 


oratory,  we  examined  the  organisms  using  a 
Zeiss  RA  Research  microscope  with  No- 
marski  differential  phase  contrast  accessories. 
We  were  surprised  to  find  that  'the  red  color- 
ation was  not  due  to  Palmella  but  to  a  fila- 
mentous prokaryote.  Careful  study  elimi- 
nated known  blue  green  algae  as  the  causal 
organism.  Further  examination  showed  the 
organism  to  be  a  bacterium  in  the  family 
Beggiatoaceae  Migula.  These  bacteria  are 
gram-negative  filaments  that  are  motile  by 
gliding  motion.  Three  genera  are  presently 
placed  in  this  family:  Beggiatoa  Trevisan, 
Vitreoscilh  Pringsheim,  and  Thioploca  Lau- 
terbom.  In  addition,  the  three  genera  Bactos- 
cilla  Pringsheim,  Flexoscilla  Pringsheim,  and 
Thiospirillopsis  Uphof  are  possible  members 
of  the  family  (Leadbetter  1974). 

The  organisms  we  collected  from  Weeping 
Rock  in  Zion  National  Park  may  be  placed  in 
the  genus  Thioploca,  based  upon  the  presence 
of  more  than  one  filament  in  the  sheaths 
(Figs.  2-4).  Furthermore,  the  filaments 
ranged  between  the  width  of  2  and  7  um, 
placing  them  in  Thioploca  ingrica  Visloukh. 
The  filaments  often  served  as  a  substrate  for 
epiphytic  algae,  particularly  Cluimaesiphon 
species  (Fig.  3). 

This  observation  of  T.  ingrica  is  interesting 
for  several  reasons.  First,  it  represents  the 
only  example  in  the  several  gardens  we  have 
studied  where  the  predominant  mucilaginous, 
matrix-forming  organism  was  a  bacterium 
rather  than  an  alga.  Second,  it  represents  the 
only  reported  occurrence  of  Thioploca  from 
intermountain  western  North  America  that 
we  are  aware  of.  Third,  Thioploca  usually  ex- 
hibits a  greenish  blue  color.  However,  the 
specimens  we  have  collected  produce  a 
bright  orange  red  color  on  the  wall  itself  and 
a  paler  orange  color  when  examined  beneath 


the  microscope.  The  nature  of  this  color  is 
unknown,  although  Beggiatoaceae  are  report- 
ed to  not  form  carotenoid  pigments.  And 
fourth,  the  presence  of  this  organism  on  the 
moist  walls  is  itself  unusual  because  all  the 
reports  to  date  that  we  are  aware  of  chron- 
icle Thioploca  species  as  inhabiting  the  upper 
layers  of  bottom  muds  of  freshwater  and 
brackish  habitats  (Maier  1974).  Specifically, 
such  organisms  have  been  collected  from 
both  oxidizing  and  reducing  environments  in 
such  muds  (Perfil'ev  et  al.  1965). 

Literature  Cited 

Clark,  R.  L.,  and  S.  R.  Rushforth.  In  review.  A  study 
of  the  algal  flora  of  selected  hanging  gardens  of 
Glen  Canyon,  Utah.  Great  Basin  Naturalist  Mem- 
oirs. 

Leadbetter,  E.  R.  1974.  Beggiatoaceae  Migula.  Pages 
112-115  in  R.  E.  Buchanan,  N.  E.  Gibbons,  eds. 
Bergey's  manual  of  determinative  bacteriology, 
8th  ed.  Williams  and  Wilkins,  Baltimore. 

Maier,  S.  1974.  Thioploca  Lauterborn.  Pages  113-116  in 
R.  E.  Buchanan,  N.  E.  Gibbons,  eds.  Bergey's 
manual  of  determinative  bacteriology,  8th  ed. 
Williams  and  Wilkins,  Baltimore. 

Perfil'ev,  B.  V.,  D.  R.  Gabe,  A.  M.  Galperina,  V.  A. 
Rabinovich,  a.  a.  Sapotnitskii,  E.  E.  Sherman, 
AND  E.  P.  Troshanov.  1964.  Applied  capillary 
microscopy:  The  role  of  microorganisms  in  the 
formation  of  iron-manganese  deposits.  Izadatels- 
tov,  Akad  Nauk  SSSR,  Savarenskii  Laboratory  for 
Hydrogeological  Problems.  Moscow.  In  Russian 
(Transl.  Consultants  Bureau  Enterprise  Inc.  1965 
New  York). 

Rushforth,  S.  R.,  L.  L.  St.  Clair,  T.  A.  Leslie,  K.  H. 
Thorne,  and  D.  S.  Anderson.  1976.  The  algal 
flora  of  two  hanging  gardens  in  southeastern 
Utah.  Nova  Hedwigia  27:231-323. 

Welsh,  S.  L.,  and  G.  Moore.  1965a.  Plants  of  Canvon- 
lands  National  Park.  Part  I.  The  Needles  Region. 
Proc.  Utah  Acad.  Sci.  42(1):  160. 

Welsh,  S.  L.,  and  G.  Moore.  1965li.  Plants  of  Canyon- 
lands  National  Park.  Part  II.  The  Islands  in  the 
Sky  Region.  Proc.  Utah  Acad.  Sci.  42(1):160-161. 


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TABLE  OF  CONTENTS 

Some  aspects  of  succession  in  the  spruce-fir  forest  zone  of  northern  Utah.  David  J. 

Schimpf,  Jan  A.  Henderson,  and  James  A.  MacMahon  1 

Utah  Flora:  Malvaceae.  Stanley  L.  Welsh  27 

Utah  Flora:  Miscellaneous  families.  Stanley  L.  Welsh  38 

The  taxonomic  status  of  the  rosy  boa  Lichamira  roseofusca  (Serpentes:  Boidae).  John 

R.  Ottley,  Robert  W.  Murphy,  and  Geoffrey  V.  Smith  59 

Hespewperla  lio^uei,  a  new  species  of  stonefly  from  California  (Plecoptera:  Perlidae). 

Richard  W.  Baumann  and  Bill  P.  Stark  63 

Reproduction  in  three  sympatric  lizard  species  from  west-central   Utah.  John  B. 

Andre  and  James  A.  MacMahon  68 

Haplopappus  alpinus  (Asteraceae):  A  new  species  from  Nevada.  Loran  C.  Anderson  ..         73 

Miscellaneous  plant  novelties  from  Alaska,  Nevada,  and  Utah.  Stanley  L.  Welsh  and 

Sherel  Goodrich  78 

New  genera  and  new  generic  synonvmv  in   Scolytidae   (Coleoptera).   Stephen   L. 

Wood [ '.....'. '. 89 

The  bacterium  Thioplova  ingrica  on  wet  walls  in  Zion  National  Park,  Utah.  Samuel 

R.  Rushforth,  Sheril  D.  Burton,  Jeffrey  R.  Johansen,  and  Judith  A.  Grimes  98 


HE  GREAT  BASIN  NATURALIST 


ume  40  No.  2 


June  30,  1980 


Brigham  Young  University 


MUS.  COMP.  ZOOU, 

NOV 


GREAT  BASIN  NATURALIST 

Editor.  Stephen  L.  Wood,  Department  of  Zoology,  Brigham  Young  University,  Provo,  Utah 

84602. 
Editorial  Board.   Kimball  T.  Harper,  Botany;  Wilmer  W.  Tanner,   Life   Science  Museum; 

Stanley  L.  Welsh,  Botany;  Clayton  M.  White,  Zoology. 
Ex  Officio  Editorial  Board  Members.  A.  Lester  Allen,  Dean,  College  of  Biological  and  Agricul- 
tural Sciences;  Ernest  L.  Olson,  Director,  Brigham  Young  University  Press,  University 
Editor. 

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cover. 


5-80  650  45971 


ISSN  0017-3614 


The  Great  Basin  Naturalist 


Published  at  Provo,  Utah,  by 
Brigham  Young  University 

ISSN  0017-3614 


Volume  40 


June  30,  1980 


No.  2 


FEEDING  ECOLOGY  OF  GILA  BORAXOBIUS  (OSTEICHTHYES:  CYPRINIDAE) 
ENDEMIC  TO  A  THERMAL  LAKE  IN  SOUTHEASTERN  OREGON' 

Jack  E.  Williams'  and  Cynthia  D.  W'illianis' 


.\bstr.\ct.—  (w/a  hoiaxchiit.s  is  a  dwarf  species  of  cvprinid  endemic  to  a  thermal  lake  in  southeastern  Oregon. 
Despite  a  relatively  depauperate  fauna  and  flora  in  the  lake,  24  food  items  were  foimd  in  intestines  of  G.  horaxobins. 
Ten  of  the  24  foods,  including  six  insects,  were  of  terrestrial  origin.  The  relative  importance  of  food  items  fliictuated 
seasonally.  Diatoms,  chironomid  larvae,  microcrustaceans,  and  dipteran  adults  were  the  primary  foods  during  spring. 
In  summer,  diatoms  decreased  and  terrestrial  insects  increased  in  importance.  During  autumn  important  foods  were 
terrestrial  insects,  chironomid  larvae,  and  diatoms.  Diatoms  and  microcrustaceans  increased  in  importance  during 
winter.  Chironomid  larvae  were  of  importance  in  winter,  when  the  importance  of  terrestrial  food  items  decreased 
.substantially.  Similar  food  habits  were  observed  between  juveniles  and  adults,  except  that  adults  consumed  more 
gastropods  and  diatoms  and  juveniles  consumed  more  copepods  and  terrestrial  insects.  Gila  howxobius  feeds  op- 
portunistically with  individuals  commonly  containing  mostly  one  food  item.  Fish  typically  feed  by  picking  foods 
from  soft  bottom  sediments  or  from  rocks.  However,  fish  will  feed  throughout  the  water  column  or  on  the  surface  if 
food  is  abundant  there.  Gila  boraxohiiis  feeds  throughout  the  day,  with  a  peak  in  feeding  activity  just  after  sunset.  .\ 
daily  ration  of  H.l  percent  boch'  weight  was  calculated  for  the  species  during  June.  .\  comparison  of  food  habits 
among  G.  boraxobius  and  populations  of  G.  alvordensis  during  the  summer  shows  that  all  are  opportimistic  in  feed- 
ing, but  that  G.  boraxobius  relies  more  heavily  on  terrestrial  foods. 


The  Borax  Lake  chub,  Gilo  boraxobius,  is  a 
rare  species  of  cvprinid  fish  endemic  to  a 
thermal  lake  in  the  Alvord  Basin  of  south- 
eastern Oregon.  The  restricted  habitat  of  G. 
boraxobius  is  threatened  bv  geothermal 
energy  development  in  the  Alvord.  In  recog- 
nition of  this  problem,  the  species  is  listed  as 
"threatened"  by  the  Endangered  Species 
Committee  of  the  American  Fi.sheries  Society 
(Deacon  et  al.  1979)  and  is  currently  on  the 
protected  list  of  the  Oregon  Department  of 
Fish  and  Wildlife. 

Gila  boraxobius  was  recently  described  and 
has  been  diagnosed  as  a  dwarf  relative  of  G. 
alvordensis  (Williams  and  Bond,  in  press). 
Typical  adult  size  of  Gila  boraxobius  is  33-50 
mm  standard  length  (SL).  Individuals  longer 
than  55  mm  SL  are  rare. 

Becau.se   no   life   history   information    was 


known  concerning  this  species,  a  study  was 
conducted  to  determine  its  food  habits  rela- 
tive to  sea.sons,  fish  length,  diel  feeding  chro- 
nology, and  food  habits  of  other  Alvord  Basin 
fishes  of  the  genus  Gila. 

Methods 

Feeding  ecology  of  Gila  boraxobius  was  in- 
vestigated by  analyzing  intestinal  contents  of 
fish  collected  monthly  from  March  1978  to 
June  1979.  Fish  were  collected  from  the 
southwest  one-quarter  of  Borax  Lake  using  a 
3  X  5  mm  me.sh  seine  approximately  3  m  in 
length.  Specimens  were  preserved  in  10  per- 
cent formalin  and  transferred  to  45  percent 
isopropanol  after  one  week.  Standard  length 
of  specimens  was  measured  to  the  nearest  0.1 
mm  with  dial  calipers.  After  blotting  fish  dry 
on  paper  towels,  wet  weight  was  measured  to 


'Technical  Paper  5335,  Oregon  .Agricultural  Experiment  Station. 

■Department  of  Fisheries  and  Wildlife.  Oregon  State  University.  Corvallis,  Oregon  97331. 


101 


102 


Great  Basin  Naturalist 


Vol.  40,  No.  2 


the  nearest  0.01  g.  Gila  boraxohius,  like  other 
cvprinids,  has  no  stomach;  therefore,  contents 
of  the  intestine,  from  esophagus  to  anus,  were 
removed  and  examined  under  dissecting  mi- 
croscopes. Percent  frequency  of  occurrence, 
mean  number  per  intestine,  mean  percent 
volume,  and  a  value  of  relative  importance 
are  reported  for  each  food  item.  Percent  fre- 
quency of  occurrence  is  defined  as  the  num- 
ber of  intestine  samples  in  which  one  or  more 
of  a  given  food  item  is  found  expressed  as  a 
percentage  of  all  nonemptv  intestines  exam- 
ined (Windell  and  Bo  wen  1978).  The  total 
number  of  a  given  food  item  observed  in  the 
intestines  divided  by  the  number  of  nonemp- 
tv intestines  examined  is  the  mean  number 
per  intestine.  Mean  percent  volume  is  de- 
fined as  the  total  volume  estimates  for  a  giv- 
en food  item  divided  by  the  number  of  non- 
empty intestines  examined.  Percent  volumes 
were  derived  by  separating  the  intestine  into 
three  subsamples  and  visually  estimating  the 
percent  contribution  of  a  given  food  item  in 
each  sample.  The  percent  contribution  of 
each  subsample  to  the  contents  of  the  entire 
intestine  was  estimated  so  that  the  volume  of 
a  <£iven  food  item  relative  to  all  intestinal 
contents  could  be  made.  Percent  frequency 
of  occurrence,  mean  number  per  intestine, 
and  mean  percent  volume  each  contain  a  bias 
which  limits  their  usefulness  when  used  sepa- 
rately (Windell  and  Bowen  1978).  For  ex- 
ample, percent  frequency  of  occurrence 
overemphasizes  the  importance  of  small  food 
items  that  may  be  ingested  frequently  but 
have  a  small  impact  on  the  volume  of  food  in 
the  intestine.  On  the  other  hand,  mean  per- 
cent volume  overemphasizes  the  importance 
of  large  food  items,  such  as  adult  insects,  that 
may  be  consumed  infrequently  but  have  a 
large  volume.  To  offset  these  biases  against 
each  other,  an  index  of  relative  importance 
(RI)  is  reported  for  each  food  item.  The  rela- 
tive importance  index  combines  the  percent 
frequency  of  occurrence  and  mean  percent 
volume  for  food  item  a  into  an  absolute  im- 
portance index  (AI„)  as  follows: 


AT    =       P^''<-'(-'"t  frequency  of  occurrence  + 
mean  percent  volume 

RI„  =  100AI„/SAI„,  where  n  is  the  number 


of  different  food  items.  The  determination  of 
Rio  and  AI^  are  by  methods  modified  from 
George  and  Hadley  (1979). 

Invertebrate  identification  was  based  on 
Pennak  (1978). 

Food  habits  of  G.  boraxohius  are  summa- 
rized as  follows:  (1)  seasonal  variation  of 
foods  consumed,  (2)  foods  consumed  by  dif- 
ferent size  classes  of  fish,  (3)  diel  feeding 
chronology,  and  (4)  comparison  with  food 
habits  of  two  populations  of  Gila  alvordcnsis. 
Changes  in  food  habits  with  season  and  fish 
size  class  were  determined  from  monthly  col- 
lections taken  from  March  1978  through  Jan- 
uary 1979.  Monthly  collections  were  grouped 
into  seasons  as  follows:  spring  (March-May), 
summer  (June-August),  autumn  (Septem- 
ber-November) and  winter  (Decem- 
ber-January). To  compare  changes  in  food 
habits  with  fish  size,  two  size  classes  of  fish 
were  compared,  15.0  mm  to  32.9  mm  SL 
(juveniles),  and  33.0  mm  SL  or  longer 
(adults).  Gila  boraxobius  matures  at  approx- 
imately 33.0  mm  SL  (Williams  and  Bond,  in 
press).  Diel  feeding  chronology  was  deter- 
mined from  collections  made  at  3-hour  inter- 
vals during  a  24-hour  period  in  June  1979. 
Fullness  of  the  intestine  was  determined  ac- 
cording to  the  formula  defined  by  Hureau 
(1969  in  Berg  1979)  as  follows: 

_    weight  of  ingested  food  X  100  percent 
weight  of  fish 

(Ir  =  L'indice  de  repletion  =  fullness  index). 
The  daily  ration  of  food  for  G.  boraxobius 
was  estimated  from  diel  trajectories  of  the 
fullness  of  the  intestine.  This  estimate  was  de- 
rived by  the  following  formula  proposed  by 
Bajkov  (1935)  and  modified  by  Darnell  and 
Meierotto  (1962)  and  Eggers  (1977): 

Rt  =  24Sa 

where  Rj  is  the  daily  ration,  S  is  the  average 
weight  of  intestinal  contents  expressed  as 
percent  of  body  weight  during  the  24-hour 
period,  and  a  is  the  intestinal  evacuation 
rate.  An  intestinal  evacuation  rate  of  0.2  hr' 
is  assumed  (Eggers  1977). 

Intestinal  contents  of  Gila  alvordcnsis  col- 
lected from  Thousand  Creek,  Nevada,  on  13 
June  1978  and  G.  alvordensis  collected  from 
Serrano  Pond,  Oregon,  on  6  August  1977 
were  compared  to  those  of  G.  boraxobius  col- 


June  1980 


Williams,  Williams:  Borax  Lake  Chub 


103 


lected  during  the  summer  of  1978.  Methods 
of  collection  and  food  habits  analysis  for  pop- 
ulations of  G.  alvordensis  were  the  same  as 
those  used  for  G.  boraxobius. 

Study  Area 

Fish  were  collected  from  Borax  (  =  Hot) 
Lake,  Serrano  Pond,  and  Thousand  Creek,  all 
in  the  Alvord  Basin  of  southeastern  Oregon 
and  northwestern  Nevada.  The  Alvord  is  an 
endorheic  basin  of  semiarid  climate,  sur- 
rounded by  fault-block  mountain  ranges.  Bo- 
rax Lake'(T37S,  R33E,  Sec.  14,  Harney 
County,  Oregon)  is  a  4.1  ha  natural  lake  that 
receives  water  from  several  thermal  springs 
in  the  lake  bottom.  Water  temperature  in 
Borax  Lake  is  typically  29-32  C,  with  ex- 
tremes of  35  C  or  greater  near  spring  sources 
and  17  C  near  the  lake  margin  during  winter. 
The  lake  is  relatively  shallow,  with  a  soft,  sil- 
tv  bottom  interspersed  with  rocks  and  hard 
outcroppings.  The  water  is  transparent,  and 
aquatic  vegetation  is  limited  to  a  few  areas 
along  the  lake  margin.  Sodium  is  the  princi- 
pal cation,  whereas  bicarbonate,  sulfate,  and 
chloride  are  the  major  anions  in  Borax  Lake 
(Mariner  et  al.  1974).  Specific  conductance  of 
the  water  is  2410.  Serrano  Pond  (T36S, 
R33E,  Sec.  1,  Harney  County,  Oregon)  is  a 
0.1  ha  reservoir  that  receives  water  from  a 
cool  spring  approximately  60  m  distant.  Wa- 
ter temperature  is  usually  16-21  C  during  the 
summer.  The  bottom  of  this  shallow  pond  is 
primarily  silt.  The  water  is  turbid,  and  aquat- 
ic vegetation  is  abundant.  Thousand  Creek 
(collection  site  at  T46N,  R28E,  Sec.  34, 
Humboldt  County,  Nevada)  is  a  small,  shal- 
low stream  rarely  exceeding  2  m  in  width. 
The  creek  often  becomes  intermittent  during 
summer  months.  Water  temperature  varies 
from  15-27  C  during  the  summer,  fluctuating 
rather  closely  with  air  temperature.  The  bot- 
tom is  a  silt  and  gravel  mix.  The  water  is  tur- 
bid, and  aquatic  vegetation  is  moderately 
abundant. 

Results 

Seasonal  Variation  in  Foods  Consumed 

Twenty-four  different  food  items  were 
found  in  intestines  of  Gila  boraxobius  during 


this  study  (Tables  1-4).  Ten  of  the  24  food 
items  were  encountered  in  all  seasons.  Many 
food  items  fluctuated  seasonally  in  occur- 
rence; however,  some  insects,  especially  chi- 
ronomid  larvae,  diatoms,  and  micro- 
crustaceans,  were  of  importance  throughout 
the  year.  During  the  spring,  algae,  chirono- 
mid  larvae,  copepods,  dipteran  adults,  and 
ostracods  were  the  predominant  food  items 
(Table  1).  Algae,  which  was  composed  almost 
wholly  of  diatoms,  was  the  most  frequently 
ingested  food  during  the  spring,  occurring  in 
over  one-half  of  the  intestines.  Some  of  the 
diatoms  were  secondarily  ingested  with  mi- 
crocmstaceans;  however,  in  some  individuals 
diatoms  accounted  for  70-80  percent  of  the 
intestinal  contents  by  volume.  The  high  vol- 
ume suggests  that  diatoms  are  not  exclusively 
the  result  of  secondary  ingestion  but  are  a 
preferred  food  item  for  most  fish.  The  most 
common  diatom  observed  in  intestines  was  a 
benthic  species,  Denticida  thermalis.  Nov- 
iciila  sp.,  Synedra  sp.  Achnanthes  lanceolata, 
and  A.  minutissima  were  observed  in  lesser 
numbers.  Both  Achnanthes  species  are  ben- 
thic, whereas  the  Navicula  and  Synedra  spe- 
cies could  be  benthic  or  planktonic  forms. 
Adult  dipterans  accounted  for  the  highest 
mean  percent  volume,  nearly  20  percent,  of 
all  food  items  during  spring.  Several  fish  fed 
on  dipteran  adults  exclusive  of  other  foods. 
During  May  1978  dipteran  adults  were  heav- 
ily utilized,  appearing  in  19  of  23  intestines 
examined.  During  the  summer,  diatoms  were 
less  frequently  encountered  in  intestines  and 
comprised  a  smaller  mean  percent  volume 
than  in  spring.  Chironomid  larvae,  copepods, 
dipteran  adults,  and  gastropods  were  the 
most  important  food  items  in  summer  (Table 
2).  Gila  boraxobius  utilized  more  terrestrial 
insects  and  spiders  and  fewer  micro- 
crustaceans  and  diatoms  in  summer  than  in 
spring.  Terrestrial  insects  and  spiders  ac- 
counted for  approximately  31  percent  mean 
volume  of  foods  consumed  during  summer 
compared  to  approximately  21  percent  in 
spring.  During  autumn  terrestrial  insects,  chi- 
ronomid larvae,  and  diatoms  were  the  princi- 
pal food  items  (Table  3).  In  winter  Gila 
boraxobius  relied  more  heavily  on  autoch- 
thonous food  items,  utilizing  primarily  dia- 
toms, ostracods,  copepods,  chironomid  larvae 
and  cladocerans  (Table  4).  Terrestrial  insects. 


104 


Great  Basin  Naturalist 


Vol.  40,  No.  2 


which  were  of  importance  in  spring,  summer, 
and  autumn,  seldom  appeared  in  intestines  of 
G.  boraxobius  during  winter,  when  they  con- 
tributed only  2  percent  mean  volume.  Food 
items  consumed  in  autumn  and  winter  were 
less  diverse  than  in  other  seasons.  Sixteen 
food  items  were  observed  in  fish  collected  in 
autumn  and  winter,  20  food  items  during 
spring,  and  21  during  summer.  Aquatic  in- 
sects were  important  foods  throughout  the 
year,  comprising  mean  volumes  of  approx- 
imately 19  percent,  23  percent,  16  percent, 
and  13  percent  in  spring,  summer,  autumn, 
and  winter,  respectively.  The  primary  con- 
tributor to  these  high  values  were  chirono- 
mid  larvae,  which  consistently  exhibited  a 
high  relative  importance.  Chironomid  pupae 
and  Odonata  nymphs  were  also  consumed 
throughout  the  year  but  were  of  much  less 


importance.  Coleoptera  larvae  and  aquatic 
Coleoptera  adults  were  utilized  to  lesser  de- 
grees seasonally.  The  increased  consumption 
of  copepods,  ostracods,  and  cladocerans  in 
the  winter  was  dramatic.  These  micro- 
CRistaceans  comprised  approximately  35  per- 
cent mean  volume  of  intestines  during  win- 
ter, but  only  16.5  percent,  12  percent  and  4.5 
percent  in  spring,  summer,  and  autumn,  re- 
spectively. Large  amounts  of  inorganic  debris 
were  found  in  intestines  throughout  the  year. 
Tliis  was  probably  ingested  accidently  while 
the  fish  were  feeding  on  bottom  organisms. 
Many  important  foods  in  Borax  Lake,  such  as 
insect  larvae,  gastropods,  diatoms,  and  prob- 
ably many  small  invertebrates,  are  benthic. 
Observations  in  Borax  Lake  and  in  aquaria 
show  that  G.  boraxobius  feeds  primarily  by 
rooting  around  in  bottom  material  and  pick- 


Table  1.  Contents  of  71  intestines  oi  Cihi  boraxobius  collected  during  the  spring  of  1978.  ND  =  no  data. 


Item  ingested 


Percent 

frequency 

of  occurrence 


Mean  number 
per  intestine 


Mean  percent 
volume 


RI 


.\lgae' 

C'.astropods- 

C.astropod  eggs 

Haplotaxid  oligochaetes 

Harpacticoid  copepods 

Ostracods 

Cladocerans 

Plant  seeds 

Higher  plants 

Fish  scales 

Araneae 

Insect  eggs 

Unidentified  insects' 

Tekrestrial  i\sec;ts 
CoUemhola 
Thysanoptera  adults 
Hemiptera  adults 
Coleoptera  adults 
Hymcnoptera  adults 
Diptera  adults 

Asiatic:  insects 
Chironomid  larvae 
C;hironomid  pupae 
Odonata  nvmphs 
Klmid  larvae 
C'oleoptera  adults 

In()Iu;amc:  dehhis 


56.14 

ND 

12.67 

15.70 

22.81 

1.07 

6.69 

6.73 

0.00 

0.00 

0.00 

().(M) 

1.75 

0.11 

0.32 

0.47 

49.12 

44.95 

10.38 

13..57 

40.35 

8.39 

5.91 

10..55 

15.79 

0.21 

0.17 

3.64 

5.26 

0.11 

0.06 

1.21 

12.28 

ND 

1.72 

3.19 

3.51 

0.11 

0.04 

0.81 

3.51 

0.04 

0.34 

0.88 

10.53 

12.26 

2.15 

2.89 

26.32 

ND 

2.50 

6..57 

0.00 

().(H) 

0.(X) 

0.00 

3.51 

0.04 

0..39 

0.89 

0.00 

0.00 

0.00 

0.(K) 

1.75 

0.02 

0.21 

0.45 

3.51 

0.04 

0.49 

0.91 

35.09 

1.93 

19.49 

12.45 

47.37 

12.30 

13.71 

13.93 

7.02 

0.14 

2.32 

2.13 

3.51 

0.04 

0.91 

1.01 

7.02 

0,39 

1.76 

2.(X) 

0.00 

{).()() 

().(K) 

0.(X) 

61.40 

ND 

17.48 

- 

Total 


99.71 


'Mostly  diatoms 

'Mostly  PUinorhulla,  rarely  Phy.m 

'Mostly  terrestrial  forms 


June  1980 


Williams,  Williams:  Borax  Lake  Chub 


105 


ing  up  food  items.  However,  if  Ijenthic  food 
items  are  scarce,  or  if  other  foods  are  abun- 
dant, C.  homxohius  will  readilv  feed  on  ma- 
terials drifting  through  water  column  or  on 
the  surface.  Thus,  during  the  summer  some 
G.  boraxohius  readilv  switched  to  ingestion  of 
terrestrial  invertebrates.  This  resulted  in  the 
lowest  mean  percent  volume  of  inorganic 
debris  ingested  for  any  season.  The  shoreline 
of  Borax  Lake  provides  habitat  for  many  ter- 
restrial invertebrates  that  can  enter  Borax 
Lake.  Terrestrial  invertebrates  are  scarce 
during  winter,  reducing  the  likelihood  of 
their  being  a  primary  food  source  at  that 
time. 

Gila  boraxohius  is  often  a  highly  exploitive 
omnivore,  feeding  almost  entirelv  on  one 
food  source.  For  example,  examination  of  in- 
testines of  fish  collected  during  May  1978 
disclosed  the  following  (percent  volumes  of 

Tahle  2.  C'ontents  of  70  intestines  of  Cila  boraxob'uts  colh 


the  food  item  are  given  in  parentheses):  one 
individual  contained  32  gastropods  (84  per- 
cent volume),  a  second  fish  contained  14 
adult  dipterans  (98  percent  volume),  a  third 
contained  775  copepods  (79  percent  volume), 
a  fourth  contained  340  first  instar  chironomid 
larvae  (69  percent  volume),  and  a  fifth  con- 
tained 485  insect  eggs  (64  percent  volume)! 
Although  the  preceding  is  somewhat  unusual, 
many  fish  were  found  with  one  food  item 
dominating  their  intestinal  contents. 

Foods  Consumed  by 
Different  Size  Classes  of  Fish 

To  study  the  effect  of  fish  size  and  age  on 
foods  consumed,  we  compared  intestinal  con- 
tents of  juvenile  and  adult  Gila  boraxohius. 
Overall  food  habits  of  juveniles  and  adults 
were  similar.  Both  consumed  large  amounts 


:ted  di 


tile  summer  of  1978.  ND  =  no  data. 


Item  ingested 


Percent 

frequency 

of  occurrence 


Mean  number 
per  intestine 


Mean  percent 
volinne 


RI 


■\lgae' 
Gastropods- 
Gastropod  eggs 
Haplotaxid  oligochaetes 
Harpacticoid  copepods 
Ostracods 
Cladocerans 
Plant  seeds 
Higher  plants 
Fish  scales 
Araneae 
Insect  eggs 
Unidentified  insects' 

Terrestrial  insects 
Collembola 
Thvsanoptera  adults 
Hemiptera  adults 
Coleoptera  adults 
H\  nienoptera  adults 
Diptera  adults 

Asiatic:  insects 
C!hironoinid  larvae 
(Ihironomid  pupae 
Odonata  nvmphs 
Elmid  larvae 
Coleoptera  adults 

Inorganic  debris 


31.15 

ND 

6.42 

7.42 

32.79 

1.07 

9.04 

8.26 

1.64 

0.66 

1.56 

0.63 

1.64 

0.08 

0..30 

0.38 

47.54 

33.30 

8.45 

11.06 

32.79 

2.87 

3.03 

7.08 

8.20 

0.38 

0.31 

1.68 

13.11 

0.25 

0.49 

2.69 

0.00 

0.00 

0.00 

0.(X) 

().(X) 

0.00 

0.00 

0.(X) 

14.75 

0.25 

3.21 

3..55 

3.28 

0.69 

0.15 

0.68 

37.70 

ND 

4.50 

8.34 

6.56 

0.16 

0.23 

1.34 

9.84 

0.13 

0.45 

2.03 

0.00 

0.00 

0.00 

0.00 

26.23 

0.70 

5.94 

6.36 

18.03 

0.21 

2.27 

4.01 

39.34 

1.62 

14.31 

10.60 

.57.38 

4.10 

13.79 

14.06 

9.84 

0.11 

1.12 

2.17 

1.64 

0.03 

0.49 

0.42 

11.48 

0.23 

0.85 

2.44 

18.03 

0.31 

6.29 

4.80 

.54.10 

ND 

15..30 

- 

Total 


98.50 


'Mostly  diatoms 

'Mostly  Planorhulla.  rarely  Phy.sa 

'Mostly  terrestrial  forms 


106 


Great  Basin  Naturalist 


Vol.  40,  No.  2 


of  terrestrial  insects  in  spring,  summer,  and 
autumn,  and  algae  (mostly  diatoms)  and  mi- 
crocm-staceans  in  winter  (Fig.  1).  Aquatic  in- 
sects, primarily  chironomid  larvae,  were  im- 
portant to  juveniles  and  adults  throughout 
the  year.  Despite  the  overall  similarity  of 
food  habits  between  juveniles  and  adults, 
some  differences  were  noted.  Elmid  (Coleop- 
tera)  larvae  were  consumed  almost  exclusi- 
vely by  adults.  Elmid  larvae  were  found  in 
intestines  of  15  adults  but  only  one  juvenile. 
More  terrestrial  insects  were  consumed  by 
juveniles  than  adults  except  during  winter, 
when  terrestrial  insects  were  relatively  unim- 
portant to  both  groups.  The  relatively  large 
size  of  many  terrestrial  insects,  such  as  the 
commonly  consumed  muscoid  fly  adults,  did 
not  deter  their  ingestion  by  juvenile  Gila 
horaxohius.    Adults   consumed   more   gastro- 


pods than  did  juveniles  during  all  seasons.  In- 
testines of  adults  averaged  8.5  percent  mean 
volume  of  gastropods  during  the  year,  where- 
as intestines  of  juveniles  averaged  2.3  per- 
cent. Adults  also  consumed  more  diatoms 
than  did  juveniles.  Increased  relative  con- 
sumption of  diatoms  by  adults  was  primarily 
evident  in  summer  and  autumn,  when  adults 
consumed  9.3  percent  and  6.9  percent  mean 
volume  of  diatoms,  respectively;  however, 
juveniles  consumed  0.1  percent  and  1.3  per- 
cent mean  volume,  respectively.  The  small 
volume  of  gastropods  and  diatoms  ingested 
by  juveniles  was  compensated  for  by  inge- 
stion of  large  numbers  of  copepods.  Intestines 
of  juveniles  averaged  13.4  percent  mean  vol- 
ume of  copepods  during  the  year,  whereas 
adults  averaged  4.2  percent  mean  volume  of 
copepods. 


Table  3.  Contents  of  57  intestines  of  C.iUi  horaxohius  collected  during  the  autumn  of  1978.  ND  =  no  data. 


Item  ingested 


Percent 

frequency 

of  occurrence 


Mean  number 
per  intestine 


Mean  percent 
volume 


RI 


Algae' 
Gastropods- 
Gastropod  eggs 
Haplotaxid  oligochaetes 
Harpacticoid  copepods 
Ostracods 
Cladocerans 
Plant  seeds 
Higher  plants 
Fish  scales 
Araneae 
Insect  eggs 
Unidentified  insects' 

Terrestrial  insects 
Collembola 
Thysanoptera  adults 
Hemiptcra  adults 
Coleoptera  adults 
Hymenoptera  adults 
Diptera  adults 

AgUATIC  INSECTS 

Chironomid  larvae 
Chironomid  pupae 
Odonata  nymphs 
Elmid  larvae 
Coleoptera  adults 

I.NORGANIC  DEBRIS 


50.00 

ND 

5.03 

12.63 

10.87 

0.50 

4.04 

3.42 

0.00 

0.00 

0.00 

0.00 

0.00 

0.00 

0.00 

0.00 

34.78 

25.46 

4.22 

8.95 

10.87 

0.50 

0.15 

2.53 

0.00 

0.00 

0.00 

0.00 

39.13 

1,57 

0..59 

9.12 

2.17 

ND 

0.07 

0.51 

0.00 

0.00 

0.00 

0.00 

0.00 

0.00 

0.00 

0.00 

2.17 

0.04 

0.04 

0.51 

67.39 

ND 

19.54 

19.95 

0.00 

0.00 

0.00 

0.00 

21.74 

0.46 

1.69 

5.38 

13.04 

0.22 

2.55 

3.58 

4.35 

0.04 

0.22 

1.05 

8.70 

0.11 

l.,54 

2.35 

19.56 

0.22 

3..52 

5.30 

71.74 

5.43 

12.00 

19.22 

13.04 

0.65 

3.15 

3.72 

6.52 

0.07 

1.29 

1.79 

0.00 

0.(K) 

0.(X) 

0.00 

().(X) 

0.(X) 

0.(K) 

0.00 

86.96 

ND 

40.34 

- 

Total 


99.94 


'Mostly  diatoms 

'Mostly  Planorhulla,  rarely  Physa 

'Mostly  terrestrial  forms 


June  1980 


Williams,  Williams:  Borax  Lake  Chub 


10' 


Diel  Feeding  Chronology 

Feeding  chronologv  and  daily  ration  were 
determined  bv  the  relative  weight  of  material 
ingested  bv  CUla  honixohiiis  collected  during 
a  24-hour  period  in  June  1979  (Fig.  2).  The 
average  weight  of  fish  was  1.21  g.  Gila  horax- 
ohius  fed  throughout  the  day  with  peak  feed- 
ing activity  shortly  after  sunset.  Minimal 
feeding  activities  occurred  after  sunrise.  An 
increase  in  feeding  activity  after  sunset  has 
been  observed  in  Gila  hicolor  (Snyder  1917). 
The  average  weight  of  ingested  material  in 
intestines,  as  determined  from  1800-1500 
hours,  was  2.32  percent  of  body  weight.  This 
average  weight  of  ingested  material  (S)  was 
used  to  determine  the  daily  ration  (Rj)  as  fol- 
lows: 


Rt  =  24Sa  =  24(2.32)(.2)  =  11.14 

By  the  above  method  we  calculated  that  G. 
horaxohius  ingested  11.14  percent  of  their 
body  weight  daily.  This  estimate  is  larger 
than  most  reported  by  researchers  for  other 
species.  Brett  (1979)  summarized  in- 
vestigations made  by  various  researchers  who 
calculated  daily  rations  that  were  typically 
2-5  percent  of  body  weight.  Several  studies 
have  noted  increased  relative  ration  with  in- 
creased temperature  (e.g.,  Brett  et  al.  1969, 
Kinne  1960,  Stauffer  1973)  and  with  smaller 
fish  size  (e.g.,  Brett  1971,  Brett  and 
Shelbourn  1975,  Elliott  1975).  Brett  (1979) 
reported  that  temperature  and  fish  size  were 
of  greatest  importance  in  determining  ration 
size.  The  dwarf  size  of  G.  horaxohius  and  its 


Table  4.  Contents  of  62  intestines  oi  Gila  hoiaxobius  collected  during  the  winter  of  1978-79.  ND  =  no  data. 


Item  ingested 


Percent 

frequency 

of  occurrence 


Mean  nmuber 
per  intestine 


Mean  percent 
volume 


RI 


Algae' 
Gastropods- 
Gastropod  eggs 
Haplotaxid  oligochaetes 
Harpacticoid  copepods 
Ostracods 
Cladocerans 
Plant  seeds 
Higher  plants 
Fish  scales 
Araneae 
Insect  eggs 
Unidentified  insects' 

Terrestrial  i\sec:ts 
C'ollembola 
Thvsanoptera  adults 
Ileniiptera  adults 
Goleoptera  adults 
Hynienoptera  adults 
Diptera  adults 

AyiATic:  insects 
('hironomid  larvae 
(;hir()nomid  pupae 
Odouata  nymphs 
Elmid  larvae 
Goleoptera  adults 

I.N()R{;ank:  debris 


94.44 

ND 

18.81 

22.05 

18.52 

0.54 

3.88 

4.36 

0.00 

0.00 

0.00 

0.(K) 

5.56 

0.91 

2.21 

1,51 

75.93 

16.44 

9.96 

16.72 

77.78 

21.65 

20.58 

19.15 

55.56 

6.52 

4.07 

11.61 

3.70 

0.04 

0.02 

0.72 

5.56 

ND 

0.35 

1.15 

1.85 

0.04 

0.03 

0.37 

0.00 

0.00 

().(X) 

(),(K) 

0.00 

0.00 

(),()() 

0.00 

9.26 

ND 

1.49 

2.09 

1.85 

0.04 

0.33 

0.42 

0.00 

0.00 

0.00 

0.(X) 

0.00 

0.00 

0.(K) 

0.00 

0.00 

0.00 

0.(K) 

0.(K) 

0.00 

0.00 

().(K) 

0.00 

1.85 

0.02 

0.15 

0.39 

70.37 

5.15 

10.94 

15.83 

1.85 

0.02 

0.22 

0.40 

5.56 

0.07 

0.77 

1.23 

9.26 

0.11 

0.84 

1.97 

0.00 

0.00 

().()() 

().(M) 

90.74 

ND 

25.37 

- 

Total 


100.02 


'Mostly  diatoms 

■Mostly  PkmorhuUa.  rarely  Physa 
Mostly  terrestrial  forms 


108 


Great  Basin  Naturalist 


Vol.  40,  No.  2 


gastropod 


gastroDod 


copepod 


algae 


est  racod 


copepod 


algae 


debris 


ostracod 


other 


debris 


other 


terrestrial      insect 


JUVEN I LES 


ADULTS 


SPRING 


gastropod  2.5 


0.1  algae 


copepod 


ostracod 


other 


gastropod 


ebris 


aquatic 
insect 


other 


teirest  rial     i  nsec  t 


JUVENI LES 


ADULTS 


SUMMER 


Fig.  1.  Comparison  of  food  habits  between  128  juvenile  and  1.32  adult  Cila  Inmixohitis.  Mean  percent  volume  of 
food  items  are  given  in  circle. 


June  1980 


Williams,  Williams:  Borax  Lake  Chub 


109 


copepod 


terrestrial      insect 


debris 


copepod         gastropod 


istracod    0.2 


debris 


0.9   other 


JUVENILES 


ADULTS 


AUTUMN 


0.9   gastropod 

1 


copepo 


alqae 


est  racod 


gast  ropod 


copepo 


ost  racod 


debris 


ther 
aquatic    insect  i 

1.0    terrestrial       insect 


debris 


f  other 

2-^    terrestrial       insect 


JUVENI LES 


ADULTS 


Wl  NTER 


Fig.  1  continued. 


110 


Great  Basin  Naturalist 


Vol.  40,  No.  2 


habitation  in  thermal  spring  waters  contrib- 
uted to  the  large  daily  ration  found  in  this 
species.  Many  researchers  determined  daily 
ration  using  prepared  foods  that  were  per- 
haps more  nutritious  than  material  ingested 
by  Gila  boraxobiiis,  which  included  such  un- 
digestible  items  as  insect  exoskeletons,  gastro- 
pod shells,  and  inorganic  debris.  The  pres- 
ence of  large  amounts  of  undigestible 
material  increases  S  because  the  fish  must  in- 
gest a  larger  volume  of  material  to  get 
enough  calories.  This  results  in  a  larger  daily 
ration  than  would  be  calculated  if  fish  con- 
sumed only  digestible  foods. 


Food  Habits  of  Gila  ahordensis 

Ten  food  items  were  foimd  in  intestines  of 
Gila  alvordensis  collected  from  Thousand 
Creek,  Nevada,  in  June  1978  (Table  5).  Of 
the  ten  food  items,  chironomid  larvae,  cla- 
docerans,  copepods,  and  ostracods  were  of 
greatest  importance.  Chironomid  larvae  oc- 
curred in  all  intestines  examined  and  ac- 
counted for  approximately  26  percent  mean 
volume  of  intestines.  Microcrustaceans  com- 
prised almost  45  percent  mean  volume  of  in- 
testines. Diatoms  accounted  for  5  percent 
mean  volume  of  intestines.  No  terrestrial  in- 
sects were  ingested  by  G.  alvordensis  from 
Thousand  Creek. 

Eleven  food  items  were  found  in  intestines 
of  Gila  alvordensis  collected  from  Serrano 
Pond,  Oregon,  in  August  1977  (Table  6).  Of 
the  eleven  food  items,  chironomid  larvae,  di- 
atoms, cladocerans,  and  ostracods  were  of 
greatest  importance.  Chironomid  larvae  oc- 
curred in  over  three-quarters  of  the  intestines 
examined  and  accounted  for  approximately 
50  percent  mean  volume  of  all  intestines.  Di- 
atoms occurred  in  one-half  of  the  intestines 
and  accounted  for  almost  23  percent  mean 
volume  of  all  intestines.  Microcrustaceans 
comprised  approximately  17  percent  mean 
volume  of  intestines.  No  intestines  contained 
terrestrial  insects.  Gila  alvordensis  from  Ser- 
rano Pond  were  highly  opportimistic  feeders. 
Eighty-nine  percent  of  the  fish  from  Serrano 
Pond  with  food  in  their  intestines  contained 
one  item  that  accounted  for  more  than  50 
percent  of  their  intestinal  volume.  Thirty- 
nine  percent  of  fish  contained  one  food  item 


that  comprised  90  percent  or  more  of  in- 
testinal volume.  This  exploitive  feeding  was 
focused  on  chironomid  larvae,  cladocerans, 
or  algae.  One  intestine  was  exclusively  filled 
with  2570  cladocerans.  Such  exploitive  feed- 
ing was  not  noted  in  Gila  alvordensis  from 
Thousand  Creek. 

A  comparison  of  foods  of  Gila  alvordensis 
collected  during  June  and  August  with  foods 
of  Gila  boraxobiiis  collected  during  the  sum- 
mer shows  several  differences.  Terrestrial  in- 
sects were  important  foods  for  Gila  borax- 
obiiis during  the  summer  but  were  absent 
from  intestines  of  Gila  alvordensis.  Gila 
boraxobiiis  also  consumed  larger  quantities  of 
other  terrestrial  foods,  such  as  spiders  and  in- 
sect eggs,  than  did  G.  alvordensis.  Intestines 
of  G.  alvordensis  from  Thousand  Creek,  and 
to  a  lesser  extent  those  from  Serrano  Pond, 
contained  much  larger  amounts  of  micro- 
ciTistacea  than  did  intestines  of  G.  boraxobiiis 
during  the  summer.  Diatoms  were  a  major 
food  item  of  fish  in  Serrano  Pond  during  the 
simimer,  but  not  of  fish  in  Thousand  Creek  or 
of  Gc  boraxobiiis.  Gila  boraxobiiis  consumed 
a  larger  number  of  food  items  than  either 
population  of  G.  alvordensis.  This  is  due  to 
the  greater  opportimism,  including  the  use  of 
terrestrial  foods,  exhibited  by  G.  boraxobiiis. 
A  larger  sample  size  may  also  contribute  to 
the  greater  diversity  of  foods  utilized  by  G. 
boraxobiiis. 


Discussion 

Gila  boraxobiiis  and  G.  alvordensis  have 
tentatively  been  included  with  G.  bicolor  in 
the  subgenus  Siphateles  (Hubbs  and  Miller 
1972,  Williams  and  Bond,  in  press).  Although 
no  life  history  information  has  previously 
been  published  for  G.  boraxobiiis  or  G.  alvor- 
densis, several  researchers  have  examined 
food  habits  of  G.  bicolor  and  concluded  that 
they  are  primarily  opportunistic  omnivores 
(Bird  1975,  Cooper  1978,  La  Rivers  1962). 
However,  differences  in  food  habits  between 
the  coarse  gill  raker  form,  G.  b.  obesa,  and 
the  form  with  numerous,  fine  gill  rakers,  G. 
b.  pectinifer,  have  been  noted.  In  describing 
habits  of  the  form  with  coarse  gill  rakers 
from  Lake  Tahoe,  Miller  (1951)  foimd  them 
to  be  primarily  benthic  feeders,  with  a  diet 


June  1980 


Williams,  Williams:  Borax  Lake  Chub 


111 


composed  of  89  percent  bottom  organisms. 
Snyder  (1917)  noted  that  the  form  with 
coarse  gill  rakers  collected  from  the  littoral 
zone  of  Lake  Tahoe  fed  on  algae,  other  plant 
material,  and  insects.  Gila  b.  pectinifer  from 
Lake  Tahoe,  with  its  numerous  gill  rakers, 
fed  almost  exclusively  on  midwater  micro- 
crustacea  (Miller  1951).  La  Rivers  (1962)  also 
reported  that  G.  b.  pectinifer  contained  many 
midwater  foods  in  their  intestines,  primarilv 
consuming    diatoms    and    microcrustaceans. 


Cooper  (1978)  reported  that  the  form  with 
numerous  gill  rakers  (although  he  referred  it 
to  G.  h.  obesa)  in  Walker  Lake,  Nevada,  fed 
mostly  on  zooplankton  and  filamentous  al- 
gae. A  population  complex  of  G.  bicolor  in 
Eagle  Lake,  California,  that  included  forms 
with  both  coarse  and  fine  gill  rakers  fed  on  a 
variety  of  foods,  including  zooplankton,  plant 
material,  insect  larvae,  and  surface  insects 
(Kimsey  1954).  There  appears  to  be  a  definite 
correlation   between   gill   raker   morphology 


Table  5.  Contents  of  21   intestines  of  Gila  uhordctisis  collected   13  June  1978  from  Thousand  Creek,  Nevada. 
ND  =  no  data. 


Item  ingested 


Percent 

frequency 

of  occurrence 


Mean  number 
per  intestine 


Mean  percent 
volume 


RI 


Diatoms 

Gastropods 

Harpacticoid  copepods 

Ostracods 

Cladocerans 

Plant  seeds 

Araneae 

AgiL-VTIC  INSECTS 

Chironomid  larvae 
Chironomid  pupae 
Coleoptera  adults 

I.NORCANIC:  DEBRIS 


36.36 

ND 

5.00 

7.91 

9.09 

0.09 

0.36 

1.81 

81.82 

6.82 

6.82 

16.94 

72.73 

15.82 

13.73 

16.53 

90.91 

27.73 

24.23 

22.01 

27.27 

0.27 

0.45 

5.30 

9.09 

0.09 

0.55 

1.84 

10().(K) 

16.73 

25.78 

24.04 

9.09 

0.09 

0.09 

1.75 

9.09 

0.09 

0.68 

1.87 

lOO.(K) 

ND 

20.95 

- 

Total 


98.64 


Table  6.  Contents  of  20  intestines  of  Gila  akordensis  collected  6  August    1977  from   Serrano  Pond,  Oregon. 
ND  =  no  data. 


Item  ingested 


Percent 

frequency 

of  occurrence 


Mean  number 
per  intestine 


Mean  percent 
volume 


RI 


Diatoms 

Harpacticoid  copepods 

Ostracods 

Cladocerans 

.Araneae 

Insect  eggs 

Unidentified  insects 

.\quatic  insects 
Chironomid  larvae 
Chironomid  pupae 
Odonata  nymphs 
Ephemeroptera  larvae 

Inorganic  debris 


50.00 

ND 

22.94 

18.16 

.33.33 

1.67 

0.36 

8.39 

38.89 

5.94 

1.70 

10.11 

55.56 

253.28 

15.34 

17.65 

5.56 

0.06 

0.22 

1.44 

5..56 

0.06 

0.05 

1.40 

5.56 

ND 

0.05 

1.40 

77.78 

20..33 

50.41 

31.92 

22.22 

0.33 

2.96 

6.27 

5.56 

0.06 

0.67 

1.55 

5.56 

0.06 

1.33 

1.72 

55..56 

ND 

3.96 

- 

Total 


99.99 


112 


Great  Basin  Naturalist 


Vol.  40,  No.  2 


and  food  habits,  tho.se  with  coarse  gill  rakers 
ingesting  more  benthic  food  organisms  and 
those  with  fine  gill  rakers  ingesting  more 
zooplankton.  Gila  boraxobius  and  G.  alvor- 
densis  possess  approximately  16  and  20  short 
gill  rakers,  respectively,  agreeing  closely  with 
the  gill  raker  morphology  of  G.  bicolor  obesa 
form.  Although  G.  boraxobius  typically  feeds 
on  benthic  organisms,  large  amounts  of  dia- 
toms, microcnistaceans,  and  terrestrial  insects 
are  ingested  sea.sonally. 

The  ingestion  of  terrestrial  insects  by  Gila 
is  not  common.  However,  several  researchers 
have  found  that  terrestrial  insects  comprised 
a  small  part  of  the  diet  of  Gila  (Cross  1978, 
Kimsey  1954,  Moyle  1976,  Sigler  and  Miller 
1963).  Terrestrial  insects  were  the  primary 
foods  of  G.  robusta  and  G.  elegaas  longer 
than  200  mm  SL  collected  from  the  Green 
River  (Vanicek  and  Kramer  1969).  Smaller  G. 
robusta  and  G.  elegans  contained  pre- 
dominantly aquatic  insect  larvae.  Juvenile 
and  adult  G.   boraxobius  consumed  large 


quantities  of  terrestrial  insects.  Several  re- 
searchers (Kimsey  1954,  Miller  1951)  have 
noted  that  as  Gila  grow,  they  switch  to  larger 
food  items;  however,  at  least  one  study  (Gra- 
ham 1961)  found  foods  of  different  sized 
groups  of  CAla  to  be  nearly  identical.  We 
found  foods  of  juvenile  and  adult  G.  borax- 
obius to  be  very  similar,  except  that  adults 
exhibited  a  greater  consumption  of  gastro- 
pods and  diatoms,  and  juveniles  consumed 
more  copepods  and  terrestrial  insects.  The 
hard  shells  and,  to  a  lesser  extent,  the  rela- 
tively large  size  of  gastropods  probably  con- 
tributed to  juveniles  avoiding  them  as  a  food 
source.  Larger  Gila  ingest  more  algae  than 
do  smaller  fish  in  studies  by  Moyle  (1976)  and 
Vanicek  and  Kramer  (1969).  Age  II  Gila  coe- 
rulea  feed  predominantly  on  filamentous  al- 
gae, whereas  algae  were  entirely  or  prac- 
tically absent  from  age  I  fish  (Moyle  1976). 
Large  adult  Gila  from  the  Green  River  con- 
sumed more  algae  than  did  smaller  fish  (Van- 
icek and  Kramer   1969).  Juvenile  G.  borax- 


Tl  ME        OF        DAY 


Fig.  2.  Feeding  throiiologv^  of  Cila  boraxobius  on  5-6  June  1979.  N  =  72. 


June  1980 


Williams,  Williams:  Borax  Lake  Chub 


113 


obitis  consumed  more  terrestrial  insects  than 
did  the  adults,  except  in  winter,  when  small 
amoiuits  of  terrestrial  insects  were  ingested 
by  both  groups.  The  reason  for  juveniles  con- 
suming large  volumes  of  terrestrial  insects  is 
unknown;  apparently  the  relatively  large  size 
of  this  food  item  is  not  a  deterrent. 

Both  juvenile  and  adult  G.  horaxoJyiiis  in- 
creased consumption  of  diatoms  and  micro- 
crustaceans  in  winter.  This  probably  in- 
dicates a  scarcity  of  food  items  during  the 
winter,  as  is  reflected  by  finding  24  percent 
fewer  food  items  in  intestines  during  winter 
than  in  summer.  A  factor  contributing  to  the 
winter  scarcity  of  food  items  is  a  decrease  in 
the  availability  of  terrestrial  foods,  causing 
concentrated  feeding  on  remaining  food 
items.  For  example,  the  Devil's  Hole  pupfish, 
Cyprinodon  diabolis,  which  inliabits  a  small 
thermal  spring,  dramatically  increases  in- 
gestion of  diatoms  during  winter  due  to  a 
scarcity  of  preferred  foods  (Minckley  and 
Deacon  1975).  Although  the  amount  of  nutri- 
tion derived  from  consuming  diatoms  is  un- 
known, we  suspect  that  the  large  con- 
sumption of  diatoms  at  certain  times  of  the 
year  by  G.  bomxobius  would  indicate  that 
some  nutritive  value  is  gained.  Arnold  (1971) 
found  that  species  of  Ci/prinodon  derived  oil 
droplets  from  ingested  diatoms,  thus  extract- 
ing nutritive  value.  A  similar  mechanism 
could  operate  in  Gila  bomxobius. 

Examination  of  summer  foods  of  both  G. 
ah'ordensis  populations  showed  differences 
from  the  summer  foods  of  G.  boraxobius. 
During  summer  Gilo  boraxobius  relied  heavi- 
ly on  terrestrial  food  items,  whereas  popu- 
lations of  G.  alvordensis  consumed  prac- 
tically no  terrestrial  foods.  Forty-three 
percent  of  food  items  consumed  by  G.  bora- 
xobius during  the  summer  were  of  terrestrial 
origin.  Surveys  of  potential  food  items  in  Bo- 
rax Lake  conducted  at  various  times  of  the 
year  found  that  all  potential  food  items  were 
utilized  by  G.  boraxobius  except  some  adult 
hemipteran  and  coleopteran  insects  that  were 
probably  too  large  to  be  ingested.  Also,  many 
hemipterans  possess  scent  glands  that  render 
them  impalatable  to  predators. 

Acknowledgments 

The  authors  are  indebted  to  Carl  E.  Bond 
for  his  guidance   during   the   course  of  this 


study  and  for  his  review  of  the  manuscript. 
Stanley  V.  Gregory  provided  identification  of 
diatoms  and  information  on  their  habitats, 
the  Oregon  Department  of  Fish  and  Wildlife 
provided  collecting  permits,  and  James  J. 
Long,  Kevin  M.  Howe,  and  Glen  DeMott  as- 
sisted with  field  collections.  Kevin  M.  Howe 
reviewed  the  manuscript  and  provided  field 
notes  of  the  Serrano  Pond  area.  This  informa- 
tion is  part  of  the  senior  author's  doctoral  dis- 
sertation at  Oregon  State  University  and  is 
prepublished  by  permission  of  the  Graduate 
School  and  the  Department  of  Fisheries  and 
Wildlife. 

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FIRST  RECORD  OF  THE  PALLID  BAT  (ANTROZOUS  PALLIDUS)  FROM  MONTANA 

Jeff  Shryer'  and  Dennis  L.  Flath' 

.\bstract.—  a  pallid  bat  {Antrozous  pallidiis)  was  taken  20  km  SE  Warren,  Carbon  County,  Montana.  This  repre- 
sents a  410  km  range  extension  and  a  new  record  for  Montana. 


The  pallid  bat  {Antrozous  pallidiis)  has  not 
been  previously  reported  from  Montana.  On 
the  basis  of  specimens  from  Grangeville  and 
Pocatello,  Idaho,  Hoffman  and  Pattie  (1968) 
suggested  that  the  species  may  occur  in 
southwestern  Montana.  Fichter  (1964)  report- 
ed A.  pallidus  from  Boise,  Idaho. 

On  25  August  1978  we  mist-netted  an 
adult  female  A.  pallidus  (Montana  Fish, 
Wildlife,  and  Parks  Collection  NG  748)  at  an 
isolated  spring  approximately  20  km  SE 
Warren,  Carbon  County,  Montana.  This  rec- 
ord extends  the  known  range  of  A.  pallidas 
approximately  410  km  northeast  of  Pocatello, 
Idaho,  and  approximately  460  km  north  of 
Dinosaur  Quarry,  Utah,  where  Knitzsch  and 
Heppenstall  (1955)  obtained  a  specimen. 

The  new  locality  is  in  an  alluvial  plain 
south  of  the  Pryor  Mountains,  at  1370  m  ele- 
vation, and  lies  within  the  25-30  cm  precipi- 
tation zone  (U.S.  Soil  Conservation  Service 
1977).  The  area  is  characterized  by  scattered 
outcrops  of  sandstone  and  .shale,  with  a  vege- 
tative community  dominated  by  western 
wheatgrass  {Agropyron  smitliii),  sagebrush 
{Artemisia  spp.),  and  saltbush  (Atriplcx  spp.). 

External  measurements  of  the  specimen 
are  as  follows:  total  length,  110  mm;  length 
of  tail,  37  mm;  length  of  hind  foot,  13  mm; 
length  of  ear,  29  mm;  and  length  of  forearm, 
59  mm.  The  dentition  displaved  substantial 
wear,  indicating  the  bat  was  old.  This  speci- 
men is  similar  to  specimens  in  the  University 
of  Montana  Zoological   Museum  from  New 


Mexico  (MSUZ  10419)  and  Arizona  (MSUZ 
12996)  in  that  it  is  of  a  similar  color— a  very 
pale  fawn.  Allen  (1864)  described  A.  p.  pal- 
lidas as  having  two  varieties  of  color,  fawn 
and  yellowish  brown.  Bailey  (1936)  described 
A.  p.  cantwelli  as  darker  than  pallidus,  with 
dark  brown  ears  and  membranes.  The  mem- 
branes of  our  specimen  are  medium  brown. 

Other  species  captured  concurrently  in- 
clude: Myotis  lucifugus  and  M.  leihii. 

We  thank  John  Ciralli  for  field  a.ssistance 
and  P.  L.  Wright  for  assistance  in  the  identi- 
fication of  the  specimen.  This  paper  is  a  con- 
tribution of  the  U.S.  Bureau  of  Land  Manage- 
ment, Montana  Nongame  Program,  and  the 
Montana  Department  of  Fish,  Wildlife,  and 
Parks,  Nongame  Program,  State  Project 
5853. 

Literature  Cited 

.\LLEN,  H.  1864.  Monograph  of  the  bats  of  North  Ameri- 
ca. Smithsonian  Misc.  Coll.  165:68-69. 

.\.\o.NYMOUS.  1977.  Average  annual  precipitation  for 
Montana  based  on  1941-1970  base.  USDA,  Soil 
Cons.  Serv.,  Portland,  Ore.  13pp. 

B.\iLEY,  V.  1936.  The  mammals  and  life  zones  of  Oregon. 
North  American  Fauna  55:390-.393. 

Fichter,  E.  1964.  The  pallid  bat  in  Idaho.  Tebiwa 
7:23-27. 

Hoffman,  R.  S.,  a.nd  D.  L.  Pattie.  1968.  .-^  guide  to 
Montana  mammals:  Identification,  habitat,  distri- 
bution and  abundance.  Univ.  Montana  Print. 
Serv.  1.33  pp. 

Krutzsch,  p.  H.,  and  C.  A.  Heppenstall.  1955.  Addi- 
tional distributional  records  of  bats  in  Utah.  J. 
Manmial.  .36:126-127. 


'U.S.  Bureau  of  Land  Management,  Lewistown  District  Office.  Lewistown,  Montana  594.57. 

'Montana  Department  of  Fish,  Wildlife,  and  Parks,  Montana  State  University,  Box  5,  Bozeman,  Montana  59717 


115 


A  CHIRACANTHIUM  SPIDER  BITE 

Dorald  M.  Allied' 
.\bstr.\ct.—  a  bite  bv  Cliimcanthium  mildci  L.  Koch  is  described. 


In  May  1979,  Mr.  Lee  Carson  of  Provo, 
Utah,  brought  a  spider  to  me  which  had  bit- 
ten him  on  the  right  index  finger.  It  was  sub- 
sequently identified  by  Dr.  Wilhs  J.  Gertsch 
as  a  female  Chiracanthium  niildei  L.  Koch. 

Mr.  Carson  was  in  the  process  of  placing  a 
pair  of  Ribbers  over  his  shoes  to  work  in  his 
garden.  The  mbbers  were  kept  in  his  closed 
garage,  and  had  been  worn  a  few  days  pre- 
viously. As  he  inserted  his  fingers  into  one  of 
the  rubbers,  he  felt  a  sudden  pain  at  the  tip 
of  his  finger.  Examination  disclosed  the  spi- 
der with  a  web  and  cocoon  in  a  rubber. 

Within  3  to  4  seconds  after  the  bite,  the 
finger  began  to  ache  severely.  The  pain  soon 
extended  to  his  upper  arm  where  it  remained 
for  2  to  3  hours,  although  it  was  most  con- 
centrated in  the  finger.  He  described  the  sen- 
sation as  a  painful  "tingling"  similar  to  what 
one  experiences  when  his  elbow  "crazy- 
bone"   is  bumped.   No  nausea,  headache. 


swelling,  or  numbness  was  experienced.  No 
evidence  of  inflammation  or  necrosis  oc- 
curred at  the  site  of  the  bite,  and  the  cheli- 
ceral  punctures  healed  rapidly.  Pain  in  the 
finger  and  arm  disappeared  after  about  4 
hours. 

Dr.  Gertsch  kindly  supplied  information  on 
the  spider  and  its  ecology.  The  species  appar- 
ently was  introduced  into  the  United  States, 
became  a  typical  house  spider  in  the  New 
York  and  Boston  area,  and  subsequently 
spread  to  other  areas  of  the  U.S.  As  early  as 
1930  Dr.  Gertsch  collected  it  at  Salina,  Utah. 
It  is  known  from  southern  California,  and 
Kaston  (1972,  "How  to  Know  the  Spiders,  p. 
221)  indicated  its  distribution  as  New  Eng- 
land, New  York,  New  Jersey,  Alabama,  Mis- 
souri, and  Utah.  Gertsch  further  stated  that 
the  genus  is  reputed  to  have  a  venom  of  in- 
termediate potence,  and  that  of  the  Eu- 
ropean species  is  said  to  be  next  to  their  Lat- 
rodectus  in  severity. 


'Department  of  Zoology,  Brigham  Young  University,  Provo,  Utah  84602. 


116 


IDENTITY  OF  NARROW-LEAVED 
CHRYSOTHAMNUS  VISCIDIFLOR US  (ASTERACEAE)' 

Loraii  ('.  Aiidersoir 


.\bstr.\ct.—  Two  ijroups  of  glabrous,  narrow-leaved  Chiysotliaiiinits  visciclifloius  (Asteraceae)  are  perceived,  and 
appropriate  taxononiie  combinations  are  made,  i.e.,  C.  v.  ssp.  vmidiflorus  var.  stcnofjlttjlhis  and  C.  v.  ssp.  axillaris. 
The  two  are  fairl\'  distinct  ideograph icallv,  and  thev  can  be  separated  l)v  floral  niorpliologv.  .\  key  to  all  species  of 
section  Chrtjsothanntus  (to  which  C.  liscidiflorus  belongs)  is  given. 


Classification  of  intra.specific  variants  of 
Chrysothamnus  liscidiflorus  has  been  prob- 
lematic, in  part,  becau.se  floral  features  seem- 
ingly lacked  sufficient  distinctions.  Hence, 
vegetative  aspects  such  as  stature,  vesture, 
and  leaf  dimensions  have  been  used.  Experi- 
mental -Studies  (Anderson  1964)  have  demon- 
strated that  in  many  in.stances  plant  height, 
leaf  twisting,  and  leaf  width  are  expressions 
of  differing  edaphic  conditions,  droughtiness, 
or  other  environmental  parameters  and 
thereby  complicate  taxonomic  resolution. 

Study  of  interpopulational  variation  and 
the  distribution  of  the  narrow-leaved,  yellow 
rabbitbmsh  (C.  viscidiflorus  ssp.  stenophyUus) 
reveals  that  there  are  two  different  taxa  rep- 
resented. One  is  found  sporadically  through 
the  northern  latitudes  of  the  western  United 
States  and  into  southern  California  (open  cir- 
cles in  Fig.  1).  The  other  taxon  (stars.  Fig.  1) 
is  found  further  .south  and  is  more  generally 
distributed,  i.e.,  has  a  more  pronounced 
"range  integrity."  In  south-central  Nevada, 
Beatley  (1976)  reports  it  is  the  common 
Chrysothamnus  of  basin  floors  and  foothills, 
especially  in  volcanic  areas  and  on  disturbed 
sites,  usually  below  5500  ft.  Examination  of 
populations  at  possible  type  locality  sites 
(type  specimens  labeled  either  West  Hum- 
boldt Mountains  or  Huntington  Valley)  for 
ssp.  StenophyUus  shows  that  the  narrow- 
leaved  plants  represent  extremes  of  the 
broader-leaved  ssp.  viscidiflorus. 

I  have  concluded  from  field  observations, 
garden  culture,  and  herbarium  studies   that 


the  northern  elements,  which  include  the 
tvpe  collection  of  C.  i".  ssp.  stenophyUus,  are 
actually  environmentally  induced  variants  of 
ssp.  liscidiflorus.  Although  quadrinomials  are 
cumbersome,  the  following  nomenclatural 
combination  more  appropriately  identifies 
the  relationship  of  these  plants: 

Chrysothamnus  viscidiflorus  (Hook.)  Nutt. 
ssp.  viscidiflorus  var.  stenophyUus  (Gray)  L. 
C.  Anderson,  comb.  nov. 

Basionym:  Bigelovia  douglasii  Gray  var. 
stenophylla  Gray.  Proc.  Am.  Acad.  Sci. 
8:646,  1873.  W. '  Humboldt  Mtns,  Nevada, 
Watson  566  (GH,  holotype;  NY,  US,  iso- 
types). 

Synonymy:  Chrysothamnus  pumilus  Nutt. 
var.  varus  A.  Nels.  Bot.  Gaz.  28:  375,  1899. 
Centennial  Valley,  Wyoming,  Nelson  1847 
(RM,  holotype;  GH,  NY,  i.sotypes). 

Tlie  .southern  elements  that  had  previously 
been  referred  to  ssp.  stenophyUus  warrant 
subspecific  recognition.  Their  narrow-leaved 
characteristic  is  independent  of  environmen- 
tal conditions.  These  plants  are  diploids; 
broad-leaved  forms  of  ssp.  viscidiflorus  that 
grow  in  the  same  region  are  tetraploids  or 
hexaploids  (Anderson,  1966,  1971).  The  only 
available  name  for  these  narrow-leaved 
plants  is  C.  axillaris.  In  1964,  I  noted  that  C. 
axillaris  was  not  specifically  distinct  from  C. 
viscidiflorus,  and  the  name  was  synonomized 
luider  .ssp.  stenophyUus.  Munz  (1968),  in  re- 


'This  shidy  was  supported  by  .National  Science  Foundation  Grant  DEB  76-10768.  Ken  VVonilile  helped  with  ijraphics. 
•Department  of  Biological  Science,  Florida  Stale  University,  Tallahassee,  Florida  32306. 


117 


118 


Great  Basin  Naturalist 


Vol.  40,  No.  2 


ferring  to  my  studies,  stated  I  had  made  C. 
uxillaris  a  suljspecies  of  C.  viscidiflonis.  The 
statement  was  inaccurate  at  that  time— but 
prophetic.  The  combination  is  now  made: 

Chrysothamnus  viscidiflonis  (Hook.)  Nutt. 
ssp.  axillaris  (Keck)  L.  C.  Anderson,  comb.  & 
Stat.  nov. 

Basionvm:  Chri/sothamntis  axiUaris  Keck. 
Ahso  4:103,  1958.'  Deep  Springs  Valley,  Cali- 
fornia, Ferris  6924  (NY,  holotype;  DS,  LL, 
POM,  isotypes). 

Keck  (1958),  in  describing  C.  axiUaris,  re- 
lated it  to  C.  albidus  and  more  distantly  to  C. 
greenei.  Actually,  ssp.  axillaris  is  not  close  to 
C.  albidus  in  relationship,  but  it  is  to  C. 
greenei.  In  fact,  the  type  collection  of  ssp.  ax- 
illaris with  very  acute  phyllaries  suggests 
some  intergradation  with  C.  greenei.  Keck 
stated  that  the  latter  was  known  only  from 
eastern  Nevada  and  eastward,  but  it  does  ex- 


tend through  southern  Nevada  into  Inyo 
County,  California.  At  its  western  limit  in 
California  and  also  in  northeastern  Arizona, 
C.  greenei  intergrades  somewhat  with  C.  vis- 
cidiflorus.  The  feature  of  vertically  aligned 
phyllaries  noted  by  Keck  (1958)  is  not  con- 
sistent for  ssp.  axillaris  (Anderson  1964). 

Although  ssp.  axillaris  and  var.  steno- 
plnjllus  are  fairly  distinctive  habitally,  they 
could  not  be  "keyed  out"  easily  vinless  refer- 
ence was  made  to  geographic  distribution 
(Fig.  1).  Floral  morphology  was  studied  in 
search  of  additional  distinguishing  features. 
Methods  are  those  used  earlier  (Anderson 
1964).  Detailed  floral  data  and  a  list  of  speci- 
mens examined  are  on  file  at  FSU.  Significant 
comparisons  are  graphically  illustrated  in 
Figure  2. 

Involucral  width  in  Chrysothamnus  is  gen- 
erally strongly  correlated  with  flower  num- 
ber because  more  flowers  per  head  require  a 
broader  receptacle.  The  pattern  is  evident  in 


general    range   for    C.  v. 
sep.    viscidiflorus 

C.  V.    ssp.    viscidiflorus 
var.    stenophyllus 

C.  V.    ssp.    axillaris 


Fig.  1.  Range  of  Clirysolhamnu.^  viscidiflonis  ssp.  viscidiflonis  and  ssp.  axillahs.  Range  for  ssp.  viscidiflonis  nearly 
equals  that  of  the  species;  ssp.  tanceolatus  extends  into  southern  British  Columbia  and  north-central  New  Mexico. 
Distribution  of  ssp.  (ixillaiis  (stars)  is  fairly  general  through  southern  parts  of  Utah  and  Nevada  and  adjacent  regions, 
whereas  that  of  .ssp.  viscidiflonis  var.  stenophijllus  (open  circles)  is  sporadic,  but  mostly  north  of  ssp.  axillaris. 


June  1980 


Anderson:  Chrysothamnus 


119 


C.  viscidiflorus  ssp.  viscidiflorus,  wherein 
plants  averaging  10.8  (with  up  to  12)  flowers 
per  head  have  involucral  widths  over  50  per- 
cent of  involucral  length  (Fig.  2);  these  plants 
represent  an  altitudinal  record  for  the  genus 
of  12,800  ft  in  the  White  Mountains  of  Cali- 
fornia. Previous  descriptions  of  C.  t/,s- 
cidiflonis  listed  flower  number  as  about  5 
(Hall  and  Clements  1923).  Nearly  all  plants 
of  ssp.  viscidiflorus  with  high  flower  number 
come  from  altitudes  over  10,000  ft.  Most 
populations  of  the  subspecies,  including  var. 
stenophyUus,  have  4-6(7)  flowers  per  head 
with  proportionately  narrower  involucres. 

Heads  of  ssp.  axillaris  depart  from  the  bas- 
ic correlation  of  flower  number— involucral 
width /length  ratio.  Although  they  average 
fewer  than  5  flowers  per  head,  the 
width /length  ratio  for  the  involucre  is  high 
(Fig.  2).  Tlius,  ssp.  axilluris  and  var.  steno- 
phijllus  can  be  distinguished  by  floral  fea- 
tures as  well  as  geographically. 

Taxonomic  interpretation  of  Chryso- 
thamnus section  Chrysothamnus  (to  which  C. 
viscidiflorus  belongs)  has  been  altered  consid- 
erably since  Hall's  monograph  (Hall  and  Cle- 
ments 1923).  Chrysothamnus  vaseyi  and  C. 
molestus  (C.  viscidiflorus  var.  tnolestus)  have 
been  transferred  to  section  Fulchelli  (Ander- 
son 1970)  and  C.  gramineus  to  Petradoria 
(Anderson  1963).  Additional  species  have 
been  recognized  in  the  section.  A  key  to  sec- 
tion Chrysothamnus  as  currently  understood 
is  presented  here. 


-/At- 


Fig.  2.  Correlation  of  involucral  shape  and  flower 
number  in  heads  of  Cltrysothamntis  viscidiflorus  ssp.  vis- 
cidiflorus (closed  circles,  var.  viscidiflorus;  open  circles, 
var.  stcnoplnjilus)  and  ssp.  axillaris  (stars).  Vertical  axis 
represents  the  involucral  width /length  relationship  ex- 
pressed as  percent;  the  horizontal  axis  is  average  flower 
number  per  head.  Note  that  ssp.  axillaris  departs  from 
the  general  correlation  in  ssp.  viscidiflorus  and  other 
members  of  the  genus. 


1.  Flowers  white;  leaves  terete,  strongly  punctate C.  albidus  (Jones)  Greene 

—  Flowers  yellow;  leaves  planate  or  involute,  not  punctate  2 

2(1).         Flowers  2-3(4);  style  branches  included  in  erect  corolla  lobes;  plants  mostly 

less  than  1.5  dm  tall C.  hurnilis  Greene 

—  Flowers  (3)4-6;  styles  exerted  beyond  spreading  corolla  lobes;  plants  often  over 

2  dm  tall 3 

3(2).         Style  appendages  long  (40-70  percent  of  style  branch);  leaves  never  twisted  or 

involute;  tall  shRibs  4 

—  Style  appendages  short   (30-45  percent  of  style  branch);   leaves  frequently 
twisted  or  involute 5 

4(3).         Leaves  lanceolate;  achenes  densely  pubescent  C.  linifolius  Greene 

—  Leaves  spatulate  to  oblanceolate;  achenes  sparsely  pubescent  

C.  spathulatus  L.C.Anders. 

5(3).         Phyllaries  acuminate-cuspidate;  leaves  1-2  mm  wide C.  greenei  (Gray)  Greene 

—  Phyllaries  obtuse  to  acute;  leaves  1-10  mm  wide  (C.  viscidiflorus)  6 


120 

6(5). 

7(6). 
8(7). 

9(7). 

10(9). 


Great  Basin  Naturalist 


Vol.  40,  No.  2 


Leaves  planate,  glabrous;  flowers  3.5-4(4.5)  mm  long 

C.  V.  ssp.  planifolius  L.C.Anders. 

Leaves  twisted  or  pubescent  or  flowers  longer 7 

Upper  stems,  frequently  leaves,  hairy  8 

Stems  glabrous;  leaves  only  ciliate  9 

Stems  and  leaves  pubenilent;  leaves  1-2(4)  mm  wide 

C.  V.  ssp.  puberiilus  (D.C.Eat.)  H.&C. 

Stems  hispid  near  inflorescence;  leaves  over  2  mm  wide,  hirsute  or  glabrous  

C.  t;.  ssp.  lanceolatiis  (Nutt.)  H.&C. 

Leaves  ±  1  mm  wide;  flowers  3-4(5);  involucres  somewhat  turbinate  

C.  r.  ssp.  axillaris  (Keck)  L.C.Anders. 

Leaves  1-10  mm  wide;  if  1  mm,  flowers  4  or  more  and  involucres  narrowly 
cylindric 10 

Leaves  more  than  1.5  mm  wide;  plants  up  to  1  m  tall 

C.  V.  ssp.  viscidiflorus  var.  viscidiflorus 

Leaves  1-1.5  mm  wide;  plants  mostly  less  than  3  dm  tall  

C.  V.  ssp.  viscidiflorus  var.  stenophyllus  (Gray)  L.C.Anders. 


Names  often  applied  in  C.  viscidiflorus, 
but  not  representative  as  distinct  subspecies, 
include:  (1)  elegans,  usually  misapplied  to 
certain  forms  of  ssp.  puberulus  with  bracts 
with  enlarged  green  tips,  but  the  type  speci- 
men does  not  have  such  bracts  and  is  clearly 
part  of  ssp.  lanceolatus;  (2)  pumilus,  low 
form  that  is  part  of  ssp.  viscidiflorus;  (3)  torti- 
folius,  environmental  variant  with  strongly 
twisted  leaves,  part  of  ssp.  viscidiflorus;  and 
(4)  latifolius,  wide-leaved  plants  from  north- 
ern Nevada  that  could  be  considered  a  varie- 
ty of  ssp.  viscidiflorus;  however,  not  all  wide- 
leaved  plants  of  the  subspecies  would  belong 
to  that  variety. 

A  conceptual  distinction  between  sub- 
species and  variety  exists  in  Chnjsothamnus 
for  me.  Subspecies  is  applied  to  groups  of 
populations  with  pronounced  geographical 
and  fairly  distinct  morphological  limits.  The 
variety  can  be  applied  in  two  ways.  It  may 
represent  a  sporadic  but  rather  distinctive 
morphotype  within  a  given  subspecies,  such 
as  ssp.  viscidiflorus  var.  stenophijllus,  and  it 
could  possibly  be  applied  to  such  variation  in 
species  where  subspecies  are  not  recognized, 
such  as  C.  grcenci  var.  filifolius  for  the  nar- 
row-leaved variant.  The  second  application 
(my  preferred  usage)  of  variety  would  be  for 
elements  of  a  subspecies  that  have  some  fair- 
ly consistent  morphological  distinction  and 
also  have  relatively  sharp  geographic  limits. 


but  are  clearly  subordinate  to  the  subspecies. 
An  example  would  be  ssp.  viscidiflorus  var. 
latifolius— a  the  combination  were  made. 
Publication  of  additional  quadrinomials  as 
needed  to  clarify  relationships  in  the  genus 
will  be  part  of  my  upcoming  monograph. 


Literature  Cited 

Anderson,  L.  C.  1963.  Studies  on  Pctradoria  (Conipos- 
itae):  Anatoniv,  cvtologv,  taxonoiiiv.  Trans.  Kans. 
Acad.  Sci.  66:632-684."' 

1964.   Taxononiic  notes  on  the  CIinjsotlKininus 

viscicliflonis  complex  (Astereae,  Conipositae). 
Madrono  17:222-227. 

1966.  Cytotaxonomic  studies  in  Clinisothiimnus 

(Astereae,  Conipositae).  Amer.  J.  Bot. 
53:204-212. 

1970.  Floral  anatomy  of  Chrysotli(iiuniis  (As- 
tereae, Conipositae).  Sida  3:466-503. 

1971.  Additional  chromosome  counts  in  Chnjso- 

tlunnntts  (Asteraceae).  Bull.  Torrev  Bot.  Club 
98:222-225. 

Be.\tley,  J.  C.  1976.  Vascular  plants  of  the  Nevada  Test 
Site  and  central-southern  Nevada.  Energy  Res.  & 
Dev.  ,\dmin.,  Nat'l  Tech.  Inf.  Service, 
Springfield,  Va. 

Hall,  H.  M.,  .\nu  F.  E.  Clements.  1923.  The  phyloge- 
netic  method  in  taxonomy:  The  North  American 
species  of  Artemisia,  Chnjsoihamnus,  and  Atri- 
plcx.  Carnegie  Inst.  Publ.  326:l-.355. 

K.ec:k,  D.  D.  1958.  Taxononiic  notes  on  the  California 
flora.  Aliso  4:101-114. 

Munz,  p.  a.  1968.  Supplement  to  A  California  Flora. 
I'niv.  Calif.  Press,  Berkeley. 


RIBULOSE  DIPHOSPHATE  CARBOXYLASE  ACTIVITIES  IN  COLD-RESISTANT 

COMMON  MALLOW,  MALVA  NEGLECTA  WALLR.  AND  A  COLD-SENSITIVE 

TOMATO,  LYCOPERSICON  ESCULENTUM  L.,  ACE  55  VAR. 

W  illiam  H.  AiidfiSL-ii'  and  Jack  D.  Brotlicrson' 

Abstfl^ct.—  Coiiinioii  mallow  (Malta  nc^hcta  W'allr. )  and  tomato  (Li/copcisicon  csciiUnttim  L.  var.  Ace  55)  were 
compared  as  to  certain  characteristics:  C02  fixation  properties,  ribulose  diphosphate  carboxyl  activities,  (RiiDPCase) 
photosynthesis,  respiration,  and  compensation  points.  Significant  differences  in  these  factors  were  observed  in  all 
cases  except  dark  respiration.  Mallow  enzyme  (RuDPCase)  activities  were  higher  per  unit  of  enzyme  than  those  of 
tomato.  The  Mallow  RuDPCase  exhibited  slightly  higher  activity  at  5  to  25  C.  Mallow  leaves  retained  their  capacity 
for  photosynthesis  and  respiration  after  long  periods  of  exposure  to  subfreezing  temperature.  The  cold  adapted  mal- 
low had  a  higher  C(%  compensation  point,  suggesting  a  lower  efficiency  for  CO2  fixation.  The  results  suggest  that 
cold  acclimation  in  common  mallow  affects  photosynthesis  but  has  little  effect  on  respiration. 


Several  physiological  factors  are  associated 
with  the  development  of  resistance  to  winter 
injury  in  plants.  Qualitative  and  quantitative 
changes  in  protein,  carbohydrate,  and  lipid 
contents  have  been  observed  during  cold  ac- 
climation (Roberts  1969,  Cerloff  et"  al.  1967, 
Hochachka  and  Somaro  1968,  Zeller  1951). 
However,  the  in  vivo  features  of  observed 
biochemical  and  physiological  alterations  as- 
sociated with  cold  acclimation  in  specific  in- 
stances are  not  clear.  In  particular,  very  little 
is  known  about  the  intracellular  mechanisms 
of  freezing  resistance  in  broad-leaved  plants 
that  remain  conspicuously  green  and  meta- 
bolically  active  throughout  the  winter 
months  of  cool  temperature  regions. 

Common  mallow,  Malva  neglecta  Wallr., 
is  an  example  of  a  broad-leaved  plant  that  of- 
ten remains  green  and  succulent  throughout 
the  winter  in  north-temperature  regions.  Its 
green  leaves  can  tolerate  subfreezing  temper- 
atures without  visible  evidence  of  injury.  It 
appears  that  common  mallow  is  capable  of 
surviving  winter  cold  by  some  mechanism 
other  than  dormancy,  because  the  plant  re- 
tains the  capacity  for  photosynthesis  and  rel- 
atively high  respiration  rates  when  favorable 
conditions  are  present. 

These  observations  have  prompted  an  in- 
vestigation of  certain  photosynthetic  charac- 


teristics and  CO2  fixation  properties  in  win- 
ter-hardened mallow.  This  paper  reports  the 
activity  of  purified  ribulose  diphosphate  car- 
boxylase (RuDPCa.se)  and  the  capacity  of 
whole  leaves  to  fix  COo  from  cold-accli- 
mated, field-grown  mallow  and  from  green- 
house-grown mallow  and  tomato. 

Materials  and  Methods 

Plant  nuitcriah:  Common  mallow  is  a  per- 
ennial weed  characteristic  of  cultivated 
ground,  gardens,  yards,  and  waste  places 
throughout  the  United  States.  Introduced 
from  Eiuope,  the  weed  belongs  to  the  same 
plant  family  as  cotton,  hollyhocks,  rose  of 
sharon,  and  the  weeds  known  as  velvet-leaf 
and  flower-of-the-hour.  This  family  (Mal- 
vaceae) has  flowers  which  contain  a  tube  of 
stamens  siurounding  the  pistil  and  a  ring  of 
seeds  centered  in  persistent  floral  parts  remi- 
niscent of  a  small  flat  cheese  (thus  one  of  the 
plant's  common  names,  "cheese  weed").  The 
plant's  long  tap  root  and  its  wide  distribution 
in  relation  to  habitat  occupation  indicates  a 
wide  ecological  amplitude  in  regard  to  envi- 
ronmental stress  factors. 

The  garden  tomato,  Lycopersicon  esculen- 
tiim  L.,  variety  Ace  55,  cannot  tolerate  sub- 
freezing  temperatiues.  Tomatoes  are  warm 


'Department  of  Botany  and  Range  Science,  Bni;liam  Young  University.  Provo.  Utah  84602. 


121 


122 


Great  Basin  Naturalist 


Vol.  40,  No.  2 


season  plants;  the  Ace  55  variety  yields  very 
well  under  high  day  and  cool  night  temper- 
ature regimes. 

Method  of  sampling  plants  for  measure- 
ment of  photosynthesis  and  respiration  rates: 
Plants  of  mallow  and  tomato  were  grown  in 
the  greenhouse  at  25  C  (76  F)  day  and  20  C 
(68  F)  night  temperatures.  Mallow  plants 
were  also  grown  in  the  field  near  Provo, 
where  they  were  exposed  to  subfreezing  tem- 
peratures. Plant  samples  were  taken  from 
greenhouse  and  field  areas  during  January. 
When  harvested,  plants  were  collected 
whole,  petiole  ends  were  cut  under  water, 
and  then  they  were  placed  in  controlled  envi- 
ronment chambers  with  the  cut  ends  remain- 
ing immersed  in  water.  Only  deep  green  suc- 
culent growth  was  harvested.  Photosynthetic 
and  respiration  measurements  were  then 
made  repeatedly  as  described  below. 

COo  fixation  methods:  Rates  of  net  photo- 
synthesis (APS),  dark  respiration  (DR),  and 
CP2  compensation  points  (CP)  were  deter- 
mined in  mid-January  on  excised  shoots  cut 
under  water.  Analysis  was  made  utilizing  a 
Beckman  IR-215  infrared  gas  analyzer  and  a 
plexiglas  controlled-environment  chamber. 
Apparent  photosynthesis  (APS)  was  deter- 
mined by  the  time  required  for  the  closed 
system's  CO2  content  to  decrease  from  315  to 
275  jul  per  1  of  air.  Dark  respiration  (DR)  was 
determined  by  the  time  for  the  closed  sys- 
tem's CO2  content  to  return  from  275  to  315 
jtxl  per  1.  The  CO2  compensation  point  (CP) 
was  determined  in  a  closed  system  by  allow- 
ing the  plants  to  fix  CO2  until  no  further 
change  in  CO2  concentration  occurred  in  the 
atmosphere  of  the  lighted  plexiglas  chamber 
surrounding  the  plant.  The  assimilation 
chamber  was  housed  inside  a  large  growth 
chamber  with  lighting  provided  by  8  cool- 
white  inforescent  tubes,  8  grolux  (Sylvania) 
inforescent  tubes,  and  10  25-watt  in- 
candescent globes.  The  light  was  filtered 
through  4  cm  of  water  and  provided  an  in- 
tensity of  6.05  X  104  ergs  per  cm  per  minute 
at  leaf  height.  This  light  intensity  has  been 
indicated  to  be  saturating  for  tomato  at  315 
jul  CO2  per  1  of  air.  Chamber  parameters 
were:  temperature  -23 ±.6  C;  relative  hu- 
midity—65  ±10  percent;  wind  speed— 4.0  dm 
per  minute  (3  chamber  volumes  per  minute) 
(Brewster  1971). 


Preparation  of  extracts  from  leaf  homoge- 
nates  and  enzyme  purification:  Fully  ex- 
panded leaves  were  washed  and  their  midribs 
removed  and  blotted  dry.  From  this  step  on, 
all  procedures  were  carried  out  at  5  C.  Ap- 
proximately 3.0  gm  samples  of  leaf  tissue 
were  groimd  manually  with  cold  mortar  and 
pestle  for  5  minutes,  using  5  ml  of  0.1  M 
HEPES  (N-2-hvdroxvethvlpiperzaine-N-2- 
ethanesulfonic  acid)  buffer  pH  8.00,  0.001  M 
EDTA,  0.0001  M  DTT,  0.01  M  MgClai,  0.025 
mM  NaHC03,  per  gram  fresh  weight  leaf  tis- 
sue. The  homgenates  were  centrifuged  for  10 
minutes  at  20,000  rpm  in  a  Sorvall  model 
RC-2B  centrifuge  with  the  S-34  rotor.  The 
supernatant  was  decanted  and  used  as  the 
crude  enzyme  extract.  The  crude  enzyme  ex- 
tract from  the  low  speed  centrifugation  was 
further  clarified  by  centrifugation  at  40,000 
rpm  in  a  Spinco  model  L3-50  for  5  minutes. 
The  supernatant  fraction  was  collected.  The 
RuDPCase  enzyme  was  purified  further  by 
sedimentation  of  the  extract  into  a  sucrose 
step  gradient  consisting  of  2  ml  of  5  percent, 
2  ml  30  percent  and  3  ml  50  percent  sucrose 
solution  in  HEPES  buffer  layered  in  a  centri- 
fuge tube.  The  rapidly  sedimenting  RuD- 
PCase accumvilated  in  the  50  percent  sucrose 
layer  after  sedimentation  for  12  hours  at 
25,000  rpm  in  a  Spinco  SW-25  rotor.  The  su- 
crose-enzyme solution  was  then  passed 
through  a  10  X  1.0  cm  Sephadex  G-25  col- 
umn for  further  purification. 

Measurement  of  enzyme  activities:  Deter- 
mination of  RuDPCase  activity  was  based 
upon  fixation  of  ^^COa.  into  acid  stable  prod- 
ucts. The  assay  mixture  contained  0.01  M 
HEPES-S04  buffer  (pH  8.00)  0.01  M  MgC12, 
0.001  M  DTT,  0.02  M  NaHi^COa,  and  ribu- 
lose-1,  5-bisphosphate  in  200  [i\.  The  enzyme 
(30jul)  was  added  to  initiate  the  reaction  and 
was  allowed  to  proceed  for  10  minutes.  The 
reaction  rates  were  linear  over  this  time  peri- 
od. The  enzyme  reaction  was  stopped  by  the 
addition  of  50  ju.1  of  glacial  acetic  acid.  A  100 
jul  aliquot  of  the  reaction  mixture  was  spotted 
onto  a  strip  of  Whatman  No.  1  filter  paper 
and  dried  under  the  hood.  The  sample  was 
counted  in  a  Packard  Tri-Carb  (Model  3320) 
liquid  scintillation  counter  in  toluene  scintil- 
lation fluid.  The  counted  samples  were  cor- 
rected for  machine  efficiency  and  quenching 


June  1980 


Andersen,  Brotherson:  Cold  Acclimation 


123 


and  the  values  converted  to  disintegrations 
per  minute  (dpm). 

Determination  of  specific  activity  of  RuD- 
PCase:  RuDPCase  has  been  identified  with  a 
large,  rapidly  moving  boundary  observed  in 
the  Spinco  Model  E  ultracentrifuge  known  as 
fraction  I  protein.  The  area  of  the  Schlieren 
boundary  curve  corresponding  to  fraction  I 
protein  provides  a  means  to  determine  the 
relative  concentration  of  RuDPCase  present. 
An  estimate  of  specific  activity  per  unit  of 
enzyme  can  then  be  calculated  by  comparing 
enzvme  activitv  in  a  given  extract  with  the 
area  of  the  corresponding  fraction  I  protein 
peak  (Andersen  et  al.  1970). 

Results 

Net  photosynthesis,  dark  respiration,  CO2 
compensation  point:  Table  1  shows  that  cold- 
adapted,  field-grown  mallow  exliibited  lower 
rates  of  APS  than  greenliouse-grown  mallow 
or  tomato.  However,  rates  of  DR  were  sim- 
ilar in  all  cases.  This  resulted  in  an  APS  to 
DR  ratio  for  field-grown  mallow  of  one-half 
that  for  the  greenhouse-grown  plants.  The 
CP  shows  a  significant  increase  for  field- 
grown  Mallow  over  values  for  the  green- 
house-grown plants. 

Reaction  velocities  for  carbonate  and  ri- 
hu\osedip}\osphate  substrates:  The  com- 
parative reaction  velocities  for  purified 
"cold-adapted"  mallow  and  tomato  RuD- 
PCase at  different  carbonate  substrate  con- 
centrations are  depicted  in  Figure  1.  The  ri- 
bulosediphosphate  (RuDP)  substrate  was 
maintained  at  maximum  concentration  for 
both  enzymes.  The  mallow  RuDPCase  exliib- 
ited  higher  catalytic  capacity  per  unit  of  en- 


zyme than  tomato  RuDPCase.  The  turnover 
number  at  V.^^,  (4  u  moles  of  carbonate  sub- 
strate per  200  ul  of  reaction  mixture)  for  to- 
mato (RuDPCase)  was  calculated  at  1036 
moles  carbonate  fixed  per  mole  of  enzvme 
per  minute.  The  turnover  number  for  puri- 


2 
V  10 


i    8 


<    6 


O •MALLOW 
•■TOMATO 


Ofl        1£       2.4       32        40       4B       56       64 

CONCENTRATION     HCO,"     (>i  MOLES) 

Fig.  1.  Dependence  of  RuDPCase  activity  in  pmified 
extracts  upon  HC0.3  concentrations.  RuDPCase  was  pu- 
rified from  common  mallow  and  tomato  ACE  55  var. 
The  enzyme  activities  are  based  upon  total  amount  of 
RuDPCase  enzvme  present  in  the  reaction  mixture  as 
calculated  from  the  Schlieren  curve  of  the  sedimenting 
boundaries  in  a  model  E  ultracentrifin'e. 


T.\BLE  1.     Rates  of  net  photosynthesis,  dark  respiration,  and  the  CO2  compensation  point  of  excised  plant  shoots. 
See  text  for  description  of  plant  treatments. 


Species 


Location 


.\pparent 

photosynthesis 

(ugCo2'dnr^*min"^ ) 


Mallow 
Mallow 
Tomato  (ACE  55) 


Greenhouse 

Field 

Greenhouse 


3.2 
1.5 
2.6 


Dark  respiration 
(ugCo2'dnr^Mnin"') 


L7 
1.7 
1.3 


CC^  compensation 

point  (uI.LM 


64 
86 
68 


*.\pparent  photosynthesis  {.\?S)  was  measured  bv  determining  time  required  to  lower  closed  system  COj  concentration  from  320  ul/I  to  280  ul/1  air-1. 

^'Dark  respiration' (DR)  was  measured  by  determining  the  time  required  for  a  darkened  closed  system  to  return  COj  concentration  from  280  to  320  ul/1 
air. 

'^Compensation  point  was  measured  by  allowing  photosynthesizing  plants  to  fix  CO2  from  a  closed  atmosphere  until  no  further  change  in  CO2  concentra- 
tion could  be  obser\  ed. 


124 


Great  Basin  Naturalist 


Vol.  40,  No.  2 


fied  mallow  RuDPCase  was  calculated  at 
1400  moles  of  carbonate  fixed  per  mole  en- 
zyme per  minute.  The  differences  in  V,„^^  val- 
ues for  the  two  enzymes  were  judged  to  be 
highly  significant,  based  on  the  student-t  test 
for  measuring  differences  between  paired 
variates.  The  calculated  t  value  exceeded  the 
0.001  level  of  significance.  Only  slight  differ- 
ences between  the  corresponding  Km  values 
could  be  observed. 

The  reaction  velocities  of  mallow  and  to- 
mato RuDPCase  for  different  ribulosediphos- 
phate  substrate  concentrations  are  graphed  in 
Figure  2.  The  shapes  of  the  reaction  velocity 
curves  for  tomato  and  Mallow  RuDPCse  are 
similar.  The  mallow  enzyme  exhibited  a  sig- 
nificantly higher  V^,^,  value.  Both  enzymes 
showed  substrate  inhibition  at  RuDP  substate 
concentrations  higher  than  8  u  moles  per  200 
jLil  of  reaction  mixture.  The  significance  of 
the  differences  between  the  two  reaction  ve- 
locity cmves  in  Figure  2  was  measured  by 
the  student-t  test  for  paired  variates.  The  cal- 
culated t  value  exceeded  the  0.01  level  of  sig- 
nificance. The  Km  values  for  substrate  con- 
centration at  half  maximal  velocity  were 
slightly  higher  for  tomato. 

Effect  of  temperature  on  reaction  velocity 
with  purified  RuDPCase  enzyme:  Purified  to- 
mato and  mallow  RuDPCase  enzyme  extracts 
were  compared  for  catalytic  velocities  at  re- 
action temperatures  ranging  from  4  C  to  63 
C  (Figure  3).  Purified  mallow  RuDPCase  had 
significantly  higher  catalytic  activity  per  unit 
of  enzyme  under  the  temperatine  range  of  4 
to  25  C.  The  calculated  student-t  value  for 
differences  between  paired  variates  exceeded 
the  0001  level  of  significance.  On  the  other 
hand,  purified  tomato  enzyme  showed  signifi- 
cantly higher  catalytic  capacity  in  the  38  to 
53  C  temperature  range.  The  corresponding 
calculated  student-t  value  for  paired  reaction 
rates  in  the  38  to  53  C  temperature  range  ex- 
ceeded the  0.01  significance  level.  The  gener- 
al shapes  of  the  temperature  curves  for  the 
purified  mallow  and  tomato  RuDPCase  were 
quite  similar,  however,  with  heat  denatura- 
tion  for  both  enzymes  occurring  near  53  C. 

Discussion 

Our  studies  indicate  that  respiration  and 
photosynthesis  are  differentially  effected  by 


COMCCNTRATION    OF    RuOP    (ji  MOLES) 


Fig.  2.  Dependence  of  RuDPCase  activity  in  purified 
extracts  from  mallow  and  tomato  ACE  55  var.  upon 
RuDP  concentrations.  Enzyme  activities  are  normalized 
for  equal  concentrations  of  RuDPCase. 


20    25     30    35     40    45     50    55    60    65 
TEMPERATURE    °C 


Fig.  3.  Dependence  of  RuDPCase  activity  in  purified 
extracts  from  mallow  and  tomato  ACE  55  var.  upon 
temperatine.  Enzyme  activities  are  normalized  for  equal 
concentrations  of  RuDPCase. 


June  1980 


Andersen,  Brotherson:  Cold  Acclimation 


125 


cold  acclimation  in  mallow.  The  measured 
rates  of  apparent  photosynthesis  values  in 
greenhouse-thrown  mallow  were  approx- 
imately twice  the  measured  rate  of  apparent 
photosynthesis  values  of  cold-acclimated, 
field-grown  mallow.  No  differences  were  ob- 
served for  rates  of  dark  respiration  (Table  1). 
Higher  COo  compensation  points  were  ob- 
.served  for  field-grown,  cold-adapted  mallow, 
which  suggested  a  depressed  efficiency  for 
CO2  fi.xation.  However,  because  respiration 
and  photosynthesis  measurements  were  made 
at  25  C  in  the  laboratory,  it  is  possible  that 
relative  efficiencies  of  carbohydrate  accumu- 
lation would  change  at  lower  temperatures. 
The  cold-acclimated  mallow  might  under 
such  circumstances  become  relatively  more 
efficient.  Present  evidence,  however,  in- 
dicates that  cold  acclimation  in  field-grown 
mallow  is  a  matter  of  maintaining  a  steady 
state  of  metabolic  activity  rather  than  the 
rapid  acciunulation  of  carbohydrate  reserves. 
Further  studies  are  underway  to  assess  inter- 
action of  lower  temperatures  and  carbohy- 
drate accumulation  in  cold-acclimated  mal- 
low. 

Purified  RuDPCase  from  cold-adapted 
Malva  neglecta  had  the  highest  catalytic  ca- 
pacity per  unit  of  enzyme.  The  Vmax  values 
for  carbonate  and  ribulosediphosphate  sub- 
strates were  highest  for  mallow  RuDPCase 
(Figs.  2  and  3).  The  Km  values  for  tomato 
were  only  slightly  higher.  The  Km  and  Vmax 
values  for  tomato  RuDPCase  agree  in  general 
with  corresponding  reported  values  (Ander- 
sen et  al.  1970).  Although  we  cannot  yet 
compare  in  vitro  RuDPCase  activity  to  in 
vivo  CO2  fixation  without  some  misgivings, 
our  studies  suggest  that  mallow  RuDPCa.se 
would  promote  slightly  more  rapid  CO2  fix- 
ation per  unit  of  enzyme  in  vivo.  The  lower 
Km  value  for  CO2  substrate  of  RuDPCase 
from  the  cold-adapted  mallow  would  suggest 
a  higher  photosynthetic  efficiency  for  the  in- 
tact plant.  Yet  the  higher  compensation  point 
of  these  cold-adapted  plants  indicates  that 
photosynthetic  efficiency  is  depressed  in  the 
intact  leaf.  Recent  work  by  several  in- 
vestigators clearly  implicates  RuDPCa.se  as  a 
major  contributing  factor  to  the  high  com- 
pensation points  of  the  C3  species  (Ogren 
and  Hunt   1978).  Our  study  would  indicate 


that  the  higher  compensation  point  in  the 
cold-adapted  mallow  is  due  to  some  other 
factor  in  the  photosynthetic  carbon  cycle 
than  RuDPCase.  On  the  other  hand,  since  we 
are  using  purified  enzyme  for  our  studies,  it 
is  likely  that  control  molecules  that  may  af- 
fect Km  for  CO2  fixation  of  RuDPCa.se  could 
be  removed  in  our  purification  process.  In 
any  case,  if  RuDPCase  has  a  higher  Km  for 
CO2  in  vitro,  which  would  result  in  a  higher 
compensation  point  in  the  cold-adapted  mal- 
low, the  effect  does  not  persist  through  puri- 
fication of  the  enzyme.  Therefore,  at  least  a 
change  has  not  been  detected  on  the  purified 
enzyme  that  would  affect  the  compensation 
point  and  thus  be  a  basis  for  lower  photo- 
synthetic efficiency  durmg  cold  acclimation. 

The  mallow  RuDPCase  enzyme  showed 
higher  catalytic  capacity  than  tomato  RuD- 
PCase under  temperature  ranges  of  0-25  C, 
and  the  tomato  RuDPCase  enzyme  exhibited 
higher  activity  under  temperature  ranges 
40-60  C.  This  may  be  indicative  of  Mallow's 
lower-temperature  environmental  adaptation 
and  its  COo-fixing  enzymes.  Other  studies 
have  shown  that  RuDPCase  extracts  from 
plants  of  different  climatic  regions  exliibit 
correspondingly  different  temperature  reac- 
tions (Trihame  and  Cooper  1969).  Also,  we 
have  observed  that  the  RuDPCase  activities 
in  ciiide  extracts  from  Mallow  were  con- 
sistently higher  than  RuDPCase  activities  in 
tomato  crude  extracts,  (on  a  per-gram  fresh 
weight  basis).  These  results,  along  with  the 
distribution  patterns  of  these  two  species, 
suggest  that  the  temperature  interaction  of 
the  enzyme  might  be  related  in  some  way  to 
the  different  seasonal  adaptations  of  the  two 
species. 

The  results  also  suggest  that  the  process  of 
cold  acclimation  in  mallow  affects  photo- 
synthesis and  dark  respiration  differently. 
Respiration  was  not  seriously  affected,  but 
photosynthetic  capacity  per  unit  of  leaf  area 
and  photosynthetic  efficiency  were  signifi- 
cantly reduced  (Table  1).  It  may  be  hypoth- 
esized then  that  the  processes  of  cold  accli- 
mation in  mallow  either  depresses  the  in  vivo 
activity  of  RuDPCase  or  alters  in  some  way 
other  chloroplast  functions  which  affect  the 
plant's  capacity  for  photosynthesis. 


126 


Great  Basin  Naturalist 


Vol.  40,  No.  2 


Acknowledgment 

This  research  was  supported  in  part  by  the 
National  Institute  of  Health,  Grant  GM 
17868-02. 


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RECOVERY  OF  GAMBEL  OAK  AFTER  FIRE  IN  CENTRAL  UTAH 

L.  M.  Kiinzler'  and  K.  T.  Harper' 

.\bstr.\ct.—  The  height  of  oak  (Qucrcus  gdinbelii  Nutt.)  stems  was  measured  on  several  fire  scars  within  the  Uinta 
National  Forest  and  vicinity  and  compared  with  the  height  of  oak  stems  on  adjacent,  nonhumed  areas.  A  significant 
relationship  exists  between  the  recovery  rate  of  oak  after  fire  and  elevation,  with  the  recovery  rate  being  greatest  at 
low  elevations.  \  trend  also  exists  showing  that  recoverv  tends  to  be  greater  on  south  to  westerly  exposvires  than  on 
north  to  easterly  exposures. 


Gainbel  oak  {Qucrcus  gambelii  Nutt.)  is  an 
important  species  of  the  deer  winter  ranges 
of  central  Utah,  providing  both  food  and  cov- 
er for  deer  (Alhiian  1952,  Smith  1949).  How- 
ever, because  of  its  growth  habit,  it  often 
forms  impenetrable  thickets  (Allman  1952, 
Baker  1949,  Dills  1970,  Marquiss  1972, 
McKell  1950).  This,  coupled  with  its  height, 
places  most  of  the  available  browse  out  of 
reach  of  big  game  (Plummer  et  al.  1966, 
1970).  By  treating  these  dense  stands  of  oak 
with  chemical  herbicides,  fire,  or  machinery 
to  break  them  up,  the  stands  can  be  opened 
up  and  made  available  to  browsing  animals 
(Anon.  1966,  Dills  1970,  Hallisey  et  al.  1976, 
Marquiss  1971,  1972,  Plummer  et  al.  1966, 
1970,  Price  1938).  Because  oak  stands  recov- 
er rather  rapidly  after  these  treatments,  it  is 
necessary  to  determine  a  rotation  period  for 
treatment  to  maintain  an  optimum  amount  of 
browse  for  wildlife  (McKell  1950,  Plummer 
et  al.  1966,  1970). 

Literature  Review 

In  central  Utah,  Gambel  oak  has  been  re- 
ported to  occur  in  almost  pure  stands  from 
5000  (1525  m)  to  8000  (2440  m)  feet  eleva- 
tion along  the  Wasatch  Range  (Allman  1952, 
Baker  1949,  McKell  1950).  This  area  con- 
stitutes a  large  portion  of  the  deer  winter 
range  in  the  area  (Allman  1952,  Anon.  1966, 
Dills  1970,  Hallisey  et  al.  1976,  Plummer  et 
al.  1966,  1970,  Smith  1949). 

Treatments  of  oak  using  fire,  herbicides,  or 


machinery  to  destroy  the  oak  canopy  result 
in  prolific  sprouting,  with  several  stems  re- 
placing each  preexisting  stem.  Impenetrable 
thickets  often  result  from  such  treatments 
(Allman  1952,  Baker  1949,  Dills  1970,  Mar- 
quiss 1972,  McKell  1950).  Yet,  treatments  can 
be  effective  in  improving  deer  range.  When  a 
follow-up  program  is  used,  such  as  seeding 
with  competitive  herbs  and  grasses,  the  ben- 
efits of  the  treatment  can  be  prolonged  for 
over  15  years  (Anon.  1966,  Dills  1970,  Hal- 
lisey et  al.  1976,  Marquiss  1971,  1972,  Plum- 
mer et  al.  1966,  1970,  Price  1938).  By  treat- 
ing oak,  deer  use  can  be  increased  up  to  four 
times,  but  deer  use  declines  as  the  time  from 
treatment  increases  (Anon.  1966,  Hallisey  et 
al.  1976,  Price  1938). 

Methods 

The  height  of  oak  stems  was  measured  on 
several  stands  within  the  Uinta  National  For- 
est and  vicinity.  One  half  of  these  stands 
were  located  in  oak  stands  that  had  burned  3 
to  15  years  ago.  The  other  half  of  the  stands 
were  located  in  nonburned  areas  adjacent  to 
each  burned  stand  considered.  Unburned 
stands  were  selected  so  as  to  have  the  same 
slope,  exposure,  and  elevation  as  the  burned 
stand  that  each  was  paired  with.  Mea.sure- 
ments  were  taken  along  a  100-foot  transect  at 
16-foot  intervals,  with  the  oak  stem  that  was 
nearest  to  the  point  being  measured.  Slope 
varied  from  20  to  70  percent  and  elevation 
ranged  from  5100  feet  (1555  m)  to  6800  feet 


'Department  of  Botany  and  Range  Science,  Brigham  Young  University,  Provo,  Utah  84602. 


127 


128 


Great  Basin  Naturalist 


Vol.  40,  No.  2 


(2070  m)  in  elevation  among  the  stand  pairs 
considered.  Exposure  also  varied,  with  one- 
half  of  the  stands  having  a  south  to  west  ex- 
posure and  the  other  half  having  a  north  to 
east  exposure.  A  percent  recovery  value  was 
calculated  for  each  pair  by  dividing  the  aver- 
age height  of  the  oak  in  the  fire  scar  by  the 
average  height  of  the  oak  in  the  nonburned 
area.  A  recovery  rate  was  then  calculated  by 
dividing  the  percent  recovery  value  by  the 
age  of  the  fire  scar  (Table  1).  To  minimize 
the  variation  among  recovery  rates  caused  by 
the  nuisance  factors  of  age,  elevation,  slope, 
and  site,  recovery  rates  were  also  calculated 
on  a  uniform  hill  within  the  Wallsburg  Burn 
(Table  2).  Three  transects  were  placed  in  the 
Ijumed  area  at  500-foot  (152  m)  intervals  in 
elevation. 

Results 

In  comparing  the  recovery  rates  from  the 
stands  throughout  the  Uinta  National  Forest 


and  vicinity,  a  significant  correlation  (power 
equation)  exists  between  elevation  and  oak 
recovery  rates  (r  =  -.85,  .01  <  P  <  .05,  Fig. 
1).  Using  data  collected  from  the  Wallsburg 
Burn,  a  similar  analysis  confirms  that  a  signif- 
icant correlation  (linear  equation)  exists  be- 
tween elevation  and  the  recoverv  rates  of  oak 
stems  (r  =  .99,  P  <.01,  Fig.  2)'.  Both  sets  of 
data  show  that,  as  elevation  is  increased,  the 
recovery  rates  decrease.  Although  there  is  in- 
dication that  stands  on  south  to  west  expo- 
sures have  faster  recovery  rates,  the  differ- 
ence between  the  recovery  rates  of  these 
stands  and  those  with  north  to  east  exposures 
is  not  significant  (Fig.  3). 

Discussion 

Comparing  the  results  of  the  two  sets  of 
data,  it  seems  clear  that  elevation  has  a 
strong  influence  on  oak's  rate  of  recovery  fol- 
lowing fire.  The  variation  between  recoverv 
rates  in  Figure  2  is  primarily  due  to  elevation 


Table  1.  Annual  recoverv  rates  in  percent  for  oak  on  burns  scattered  throuij;hout  the  Uinta  National  Forest  and 
nearby  areas. 


Average 

Average 

height  of 

Age  of 

height  of 

oak  in  non- 

Recoverv 

Stand 

Elevation 

burn 

oak  on  burn 

burned  area 

Percent 

rate 

# 

(m) 

Exposure 

(years) 

(cm) 

(cm) 

recovery 

(percent) 

1 

1,740 

East 

3 

77 

579 

13.3 

4.4 

2 

1,800 

North 

15 

140 

320 

43.7 

2.9 

3 

1,560 

South 

3 

145 

274 

52.8 

17.6 

4 

1,770 

South 

4 

83 

343 

24.2 

6,1 

5 

1,650 

East 

3 

84 

290 

28.9 

9.6 

6 

1,680 

West 

3 

61 

135 

45.0 

15.0 

7 

1,680 

East 

3 

83 

290 

28.5 

9.5 

8 

1.770 

West 

4 

88 

399 

22.1 

5.5 

9 

1,860 

East 

6 

116 

533 

21.7 

3.6 

10 

2,070 

West 

4 

37 

320 

11.4 

2.9 

T.\BLE  2.   Percent  recovery  values  for  oak  on  a  unih)rni  hill  in  the  Wallsburg  Burn.  This  stand  had  burned  four 
years  earlier. 


Transect 


1 

1-a 

2 

2-a 

3 

3-a 


Elevation 

Status 

Average  height 

Percent  recovery 

1,770  m 

Burned 

88.4  cm 

22.2 

1,770  m 

Unburned 

398.8  cm 

1,920  m 

Burned 

59.9  cm 

16.8 

1,920  m 

Unburned 

356.6  cm 

2.070  m 

Burned 

■36.6  cm 

11.4 

2,070  m 

Unburned 

320.0  cm 

June  1980 


KuNZLER,  Harper:  (jAmbel  Oak 


129 


alone.  Even  with  the  nuisance  variables  men- 
tioned earlier,  it  will  he  noted  that  elevation 
is  a  significant  factor  for  recovery  rates  of 
oak  (Fig.  1). 

Possible  reasons  for  the  change  in  oak  re- 
coverv  rates  with  elevation  include  the  fol- 
lowing: (1)  the  species  is  approaching  its  up- 
per elevational  limit  on  some  of  the  burns 
and,  because  of  this,  its  growth  may  be  slow- 
er; (2)  more  moisture  and  nutrients  may  be 
available  to  plants  at  the  bottom  of  slopes  be- 
cause of  precipitation's  surface  nuioff  and  at- 
tendant erosion,  nutrient  transport,  and  re- 
sultant differences  in  soil  depth  at  the  top 
and  bottom  of  the  slope;  and  (3)  the  shorter 
growing  season  at  the  higher  elevations  gives 
less  time  for  growth  there.  There  may  be  oth- 
er reasons  or  a  combination  of  reasons  for 
this  phenomenon.  In  anv  event,  the  relation- 
ship is  strong  and  has  management  implica- 
tions. 

In  the  winter,  deer  in  the  Uinta  National 
Forest  and  nearby  areas  primarily  use  south- 
and  west-facing  slopes  at  lower  elevations 
(Bruce  Giunta  and  Jordon  Pederson,  Utah  Di- 
vision of  Wildlife  Resources,  and  Juan  Spil- 
lett,  Uinta  National  Forest,  pers.  comm.). 
Such  areas  coincide  with  situations  where 
oak  recovery  is  most  rapid.  Thus,  manage- 
ment programs  to  regenerate  oakbrush  on 
deer  winter  ranges  in  our  area  may  be  short- 
lived. If  Gambel  oak  is  to  be  manipulated  to 
improve  deer  range  using  conventional  meth- 
ods in  this  area,  a  follow-up  program  that 
will  retard  oak  recovery  should  be  used.  The 


20-1 


15 


10- 


.85 


1,530m 


1,740  m  1,950  m 

Elevation 


2,130m 


Fig.  1.  The  relationship  between  recovery  rates  of 
oak  and  elevation  on  various  burns  throughout  the  Uinta 
National  Forest  and  nearbv  areas. 


significant  increase  in  time  between  major 
treatments  will  thus  minimize  management 
costs. 


Conclusions 

Elevation  is  a  significant  factor  in  affecting 
the  recovery  rate  of  oak  after  fire,  with  high- 
er elevation  stands  recovering  more  slowly. 
Recovery  takes  from  6  to  .35  years  in  this 
area,  with  a  modal  recovery  time  of  about  15 
years. 


25H 


20- 


15- 


10 


r  =    .9998 


1,770m 


1,920m 
Elevation 


2,070  m 


Fig.  2.  The  relationship  between  recovery  rate  and 
elevation  for  oak  in  the  Wallsburg  Burn  area. 

10-1 


0) 
4-1 
CD 

P^ 

>, 
U 

> 
o 
o 

0) 

Pi 


> 

<: 


5- 


s-w  n -e 

Exposure 

Fig.  .3.  Histograph  of  the  average  recovery  rate  of  oak 
stands  with  south  to  west  exposures  and  north  to  east  ex- 
posures on  burns  throughout  the  Uinta  National  Forest 
and  nearby  areas. 


130 


Great  Basin  Naturalist 


Vol.  40,  No.  2 


Acknowledgments 

The  authors  wish  to  thank  the  Uinta  Na- 
tional Forest  and  the  Utah  Division  of  Wild- 
life Resources  for  permitting  access  to  the 
study  areas.  Funding  for  this  study  was  pro- 
vided by  the  Uinta  National  Forest  (Supple- 
ment to  Cooperative  Agreement  12-11-204- 
31).  The  authors  also  express  thanks  to  Karl 
McKnight  for  help  in  the  statistical  analysis 
of  the  data. 

Literature  Cited 

Allman,  v.  p.  1952.  A  preliminary  study  of  the  vegeta- 
tion in  an  exclosure  in  the  chaparral  of  the 
Wasatch  Mountains,  Utah.  Unpublished  thesis, 
Brigham  Young  University,  Provo,  Utah.  236  pp. 

Anon.  1966.  Conversion  of  thicket  covered  areas  to  pro- 
ductive grazing  lands.  USDA,  Forest  Service,  Int. 
Reg.  Range  Imp.  Notes  11(4):  1-4. 

Baker,  W.  L.  1949.  Soil  changes  associated  with  recov- 
ery of  scrub  oak  (Querctis  gambelii)  after  fire.  Un- 
published thesis,  Univ.  of  Utah,  Salt  Lake  City, 
65  pp. 

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

Dills,  G.  G.  1970.  Effects  of  prescribed  burning  on  deer 
browse.  J.  Wildl.  Manage.  34:540-545. 


Eastmond,  R.  J.  1968.  Vegetational  changes  in  a  moun- 
tain brush  commimity  of  Utah  during  18  years. 
Unpublished  thesis,  Brigham  Yoiuig  Univ.,  Provo, 
Utah,  64  pp. 

Hallisey,  D.  M.,  and  G.  W.  Wood.  1976.  Prescribed 
fire  in  scrub  oak  habitat  in  central  Pennsylvania. 
J.  Wildl.  Manage.  40:507-516. 

Marquiss,  R.  W.  1971.  Controlling  Gambel  oak  on 
rangelands  of  southwest  Colorado.  Colo.  .'Vgri. 
Exp.  Stat.  Progress  Report  #PR71-9.  2  pp. 

1972.  Soil  moisture,  forage,  and  beef  production 

from  Gambel  oak  control  in  Southwest  Colorado. 
J.  Range  Manage.  25:146-150. 

McKell,  C.  M.  1950.  A  study  of  plant  succession  in  the 
oakbnish  (Querctis  gambelii)  zone  after  fire.  Un- 
published thesis,  Univ.  of  Utah,  Salt  Lake  City. 
74  pp. 

Plummer,  a.  p.,  D.  R.  Christensen,  and  S.  B.  Monsen. 
1966.  Highlights,  results,  and  accomplishments  of 
game  range  restoration  studies.  Utah  State  Dept. 
of  Fish  and  Game.  Publ.  No.  67-4:7-9. 

Plummer,  A.  P.,  R.  Stevens,  and  K.  R.  Jorgensen.  1970. 
Highlights,  results,  and  accomplishments  of  game 
range  restoration  studies.  Utah  State  Dept.  of 
Fish  and  Game.  Publ.  No.  70-3:26-30. 

Price,  R.  1938.  Artificial  reseeding  on  oakbrush  range  in 
Central  Utah.  USDA  Circ.  No.  458.  19  pp. 

Smith,  J.  G.  1949.  Deer  forage  observations  in  Utah.  J. 
Wildl.  Manage.  13:314-315. 


RELATIONSHIPS  AMONG  TOTAL  DISSOLVED  SOLIDS,  CONDUCTIVITY, 
AND  OSMOSITY  FOR  FIVE  ARTEMIA  HABITATS  (ANOSTRACA:  ARTEMIIDAE) 

Nicholas  C.  Collins'  and  Gray  Stirling' 

.\bstract.—  Graphs  allowing  interconversion  between  various  physical  chemical  parameters  are  presented  for  five 
Aiicmia  habitats  in  the  western  USA.  Both  the  mean  osmosity  and  its  typical  yearly  range  differ  greatly  among  habi- 
tats. Consequently,  Arteiiiici  populations  provide  an  interesting  opportimity  to  study  physiological  and  life  history 
adaptations  to  differing  degrees  of  habitat  stability. 


Populations  of  Aiiemia,  the  brine  shrimp, 
exist  in  isolated  hypersaline  environments 
throughout  most  of  the  world  (McCarraher 
1972).  Their  source  waters  span  the  entire 
natural  spectrum  of  ion  ratios  (Cole  and 
Brown  1967)  and  range  from  the  massive  and 
relatively  permanent  Great  Salt  Lake  to  tem- 
porary ponds  50  m  to  diameter  (e.g.  Broch, 
1969;  Khalaf  et  al.  1977).  Not  surprisingly, 
the  individual  populations  exhibit  morpholo- 
gical, physiological,  developmental,  and  gen- 
etic differences  that  indicate  they  are  locally 
adapted  (e.g.  D'Agostino  1965,  Clark  and 
Bowen  1976,  Glaus  et  al.  1977,  Collins  1977). 

Because  the  resting  cysts  of  these  popu- 
lations are  easy  to  collect,  transport,  store, 
and  hatch,  Artemia  populations  are  excellent 
subjects  for  comparative  studies  of  genetics 
(Barigozzi  1974,  Clark  and  Bowen  1976), 
physiology  of  ion  regulation  (e.g.,  Geddes 
1975a,  b,  c),  and  life  history  tactics  (Collins 
1977,  Glaus  et  al.  1977).  Many  such  studies 
have  involved  comparisons  of  the  perform- 
ance of  various  strains  grown  in  a  common 
medium,  usually  diluted  or  concentrated  sea- 
water.  An  alternate  approach,  which  accom- 
modates some  strains  that  will  not  grow  well 
in  sea  water,  involves  growing  them  each  in 
their  own  source  water,  but  at  a  common  os- 
motic pressure.  This  alternative  requires 
measurement  of  the  osmotic  characteristics 
of  various  dilutions  of  water  from  each 
source  lake,  a  time-consuming  process  requir- 
ing osmometers  that  are  both  expensive  and 


uncommon.  To  reduce  the  necessity  for  fu- 
ture osmometric  measurements  for  studies  of 
Artemia  populations  in  the  western  United 
States,  this  paper  presents  relationships  be- 
tween osmosity  and  more  easily  measured  pa- 
rameters for  source  waters  of  five  popu- 
lations. Data  on  pH  changes  with  source 
concentration  and  information  on  the  natural 
range  of  concentrations  for  each  source  are 
also  presented. 

Methods 

Source  waters  from  Arizona,  New  Mexico, 
Nebraska,  and  Washington,  collected  during 
1976  and  1977,  were  filtered  and  diluted  or 
concentrated  by  evaporation.  Locations  for 
each  source  are  specified  in  the  references  in 
Table  1.  Total  dissolved  solids  (TDS)  concen- 
trations were  determined  by  evaporating  five 
or  ten  ml  samples  to  a  constant  weight  at  100 
C.  Salt  scale  had  to  be  repeatedly  broken  to 
insure  completion  of  the  drying  process. 
Drying  at  temperatures  higher  than  100  G 
resulted  in  steam  explosions  within  salt  mas- 
ses that  scattered  the  salt  and  biased  the  de- 
terminations. 

Conductivity  meter  readings  were  con- 
verted to  specific  conductance  at  20  G  using 
an  NaCl  calibration  curve  based  on  Wolf, 
Brown,  and  Prentiss  (1975). 

Specific  gravity  at  20  C  was  measured 
gravimetrically  using  individually  calibrated 
50  ml  volumetric  flasks.  Each  determination 


'Department  of  Zoolog)'  and  Erindale  College,  University  of  Toronto,  3359  Mississauga  Road,  Mississauga,  Ontario  L5L  1C6. 


131 


132 


Great  Basin  Naturalist 


Vol.  40,  No.  2 


is  the  average  of  duplicate  measurements. 

Osmosity,  the  molar  concentration  of  NaCl 
having  the  same  freezing  point  or  osmotic 
pressure  as  the  measured  solution,  was  deter- 
mined with  a  Wescor  vapor  pressure  os- 
mometer. The  microvoltmeter  output  was 
calibrated  with  a  series  of  NaCl  solutions 
whose  osmotic  properties  were  assumed  to 
correspond  with  Wolf  et  al.  (1975).  Two  to 
five  determinations  were  averaged  for  each 
data  point  in  the  figures. 


Results 

For  three  of  the  five  sources  (Figs.  1-3)  os- 
mosity is  closely,  linearly  related  to  TDS. 
Where  Y  is  osmosity,  X  is  TDS,  n  is  the  num- 
ber of  measurements,  and  r  is  the  product- 
moment  correlation  coefficient,  the  relation- 
ships for  Penley  Lake  (Washington),  Green 
Pond  (Arizona),  and  Lily  Lake  (Nebraska)  are 
respectively: 


E 
u 

(/) 

o 

E 
E 


TOTAL  DISSOLVED  SOLIDS 

Fig.  1.  Relationships  among  TDS,  conductivity  (open  circles),  and  osinositv  (closed  circles)  for  Penley  Lake,  near 
Oniak,  Washington  (Broch  1969).  Numbers  below  upper  margin  are  specific  gravity  measurements  and  those  above 
lower  axis  are  pH  values  for  indicated  TDS  levels. 


June  1980 


Collins,  Stirling:  Artemia  Habitats 


133 


Y  =  .0081  X  + 

.0176 

n  =  23 

r  =  .995 

Y  =  .0160  X  + 

.0141 

n  =  19 

r  =  .995 

Y  =  .0103  X  + 

.0439 

n  =  30 

r  =  .998 

The  TDS-osmosity  relationships  for  the  other 


two  lakes  (Figs.  4,  5)  did  not  appear  to  be  lin- 
ear, and  were  drawn  by  eye.  Similarly,  the 
curvihncar  TDS-conductivity  relationships 
lor  all  lakes  were  drawn  by  eye. 


E 
a 

«/) 

o 

"I 
E 


>- 
> 

a 

o 
u 


160 

40              80 

120 

160 

1         i    1    II       1 

'     1    ' 

1         1         1 

- 

1.0390- 
1.0465  • 

1 

CD 

CO 

eo 

yv 

- 

140 

- 

/  / 

"~ 

- 

o/ 

°  / 

- 

120 

- 

GREEN  POND 

o    / 
/o 

/ 

— 

■ 

jPo           / 

100 

f    /• 

80 

- 

/• 

- 

60 

- 

• 

— 

40 

- 

— 

20 

-   [ 

— 

-/ 

9.8 

1         1         ll       1 

9.4 
ll         1 

1  1  1 

— 

3.20 


-  2.80 


-  2.40 


-  2.00      _. 

o 
E 

-11.60    r" 


-  1.20 


-  .80 


CO 

o 

C/) 

o 


0  40  80  120  160  200 

TOTAL  DISSOLVED  SOLIDS  (  g/l  ) 


Fig.  2.  Relationships  among  physical-chemical  parameters  for  Green  Pond,  near  St.  Johns,  .\rizona  (Cole  and 
Whiteside  196.5).  Legend  as  in  Figure  1. 


134 


Great  Basin  Naturalist 


Vol.  40,  No.  2 


Discussion 

Generally,  conductivity  is  the  most  con- 
venient indicator  of  osmosity  or  TDS.  The 
natural  concentrations  of  these  soiuce  lakes 
fall  well  above  the  highest  scale  of  most  exist- 
ing conductivity  meters,  but  Figs.  1-5  will  al- 
low rough  predictions  of  TDS  or  osmosity 
from  conductivity  measurements  of  samples 


diluted  to  within  the  range  of  such  meters.  In 
such  cases  the  errors  in  any  prediction  based 
on  the  graphs  will  be  multiplied  by  the  dilu- 
tion factor.  At  the  highest  source  concentra- 
tions conductivity  is  not  a  precise  predictor 
of  TDS  or  osmosity  (Figs.  4,  5).  For  such  solu- 
tions an  accurate  dilution  can  be  made  for 
conductivity  measurement,  or  TDS  can  be 
measured  directly.  Inaccuracies  in  TDS  mea- 


O 

E, 

>- 

CO 

o 

C/3 

O 


40  80  120  160  200 

TOTAL  DISSOLVED  SOLIDS  (g/l) 


240 


Fit;.  3.   Kelationships  among  pliysical-eheniical  paraiufteis  ior  Lilv  Lake,  near  Alliance,  Nebraska  (McCarraher 
1970).  Legend  as  in  Figure  L 


June  1980 


Collins,  Stirlinc;:  Arteml\  Habitats 


135 


siirements  from  water  inclusions  in  the  crys- 
talizing  salt  mass  can  be  eliminated  by  using 
a  micrometer  syringe  for  applying  precisely 
measured  small  volumes  (<  1  ml)  to  a  pre- 
weighed  filter  paper  circle.  Such  a  technique 
will  also  allow  much  faster  determinations 
tlian  the  one  we  used. 

Hydrometer    measurements    of   specific 


gravity  are  often  reported  as  a  measure  of 
concentration.  Although  this  method  appears 
to  be  quick,  straightforward,  and  suitable  for 
field  measurements,  our  experience  indicates 
even  marginally  accurate  results  require 
careful  control  of  water  temperature,  wind, 
and  cleanliness  of  the  hydrometer  that  to- 
gether preclude  most  field  measurements.  In 


160 


140 


^        120 

E 
u 

o 


E 
E 


100 


80 


60 


40 


20  - 


50           100          150          200         250         300         350 
T — ^ — T] — rpi — I 1 1 1 1 1 1 1 — I — r 


2.0 


4.0 


50     100    150    200    250    300    350    400 


TOTAL  DISSOLVED  SOLIDS  (g/l) 


CO 

o 

CO 


Fig.  4.  Relationships  among  phvsical-chemical  parameters  of  Jesse  Lake,  near  .Alliance,  Nebraska  (McCarraher 
1970).  Legend  as  in  Figure  1. 


136 


Great  Basin  Naturalist 


Vol.  40,  No.  2 


the  laboratory,  TDS  measurements  or  gravi- 
metric density  determinations  are  almost  as 
easy  as  hydrometer  readings,  if  a  con- 
ductivity meter  is  not  available. 

A  summary  of  natural  osmosity  and  TDS 
levels  from  western  Artemia  habitats  (Table 


1)  indicates  that  different  populations  expe- 
rience not  only  very  different  ion  ratios  and 
mean  osmosities,  but  also  very  different  sea- 
sonal ranges  in  source  osmosity.  Western 
North  American  Aiiemiu  populations  there- 
fore  provide   an   interesting   opportunity   to 


E 
o 

o 

"I 

E 


>- 

> 
I— 

Q 
O 


50          100          150 

200          250 

300 

225 

1 
1.0827-= 

1 

52   /     52-^ 

1      1 

-75 8- 

200 

ZUNI  SALT  LAKE 

/ 

/ 

: 

175 

■  1 

' 

^ 

150 

/    • 

/ 

- 

125 

/  / 

- 

100 

-        /  / 

- 

75 

■  /y 

- 

50 

'  1/ 

- 

25 

r 

- 

•                 8.7                    \ 

).5 

8.1 

7.9 

1       1       ill       1.  .   J- 

1    1 

1       1     li 

1       1 

4.5 


-  4.0 


3.5 


-13.0       =; 

o 
H2,5        ^ 


-  2.0 


-   1.5 


-   1.0 


-    .5 


CO 

o 

C/9 

O 


0  50  100  150  200  250         300  350 

TOTAL  DISSOLVED  SOLIDS  (g/l) 

Fig.  5.  Relationships  amonn  pliysical-cliemital  parameters  of  Zuiii  Salt  Lake,  near  Queniado,  New  Mexico  (Brad- 
bury 1971).  Legend  as  in  Figure  1. 


June  1980 


Collins,  Stirling:  Artemia  Habitats 


137 


identity  the  genetic,  physiological,  and  life 
history  characteristics  that  evolve  in  response 
to  variability  of  environmental  conditions, 
and  in  response  to  substantial  differences  in 
length  of  the  growing  season. 

Note  added  in  proof:  Recent  talks  with 
Nebraska  residents  revealed  that  the  lake  re- 
ferred to  in  this  paper  and  in  Collins  (1977) 
as  Lily  Lake  of  McCarraher  (1970,  1972)  is 
actually  an  imnained  smaller  lake  slightly 
northwest  of  the  tnie  Lily  Lake.  The  range  of 
osmosity  for  this  lake  is  unknown;  therefore 
the  figures  for  it  in  Table  1  should  be  dis- 
regarded. 

Acknowledgments 

The  National  Geographic  Society,  the  Nat- 
ural Sciences  and  Engineering  Research 
Council  of  Canada,  and  the  University  of  To- 
ronto supported  this  work  financially.  1  thank 
Mr.  Drew  Piatt  for  permission  to  sample 
Green  Pond,  Gerald  Cole,  J.  P.  Bradbiuy,  and 
Sarane  Bowen  for  information  on  sampling 
sites,  and  R.  W.  Cummins  and  J.  Svoboda  for 
access  to  the  o.smometer.  LeeAnne  Wilson 
and  John  Stoneman  provided  excellent  tech- 
nical assistance. 


Liter.\ture  Cited 

Baricozzi.  C.  1974.  Aitcinui:  a  survey  of  its  significance 
in  genetic  prolileins.  Evolutionary  Biol.  7: 
221-252. 

Bhadburv,  J.  P.  1971.  l.iniiiology  of  Zuni  Salt  Lake,  New 
Mexico.  Geol.  Soc.  Anier.  Bull.  82:  .379-398. 

Broch.  E.  S.  1969.  The  osmotic  adaptation  of  the  fairy 
slirinip  Brcinrhinccta  campestris  Lynch  to  salaine 
astatic  waters.  Linmol.  Oceanogr.  14:  48.5-492. 

Clark,  L.  S.,  and  S.  T.  Bowe.n.  1976.  The  genetics  of 
Artemia  xalina.  VIL  Reproductive  isolation.  J. 
Hered.  67:  38.5-.388. 

Claus,  C,  F.  Benijts,  a.nd  P.  Sorgeloos.  1977.  Com- 
parative study  of  different  geographical  strains  of 
the  brine  shrimp,  Artemia  salimi,  pp.  91-105.  In: 
Jaspers,  E.  (ed.).  European  Mariculture  Society 
special  publication  No.  2.  Bredene,  Belgium. 

Cole,  G.  \.,  a.nd  M.  C.  Whiteside.  1965.  Kiatuth- 
lanna— a  limnological  appraisal.  II.  Chemical  fac- 
tors and  biota.  PJateau  .38:  .36-48. 

Cole,  G.  \.  and  R.  J.  Brown.  1967.  The  chemistry  of 
Artemia  habitats.  Ecology'  48:  8.58-861. 

Collins,  N.  C.  1977.  Ecological  studies  of  terminal 
lakes— their  relevance  to  problems  in  limnology 
and  population  biology,  pp.  411-420.  In:  D.  C. 
Greer  (ed.).  Desertic  terminal  lakes.  Utah  Water 
Research  Laboratory,  Logan,  Utah. 

D".\(;osTiNo,  \.  1965.  Comparative  studies  of  Artemia 
■salina  (development  and  physiology).  Unpub- 
lished dissertation.  New  York  Univ. 

Gedde.s,  M.  C.  197.5a.  Studies  on  an  .Australian  brine 
shrimp,  Parartemia  zietziana  Sayce  (Crustacea: 
.\nostraca)— I.  Salinity  tolerance.  Comp.  Bio- 
chem.  Physiol.  51,\:  .5.53-5.59. 


Table  1.  Comparisons  of  physical  and  chemical  characteristics  of  six  Artemia  .sources  in  the  western  USA. 


Water  body 

Major  salt 

Great  Salt  Lake, 

Utah 

NaCl 

Jesse  Lake. 
Nebraska 

NaCOj 

Lily  Lake. 
.Nebraska 

NaC()3-Cl 

Green  Pond, 

NaC03-Cl 

.\rizona 

Penley  Lake, 

Na2S()4 

Washington 

Zuni  Salt  Lake, 
New  Mexico 

NaCl 

Observed^  TDS 

range  (g/1) 


"Typical"  annual 
osmosity  range^ 

(moles/1)  References 


130-300 


.52-87 


12-69 


61-112 


33-230 


17.5-.350 


2.-2.25 


..57-.90 


.16-. 76 


.98-1.8 


.24-dryness 


Handv  and  Hahl 

1966  ' 

McC:arraher  1970. 
1972 

.McCarraher  1970, 
1972 

Cole  and 
Whiteside  1965 

Broch  1969 


..35-5.3-1-  Bradbury  1971 


Range  of  values  recorded  in  literature  and  personal  observations,  .\nnual  range  is  usually  smaller. 

IncKides  only  the  period  during  which  active  Artemia  are  present.  For  some  lakes,  only  one  year's  data  are  available,  so  nothing  Ls  known  about  be- 
tween-vear  differences. 


138 


Great  Basin  Naturalist 


Vol.  40,  No.  2 


1975b.  Shidies  on  an  Australian  brine  shrimp, 

Parartemia  zietziana  Sayce  (Crustacea:  Anost- 
raca)— II.  Osmotic  and  ionic  rei;iilation.  Comp. 
Biochem.  Physiol.  Sl.A:  561-571. 

1975c.  Studies  on  an  Australian  brine  shrimp, 

Parartemia  zietziana  Sayce  (Cnistaceae:  .\nost- 
raca)— III.  The  mechanisms  of  osmotic  and  ionic 
regulation.  Comp.  Biochem.  Physiol.  51A: 
573-578. 

Handy,  A.  H.,  a.nd  D.  C.  Hahl.  1966.  Chemistry  of  the 
water  in  the  Great  Salt  Lake,  pp.  135-151.  In: 
Stokes,  VV.  L.  (ed.).  The  Great  Salt  Lake,  Guide- 
book 20.  Utah  Geol.  Soc.,  Salt  Lake  City. 

Khal.\f,  a.  N.,  M.  a.  Lattif,  H.  H.  Mangalo,  and  M. 
Salih.   1977.  A  bioecological  study  on  the  brine 


.shrimp  Artemia  salina  L.  (Anostraca:  Branchiop- 

oda)  in  two  inland  brine  temporary  ponds  in 

Iraq.  Bull.  Biol.  Res.  Center,  Bagdhad.  8:  .37-48. 
McCarraher,  D.  B.  1970.  Some  ecological  relations  of 

fairy   shrimps   in   alkaline   habitats   of  Nebraska. 

.\mer.  Midi.  Nat.  84:  .59-68. 
1972.  A  preliminary  bibliography  and  lake  index 

of  the  inland  mineral  waters  of  the  world.  F.\0 

Fisheries  Circular  No.  146.  .33  pp. 
Wolf,  A.  V.,  M.  G.  Brown,  and  P.  G.  Pre.ntiss.  197.5. 

Concentratiye   properties   of  aqueous   solutions: 

conversion  tables,  pp.  D218-D273.  In:  Weast,  R. 

C.   (ed.).   Handbook  of  chemistry  and  physics, 

56th  ed. 


SPAWNING  OF  THE  LEAST  CHUB  [lOTICHTHYS  PHLEGETHONTIS) 


riioiiKis  M.  Baugh' 


.\bstract.—  The  least  tluib,  loticlitlujs  phlf^clhonlis  (Cope),  a  relict  fisli  in  I'tali,  spawned  suecesslully  under  lab- 
oratory conditions. 


The  least  chub,  lotichthys  pJiIcgethontis 
(Cope),  is  a  small  (ca  5  cm)  relict  fish  (Hubbs 
and  Miller  1948)  found  only  in  a  few  local- 
ities in  the  western  desert  region  of  Utah.  In 
1973  the  Utah  Division  of  Wildlife  Resources 
classified  this  species  as  endangered.  There  is 
little  literature  on  /.  phlcgethontis  (Crawford 
1979,  Hubbs  and  Miller  1948,  Pendleton  and 
Smart  1954,  Sigler  and  Miller  1963),  and 
there  are  no  reports  of  this  species  spawning 
under  artificial  light  in  closed-svstem  aquaria. 

On  26  May  1979,  I  obtained  five  male  and 
five  female  least  chub  from  an  open  raceway 
at  the  Utah  Division  of  Wildlife  Resources  fa- 
cility, Logan,  Utah.  The  fish  had  originally 
been  collected  from  Leland  Harris  Spring 
and  the  associated  marsh  between  May  1977 
and  February  1978  by  Crawford  (1979).  I 
placed  the  fish  in  a  61  X  41  X  31  cm  (72.5 
liter)  aquarium  with  commercial  aquarium 
gravel  placed  over  a  subgravel  filter  to  a 
depth  of  about  5  cm.  The  water  was  con- 
stantly aerated  and  the  tank  was  densely 
planted  with  simulated,  broad-leafed  plants. 
The  fish  were  fed  a  mix  of  TetraMin^  Staple 
Food  and  Tetra  KrillflakesR  at  0630  and  fro- 
zen San  Francisco  Bay  Brand'^  brine  shrimp 
at  1630  each  day. 

Once  each  week,  for  a  two-hour  period, 
the  water  was  filtered  through  a  Vortex  Dia- 
tom'^ filter.  Also  once  each  week,  15  percent 
of  the  aquarium  water  was  drawn  off  and  re- 
placed with  an  equal  amount  of  aged  tap  wa- 
ter. One  oimce  of  Instead  Ocean*^  marine  salt 
mix  in  solution  was  added  each  week. 

On  15  October  1979,  I  added  two  29  cm 
long  strips  of  Living  World^  spawning  grass 


to  the  aquarium.  This  spawning  meditun  was 
examined  daily,  and  on  26  October  one 
length  of  the  medium  contained  five  mildly 
adhesive  eggs.  This  piece  of  medium  was  re- 
moved from  the  aquarium  and  placed  in  a  3.6 
liter  glass  jar  containing  water  from  the  aqua- 
rium. The  water  in  the  jar  was  mildly  agi- 
tated with  air.  On  27  October  one  and  on  28 
October  three  additional  eggs  were  removed 
from  the  aquarium  to  the  jar.  The  water  in 
the  jar  was  maintained  at  the  same  temper- 
ature as  that  in  the  aquarium. 

Free  swimming  larvae  were  first  noted  on 
31  October,  and  by  2  November  all  nine  eggs 
had  hatched.  The  larvae  were  able  to  adliere 
to  glass  and  plastic.  The  mechanism  of  at- 
tachment was  not  studied. 

The  following  conditions  existed  at  the 
time  of  spawning.  During  the  10  days  prior 
to  the  day  of  last-noted  egg  deposition,  the 
water  temperature  ranged  from  17.7  to  18.8 
C  and  averaged  18.2  C.  The  photoperiod  was 
14  hours  of  daylight  and  10  hours  of  darkness. 
Water  conditions  were:  pH  7.6,  total  alka- 
linity 84.1,  total  hardness  186,  CI-  683,  Ca 
48.4,  an  Mg  45.1.  Due  to  an  equipment  mal- 
fimction,  dissolved  oxygen  was  not  measured. 

Two  additional  spawnings  took  place  on  5 
Noveiuber  1979  and  8  November  1979.  Fif- 
teen eggs  were  gathered  from  the  former  and 
eight  eggs  from  the  latter  spawning.  In  addi- 
tion, several  other  spawnings  from  these  fish 
occurred  from  which  the  eggs  were  not 
taken. 

From  the  above,  it  appears  that  /.  pfilc^e- 
thontis  is  amenable  to  culture  in  closed-sys- 
tem aquaria  under  artificial  light. 


'1020  Custer  .\ venue,  Ogden,  Utah  84404. 


139 


140 


Great  Basin  Naturalist 


Vol.  40,  No.  2 


Acknowledgments 

I  thank  the  Utah  Division  of  Wildhfe  Man- 
agement, especially  Donald  Andriano,  for 
granting  Permit  s'CC-SL-919,  which  made 
this  work  possible;  and  Kent  Miller,  Utah  Di- 
vision of  Wildlife  Resources,  for  reviewing 
this  note. 


Literature  Cited 

Cr.\\vford,  M.  1979.  Reproductive  modes  of  the  least 
chub  [lotichtltijs  pJilcgetJiontis  Cope).  Unpub- 
hshed  thesis.  Utah  State  Univ.  79  pp. 

HuBBs,  C.  L.  .\ND  R.  R.  Miller.  1948.  Correlation  be- 
tween fish  distribution  and  hvdrographic  historv 
in  the  desert  basins  of  the  western  United  States. 
Bull.  Univ.  Utah,  Biol.  Ser.  19(7):  17-166. 

Pe.ndleton,  R.  C,  and  E.  W.  Sm.\rt.  1954.  A  stud\  of 
the  food  relations  of  the  least  chub,  lotichthijs 
phlegcthontis  (Cope),  using  radioactive  phos- 
phonis.  J.  Wildlife  Manag.  12(2):  226-228. 

SiGLER.  W.  F..  .\ND  R.  R.  Miller.  1963.  Fishes  of  Utah. 
Pages  82-84  in  Utah  Div.  ^\■ildlife  Resour.  Salt 
Lake  Citv. 


TRANSFERRIN  POLYMORPHISM  IN  BIGHORN  SHEEP, 
OVIS  CAXADEXSIS,  IN  COLORADO 


Patrick  W,  Roberts  .  Donald  J.  Nash  .  and  Robert  E.  Keiss' 

.\bstr\ct.—  Senmi  transferrins  were  analyzed  by  polyacrylamide  gel  electrophoresis  in  four  populations  of  Colo- 
rado bighorn  sheep.  Oti.s  canadensis  canadensis.  Transferrin  was  found  to  be  polymorphic,  with  two  alleles.  Tf  D 
and  Tf  E.  being  represented  in  each  of  the  four  populations.  Within  herds  the  phenot) pic  ratios  confonned  to  values 
predicted  h\-  the  Hardv- Weinberg  equilibrium,  .\niong  populations,  significant  differences  were  seen  with  respect  to 
phenotvpic  frequencies. 


Transferrin  polymorphisms  have  been  de- 
scribed in  a  number  of  breeds  of  domestic 
sheep  and  in  different  species  of  wild  sheep 
including  Ovis  canadensis,  O.  dalli,  and  O. 
mouflon  (Nadler  et  al.  1971).  They  reported 
three  transferrin  alleles,  Tf  B+,  Tf  D,  and  Tf 
E,  in  two  subspecies  of  bighorn  sheep,  O.  c. 
canadensis  and  O.  c.  mexicana.  In  14  speci- 
mens of  O.  c.  canadensis  from  Montana,  13 
had  the  Tf  DE  phenotype  and  one  sheep  was 
B^D.  Two  specimens  of  O.  c.  mexicana  from 
.\rizona  were  of  the  EE  phenotvpe. 

In  Colorado,  bighorn  sheep  historically 
ranged  over  much  of  the  central  and  western 
parts  of  the  state,  but  the  distribution  has 
been  fragmented  in  recent  times  (Armstrong 
1972),  and  there  are  now  more  tlian  30  dis- 
junct bands  occurring  in  the  less  accessible 
parts  of  the  higher  mountains.  A  study  was 
undertaken  to  characterize  electro- 
phoretically  demonstrable  genetic  variation 
in  several  sennu  proteins  and  in  hemoglobin 
of  several  disjimct  herds  to  determine  the  de- 
gree of  genetic  similarity  or  dissimilarity 
among  and  within  the  bands  sampled.  The 
present  stud\  is  a  report  of  transferrins  ob- 
served in  several  herds. 

Blood  samples  were  collected  from  four 
different  herds  in  Colorado.  The  designation 
of  the  herds  and  their  centers  of  distribution 
are  as  follows:  (1)  Poudre— north  slope  of 
Poudre  Canvon.  Larimer  Co..  (2)  Tarrvall— 
Tarryall  and  Kenosha  Mts.,  Park  Co.,  (3) 


Chalk  Creek— Chafee  Co.,  and  (4)  Gunnison- 
Gunnison  Co. 

Transferrins  were  analyzed  by  poly- 
acrvlamide  disc  gel  electrophoresis  using  the 
techniques  described  by  Smith  (1968).  Gels 
were  prepared  at  7  percent  (w/v)  concentra- 
tion. Senuu  samples  were  prepared  by  mak- 
ing serum  with  50  percent  sucrose  containing 
0.25  percent  brom  phenol  blue  as  a  tracking 
dve.  Electrophoresis  was  carried  out  in  tris- 
givcine  buffer  at  pH  9.5.  Twelve  senmi  sam- 
ples were  electrophoresed  for  26  minutes  at  3 
milliamps  per  gel  at  10  C. 

Samples  of  domestic  sheep  blood  of  known 
transferrin  tvpe  were  obtained  from  the  Sero- 
log\"  Laboratorv  of  Dr.  Stormont  of  the  Uni- 
versitv  of  California  at  Davis  and  were  used 
as  reference  sera. 

All  populations  were  polymorphic  for 
transferrin  phenotypes  (Table  I).  Two  herds. 
Chalk  Creek  and  Poudre.  had  three  pheno- 
t\pes  and  two  herds.  Tarr\ all  and  Gunnison, 
each  had  two  phenot>pes.  The  phenotypes 


Table  1.  Transferrin  phenotypic  frequencies  of  big- 
horn sheep,  Ovis  canadensis  canadensis,  in  Colorado 
(uiunbers  of  observations  in  parentheses). 


Herd 


Tf  DD 


TfDE 


TfEE 


Gunnison  (7) 

0.000 

0.857 

0.143 

Chalk  Creek  (16) 

0.313 

0.374 

0.313 

Tarrvall  (26) 

0.577 

0.423 

0.000 

Poudre (18) 

0.316 

0.526 

0.158 

'Department  of  Zoologj-  and  Entomolog\-,  Colorado  State  University .  Fort  Collins,  Colorado  80523. 
-Colorado  Di\ision  of  Wildlife,  Fort  Collins,  Colorado  80521. 


141 


142 


Great  Basin  Naturalist 


Vol.  40,  No.  2 


were  determined  to  correspond  to  those  pro- 
duced by  two  alleles,  Tf  D  and  Tf  E.  Allelic 
frequencies  ranged  from  0.43  to  0.79  for  Tf 
D  and  from  0.21  to  0.57  for  Tf  E.  Significant 
differences  among  herds  were  observed  for 
the  distribution  of  phenotypes.  Within  herds 
the  transferrin  frequencies  followed  a  Hardy- 
Weinberg  distribution.  The  proportion  of 
heterozygotes  was  high  in  all  populations, 
with  the  lowest  value  of  0.375  being  ob- 
served in  the  Tarryall  herd. 

Although  surveys  of  isozymes  in  natural 
populations  of  small  mammals  have  indicated 
considerable  genetic  variability,  relatively 
few  biochemical  studies  have  been  done  on 
large  mammals.  Bonnell  and  Selander  (1974) 
found  no  polymorphisms  in  24  presumptive 
loci  in  northern  elephant  seals.  Hetero- 
zygosites  of  0.04  have  been  reported  for  elk 
(Cameron  and  Vyse  1978),  0.04  for  moose 
(Ryman  et  al.  1977),  and  0.32  for  white-tailed 
deer  (Manlove  et  al.  1976).  These  species 
were  monomorphic  at  the  transferrin  locus 
except  for  white-tailed  deer,  which  had  23 
percent  heterozygosity. 

It  is  of  interest  that  the  bands  of  bighorn 
sheep  in  Colorado  retain  such  a  high  degree 
of  polymorphism,  at  least  at  the  transferrin 
locus,  although  the  populations  have  been 
relativelv  isolated  and  have  had  relatively 
small  population  numbers.  Some  recent  esti- 
mates of  population  size  include  Poudre, 


65-75,  Tarryall,  100,  and  Chalk  Creek, 
90-100.  Results  at  the  transferrin  locus  in- 
dicate that  inbreeding  within  the  herds  may 
not  be  a  major  problem,  although  surveys  of 
additional  genetic  loci  should  be  undertaken. 

Literature  Cited 

Armstrong,  D.  1972.  Distribution  of  mammals  in  Colo- 
rado. University  Kansas  Mas.  Nat.  Hist.,  Monogr. 
3:1-415. 

Bonnell,  M.  L..  and  R.  K.  Selander.  1974.  Elephant 
seals:  Genetic  variation  and  near  extinction.  Sci- 
ence 184:  908-909. 

Cameron,  D.  C,  .a.nd  E.  R.  Vyse.  1978.  Heterozygosity 
in  Yellowstone  Park  elk,  Cervus  canadensis.  Bio- 
cheni.  Genet.  16:651-657. 

Manlove,  M.  N.,  J.  C.  Avise,  H.  O.  Hillestad,  P.  R. 
Ramsey,  M.  H.  Smith,  and  D.  O.  Straney.  1975. 
Starch  gel  electrophoresis  for  the  study  of  popu- 
lation genetics  in  white-tailed  deer.  Proc.  19th 
Ann.  Conf.  S.  E..  Game  and  Fish  Comm. 
29:.392-403. 

Xadler,  C.  F.,  .\.  WooLF,  AND  K.  E.  Harris.  1971.  The 
transferrins  and  hemoglobins  of  bighorn  sheep 
{Otis  canadensis),  Dall  sheep  (Oris  dalli)  and 
mouflon  (Otis  niusimon).  Comp.  Biochem.  Phvs- 
iol.  40B:  567-570. 

Ryma.n,  N.,  G.  Beckman,  G.  Briun-Petersen,  and  C. 
Reuterwall.  1977.  Variability  of  red  cell  en- 
zymes and  genetic  implications  of  management 
policies  in  Scandinavian  moose  (Alces  alces).  He- 
reditas  85:157-162. 

Smith,  I.  (Ed.).  1968.  Chromatographic  and  electro- 
phoretic  techniques.  Vol.  2.  Zone— electro- 
phoresis. Wiley,  New  York,  524  pp. 


THE  GENUS  ERIOGONUM  MICHX.  (POLYGONACEAE)  AND  MICHEL  GANDOGER 

James  L.  Reveal' 

Abstract.—  Michel  Gandoger,  a  notorious  "splitter,"  proposed  several  new  entities  in  the  plant  genus  Eriogonum 
(Polvgonaceae)  in  a  1906  paper  published  in  Belgium.  Because  he  used  the  term  species  at  two  different  ranks,  in 
violation  of  the  International  Code  of  Botanical  Nomenclature,  manv  of  his  names  are  invalid.  Unlike  his  papers 
published  in  France,  this  one  was  apparentl\-  edited  so  that  some  names  were  validly  published  and  some  are  invalid. 
A  review  of  the  1906  Eriogonum  paper  shows  that  a  majority  of  both  specific  and  infraspecific  entities  proposed  are 
valid,  but  some  names,  long  in  use  and  assinned  to  be  valid,  are,  in  fact,  invalid.  Even  so,  most  of  his  names  are 
sviionyms.  Each  name  proposed  by  Gandoger  is  reviewed  and  a  nomenclatural  and  taxonomic  disposition  made.  Two 
new  combinations  are  made  within  E.  hitcolum  Greene,  var.  caninum  (Greene)  Reveal  and  var.  pedunrulatum  (S. 
Stokes)  Reveal. 


Michel  Gandoger  (1850-1926),  in  the 
words  of  Keck  (1958),  was  a  "French  abbe; 
author  of  'Flora  Europae'  (27  vols.);  volumi- 
nous writer;  amasser  of  a  huge  herbarium 
now  at  Lyon;  a  'splitter'  who  named  thou- 
sands of  unacceptable  species."  Gandoger  is 
also  mentioned  in  the  International  Code  of 
Botanical  Nomenclature  (Stafleu  et  al.  1978) 
as  an  example  of  Art.  33.4.  This  article  deals 
with  the  problem  of  misplaced  ranks  such  as 
the  use  of  the  term  species  as  a  rank  within  a 
species.  As  noted  in  the  code,  Gandoger  ap- 
plied the  term  species  and  used  binary  no- 
menclature for  two  categories  of  taxa  of  con- 
secutive rank,  the  higher  rank  being 
equivalent  to  that  of  species  in  contemporary 
literature,  while  he  misapplied  the  same  term 
to  a  lower  rank.  These  latter  terms  are  not 
validly  published.  In  his  1906  paper,  Gando- 
ger used  three  sets  of  ranks,  the  species,  spe- 
cies of  the  second  order  ("speciebus  secundi 
ordinis"),  and  a  rank  which  has  come  to  be 
identified  with  the  rank  of  variety  (Heller 
1907),  although  Gandoger  causally  refers  to 
this  latter  rank  as  forms  ("formis,"  "formas 
secernendas,  "  "formae  memorabiles,"  "mem- 
orantur  sequentes  formae,"  "varieas  formas," 
"modo  formae,"  etc.).  He  used  the  terms 
"varians"  and  "variabilis"  to  allude  to  the 
same  category.  As  for  the  species  of  the  sec- 


ond order,  in  addition  to  terming  them  "spe- 
ciebus secimdi  ordinis,"  he  also  used  the  term 
siibspeciebus  or  subspecies  to  refer  to  entities 
he  then  proceeded  to  treat  as  binomials  and 
to  designate  by  the  expression  sp.  n. 

The  following  year.  Heller  (1907)  present- 
ed a  "compilation"  of  the  Gandoger  paper, 
noting  that  he  was  presenting  only  "the  new 
forms  described  in  this  paper."  Heller  himself 
had  some  difficulties  with  the  names.  He 
mentioned,  after  Eriogonum  aspalathoides  (as 
it  was  called  bv  Heller),  that  this  name  was 
"perhaps  intended  as  a  variety  of  E.  fasci- 
ciilatiim,  but  the  way  the  name  is  printed 
.should  indicate  a  species."  Only  once  does 
Heller  include  Gandoger's  terminology  allud- 
ing to  species  of  a  lower  rank,  this  being  for 
the  phrases  published  under  E.  sphoerocepha- 
him,  "Inter  formas  varias  duae  sequentes,  ut 
subspecies,  praecipue  distingui  possunt." 

It  must  be  noted  that  Heller  was  merely 
presenting  Gandoger's  results,  and  he  cannot 
be  assumed  to  have  validated  any  of  the  oth- 
erwise invalid  Gandoger  names. 

Normally,  Gandoger  published  his  papers 
in  the  Bulletin  de  la  Societe  Botaniqiie  de 
France,  but  for  Eriogonum  his  choice  was  the 
Bulletin  de  la  Societe  Roijale  de  Botanique  de 
Belgique,  and,  because  of  this,  some  of  the 
names    may   be   validly    described.    Possibly 


'Department  of  Botany,  University  ot  Maryland.  College  Park.  Maryland  20742,  and  National  Museum  of  Natural  History.  Smitfisonian  Institution,  Wash- 
ington, D.C.  20560.  Research  supported  by  National  Science  Foundation  Grant  BMS75-13063.  This  is  Scientific  Article  .\2714.  Contribution  No.  5761  of  the 
Maryland  .\gricultural  Experiment  Station,  Department  of  Botany. 


143 


144 


Great  Basin  Naturalist 


Vol.  40,  No.  2 


they  may  because  of  the  way  the  editor  had 
the  paper  set,  thereby  removing  from  the 
original  manuscript  some  of  Gandoger's  ec- 
centricities. In  reviewing  Gandoger's  papers 
in  the  French  journal,  it  is  clear  that  the  ma- 
jority of  his  names  are  invalidly  published.  In 
reviewing  a  few  of  his  articles  containing 
species  from  the  United  States,  I  failed  to 
note  a  single  instance  when  a  species  name 
was  validly  published.  This  little  known  fact, 
at  least  in  the  United  States  and  perhaps  else- 
where, means  that  all  the  Gandoger  names 
must  be  carefully  checked  before  they  can  be 
accepted.  This  is  particularly  true  for  floristic 
workers. 

Finally,  it  is  sometimes  difficult  to  know 
exactly  where  the  various  sections  of  Gando- 
ger's paper  start  and  end  so  that  one  can  de- 
termine if  the  name  is  validly  published  or,  in 
fact,  is  not  because  of  a  sentence  presented 
several  pages  before.  I  have  tried  to  follow 
the  intent  of  Gandoger's  paper,  and  hope  I 
have  interpreted  each  segment  correctly. 

The  following  treatment  indicates  which 
names  are  valid,  which  are  not,  and  the  tax- 
onomic  status  of  each  name.  The  full  author 
citation  of  each  name  is  given,  and  the  num- 
ber following  the  name  is  the  page  on  which 
it  was  proposed  by  Gandoger. 

Erio^onum  abertianum  Torr.  in  Emory  var. 
rubenimiim  Gandoger,  185,  valid,  a  synonym 
of  var.  abertianum. 

Eriogonum  aberiianum  var.  neomexicanum 
Gandoger,  185,  valid,  a  synonym  of  var. 
abertianum. 

Eriogonum  arizonicum  Gandoger,  non 
Stokes  ex  Jones,  186,  valid,  a  synonym  of  E. 
pharnaceoides  Torr.  in  Emory  var.  pharna- 
ceoides. 

Eriogonum  alattim  Torr.  in  Sitgr.  var.  mac- 
douglasii  (Tandoger,  186,  valid,  a  synonym  of 
var.  mogollonense  Stokes  ex  Jones. 

Eriogonum  alatum  var.  brevifolium  Gando- 
ger, 186,  valid,  a  synonym  of  var.  alatum. 

Eriogonum  anemophiUim  (as  anemophijl- 
luni)  Greene  var.  cusickii  Gandoger,  186, 
valid,  a  synonym  of  E.  cusickii  M.  E.  Jones. 

Eriogonum  angulosum  Benth.  var.  rectipes 
Gandoger,  186,  valid,  a  synonym  of  E.  ma- 
cukitum  Heller. 

Eriogonum  angulosum  var.  patens  Gando- 
ger, 187,  valid,  a  synonvm  of  E.  maculatum 
Heller. 


Eriogonum  angulosum  var.  pauciflorum 
Gandoger,  187,  valid,  a  synonym  of  E.  ma- 
culatum Heller. 

Eriogonum  angulosum  var.  flabellatum 
Gandoger,  187,  valid,  a  synonym  of  E.  ma- 
culatum Heller. 

Eriogonum  annuum  Nutt.  var.  pauciflorum 
Gandoger,  187,  valid,  a  synonym  of  E.  an- 
nuum. 

Eriogonum  hitchcockii  Gandoger,  187, 
valid,  a  synonym  of  E.  annuum. 

Eriogonum  juncinellum  Gandoger,  187, 
valid,  a  synonym  of  E.  davidsonii  Greene. 

Eriogonum  salicorniodes  Gandoger,  187, 
valid,  a  good  species  restricted  to  clay  slopes 
in  southwestern  Idaho  and  adjacent  south- 
western Oregon.  It  is  most  closely  related  to 
E.  collinum  Stokes  ex  Jones  but  most  often 
confused  with  E.  baileyi  S.  Wats.  Synonyms 
of  this  species  include  E.  demissum  S.  Stokes 
(1936)  and  the  var.  romanum  S.  Stokes. 

Eriogonum  caespitosum  Nutt.  var.  alijs- 
soides  Gandoger,  188,  valid,  a  synonym  of  E. 
caespitosum. 

Eriogonum  nevadense  Gandoger,  188, 
valid,  a  synonym  of  E.  ochrocephalum  S. 
Wats.  var.  ochrocephalum. 

Eriogonum  elatum  Dougl.  ex  Benth.  var. 
limonifolium  Gandoger,  188,  valid,  a  syn- 
onym of  £.  elatum  var.  elatum. 

Eriogonum  elatum  var.  erianthum  Gando- 
ger, 188,  valid,  a  synonym  of  var.  elatum. 

Eriogonum  fasciculatum  Benth.  var.  oleifo- 
lium  Gandoger,  189,  valid,  a  synonym  of  E. 
fasciculatum  var.  fasciculatum. 

Eriogonum  fasciculatum  Benth.  var.  as- 
palathoides  Gandoger,  189,  valid,  a  synonym 
of  var.  fasciculatum.  As  noted  by  Heller 
(1907),  there  is  some  problem  with  this  name; 
that  is,  there  is  an  E.  prior  to  the  epithet. 
However,  based  upon  the  rest  of  the  text,  this 
is  clearly  as  printer's  error  and  should  be  list- 
ed as  a  variety  and  not  as  a  species,  as  was 
done  by  Heller  in  his  "compilation.  " 

Eriogonum  flavum  Nutt.  in  Fras.  var.  folia- 
tum  Gandoger,  189,  valid,  a  synonym  of  E. 
jamesii  Benth.  in  DC.  va.r.  flavescens  S.  Wats. 

Eriogoni4m  flavum  var.  linguifolium  Gan- 
doger, 189,  valid,  a  synonym  of  var.  flavum. 

Eriogonum  leucocladum  Gandoger,  189, 
valid,  a  synonym  of  E.  baileyi  S.  Wats.  var. 
divaricatum  (Gandoger)  Reveal. 

ErioEonum  heracleoides  Nutt.  var.  micran- 


June  1980 


Reveal:  Eriogonum  Nomenclature 


145 


thum  Gandoger,  189,  valid,  a  synonym  of  £. 
herocleoides  var.  angustifolium  (Niitt.)  Torr. 
&  Gray. 

Eriogonum  hcracleoides  var.  viride  Gando- 
ger, 190,  valid,  a  synonym  of  £.  umhellatum 
Torr.  var.  nevadense  Gandoger. 

Eriogonum  hcracleoides  var.  multiccps 
Gandoger,  190,  valid,  a  synonym  of  var.  hcra- 
cleoides. 

Eriogonum  hcracleoides  var.  utahensis 
Gandoger,  190,  valid,  a  synonvm  of  var.  hcra- 
cleoides. 

Eriogonum  hcracleoides  var.  rydbergii  Gan- 
doger, 190,  valid,  a  synonym  of  var.  hcra- 
cleoides. 

Eriogonum  jamesii  Benth.  in  DC.  var.  sim- 
plex Gandoger,  190,  valid,  a  good  variety  of- 
ten included  within  var.  jamesii;  this  phase  of 
the  species  is  restricted  to  southwestern  Kan- 
sas. 

Eriogonum  jamesii  var.  neomexicanum 
Gandoger,  190,  valid,  a  synonym  of  var. 
jamesii. 

Eriogonum  longifolium  Nutt.  var.  long- 
idcns  Gandoger,  190,  valid,  a  synonym  of  E. 
longifolium  var.  gnaphalifolium  Gandoger. 

Eriogonum  longifolium  var.  gnaphalifo- 
lium Gandoger,  190,  valid,  that  phase  of  the 
species  restricted  to  Florida;  often  called  E. 
floridanum  Small. 

Eriogonum  longifolium  var.  floridanum  (as 
floridana)  Gandoger,  190,  valid,  a  synonym 
of  £.  longifolium  var.  gnaphalifolium  Gando- 
ger. 

Eriogonum  longifolium  var.  lindheimeri 
Gandoger,  190,  valid,  the  western  phase  of 
the  species  now  best  considered  a  synonvm  of 
var.  longifolium. 

Eriogonum  longifolium  var.  caput-fclis 
Gandoger,  190,  valid,  a  synonym  of  var.  lon- 
gifolium. 

Under  the  heading,  Eriogonum  micro- 
thccum  Nutt.,  Gandoger  states  "inter  quas  se- 
quentes  altem  pro  subspeciebus  habueris" 
and  refers  to  the  following  six  entities  all 
published  with  binary  names. 

Eriogonum  macdougalii  Gandoger,  191,  in- 
valid, a  synonym  of  E.  microthecum  Nutt. 
var.  foliosum  (Torr.  &  Gray)  Reveal.  The 
name  was  used  by  Stokes  (1936)  at  the  varia- 
tal  rank  within  E.  microthecum,  where  she 
validated  the  name  as  E.  microthecum  var. 
macdougalii  S.  Stokes. 


Eriogonum  myrianthum  Gandoger,  191,  in- 
valid, a  synonym  of  E.  cffusum  Nutt.  var.  cf- 
fusum. 

Eriogonum  sputhulare  Gandoger,  191,  in- 
valid, a  synonym  of  E.  microthecum  Nutt. 
var.  laxiflorum  Hook.  The  name  was  used  by 
Stokes  (1936)  at  the  variatal  rank  within  E. 
microthecum,  where  she  validated  the  name 
as  E.  microthecum  var.  spathulure  S.  Stokes. 

Eriogonum  intricatum  Gandoger,  191,  in- 
valid, a  synonym  of  E.  microthecum  Nutt. 
var.  laxiflorum  Hook. 

Eriogonum  Iwlichrysoides  Gandoger,  192, 
invalid,  a  synonym  of  E.  cffusum  Nutt.  var. 
rosmarinioides  Benth.  in  DC.  The  name  was 
validly  published  by  Rydberg  (1931)  and  the 
citation  of  the  name  should  be  E.  helichry- 
soides  Rydb.,  Brittonia  1:  87.  1931,  without 
reference  to  Gandoger.  Rydberg  was  in- 
correct that  Gandoger  had  proposed  a  varie- 
ty, so  the  combination  attributed  to  Gando- 
ger by  Rydberg  in  synonymy,  E.  nucrothecum 
var.  helichrysoides,  must  be  attributed  to 
Rydberg  as  well. 

Eriogonum  sarothriforme  Gandoger,  192, 
invalid.  The  type  of  this  name,  collected  by 
Osterhout  at  Glenwood  Springs,  Garfield  Co., 
Colorado,  may  represent  a  distinct  taxon.  It 
belongs  to  the  E.  brevicaule  Nutt.  complex 
and  is  seemingly  a  part  of  the  polymorphic 
species  E.  lonchopln/llum  Torr.  &  Gray,  a 
taxon  that  morphologically  bridges  the  E. 
brevicaule  complex,  a  group  of  herbaceous 
perennials,  with  those  species  typified  bv  E. 
corymbosum  Benth.  in  DC,  a  series  of  shrubs 
or  subshnibs.  The  Garfield  Co.  plants  tend  to 
be  more  slender  than  typical  E.  lonchophyl- 
lum  of  southern  Colorado  and  adjacent 
northern  New  Mexico.  These  plants  also  tend 
to  resemble  some  of  the  more  robust,  but  yel- 
low-flowered, forms  of  E.  brevicaule  found  in 
Rio  Blanco  Co.,  Colorado.  Although  well 
known  to  me,  I  am  still  uncertain  what  to  do 
with  the  Glenwood  Springs  plants. 

Eriogonum  niveum  Dougl.  ex  Benth.  var. 
suksdorfii  Gandoger,  192,  valid,  a  synonym  of 
E.  niveum. 

Eriogonum  niveuni  var.  candelabrum  Gan- 
doger, 192,  valid,  a  svnonvm  of  E.  niveum. 

Eriogonum  ochrolcucum  Small  var.  macro- 
podum  Gandoger,  192,  valid,  the  basionym 
for  E.  ovalifolium  Nutt.  var.  macropodum 
(Gandoger)  Reveal. 


146 


Great  Basin  Naturalist 


Vol.  40.  No.  2 


Eriogonum  ochroleucum  var.  decahans 
Gandoger,  192,  valid  a  synonym  of  E.  ocali- 
folium  Nutt.  var.  macropodum  (Gandoger) 
Reveal. 

Under  the  heading  of  Eriogonum  ovalifo- 
lium  Nutt.,  Gandoger  proposed  a  series  of 
names  in  two  ranks,  all  of  which  he  refers  to 
by  "formas  quarum  non  paucas  saltern  pro 
speciebus  secimdi  ordinis  haberi  possimt," 
which  I  believe  he  wished  to  apply  only  to 
those  names  he  indicated  by  the  designation 
oi  sp.  n. 

Eriogonum  flavissitnum  Gandoger,  193,  in- 
valid, a  svTionym  of  E.  ovalifolium  Nutt.  var. 
anserinum  (Greene)  R.  J.  Davis.  The  name 
was  used  by  Stokes  (1936)  as  a  subspecies  of 
E.  ovalifoliwn,  where  the  name  was  validated 
as  E.  ovalifolium  Nutt.  ssp.  anserinum  S. 
Stokes. 

Eriogonum  cusickii  Gandoger,  non  M.  E. 
Jones  (1903),  193,  invalid,  a  synonym  of  E. 
strictum  var.  proliferum  (Torr.  &  Gray)  Re- 
veal. The  name  was  used  at  the  variatal  rank 
by  Stokes  (1936)  within  E.  strictum,  where 
the  name  was  validated  as  E.  strictum  Benth. 
var.  cusickii  S.  Stokes. 

Eriogonum  cusickii  Gandoger  var.  califor- 
nicum  Gandoger,  193,  invalid,  a  synonvm  of 
E.  strictum  var.  proliferum  (Torr.  6c  Grav) 
Reveal. 

Eriogonum  ovalifolium  Nutt.  var.  neva- 
dense  Gandoger,  193,  valid,  a  good  varietv  of 
the  species,  this  being  the  yellow-flowered, 
early  flowering  expression  which  I  have 
called  var.  multiscapum  Gandoger  (see  be- 
low). 

Eriogonum  ovalifolium  var.  deltoideum 
Gandoger,  valid,  193,  a  synonym  of  £.  ovali- 
folium var.  nevadense  Gandoger. 

Eriogonum  dichroanthum  Gandoger,  193, 
invalid,  a  synonym  of  E.  ovalifolium  Nutt. 
var.  nevadense  Gandoger  (see  discussion  un- 
der var.  multiscapum  below). 

Eriogonum  ovalifolium  Nutt.  var.  utahense 
Gandoger,  194,  valid,  a  synonym  of  var.  ova- 
lifolium. 

Eriogonum  ovalifolium  Nutt.  var.  multi- 
scapum Gandoger,  194,  valid,  a  synonym  of 
var.  ovalifolium.  For  several  years  I  have 
misapplied  this  name  to  the  yellow-flowered 
phase  of  the  species  (Reveal  &  Munz  196S, 
Reveal  1973,  1976).  Gandoger  based  this 
name  on  plants  gathered  by  Nelson  (4658)  at 


Cokeville,  Uinta  Co.,  Wyoming,  upon  which 
he  also  based  the  name  E.  dicJiroanthum.  An 
examination  of  this  collection  b\  me  in  1966, 
and  then  by  my  brother,  Jon  A.  Reveal,  in 
1973,  was  inconclusive  in  that  it  could  not  be 
fully  determined  if  the  flowers  of  var.  multi- 
scapum were  truly  yellow.  Gandoger  divided 
the  collection  and  moimted  each  on  a  sepa- 
rate sheet,  with  E.  dichroanthum  having 
"flores  fructusque  flavissimi."  and  the  flowers 
of  var.  midtiscapum,  to  him.  were  "'ochro- 
leuci."  The  same  collection  at  KSC  is  strictly 
yellow  flowered,  but  the  specimen  at  RM  is  a 
mixture  of  a  bright-yellowed  specimens 
matching  E.  dichroanthum  and  a  whitish  or. 
at  best,  pale  yellow-flowered  specimen  that 
matches  var.  multiscapuryi.  I  now  believe  that 
the  yellow-flowered,  early  spring  flowering 
phase  of  the  species  should  be  called  var. 
nevadense  and  the  type  of  var.  lyiultiscapum 
assigned  to  var.  ovalifolium. 

Eriogonum  ovalifolium  Nutt.  var.  cijclo- 
phijllum  Gandoger,  194,  valid,  a  synonym  of 
var.  macropodum  (Gandoger)  Reveal.  A  reex- 
amination of  the  types  of  var.  cyclophyllum 
and  var.  cerastoides  (see  below),  as  part  of 
this  study,  clearly  shows  that  these  names 
must  be  referred  to  what  I  (Reveal  1968)  had 
earlier  called  var.  macropodum.  making  a 
new  combination  for  this  name  by  transfer- 
ring it  from  E.  ochroleucum  Small  to  E.  ovali- 
folium. Previously,  these  two  names  have 
been  referred  to  var.  ovalifolium  (Hitchcock 
1964).  The  var.  cijclophyUum  is  close  to  var. 
ovalifolium  and  may  represent  one  of  the 
many  intermediate  expressions  between  the 
two  varieties.  I  retain  the  usage  of  var. 
macropodum. 

Eriogonum  ovalifolium  Nutt.  var.  ceras- 
toides Gandoger,  194,  valid,  a  synonym  of 
var.  macropodum  (Gandoger)  Reveal  (see  the 
discussion  above). 

Eriogonum  ruhidum  Gandoger.  194,  in- 
valid, a  synonym  of  E.  ovalifolium  Nutt.  var. 
depressimi  Blankinship. 

Eriogonum  ruhidum  var.  frigidum  Gando- 
ger, 194,  invalid,  a  synonym  of  E.  ovalifolium 
Nutt.  var.  depressum  Blankinship. 

Eriogonum  roseiflorum  Gandoger.  194,  in- 
valid, a  svnonvm  of  E.  ovalifolium  Nutt.  var. 
ovalifolium. 

Eriogonum  piperi  Greene  var.  ochrocepha- 
lum  Gandoger,  195,  valid,  a  synonym  of  E. 


June  1980 


Reveal:  Eriogonum  Nomenclature 


14' 


flaviim  \utt.  in  Fras.  var.  piperi  (Greene)  M. 
E.  Jones. 

Eriogonum  piperi  var.  longifloruni  Gando- 
ger,  195,  valid,  a  synonym  of  E.  flavum  Xutt. 
in  Fras.  var.  piperi  (Greene)  M.  E.  Jones. 

Under  Eriogonum  pohjanthum  Benth.,  now 
better  known  as  E.  umbellatum  Torr.  var. 
pohjanthum  (Benth.  in  DC.)  M.  E.  Jones, 
Gandoger  introduces  two  validly  described 
species  simply  stating  that  "species  duae  se- 
quentes  huic  sunt  affines." 

Eriogonum  marginale  Gandoger,  195, 
valid,  a  synonym  of  E.  umbellatum  Torr.  var. 
aureum  (Gandoger)  Reveal. 

Eriogonum  glaherrimum  Gandoger,  195. 
valid,  the  basionym  of  E.  umbellatum  Torr. 
var.  glaberrimum  (Gandoger)  Reveal. 

Eriogonum  glaberrimum  var.  aureum  Gan- 
doger, 195,  valid,  the  basionym  of  E.  um- 
bellatum Torr.  var.  aureum  (Gandoger)  Re- 
veal. 

Eriogonum  pohjcladon  Benth.  var.  mexi- 
canum  Gandoger,  196,  valid,  a  synonym  of  E. 
pohjcladon. 

Eriogonum  polycladon  Benth.  var.  crispum 
Gandoger,  196,  valid,  a  synonym  of  E.  poly- 
cladon. 

Eriogonum  racemosum  Xutt.  var.  sagitta- 
tum  Gandoger,  196,  valid,  a  synonym  of  E. 
racemosum. 

Eriogonum  racemosum  Xutt.  var.  cordi- 
folium  Gandoger,  196,  valid,  a  svnonvm  of  E. 
racemosum. 

Eriogonum  reniforme  Torr.  &  Frem.  var. 
asarifolium  Gandoger,  196,  valid,  a  svnonvm 
of  E.  pusillum  Torr.  6f  Gray. 

Eriogonum  praebens  Gandoger,  196,  valid, 
a  s\nonym  of  E.  bailcyi  S.  Wats.  var.  diva- 
ricatum  (Gandoger)  Reveal. 

Eriogonum  praebens  var.  diiaricatum  Gan- 
doger, 196,  valid,  the  basionym  of  E.  baileyi 
S.  Wats.  var.  divaricatum  (Gandoger)  Reveal. 

Under  the  heading  Eriogonum  sphaero- 
cephalum  Dougl.  ex  Benth..  Gandoger  states 
"inter  formas  varias  duae  sequentes.  ut  sub- 
species, praecipue  distingui  possunt." 

Eriogonum  cupreum  Gandoger,  196.  in- 
valid, a  synonym  of  E.  umbellatum  Torr.  var. 
umbellatum. 

Eriogonum  halimioides  Gandoger,  197,  in- 
valid. This  name  was  validated  by  Stokes 
(1936)   as   E.   sphaerocephalum    var.    hali- 


mioides S.  Stokes  without  reference  to  Gan- 
doger. 

Eriogonuin  subalpinum  Greene  var.  arach- 
noideum  Gandoger,  197.  valid,  a  synonym  of 
E.  umbellatum  Torr.  var.  dichrocephalum 
Gandoger. 

Eriogonum  subalpinum  var.  vulcanicum 
Gandoger,  197,  valid,  a  synonymy  of  £.  um- 
bellatuin  Torr.  var.  majus  Hook. 

Eriogonum  subalpinum  var.  stenophyllum 
Gandoger,  197,  valid,  a  synonym  of  E.  um- 
bellatum  Torr.  var.  majus  Hook. 

Eriogonum  subalpinum  var.  subnivale 
Gandoger,  197,  valid,  a  synonym  of  E.  um- 
bellatum Torr.  var.  majus  Hook. 

Eriogonum  tenellum  Torr.  var.  grandi- 
florum  Gandoger,  197,  valid,  a  synonym  of  E. 
microthecum  Nutt.  var.  laxiflorum  Hook. 

Eriogonum  tenelhw}  var.  sessiiflorum  Gan- 
doger, 198,  valid,  a  synonym  of  E.  micro- 
thecum Xutt.  var.  laxiflorum  Hook. 

Eriogonum  tenellum  var.  erianthum  Gan- 
doger, 198,  valid,  a  synonym  of  E.  micro- 
thecum Xutt.  var.  ambiguum  (M.  E.  Jones) 
Reveal  in  Mimz. 

Eriogonum  thurberi  Torr.  var.  parishii  Gan- 
doger. 198,  valid,  a  svnonvm  of  £.  thurberi. 

Eriogonum  thurberi  var.  acutangulum  Gan- 
doger, 198.  valid,  a  synonvm  of  E.  macula- 
turn  Heller. 

Eriogonum  tlnjmoides  Benth.  in  DC.  var. 
pallens  Gandoger,  198,  valid,  a  synonym  of 
£.  thymoides. 

Eriogonum  umbellatum  Torr.  var.  cran- 
dallii  Gandoger,  198,  valid,  a  synonym  of  var. 
umbellatum. 

Eriogonum  umbellatum  var.  chrysanthum 
Gandoger,  198,  valid,  a  synonym  of  E.  um- 
bellatum var.  stellatum  (Benth.)  M.  E.  Jones. 

Eriogonum  umbellatum  var.  nevadense 
Gandoger,  198,  valid,  a  good  variety  referr- 
ing to  that  phase  of  the  species  found  in  the 
Sierra  Xevada  of  California  northward  into 
Oregon  and  eastward  into  uestern  Xevada 
that  has  been  routinelv  called  var.  umbella- 
tum (Reveal  &  Munz  1968;  see  Howell  1976). 

Eriogonum  umbellatum  var.  cladophorum 
Gandoger,  198,  valid,  a  synonym  of  var.  um- 
bellatum. 

Eriogonum  umbellatum  var.  dichro- 
cephalum Gandoger,  199,  valid,  a  good  varie- 
ty applied  to  that  phase  previously  called  E. 


148 


Great  Basin  Naturalist 


Vol.  40,  No.  2 


umbeUatum  var.  aridum  (Greene)  C.  L. 
Hitchc.  (Hitchcock  1964). 

Eriogonwn  umhellatum  var.  californicuni 
Gandoger,  199,  valid,  a  synonym  of  E.  um- 
beUatum var.  nevadense  Gandoger. 

Eriogonum  vimineiim  Dougl.  ex  Benth. 
var.  rigescens  Gandoger,  199,  valid,  a  synon- 
ym of  E.  vimeneum. 

Eriogonum  vimineum  Dougl.  ex  Benth. 
var.  califomicum  Gandoger,  199,  valid,  a  syn- 
onym of  E.  luteohim  Greene  var.  caninum 
(Greene)  Reveal,  comb,  nov.,  based  on  E.  vi- 
mineum var.  caninum  Greene,  Fl.  Francisc. 
150.  1891.  The  type  of  var.  califomicum  is 
somewhat  intermediate  between  var.  luteo- 
lum  and  var.  caninum,  being  closer  to  the 
latter  than  the  former.  I  have  long  recog- 
nized the  caninum  phase  (Reveal  &  Munz 
1968)  as  distinct  from  E.  vimineum,  but  J.  T. 
Howell,  who  has  considered  the  expression 
only  as  a  variant,  has  pointed  out  in  our  con- 
versation that  he  had  observed  a  large  series 
of  intermediate  populations,  as  had  I,  which 
held  the  Mt.  Tamalpais  plant,  var.  caninum, 
well  within  the  boundaries  of  what  he  called 
E.  vimineum.  This  latter  expression,  however, 
proved  to  be  E.  luteohim  rather  than  E.  vimi- 
neum, and  I  am  now  following  Howell's 
(1970)  taxonomic  disposition  of  this  local  en- 
demic. In  addition  to  this  variant  of  E.  luteo- 
lum,  the  Sierra  Nevada  plant  I  recognized  as 
a  distinct  species  previously  (Reveal  1970) 
should  be  included  within  this  species  as  well. 
Thus,  I  propose  E.  luteohim  var.  peduncula- 
tum.  (S.  Stokes)  Reveal,  stat.  &  comb,  nov., 


based  on  E.  pedunculatum  S.  Stokes,  Leafl. 
W.  Bot.  2:  48.  1937. 

Eriogonum  vimineum  var.  oregonense  Gan- 
doger, 199,  valid,  a  synonym  of  E.  vimineum. 

Eriogonum  restioioidcs  Gandoger,  199, 
valid,  a  synonym  of  E.  baileyi  S.  Wats.  var. 
bailey  i. 

Literature  Cited 

Gandoger,  M.  1906.  Le  genre  Eriogonum  (Poly- 
gonaceae).  Bull.  Soc.  Roy.  Bot.  Belgique  42: 
18.3-200. 

Heller,  A.  A.  1907.  Compilations.  Miihlenbergia  3: 
a3-96. 

Hitchcock,  C.  L.  1964.  Eriogonum.  Univ.  Wash.  Publ. 
Biol.  17(2):  104-138. 

Howell,  J.  T.  1970.  Marin  flora.  2d  ed.  Berkeley:  Univ. 
California  Press.  366  pp. 

1976.  Eriogonum  notes  VII.  Mentzelia  1:  17-22. 

Keck,  D.  D.  1959.  "Abbreviations  of  authors'  names." 
Pages  1551-1576  in  P.  A.  Munz,  &  D.  D.  Keck, 
eds.  A  California  flora.  Berkeley:  Univ.  California 
Press. 

Reveal,  J.  L.  1968.  Some  nomenclatural  changes  in 
Eriogonum  (Polygonaceae).  Taxon  17:  531-533. 

1973.  Eriogonum  (Polvgonaceae)  of  Utah.  Phvto- 

logia  25:  169-217. 

1976.  Eriogonum  (Polvgonaceae)  of  Arizona  and 

New  Mexico.  Phytologia  34:  409-484. 

Reveal,  J.  L.,  and  P.  A.  Munz.  1968.  "Eriogonum." 
Pages  33-72  in  P.  A.  Munz,  ed.  Supplement  to  A 
California  flora.  Berkeley:  Univ.  California  Press. 

Rydberg,  p.  a.  19.31.  Taxonomic  notes  on  the  flora  of 
the  prairies  and  plains  of  central  North  .\merica. 
Brittonia  1:  79-104. 

Stafleu,  F.  a.,  et  al.  1978.  International  code  of  bot- 
anical nomenclatme.  Regnum  Veg.  97:  1-457. 

Stokes,  S.  G.  1936.  The  genus  Eriogonum,  a  preliminary 
studv  based  on  geographic  distribution.  San  Fran- 
cisco: J.  B.  Nebiett.  124  pp. 


PARASITES  FROM  TWO  SPECIES  OF  SUCKERS  (CATOSTOMIDAE) 
FROM  SOUTHERN  UTAH 


J.  Craig  Biieiiholt'  and  Richard  A.  Heckmann 

.\bstr\c:t.— Twenty  Ciitostoiims  latipiiiuis  and  50  Catostomiis  lUscohohs  from  La  Verkin  Creek  and  the  Fremont 
River  in  southern  Utah  were  collected  and  surveyed  tor  parasites.  Data  from  the  survey  indicated  that  83  percent  of 
the  fish  were  infected  with  at  least  one  parasite,  with  the  fish  from  La  Verkin  Creek  harboring  more  parasites. 
Twelve  genera  and  12  species  of  parasites  were  identified  from  these  fish.  .\  monogenetic  trematode,  G\jwd(trtiilus 
dedans,  which  was  found  in  90  percent  of  the  fish,  was  the  most  common  parasite.  Comments  are  included  on  habi- 
tat and  host  variations  for  the  parasitofauna  from  suckers  taken  from  the  two  locations. 


A  survey  of  the  parasites  of  the  cato,sto- 
mids,  Catostomiis  latipinnis  and  C.  discoboUs, 
was  conducted  at  La  Verkin  Creek,  southern 
Utah,  and  the  Fremont  River  near  Hanks- 
ville,  Utah.  The  objectives  of  this  survey  were 
to  provide  a  hst  of  parasites  for  C.  discoboUs 
and  C.  latipinnis  in  La  Verkin  Creek  and 
Fremont  River  and  to  correlate  water  param- 
eters and  benthos  from  these  streams  with 
parasite  loads.  Both  streams  contain  well-e.s- 
tablished  populations  of  the  listed  suckers.  An 
exhaustive  survey  of  parasites  can  explain  the 
source  or  reservoir  of  serious  pathogens  for 
endangered  species  and  commercially  impor- 
tant fish. 

Catostomids  are  found  exclusively  in  North 
America,  excluding  two  or  three  Asiatic  spe- 
cies (Pflieger  1975).  Catostomiis  discoboUs  is 
found  in  Idaho,  Utah,  and  Nevada  in  the  fol- 
lowing drainages:  Colorado  River  above  the 
Grand  Canyon,  upper  Snake  River,  Bear  Riv- 
er, and  Weber  Lake  outflows.  Catostomiis 
latipinnis  is  unique  to  the  Colorado  River 
drainage  (Eddy  1959).  Information  con- 
cerning the  life  history  of  these  two  .species  is 
limited.  Catostomid  levels  in  both  study  areas 
for  this  project  are  maintained  by  resident 
sucker  populations. 

Both  of  the  streams  selected  in  this  survey 
are  unstable  desert  streams.  Much  of  the  sub- 
strate is  sand  which  shifts  and  prevents  deep 
pools  from  forming.  Flash  floods  can  disrupt 


and  completely  change  the  nature  of  the 
streams  and  change  the  macroinvertebrate 
population.  Because  of  this,  fish  species,  e.g., 
salmonids  and  centrarchids,  that  cannot  with- 
stand the  instability  of  the  stream  and  the 
consequent  change  in  macroinvertebrate  food 
source  are  not  found  extensively  in  these  two 
streams.  The  ichthyofauna  found  in  the  study 
area  of  the  Fremont  River  are:  Longnose 
dace,  RJiinichthys  cataractae;  speckled  dace, 
R.  osciiliis;  leatherside  chub,  Gila  copei;  blue- 
head  sucker,  C.  discoboUs;  and  flannelmouth 
sucker,  C.  latipinnis  (Heckmann  1976).  The 
speckled  dace  and  leatherside  chub  are  omni- 
vores  that  feed  on  aquatic  plants,  insects,  and 
cmstaceans.  The  flannelmouth  suckers  are 
herbivores  which  feed  on  algae,  diatoms, 
parts  of  higher  plants,  and  seeds.  The  blue- 
head  sucker  is  a  bottom  feeder  which  scrapes 
algae  and  other  organisms  from  rocks  (Sigler 
and  Miller  1963).  The  largest  fish  found  in 
the  Fremont  River  is  the  flannelmouth  sucker 
and  the  smallest  is  the  speckled  dace.  None 
of  the  fish  found  in  the  sample  area  are  pisci- 
vorous. All  fish  feed  either  on  aquatic  in- 
vertebrates or  plant  material.  Fi,sh  predators 
mav  include  birds  and  small  mammals. 

Fish  .species  inhabiting  the  survey  site  at 
La  Verkin  Creek  are:  speckled  dace,  R.  os- 
cuUis;  Virgin  River  spinedace,  Lcpidomeda 
mollispinis;  red  shiner,  \otropiis  liitrcnsis; 
woundfin  minnow,  Plagopterus  argentissimus; 


'Department  of  Zoolog\',  Brigham  Young  University,  Provo,  Utah  84602. 


149 


150 


Great  Basln  Naturalist 


Vol.  40,  No.  2 


flannelmouth  sucker,  C.  kitipinnis;  bluehead 
sucker,  C.  discoholis  or  desert  sucker,  C. 
clarki;  and  rainbow  trout,  Salmo  gairdneri 
(Winget  and  Baumann  1977).  La  Verkin 
Creek,  in  comparison  to  the  Fremont  River, 
is  more  stable  and  less  turbid,  resulting  in  the 
presence  of  riffles,  pools,  and  some  holes 
1-1.5  m  deep.  Because  of  the  difference  in 
stream  conditions,  rainbow  trout  are  planted 
by  the  Utah  Division  of  Wildlife  Resources  in 
small  numbers.  Two  species  of  fish,  L.  moUis- 
pinis  and  P.  argentissimus,  are  considered  en- 
dangered. The  woimdfin  minnow,  rainbow 
trout,  and  Virgin  River  spinedace  are  consid- 
ered carnivores  feeding  mainly  on  in- 
vertebrates. Catostoriius  discoholis  and  N.  lut- 
rensis  are  considered  bottom-dredging 
detritovores.  Catostoinus  kitipinnis  and  R.  os- 
culus  are  selective  omnivores  (Winget  and 
Baumann,  1977).  The  top  carnivore  in  a  tro- 
phic scheme  would  be  S.  gairdneri  because  it 
may  feed  on  smaller  fish.  Direct  competition 
"s  virtually  eliminated  because  those  species 
vith  similar  feeding  habits  have  different 
habitat  preferences  or  specific  food  prefer- 
ences (Winget  and  Baumann  1977). 

Parasites  of  catostomids  other  than  C.  lati- 
pinnis  and  C.  discoholis  have  been  studied  by 
researchers  in  the  United  States  and  Canada. 
Hoffman  (1967)  lists  known  parasites  for  12 
species  of  catostomids.  Other  surveys  have 
been  conducted  bv  Voth  and  Larson  (1968), 
Amin  (1969),  Threlfall  and  Hanek  (1970), 
Amin  (1974),  White  (1974),  Mackiewicz 
(1963),  Price  and  Arai  (1967),  Dechtiar 
(1969),  Daly  and  De  Giusti  (1971),  Clifford 
and  Facciani  (1972),  Hatha wav  and  Herlev- 
ich  (1973),  Schell  (1974),  andHayunga  and 
Grey  (1976).  The  most  widely  studied  ca- 
tostomids are  white  suckers,  C.  cornmersoni, 
and  longnose  suckers,  C.  catostomiis.  These 
surveys  deal  primarily  with  metazoan  para- 
sites, and  little  information  concerning  the 
protozoan  parasites  is  included. 


M.\TERIALS  AND  METHODS 

Through  the  use  of  electrofishing,  18 
flannelmouth  suckers  and  40  i)luehead  suck- 
ers were  collected  from  La  Verkin  Creek 
near  the  Toquerville  cemeterv,  southern 
Utah.  Two  flannel-mouth  suckers  and  10 


bluehead  suckers  were  collected  from  the 
Fremont  River  one  mile  west  of  Capitol  Reef 
National  Park,  near  Hanksville,  Utah.  The 
fish  were  transported  to  Brigham  Young  Uni- 
versity in  iced  holding  tanks.  Limited  num- 
bers of  fish  were  obtained  due  to  collecting 
restrictions. 

Each  fish  was  checked  for  parasites.  The 
suckers  were  euthanized  by  a  blow  to  the 
head  before  being  weighed  and  measured 
(Table  1).  Following  macroscopic  exam- 
ination, scrapings  of  the  surface,  gills,  medial 
area  of  the  opercula,  and  eyes,  were  exam- 
ined for  parasites.  Because  the  blood  vessels 
were  niptiued,  gill  scrapings  were  used  to 
check  for  hemoflagellates.  Intestine,  liver, 
and  gall  bladder  were  excised  and  examined 
for  endoparasites.  The  presence  of  metacer- 
cariae  inhabiting  the  liver  was  checked  by 
pressing  a  piece  of  the  organ  between  two 
glass  slides  and  examining  it  without  magnifi- 
cation. 

Protozoans  were  either  air  dried  or  pre- 
served in  10  percent  formalin.  Permanent 
preparations  of  monogenetic  trematodes 
were  made  with  Turtox  mounting  and  stain- 
ing medium  (nonresinous  stain  mountant 
CMC-S).  Leeches  were  also  fixed  in  formalin 
and  all  were  identified  tlirough  the  use  of 
keys  listed  in  Hoffman  (1967). 

Cestodes  were  placed  directly  into  AFA 
fixative  to  prevent  total  relaxation.  Digenetic 
trematodes  were  placed  in  95  C  water  to 
promote  relaxation  and  then  placed  in  AFA 
fixative.  Cestodes  and  digenetic  trematodes 
were  stained  with  Semichon's  carmine  for  12 
hours  and  then  destained  in  changes  of  acid 
alcohol  to  improve  color  contrast.  After  des- 
taining,  the  specimens  were  dehydrated  in  95 
percent  and  100  percent  ethyl  alcohol  for 
one  hoiu'  each.  Once  dehydrated,  specimens 
were  cleared  in  xylene  and  then  mounted 
with  Permount  on  glass  slides.  Morphological 
characteristics  given  in  Hoffman  (1967)  were 
used  for  identification  of  trematodes  and 
nematodes.  Preliminary  identification  of  the 
caryophyllid  tapeworms  was  confirmed  by 
John  S.  Mackiewicz  (State  University  of  New 
York  at  Albany). 

Water  chemistry  and  macroinvertebrate 
data  were  obtained  from  studies  by  Heck- 
mann  (1976),  Winget  and  Reichert  (1976), 
and  Winget  and  Baumann  (1977). 


June  1980 


Breinholt,  Heckmann:  Fish  Parasites 


151 


Results 

Data  from  the  examination  of  40  bluehead 
and  18  flannelmouth  suckers  from  La  Verkin 
Creek  in  southern  Utah  indicated  that  55 
suckers  harbored  at  least  one  species  of  para- 
site. Thirty-seven  of  40  bluehead  suckers 
were  infected  and  all  18  flannelmouth  suck- 
ers harbored  parasites  (Table  2).  Postmortem 
examination  of  12  suckers  from  the  Fremont 
River  revealed  one  of  10  bluehead  suckers 
and  2  of  2  flannelmouth  suckers  were  para- 
sitized (Table  2). 

Twelve  genera  and  12  species  of  parasites 
were  identified  from  fish  from  La  Verkin 
Creek.  The  most  frequently  encountered  par- 
asite was  Gyrodactylus  elegans,  a  mon- 
ogenetic  trematode  which  was  found  in  52 


Table   1.  Weights  and  measurements  of  fish  exam- 
ined. 


Species 

Weight 

Length 

No. 

offish 

Stream 

(gm) 

(TL)  (cm) 

1 

C.  discobolis 

La  Verkin  Creek 

45 

17 

2 

43 

14 

3 

38 

16 

4 

33 

15 

5 

33 

14 

6 

55 

18 

7 

65 

19 

8 

50 

16 

9 

40 

16 

10 

25 

13 

11 

35 

16 

12 

50 

16 

13 

60 

18 

14 

85 

21 

15 

90 

21 

16 

45 

17 

17 

20 

13 

18 

40 

16 

19 

40 

16 

20 

40 

17 

21 

15 

11 

22 

45 

17 

23 

30 

14 

24 

35 

15 

25 

25 

13 

26 

55 

17 

27 

20 

13 

28 

35 

Ifi 

29 

30 

Hi 

30 

45 

17 

31 

20 

13 

.32 

25 

14 

33 

25 

14 

34 

30 

15 

35 

25 

15 

(90  percent)  of  the  fish.  Other  monogenetic 
trematodes  recovered  were  Octomacrwn  lan- 
ceatinn,  found  in  one  (2  percent)  fish,  and 
PcUucidhaptor  alahauius,  foimd  in  six  (10 
percent)  of  the  fish.  Metacercariae  of  two 
digenetic  trematodes  were  also  recovered. 
Neascus  sp.  was  found  in  25  (43  percent)  fish 
and  Clinostomum  marginatum  was  found  in 
two  (3  percent)  fish.  Cystidicola  sp.,  a  nema- 
tode, was  found  in  one  (2  percent)  fish  and 
Monobothrium  hunteri  and  Isoglaridacris  hex- 
acotyle,  both  caryophyllid  cestodes,  were 
found  in  29  (50  percent)  and  28  (48  percent) 
fish,  respectively.  Three  protozoans  were  re- 
covered from  the  fish.  Myxosoma  sp.  was 
found  in  11  (19  percent)  fish,  Myxidiiim  sp.  in 
four  (7  percent),  and  Tridiondina  sp.  was 
found  in  20  (34  percent)  of  the  fish  examined 
(Table  3). 

Table  1  continued. 


No. 


Species 
of  fish 


Weight 
Stream  (gm) 


Length 

(TL)  (cm) 


36      C.  discobolis 

37 

38 

.39 

40 

41         C.  latipinnis 

42 

43 

44 

45 

46 

47 

48 

49 

.50 

51 

52 

53 

54 

55 

56 

57       " 

58 

59         C  discobolis 

60 

61 

62 

63 

64 

65 

66 

67 

68 

69  C.  latipinni.s 

70 


La  Verkin  Creek   30  15 

45  17 

15  11 

20  13 

20  13 

115  24 

180  27 

185  27 

230  30 

205  29 

260  32 

210  30 

125  25 

185  29 

260  31 

225  24 

110  24 

195  23 

130  25 

170  28 

125  24 

105  24 

85  22 

Fremont  Hiver   20  14 

10  11 

35  16 

35  17 

20  14 

30  16 

5  10 

25  15 

10  11 

30  15 

5.30  40 

350  37 


152 


Great  Basin  Naturalist 


Vol.  40,  No.  2 


Differences  in  protozoan  parasite  load 
from  the  two  species  of  fish  taken  from  La 
Verkin  Creek  are  as  follows:  Myxosoma  sp.. 
25  percent  bluehead  suckers  and  6  percent 
flannelmouth  suckers;  Myxidium  sp.,  0  per- 
cent bluehead  suckers  and  22  percent 
flannelmouth  suckers;  Trichodina  sp.,  80  per- 
cent bluehead  suckers  and  22  percent 
flannelmouth  suckers  were  infected.  For  the 
metazoan  parasites,  G.  elegans  was  fomid  in 
90  percent  of  the  bluehead  suckers  and  89 
percent  of  the  flannelmouth  suckers,  Octoma- 
cntm  lanceatum  and  P.  alahamus  were  found 
exclusively  on  bluehead  suckers,  3  percent 
and  6  percent,  respectively.  Neascus  sp.  was 
found  on  28  percent  of  the  bluehead  suckers 


and  78  percent  of  the  flannelmouth  suckers, 
but  C.  marginatum  was  found  only  in  11  per- 
cent of  the  flannelmouth  suckers.  The 
flannelmouth  suckers  have  a  higher  incidence 
of  both  species  of  caryophyllid  cestodes. 
MonobotJirium  hunteri  was  found  in  38  per- 
cent of  the  bluehead  suckers  and  78  percent 
of  the  flannelmouth  suckers,  and  /.  hexacotyJe 
was  found  in  33  percent  of  the  bluehead 
suckers  and  83  percent  of  the  flannelmouth 
suckers.  The  nematode  Cystidicola  sp.  and 
the  leech  Piscicola  sp.  were  symbiotic  exclusi- 
vely to  the  bluehead  suckers.  Three  percent 
of  the  fish  were  infected  with  each  of  these 
two  parasites. 

One  species  of  parasite  was  recovered  from 


T.\BLE  2.  Number  and  percentage  of  bluehead  and  flannelmouth  suckers  parasitized  from  La  Verkin  Creek  and 
the  Fremont  River. 


Host  species 


Total  fish 
sampled 


Total  fish 
parasitized 


La  Verkin  Creek 
fish  parasitized* 

37  (  93%) 
18  (100%) 

.55  (  95%) 


Fremont  River 
fish  parasitized^ 


Bluehead  sucker 
Flannelmouth  sucker 

Total 


50 
20 


38  (  76%) 
20  (100%) 

58  (  83%) 


1  (  10%) 

2  (100%) 

3  (  2,5%) 


*58  fish  examined  from  La  Verkin  Creek  (40  bluehead  and  18  flannelmouth  suckers). 
''12  fish  examined  from  the  Fremont  River  1 10  bluehead  and  2  flannelmouth  suckers). 


Table  3.  Parasites  identified  from  58  suckers  from  La  Verkin  Creek. 


Parasite  species 


Number  and 
percentage  of 

Species 

of  fish 

fish  positive 

Bluehead 

Flannelmouth 

11  (19) 

10  (25%) 

1  (  6%) 

4(   7) 

0  (  0%) 

4  (22%) 

20  (34) 

16  (80%) 

4  (22%) 

52  (90) 

.36  (90%) 

16  (89%) 

1(  2) 

1  (  .3%) 

0  (  0%) 

6(10) 

6  (15%) 

0(  0%) 

25  (43) 

11  (28%) 

14  (78%) 

2  (  3) 

0(  0%) 

2(11%) 

29(50) 

15  (38%) 

14  (78%) 

28  (48) 

13  (3.3%) 

15  (8,3%) 

1(  2) 

1  (  3%) 

1  (  0%) 

1(  2) 

1  (  3%) 

0  (  0%) 

Protozoans 
Myxosoma  sp. 
Myxiditnn  sp. 
Trichodina  sp. 

Trematodes 

Gijrodactijlufi  elegans 
Octomacrum  lanceatutn 
Pellucidhaptor  alahamus 
Postodiphstom am  mit^ im iim 
Clinostomiim  marginatum 

Cestodes 

Monobothritim  hunteri 
Isoglaridacris  hexacotyle 

Nematodes 
Cystidicola  sp. 

Leeches 
Piscicola  sp. 


June  1980 


Breinholt,  Heckmann:  Fish  Parasites 


153 


the  fish  examined  from  the  Fremont  River 
(Table  4).  Gyrodactyhis  elegans  was  identi- 
fied in  25  percent  of  the  fish  examined.  Ten 
percent  of  the  bluehead  suckers  were  in- 
fected with  this  inonogentic  trematode,  and 
100  percent  of  the  flannelmouth  suckers 
were  infected. 

Table  5  lists  the  preferred  tissue  in  the  host 
for  each  parasite.  These  parasites  were  found 
in  onlv  five  areas  of  the  fish.  Eight  of  the  par- 
asite species  were  fomid  on  the  external  sur- 
face and  only  four  were  found  in  more  than 
one  area. 


Discussion 

There  are  habitat  and  host  variations  for 
the  parasitofauna  from  suckers  taken  from  La 
V'erkin  Creek  and  Fremont  River.  The  ca- 
tostomids  from  La  V'erkin  Creek  were  more 


heavily  parasitized,  both  in  the  number  of 
fish  infected  and  in  the  number  of  species  en- 
countered, than  were  the  fish  from  Fremont 
River.  Explanation  for  these  differences  may 
be  attributed  to  many  factors,  such  as  water 
qualitv  and  macroinvertebrates.  Water  chem- 
istr\  for  the  two  streams  was  found  to  be  sim- 
ilar except  during  spring  rvmoff. 

Oligocheates,  which  usually  act  as  the  in- 
termediate hosts  for  caryophyllid  tapeworms 
(Mackiewicz  1972),  occur  in  both  streams. 
Recent  studies  listed  14,203  (Winget  and 
Baumann  1977)  and  882  (Heckmann  1976) 
oligocheates  per  m^  in  La  Verkin  Creek  and 
Fremont  River,  respectivelv,  where  the  fish 
for  this  studv  were  obtained.  The  number  of 
oligocheates  should  not  have  caused  the  dif- 
ference in  parasite  load  because  infected 
worms  would  have  been  ingested  by  fish 
from  both  streams.  Milbrink  (1975)  correlates 


Table  4.   Parasites  identified  from  12  suckers  from  the  Fremont  River. 


Number  and 
percentage  of 
fish  positive 

Species 

offish 

Parasite  species 

Bluehead 
suckers 

Flannelmouth 
suckers 

Trematodes 

Gyrodactyhis  elegans 

3  (25) 

1  (10%) 

2  (100%) 

Table  .5.   Location  in  host  of  parasites  found  in  fish  from  La  Verkin  Creek  and  the  Fremont  River. 


Parasite  species 


Surface 


Gills 


Operculimi        Intestine       Gall  bladder 


Protozoans 
Mijxosotna  sp. 
MyxidiuDi  sp. 
Trichodiud  sp. 

Trematodes 

Gyrodactyhis  ch-gans 
Octomacrum  hmccatiiiu 
PcUucidJiaptor  ahihamus 
Chnostomuin  mart^inatiiui 

Cestodes 

Monohothriuin  hunteri 
Isogla  ridac ris  h cxacoty Ic 

Nematodes 
Cystidicola  sp. 

Leeches 
Piscicola  sp. 

=  present 
o  =  not  present 


154 


Great  Basin  Naturalist 


Vol.  40,  No.  2 


the  caryophyllid  worm  burden  of  fish  with 
the  number  of  infective  oligocheates  con- 
sumed. If  the  Fremont  River  contained  in- 
fected oHgocheates,  some  of  the  fish  sampled 
should  have  been  infected. 

The  geographical  location  of  the  two 
streams  may  have  caused  the  difference  in 
parasite  load.  Parasites  can  be  found  in  one 
area  but  not  in  another  even  though  both 
have  similar  aquatic  characteristics.  Myxo- 
soma  cerebralis,  a  myxosporidan  parasite 
which  caused  whirling  disease  in  trout,  has 
been  reported  in  eight  states  (American  Fish- 
eries Society,  1974).  Whirling  disease  has  not 
spread  to  the  other  states  even  though  suit- 
able habitats  exist.  Diplostomum  spathacewn, 
the  eye  fluke  of  fish,  has  been  reported  in 
some  areas  of  Utah  but  not  others  (Palmieri, 
Heckmann,  and  Evans  1976). 

Most  parasites  have  some  effect  on  the 
health  of  the  host  (Olsen  1974).  The  fish  sam- 
pled from  the  Fremont  River  were  infected 
with  only  one  species  of  parasite,  Gijro- 
dactylus  elegans,  and  the  incidence  of  that 
parasite  was  low  in  comparison  to  infected 
fish  from  La  Verkin  Creek.  The  most  com- 
mon parasite  found  on  the  fish  from  La  Ver- 
kin Creek  is  G.  elegans.  This  organism  was  on 
the  surface  and  occasionally  in  gill  scrapings. 
Large  numbers  of  G.  elegans  can  cause  dam- 
age to  the  fish  by  physical  blockage  of  the 
gill  surface,  thus  interfering  with  the  gas  ex- 
change area  (Hoffman  1967).  Other  mon- 
ogenetic  trematodes,  Octomacrum  lanceatum 
and  Pellitcidhaptor  alahamiis,  and  the  pro- 
tozoan, Trichodina  sp.,  are  capable  of  causing 
similar  problems  in  the  fish.  These  parasites 
were  not  foimd  in  great  enough  quantities  to 
pose  a  threat  at  the  present  time.  The  two 
myxosporidans,  Myxosonia  sp.  and  Myxidium 
sp.,  are  capable  of  encysting  and  destroying 
tissue.  However,  no  cysts  were  found.  Thus, 
it  is  assumed  that  these  myxosporidans  are 
causing  little  damage  to  the  fish. 

The  fact  that  no  hemoflagellates  were 
found  dining  the  course  of  this  study  does  not 
disprove  their  existence  in  these  fish,  because 
some  of  these  parasites  have  seasonal  fluctua- 
tion. 

Only  one  nematode,  Cystidicola  sp.,  was 
recovered  from  all  fish  examined  in  the  sur- 
vey. This  round  worm  is  not  detrimental  to 
the  fish  unless  it  is  found  in  high  numbers 


(Hoffman  1967).  The  other  intestinal  hel- 
minths, Monobothrium  hunteri  and  Isoglari- 
dacris  hexacotyle,  are  adult  cestodes  that  usu- 
ally cause  little  damage  to  the  definitive  host. 
High  numbers  (200  plus)  result  in  mechanical 
blockage  or  cause  nutritional  deficiencies 
(Mackiewicz  1972).  The  adult  worms  adhere 
to  the  intestinal  lining  by  means  of  suckers. 
There  is  little  intestinal  damage  by  individual 
cestodes  because  the  scolex  is  unarmed. 

The  metacercariae  of  Neascus  sp.  and  Cli- 
nostomwn  marginatum  were  recovered  from 
suckers  from  La  Verkin  Creek.  For  these 
trematodes,  the  metacercarial  stage  is  usually 
encysted  in  the  second  intermediate  host  and 
does  not  cause  damage  through  migration 
(Hoffman  1967).  Unless  the  metacercaria  is 
encysted  in  vital  organs,  such  as  a  parasite  in 
the  eye  lens  (D.  spathaceum),  it  does  not 
present  a  pathogenic  health  problem  to  the 
fish.  (Neascus  sp.  was  observed  encysted  in 
the  fins  and  gills.  The  cysts  found  in  the  gills 
were  not  numerous  and  did  not  appear  to  in- 
terfere with  gas  exchange  for  the  fish. 

The  leech,  Piscicola  sp.,  did  not  present  a 
current  problem  to  the  fishing  that  di^ly  one 
specimen  was  recovered.  Leeches  are  period- 
ic feeders  and  should  not  attach  permanently 
to  the  host.  The  major  problem  with  leeches 
on  fish  is  due  to  large  numbers  on  one  host  or 
tlie  transmitting  by  hemoflagellates  (l:)lood 
parasites)  (Hoffman  1967).  No  blood  parasites 
were  found  during  this  survey. 

None  of  the  fish  expired  during  the  journey 
from  their  natural  habitat,  approximately  400 
km,  to  holding  tanks.  Thus,  it  is  assumed  that 
the  effects  of  all  the  parasites  on  the  suckers 
were  not  evident  when  the  fish  were  placed 
under  stress  of  capture  and  transportation. 

Limited  host  specificity  is  demonstrated  by 
the  parasites  recovered  in  this  survey.  Most 
of  the  parasites  encountered  have  been  re- 
ported in  other  species  of  fish  (Hoffman 
1967).  Species  of  Myxidium,  Myxosoma, 
Trichodina,  Cystidicola,  and  Piscicola  have  all 
been  reported  in  trout.  The  parasitic  species 
found  in  salmonids  may  be  different  than 
those  found  in  the  suckers.  Gyrodactylus  ele- 
gans and  P.  minimum  have  also  been  report- 
ed in  salmonids.  Octonuurum  lanceatum  has 
been  reported  in  the  catostomids,  Catostomus 
teres,  C.  commersoni,  C.  macrocheilus,  and 
Erimyzon  secetta,  the  cyprinids,  Mylocheilus 


June  1980 


Breinholt,  Heckmann:  Fish  Pahasites 


155 


caurinus  (peamouth),  and  Notropis  corntitits 
(common  shiner).  FclhicitUuiptor  dldluiiniis 
has  been  reported  in  Ictiohus  htihalus,  the 
smalhnoiith  buffalo  (Chien  and  Rogers  1970). 
The  caryophyllid  tapeworms.  A/,  hunteri  and 
/.  hcxncoti/lc,  common  to  C.  discobolis  and  C. 
Uitipinnis,  have  both  been  reported  from  oth- 
er catostomids  (Hoffman  1967).  Because  the 
parasites  recovered  in  this  study  have  been 
reported  in  other  species  of  fish,  it  is  possible 
that  these  parasites  may  infect  game  fish  or 
commerciallv  cultured  fish.  In  the  case  of  the 
listed  digenetic  trematodes,  infected  birds 
can  fly  from  one  body  of  water  to  another 
and  "seed"  other  streams  and  ponds.  Also, 
currents  can  carrv  infected  fish  and  other  in- 
termediate hosts  downstream  to  contaminate 
the  lower  drainage  system.  Thus,  potential 
infections  of  other  fish  in  the  same  stream 
could  threaten  endangered  species  such  as 
the  woundfin  minnow  and  the  Virgin  River 
spinedace,  which  are  also  found  in  La  Verkin 
Creek. 

The  identity  of  one  of  the  hosts  from  La 
Verkin  Creek  is  doubtful.  Originally,  it  was 
classified  as  a  chiselmouth  sucker  (Sigler  and 
Miller  1963).  Later  studies  considered  this  ca- 
tostomid  a  desert  sucker  (C.  clarki),  which  is 
still  a  valid  species  (Bailey  et  al.  1970).  Then, 
with  the  taxonomic  revision  of  some  of  the 
members  of  the  catostomid  family,  Pan- 
tosteus  delphinius,  the  bluehead  sucker,  and 
P.  virescens,  the  green  sucker,  were  combined 
to  form  C.  discobolis  (Bailey  et  al.  1970).  Af- 
ter this  revision  some  investigators  have  con- 
sidered the  fish  as  C.  discobolis.  Because  of 
the  anatomical  similarities  between  C.  dis- 
cobolis and  C.  clarki  and  the  activities  and 
feeding  habits,  the  two  could  be  considered 
similar.  It  may  also  be  concluded  that  they 
could  harbor  similar  parasites  even  if  they 
are  two  distinct  species. 

Of  the  two  suckers,  C.  latipinnis  is  more 
selective  in  its  feeding  habits  than  is  C.  dis- 
cobolis. Winget  and  Baumann  (1977)  report- 
ed stomach  contents  of  the  flannelmouth 
sucker  contained  seeds,  identifiable  plant 
matter,  and  dipteran  larvae;  stomachs  of  the 
bluehead  sucker  contained  detritus,  uniden- 
tifiable plant  matter,  and  very  few  macroin- 
vertebrates.  The  difference  in  feeding  habits 
of  these  two  fish  is  probably  the  reason  for 
the   difference   in   resident   parasite   species. 


The  parasites  that  these  two  fish  have  in 
common  may  be  due  to  the  ingestion  of  a 
common  intermediate  host.  Even  though  C. 
latipinnis  is  more  selective  in  its  feeding 
habits,  it  still  would  ingest  detritus  and  other 
material  due  to  its  feeding  technique. 

The  parasites  identified  in  this  survey  are 
not  unexpected.  Although  no  parasitic  sur- 
veys of  C.  discobolis  and  C.  latipinnis  have 
been  reported,  Mi/xosorna  sp.,  Myxidiian  sp., 
Trichodina  sp.,  G.  elegans,  O.  lanceatum,  P. 
alaJianius,  P.  mininiitm,  C.  marginatum,  I. 
Jiexacotyle,  M.  hunteri,  Cystidicola  sp.,  and 
Piscicola  sp.  have  all  been  reported  from 
suckers  (Hoffman  1967). 


Literature  Cited 

American  Fisheries  Society.  1974.  Suggested  procedures 
for  the  detection  of  certain  infectious  diseases  of 
fishes.  U.S.  Department  of  the  Interior,  Washing- 
ton. D.C. 

Amin,  O.  M.  1969.  Helminth  fauna  of  suckers  (Catosto- 
midae)  of  the  Gila  River  system,  Arizona.  II.  Five 
parasites  from  Ccito.'itomus  spp.  Amer.  Midi.  \at. 
82(2):  429-443. 

1974.   Intestinal  helminths  of  the  white  sucker, 

Catostomus  coninicrsoni  (lacepede)  in  southeast 
Wisconsin.  Proc.  Helminthol.  Soc.  Wash. 
41(l):81-88. 

Bailey,  R.  M.,  J.  E.  Fitch,  E.  S.  Herald,  E.  A. 
Lachner,  C.  C  Lindsey,  C.  R.  Robins,  and  W. 
B.  Scott.  1970.  .\  list  of  common  and  scientific 
names  of  fishes  from  the  United  States  and  Clan- 
ada.  .\merican  Fisheries  Society  Special  Pub- 
lication No.  6.  Washington,  D.C. 

Chien,  S.  M.,  and  W.  Rogers.  1970.  Four  new  species 
of  monogenetic  trematodes.  genus  PelliicicUuiptor. 
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Clifford,  T.  S.,  and  S.  Facciani.  1972.  Philoinetra 
nodulosu  in  Wvoming  white  suckers.  Prog.  Fi.sh- 
Cult.  .34(4):23.5-2.36.  ' 

Daly,  J.  J.,  a.nd  D.  L.  De(;ivsti.  1971.  Tn/pdnosoiiia  ca- 
tosiomi  n.  sp.  from  the  white  sucker  Cdtostoiiiiis 
cominersoni.  J.  Protozool.  18(.3):414-417. 

Dechtiar,  .\.  O.  1969.  Two  new  species  of  monogenetic 
trematodes  (Treniatoda:  Monogcnca)  from  nasal 
cavities  of  catostomid  fishes.  J.  Fish.  Wvs.  Board 
Can.  26(4):86.5-869. 

Eddy,  S.  1969.  The  freshwater  fishes.  Win.  C  Brown  C. 
Puhl.,  Dubuque,  Iowa. 

IL\THAWAY,  R.  p.,  and  J.  C.  Herlevich.  1973.  Gyro- 
dactyhis  stahlcri  sp.  n.  with  new  host  and  locality 
records  for  species  of  Giircddiliihis.  J.  Parasitol. 
.59(2):801-802. 

Hayunga,  E.  C,  and  .\.  J.  Grey.  1976.  Cystobracluis 
meyeri  sp.  n.  (Hirudinea:  Piscicolidae)  from  Ca- 
tostomtis  commersoni  Lacepede  in  North  Ameri- 
ca. J.  Parasitol.  62(4):621-627. 


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Heckmann.  R.  a.  1976.  Aquatic  habitat  evaluation  of 
the  Fremont,  Muddy,  and  Dirt\  Devil  rivers  and 
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Corporation  contract  .\o.  P.  O.  ESD- 151-76. 

HoFFM.w.  G.  L.  1967.  Parasites  of  North  American 
freshwater  fishes.  University  of  California  Press, 
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Mackiewicz,  j.  S.  1963.  Monobothhum  httntcri  sp.  n. 
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Price,  C.  E.,  .and  H.  P.  .\r.\i.  1967.  The  monogenean 
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ScHELL,  S.  C.  1974.  Two  new  genera  and  three  new^  spe- 
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Threlfall,  W.,  a.nd  G.  Hanek.  1970.  Metazoan  para- 
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White.  G.  E.  1974.  Parasites  of  the  common  white  suck- 
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WiNGET,  R.  N.,  AND  R.  W.  Bal'm.a.nn.  1977.  Virgin  River, 
Utah-.\rizona-Nevada,  aquatic  habitat,  fisheries, 
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WiNGET,  R.  -M.,  A.ND  .M.  K.  Reichert.  1976.  .\quatic 
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77. 


SOIL  WATER  WITHDRWVAL  AND  ROOT  DISTRIBUTION  UNDER 
GRUBBED,  SPRAYED,  AND  UNDISTURBED  BIG  SAGEBRUSH  VEGETATION 

David  L.  Stiirges' 

.Vbstract.—  Seasonal  depletion  by  vegetation  where  sagebrush  was  selecti\elv  removed  bv  gnibbing  and  where 
sagebrush  was  sprayed  with  2,4-D  was  33  and  12  percent  less,  respectively,  than  that  for  undisturbed  big  sagebrush 
vegetation  in  the  surface  122  cm  of  soil.  Differences  were  located  primarilv  below  61  cm  in  vegetation  gnibbcd  the 
previous  fall  and  below  91  cm  in  vegetation  sprayed  three  years  previously.  Total  root  weights  under  gnibbed  and 
sprayed  vegetation  were  29  and  16  percent  less,  respectively,  than  for  unclisturbed  big  sagebrush  vegetation.  Total 
herbaceous  production  by  grubbed  and  sprayed  vegetation  was  69  and  43  percent  less,  respectiveh,  than  production 
l)\  undisturbed  vegetation. 


Big  sagebrush  [Artemisia  trident  at  a)  is 
commonly  controlled  with  herbicides,  me- 
chanical methods,  or  fire  to  increase  livestock 
forage  production.  Pheno.xy  herbicides  such 
as  2,4-D  damage  forbs  as  well  as  sagebrush, 
so  that  the  net  effect  of  spraying  is  to  favor 
grass  productivity.-  Burning  or  mechanical 
sagebrush  control  techniques,  however,  do 
not  selectively  favor  grasses.  Herbaceous  pro- 
duction commonly  doubles  or  triples  bv  the 
.second  or  third  year  after  sagebnish  removal. 

The  shift  from  a  shrub  to  a  herbaceous- 
dominated  vegetation  produces  other  ecolog- 
ic  and  hydrologic  changes.  This  studv  was 
made  to  quantify  differences  in  the  soil  water 
regime  and  in  root  biomass  between  undis- 
turbed big  sagebnish  vegetation  and  (a)  her- 
baceous vegetation  three  years  after  spraying 
with  2,4-D  and  (b)  herbaceous  vegetation 
from  which  only  big  sagebnish  was  removed 
by  mechanical  means  the  previous  fall.  Infor- 
mation about  herbaceous  productivity  was 
also  collected. 

LlTER.\TL  RE  ReVIEW 

Ghanges  in  the  soil  water  regime  after 
sagebrush  control  are  strongly  influenced  by 


rooting  characteristics  of  sagebnish  and  her- 
baceous species.  Roots  of  basin  big  sagebnish 
(A.  t.  sub.  tridcntata)  and  mountain  big  sage- 
brush (A.  t.  vaseyana)  commonly  extend 
about  2  m  deep  and  have  a  maximum  lateral 
spread  from  the  trunk  of  1.5  m  (Goodwin 
1956,  Cook  and  Lewis  1963,  Tabler  1964, 
Hull  and  Klomp  1974,  Sturges  and  Trlica 
1978).  Most  roots  are  in  surface  soil  where 
maximum  spread  occurs.  About  60  percent  of 
total  root  length  (Tabler  1964)  and  85  per- 
cent of  total  root  system  weight  were  present 
in  the  surface  61  cm  of  soil,  with  only  about 
4  percent  in  soil  below  91  cm  (Sturges  and 
Trlica  1978). 

The  principal  soil  water  reservoir  utilized 
by  isolated  mountain  big  sagebnish  plants  ex- 
tended 0.9  m  laterally  from  the  trunks  and 
0.9  m  deep  (Sturges  1977b).  Tlie  plants  utiliz- 
ed water  from  surface  soil  adjacent  to  the 
trunk  early  in  the  growing  season,  but  use- 
zones  shifted  outward  and  downward  later  in 
the  summer  as  water  adjacent  to  the  trunk 
was  depleted.  .Appreciable  water  uptake  was 
detectable  until  early  in  .\ugust. 

Tabler  (1968)  and  Sturges  (1977a)  found 
that  seasonal  soil  water  withdrawal  was  re- 
duced  after   spraying   sagebnish    vegetation 


'Rocky  Mountain  Forest  and  Range  Experiment  Station,  Laramie.  Wyoming  82070.  Central  headquarters  is  at  Fort  Collins  in  cooperation  with  Colorado 
State  University;  research  reported  here  was  conducted  at  the  station's  Research  Work  Unit  at  Laramie,  in  cooperation  with  the  University  of  Wyoming. 
Portions  of  the  research  were  supported  by  the  Bureau  of  l^nd  Management.  U.S.  Department  of  the  Interior. 

This  article  reports  research  involving  pesticides.  It  neither  contains  recommendations  for  their  use  nor  implies  that  the  uses  discussed  here  have  been 
registered.  .\ll  uses  of  pesticides  must  be  registered  by  appropriate  state  and/or  federal  agencies  before  they  can  be  recommended.  Use  all  pesticides  selecti- 
vely and  carefullv,  read  and  follow  the  directions  on  the  label. 


157 


158 


Great  Basin  Naturalist 


Vol.  40,  No.  2 


with  2,4-D  on  sites  with  deep  soils  that  were 
fully  recharged  by  snowmelt.  This  reduction 
was  located  almost  entirely  below  91  cm  as 
depletion  -of  surface  soil  water  by  sprayed 
vegetation  sometimes  exceeded  depletion  by 
untreated  vegetation.  Water  depletion  in  sur- 
face soil  increased  the  first  few  years  after 
treatment  as  herbaceous  vegetation  respond- 
ed to  release  from  sagebrush  competition. 
Other  studies  also  detected  an  increasing 
moisture  draft  from  surface  soil  with  time 
(Hyder  and  Sneva  1956,  Cook  and  Lewis 
1963,  Shown  et  al.  1972). 

Herbaceous  production  was  measured  in 
most  soil  moisture  studies.  Grass  production 
doubled  the  year  after  spraying  mountain  big 
sagebrush  and  was  2.6  times  higher  than  un- 
treated vegetation  three  years  after  treat- 
ment (Sturges  1977a).  Shown  et  al.  (1972) 
found  that  usable  forage  production  in- 
creased 300  percent  compared  to  pre- 
treatment  conditions  once  a  planted  grass 
stand  became  established.  Hyder  and  Sneva 
(1956)  found  the  increase  in  grass  production 
to  be  the  same  whether  big  sagebrush  was 
controlled  by  spraying  or  by  grubbing.  Total 
herbaceous  production  increased  the  most 
where  sagebrush  was  grubbed,  because  forbs 
were  damaged  by  the  spray. 


Study  Area 

The  study  was  performed  at  the  Stratton 
Sagebnish  Hydrology  Study  area  29  km  west 
of  Saratoga  in  south-central  Wyoming.  The 
experimental  site  is  at  an  elevation  of  2,225 
m  and  lies  on  a  north-facing  slope  in  a  mod- 
erate snow  catchment  zone.  Annual  precipi- 
tation is  about  500  mm,  with  two-thirds  of 
the  total  falling  as  snow.  Precipitation  be- 
tween 1  June  and  30  September  averages  114 
mm.  Sufficient  snow  usually  accumulates  to 
completely  recharge  the  soil  mantle.  Soils  de- 
veloped in  place  from  sandstone  and  belong 
to  the  Argic  Cryoboroll  great  soil  subgroup. 
A  dense  stand  of  mountain  big  sagebrush,  un- 
derlain by  a  productive  understory  of  bunch 
grasses-primarily  Idaho  fescue  (Festuca  ida- 
Jwensis),  bluegrass  (Poa  spp.)  and  needle- 
grasses  {Stipa  spp.)-was  present  before  study 
initiation.  The  site  had  been  grazed  by  sheep, 
but  no  grazing  occurred  during  the  study. 


Methods 

Work  began  in  1968  with  a  study  that  uti- 
lized 14  0.4-ha  experimental  units  arranged 
in  seven  blocks  to  determine  how  the  soil  wa- 
ter regime  would  be  affected  by  spraying  big 
sagebnish  (Sturges  1977a).  One  experimental 
unit  within  each  block  was  sprayed  with  2,4- 
D  in  1970;  the  other  unit  remained  un- 
treated. Experimental  units  from  three  of  the 
seven  blocks  were  used  in  the  current  study. 
In  October  1972,  smaller  plots  23  m  long  and 
10  m  wide  were  established  on  either  side  of 
the  common  border  between  sprayed  and  un- 
sprayed  vegetation  (Fig.  1).  These  plots  were 
used  to  obtain  soil  cores  and  to  create  the 
grubbed  sagebrush  vegetative  condition. 
Sagebnish  was  grubbed  from  four  circular 
areas  6.1  m  in  diameter  by  cutting  plants  at 
or  slightly  below  the  ground  surface.  Grubb- 
ing was  done  in  the  fall  of  1972,  when  vege- 
tation was  dormant,  thereby  minimizing 
damage  to  residual  herbaceous  vegetation 
and  insuring  that  herbaceous  vegetation 
would  be  as  comparable  as  possible  to  that 
within  the  undisturbed  sagebnish  stand  when 
study  measurements  began  the  following 
spring. 

Soil  Water  Measurements 

Soil  water  content  was  measured  with  a 
neutron-scattering  soil  moisture  meter  at  four 
randomly  located  access  tubes  on  each  exper- 
imental unit.  Access  tubes  within  grubbed 
vegetation  were  installed  at  the  center  of 
each  cleared  circle  in  October  1972.  If  one 
assumes  that  big  sagebnish  has  a  maximum 
lateral  root  spread  of  1.5  m,  these  tubes  were 
surrounded  by  a  volume  of  soil  at  least  1.5  m 
in  radius  devoid  of  live  sagebrush  roots. 

Moisture  measurements  began  31  May 
1973  upon  completion  of  snowmelt  and  con- 
tinued at  biweekly  intervals  until  19  Septem- 
ber 1973,  when  vegetation  was  dormant. 
Measurements  were  taken  at  eight  depths: 
15,  30,  46,  61,  76,  91,  107,  and  122  cm.  The 
manufacturer-supplied  calibration  curve  re- 
lating field  neutron  count  (expressed  as  a  per- 
centage of  shield  count)  to  volume  moisture 
content  was  applied  to  all  data  except  that 
collected  at  15  cm.  Here,  a  correction  was 
made  for  escape  of  neutrons  into  the  atmo- 


June  1980 


Sturges:  Big  Sagebrush 


159 


sphere  using  a  polyethylene  shield  technique 
similar  to  Pierpoint's  (1966). 

Root  Weights 

Soil  cores  for  sampling  root  weight  were 
obtained  at  four  random  locations  within  the 
small  plots  that  straddled  the  common  border 
between  sprayed  and  midisturbed  sagebrush 
vegetation.  The  cores  were  collected  in  Sep- 
tember following  the  final  soil  water  mea- 
surement (Fig.  2).  Each  core  was  7.6  cm  in 
diameter  and  122  cm  long.  The  cores  were 
collected  in  15-cm  increments  using  the  de- 
vice described  by  Brown  and  Thilenius 
(1977).  Each  sample  site  in  grubbed  vegeta- 
tion was  located  within  2.4  m  of  an  access 
tube,  a  minimum  of  0.6  m  from  the  surround- 
ing sagebrush  cover.  Soil  cores  were  placed 


in  plastic  bags  and  frozen  on  the  day  of  col- 
lection. After  thawing,  core  segments  were 
individually  washed  in  a  core-washing  ma- 
chine (Brown  and  Thilenius  1976)  to  isolate 
root  matter  from  soil.  Roots  were  oven  dried 
for  24  hours  at  70  C  and  weighed  on  an  ana- 
lytical balance.  It  was  not  possible  to  dis- 
tinguish between  live  and  dead  roots,  but 
woody  sagebrush  roots  from  cores  taken 
within  grubbed  vegetation  were  discarded 
before  samples  were  weighed. 

Herbaceous  Production 

Above-ground  herbaceous  productivity 
was  measured  by  clipping  12  randomly  lo- 
cated plots  within  each  experimental  unit  as 
grasses  matured  in  mid-July.  In  grubbed  veg- 
etation, three  production  plots  were  placed 


Undisturbed  sagebrush 
vegetation 


Sprayed  sagebrush 
vegetation 


10  m 


23nn 


64  nn 


64  nn 


X      Access  tube  on  0.4  ha  plot 
fj    Circular  area  where  sagebrush  grubbed 
+      Access  tube  in  grubbed  vegetation 


Fig.  1.  The  experimental  design  for  one  block  showing  the  0.4-ha  experimental  units  of  undisturbed  and  sprayed 
sagebrush  vegetation  and  smaller  plots  where  sagebrush  was  grubbed.  Soil  moisture  data,  soil  cores,  and  herbaceous 
productivity  information  for  the  grubbed  treatment  were  obtained  on  the  small  plot. 


160 


Great  Basin  Naturalist 


Vol.  40,  No.  2 


at  random  within  2.4  m  of  access  tubes.  Veg- 
etation was  harvested  to  a  1  cm  stubble 
height  from  plots  30.5  cm  wide  and  61  cm 
long.  Vegetative  matter  was  separated  into 
grass,  forb,  or  sagebnish  components  and 
placed  in  paper  bags  when  harvested.  Only 
leaves  and  herbaceous  stem  material  were  in- 
cluded with  sagebnish  herbage.  Vegetation 
samples  were  subsequently  dried  at  105  C  for 
24  hours  and  weighed. 

Selected  big  sagebrush  and  productivity 
characteristics  were  measvired  in  1969  on  the 
0.4-ha  experimental  units,  the  year  before 
spraying  (Table  1).  No  statistically  significant 
differences  before  treatment  were  present. 
Big  sagebrush  contributed  76  percent  of 
aboveground  herbaceous  production  while 
grasses  contributed  20  percent  and  forbs  4 
percent.  About  one-third  of  the  area  was  cov- 
ered by  the  live,  leafy  portion  of  the  sage- 
bnish canopy.  Sagebrush  plants  had  an  aver- 
age height  of  34  cm  and  an  average  crown 
area  of  7  dm-. 

Statistical  Analysis 

Soil  water  withdrawal  and  root  weight  dif- 
ferences among  the  three  vegetative  condi- 
tions were  tested  for  statistical  significance 
by  variance  analysis  utilizing  a  split-plot  de- 
sign. Experimental  units  (whole  units)  were 
arranged  in  three  randomized 'blocks,  and  the 
eight  measurement  depths  served  as  subunits. 
Analyses  were  based  on  average  plot  values 
determined  from  the  four  replicated  mea- 
surements on  the  plot.  Variables  analyzed 
were  the  change  in  soil  water  content  be- 
tween successive  sampling  dates,  the  seasonal 
change  in  soil  water  content,  and  root 
weight.  Herbaceous  productivity  data  were 
analyzed  with  a  randomized  block  design. 

Results 

Soil  Water  Depletion 

Soil  under  undisturbed  and  sprayed  vegeta- 
tion was  completely  recharged  by  snowmelt 
on  the  first  measurement  date,  but  only  to  61 
cm  under  grubbed  vegetation  (Fig.  3).  At  the 
end  of  summer,  water  content  in  the  surface 
46  cm  of  .soil  was  similar  for  all  treatments. 
Below  46  cm,  progressively  more  water  re- 


mained in  soil  under  grubbed  vegetation 
compared  to  undisturbed  sagebnish  vegeta- 
tion, but  appreciable  differences  between 
sprayed  and  undisturbed  vegetation  were 
present  only  below  91  cm. 

Seasonal  water  withdrawal  by  undisturbed, 
sprayed,  and  grubbed  vegetation  was  24.3, 
21.4,  and  16.2  cm  of  water,  respectively,  in 
the  surface  122  cm  of  soil.  These  differences 


Table  1.  Characteristics  of  vegetation  on  plots  as- 
signed to  sprav  and  undisturbed  treatments  in  1969,  one 
vear  before  2,4-D  was  applied. 


Sagebrush 

Undisturbed 

Sprayed 

Height  (cm) 

30° 

37 

Canopy  area  (dm') 

6 

8 

Canopy  intercept  (%) 

31 

32 

Density  (number/ha) 

57,(X)0 

52,000 

Hebbac;eous  production 

(kg /ha) 

Sagebrush 

926 

1095 

Grass 

249 

290 

Forb 

53 

60 

Total 

1228 

1445 

"Differences  between  treatment  means  were  not  significant  for  any  mea- 
surement parameter  at  the  0.05  level  of  probability. 


tig.  2.  Root  samples  were  obtained  using  a  core  sam- 
pler driven  into  the  soil  to  a  122-cm  depth  by  15-cm  in- 
crements. 


June  1980 


Sturges:  Big  Sagebrush 


161 


were  significant  at  the  0.01  probability  level. 
Treatment  differences  did  not  accrue  uni- 
formly through  the  soil  mantle,  but  were  con- 
centrated at  deeper  soil  depths  (Fig.  4).  Be- 
tween 91  and  122  cm,  depletion  by  grubbed 
and  sprayed  vegetation  was  31  and  66  per- 
cent, respectively,  of  depletion  by  undis- 
turbed sagebnish  vegetation. 

Treatment   soil   water   withdrawal    differ- 


ences between  consecutive  measurement 
dates  were  significant  (p<0.05)  only  between 
25  June  and  10  July.  The  treatment  x  depth 
interaction  term  was  significant  during  five 
of  the  eight  measurement  intervals,  though, 
indicating  that  the  three  vegetative  condi- 
tions were  utilizing  water  differently  from 
within  the  soil.  For  example,  most  of  the  dif- 
ference in  depletion  below  91  cm  between 


1 20 

CL 

.^  0 

D 
Q 


Undisturbed 

Sprayed 

Grubbed 


76  cm 


91cm 


46  cm 


107cm 


61cm 


122  cm 


-J I 1 i_ 


ilLlJI IlL 


15    31    15   30 


li    .J  .1 


I  nil.  I 


-Id L 


5   31     15    31    15   30     15    31    15   30 


I  III..  I    .1  I    1. L 


5    31    15    31    15  30 


May      June       July       Aug.     Sept.  May     June       July       Aug.     Sept. 


Fig.  .3.   Daily  precipitation  and  soil  water  content  in  the  surface   122  cm  of  soil  for  undisturbed,  sprayed,  and 
grubbed  .sagebnish  vegetation  in  the  197.3  growing  season. 


162 


Great  Basin  Naturalist 


Vol.  40,  No.  2 


sprayed  and  undisturbed  vegetation  devel- 
oped after  25  July.  Sagebrush  remained  phys- 
iologically active  through  the  summer  and 
flowered  about  1  September,  so  that  appre- 
ciable water  usage  continued  all  summer. 
Most  grass  and  forb  species  had  matured  and 
set  seed  by  early  August,  thus  reducing  the 
need  for  water  by  grubbed  and  sprayed  vege- 
tation. 

Root  Weights 

The  average  weight  of  roots  obtained  from 
soil  cores  extending  122  cm  deep  was  12.2, 
10.2,  and  8.7  g  under  undisturbed,  sprayed, 
and  grubbed  vegetation,  respectively.  Nei- 
ther the  treatment,  nor  the  depth  x  treatment 
interaction  term  was  statistically  significant. 
Varying  quantities  of  dead  but  undecayed 
root  matter  and  other  organic  debris  were  in- 
cluded with  sample  material  and  could  not 
be  separated  from  live  roots.  Inclusion  of  ex- 
traneous matter  probably  accounted,  in  part, 
for  the  low  statistical  sensitivity  of  root  mea- 
surements. 

Most  of  the  weight  of  roots  was  located  in 
surface  soil  (Fig.  4).  Material  from  the  surface 
15  cm  of  soil  ranged  from  36  percent  of  total 
root  weight  in  imdisturbed  sagebRish  vegeta- 
tion to  54  percent  of  total  root  weight  in 
sprayed  vegetation.  Conversely,  only  1  to  2 
percent  of  root  weight  for  each  treatment 
came  from  the  deepest  sampling  depth. 

Herbaceous  Production 

Herbaceous  production  of  undisturbed 
sagebrvish  vegetation  was  about  a  third  great- 
er in  1973  than  in  1969,  but  composition  of 
vegetation  was  similar  both  years.  Treatment 
differences  within  sagebrush,  grass,  and  total 
production  herbage  classes  were  highly  sig- 
nificant (Table  2).  The  response  by  sprayed 
vegetation  the  third  year  after  treatment  was 
typical  to  that  reported  from  other  locations. 
Grass  production  was  2.6  times  greater  than 
production  in  imdisturbed  sagebmsh  vegeta- 
tion, but  forb  production  was  still  depressed 
below  pretreatment  levels.  Total  herbaceous 
production  by  sprayed  vegetation  was  only 
57  percent  as  large  as  production  by  undis- 
turbed sagebnish  vegetation,  the  increase  in 
grass  production  not  compensating  for  loss  of 
sagebrush. 


Grass  production  increased  27  percent 
where  sagebrush  was  grubbed  the  previous 
fall,  but  the  increase  was  not  statistically  sig- 
nificant (Table  2).  Total  production  was  31 
percent  as  high  as  that  by  undisturbed  vege- 
tation because  of  the  loss  of  sagebrush. 


Discussion  and  Conclusions 

This  study  indicates  the  soil  water  regime 
in  the  surface  91  cm  of  soil  is  unaffected  by 
sagebrush  control  once  herbaceous  vegeta- 
tion responds  to  release  from  sagebrush  com- 
petition. However,  below  91  cm,  substantial 
reductions  in  seasonal  withdrawal  can  occur 
as  reported  by  Tabler  (1968)  and  Sturges 
(1977a).  The  overall  reduction  in  soil  water 
depletion  caused  by  grubbing  sagebrush  com- 
pares closely  with  that  detected  on  the  same 
0.4-ha  experimental  units  in  1970  when  sage- 
bnish was  sprayed.  Grubbing  decreased  sea- 
sonal water  withdrawal  33  percent  in  this 
study,  and  spraying  reduced  withdrawal  from 
the  surface  137  cm  of  soil  37  percent  (from 
the  spray  date  on  22  June  through  30  Sep- 
tember). The  year  after  spraying,  a  17  per- 
cent difference  in  seasonal  withdrawal  was 
observed  with  grass  production  doubling  in 
response  to  sagebrush  removal. 

Reductions  in  moisture  withdrawal  are  re- 
lated to  decreased  aboveground  herbaceous 
productivity  of  treated  vegetation.  Produc- 
tivity in  grubbed  and  sprayed  vegetation  was 
31  and  57  percent  as  large,  respectively,  as 
that  of  undisturbed  vegetation.  Development 
of  vegetation  in  years  immediately  following 
sagebrush  control  also  influenced  water  with- 
drawal patterns.  Seasonal  depletion  under 
gnibbed  vegetation  was  less  than  that  of  un- 
disturbed sagebnish  vegetation  at  all  depths, 
but  appreciable  differences  existed  only  be- 


Table  2.  Aboveground  herbaceous  production 
(kg/ha)  by  undisturbed,  sprayed,  and  grubbed  vegeta- 
tion in  1973. 


Treatment    Sagebnish 

Grass 

Forb 

Total 

Undisturbed       °120P 
Spraved                      1^ 
C;  rubbed                    (1*^ 

347* 
918^ 
442'' 

86* 
Ifif' 
67* 

1634* 
93^ 
509^" 

"Treatment  means  having  different  letters  within  a  column  are  signifi- 
cantly different  at  the  0.05  level  o^probability. 


June  1980 


Sturges:  Big  Sagebrush 


163 


o 
o 
cr 


0-15       15 


,;yyw^.i...»,.i„.iiiiii„„„„„„„„y,^.. . 

^;M.  npot  weight  •mi       Depletion  Undisturbed 

_|  Sprayed 


t i 


Grubbed 


15-30    30 


30-46    46 


E     46-61       61 
o 


IP 

Q. 

Q 


61-76     76 


76-91      91 


91-107    107 


107-122    122 


I  2  3 

Root  weight  (g) 
Seasonal  water  depletion/15  cm  soiKcnn] 


Fig.  4.  Seasonal  water  depletion  and  weight  of  roots  in  soil  cores  7.6  cm  in  diameter  and   122  cm  long  unde 
undisturbed,  sprayed,  and  grubbed  sagebrush  vegetation. 


164 


Great  Basin  Naturalist 


Vol.  40.  No.  2 


low  61  cm.  Sprayed  vegetation,  unlike 
grubbed  vegetation,  had  fnlh-  responded  to 
release  from  sagebrush  competition  and 
depletion  did  not  become  appreciabK  less 
tlian  that  of  imdistiu-bed  vegetation  mitil  a 
91-cm  depth  was  reached.  Reductions  in 
treatment  effect  through  time  within  soil 
60-90  cm  deep  were  described  by  Hyder  and 
Sneva  (1955),  Cook  and  Lewis  (1963),  and 
Shown  et  al.  (1972). 

The  reduction  in  seasonal  water  use  and  in 
root  weight  caused  b\'  treatments  are  similar 
when  e.vpressed  as  a  percentage  of  values  for 
undisturbed  vegetation.  Seasonal  depletion 
was  33  and  12  percent  less  for  grubbed  and 
spraved  vegetation,  respectively,  and  root 
weights  were  29  and  16  percent  smaller  on 
these  same  treatments.  Similar  agreement  be- 
tween depletion  and  root  weight  did  not  exist 
for  individual  measurement  depths  (Fig.  4). 
Thus,  root  weight  measurements  do  not  veri- 
f\'  or  refute  the  hypothesis  that  root  devel- 
opment by  herbaceous  species  in  the  surface 
90  cm  of  soil  subsequent  to  sagebrush  remov- 
al accoimts  for  increases  in  moisture  use  from 
this  zone.  Measurement  of  root  length,  rather 
than  root  weight,  probably  would  have  pro- 
vided a  better  measure  of  potential  moisture 
draft  because  of  the  differences  in  morpho- 
logy of  grass  and  sagebrush  roots. 

Comparisons  of  seasonal  moisture  change 
and  root  weight  with  depth  does  indicate 
that  deep  roots  are  extremely  important  in 
extracting  soil  water,  even  though  they  com- 
prised a  small  part  of  root  weight  in  soil 
cores  (Fig.  4).  Summer  precipitation  is  usual- 
ly ineffective  in  replenishing  soil  water  levels 
in  the  sagebrush  zone,  so  that  deeper  soil  be- 
comes an  important  water  reservoir  when 
surface  soil  dries.  A  progressive,  downward 
shift  of  major  water  use  zones  in  August  was 
especially  evident  for  undisturbed  sagebrush 
vegetation  (Fig.  3). 

Results  of  this  and  other  soil  water  deple- 
tion studies  indicate  that  control  of  big  sage- 
bru.sh  with  methods  that  do  not  destroy  all 
vegetation  on  lands  with  an  adequate  popu- 
lation of  herbaceous  species  has  a  relatively 
small  effect  upon  the  soil  water  regime. 
Changes  in  the  soil  water  regime  can,  at 
most,  result  in  small  increases  of  streamflow. 


This  response  will  onl\  t)ccur  on  lauds  where 
soils  are  deeper  than  90  cm  and  soil  water 
recharge  exceeds  that  retiuired  to  fulK  wet 
the  soil  mantle.  The  maxinuuu  reduction  in 
depletion  will  usually  occur  in  the  treatment 
\ear  because  of  productivity  increases  bv 
herbaceous  species  in  xears  immediatelv  after 
treatment.  Consecjuentlv.  justification  for  big 
sagebiiish  control  must  rest  on  the  benefits 
derived  from  shifting  site  resources  to  species 
more  desirable  than  sagebrush  from  a  given 
land  management  perspective. 

Literature  Cited 

Brown,  G.  R.,  and  J.  F.  Thilenius.  1976.  A  low-cost 
machine  for  separation  of  roots  from  soil  mate- 
rial. J.  R;mge  Manage.  29;506-507. 

1977.  .\  tool  and  method  for  extracting  plant- 
root-soil  cores  on  remote  sites.  J.  Range  Manage. 
30:72-74. 

Cook,  C.  W.,  .\nd  C.  E.  Lewis.  1963.  Competition  be- 
tween big  sagebrush  and  seeded  grasses  on  foot- 
hill ranges  in  Utah.  J.  Range  Manage. 
16:245-250. 

Goodwin.  D.  L.  1956.  .\iitecological  studies  oi  Artemisia 
tridentata  Nutt.  Unpublished  dissertation.  Wash- 
ington State  Univ.,  Pullman.  79  pp. 

Hi  LL,  A.  C,  Jr.,  .and  G.  J.  Klomp.  1974.  Yield  of  crested 
wheatgrass  under  four  densities  of  big  sagebnish 
in  southern  Idaho.  U.S.  Dept.  Agric.  Tech.  Bull. 
No.  1483.  38  pp. 

Hyder,  D.  N.,  a.nd  F.  A.  Sneva.  1956.  Herbage  response 
to  sagebrush  spraying.  J.  Range  Manage.  9:34-.38. 

Pierpoi.nt,  G.  1966.  Measuring  surface  soil  moisture 
with  the  neutron  depth  probe  and  a  surface 
shield.  Soil  Sci.  101:189-192. 

Shown.  L.  M.,  G.  C.  Lusby,  and  F.  A.  Branso.n.  1972. 
Soil-moisture  effects  of  conversion  of  sagebrush 
cover  to  bunchgrass  cover.  Water  Resour.  Bull. 
8:1265-1272. 

Sti  rc;es,  D.  L.  1977a.  Soil  moisture  response  to  spraying 
big  sagebnish:  A  seven-year  study  and  literature 
interpretation.  USD.\  For.  Serv.  Res.  Pap.  R.\i- 
188.  12  pp.  Rocky  Mt.  For.  and  Range  Expt.  Stn.. 
Fort  Collins,  Colo. 

19771).  Soil  water  withdrawal  and  root  character- 
istics of  big  sagebrush.  Am.  Midi.  Nat. 
98:257-274. 

SriRCES,  D.  L.,  AND  .M.  J.  Trlica.  1978.  Root  weights 
and  carbohydrate  reserves  of  big  sagebrush.  Ecol- 
ogy 59:1282-1285. 

Tabi.er,  R.  D.  1964.  The  root  system  oi  Arteniisia  triden- 
tata    at     9,500     feet     in     Wyoming.     Ecology' 
45:633-636. 
.  1968.  .Soil  moisture  response  to  spraying  big  sage- 
brush with  2,4-D.  J.  Range  Manage.  21:12-15. 


SWARMING  OF  THE  WESTERN  HARVESTER  ANT. 
POGOXOMYRMEX  OCCIDEXTALIS 

Dorald  M.  Allred' 


,\bstract.—  The  swarming  and  mating  of  harvester  ants  was  observed  in  Utah  in  July  1979.  Workers  groom  the 
alate  forms  outside  the  mound  before  swarming  occurs  and  are  highly  aggressive  in  protecting  them.  Mating  pairs 
apparently  are  not  disturbed  by  other  ants.  Fertile  females  likely  use  moving  vehicles  that  extend  their  dispersal. 


Few  detailed  accounts  of  the  swarming  ac- 
tivities of  harvester  ants  are  existent  in  the 
literature.  Wheeler  (1910)  reported  the 
swarming  of  harvester  ants  in  the  desert 
along  the  Colorado  River.  Michener  (1942, 
1948)  noted  swarming  and  mating  of  Pogono- 
mymiex  califomicus  and  P.  barbatus.  Strandt- 
mann  (1942)  recorded  the  mating  activities  of 
P.  Comanche,  and  Chapman  (1957)  reported 
elevational  swarming  of  P.  occidentalis  on 
mountain  tops  in  five  states. 

During  the  latter  part  of  June  and  all  of 
July  in  1979,  I  traveled  extensively  over  Utah 
collecting  ants.  In  most  cases  when  harvester 
ants  were  taken,  I  partially  excavated  each 
mound  from  which  I  collected  to  determine 
the  presence  of  immature  and  winged  forms. 
Although  winged  males  and  females  were 
present  in  the  majority  of  the  moimds  during 
this  period,  swarming  was  not  seen  until  the 
latter  part  of  July. 

On  July  23  I  stopped  at  about  11:30  a.m. 
(Mountain  Daylight  Time)  to  collect  from  an 
area  of  abimdant.  large  mounds  one  mile 
west  of  Elberta.  Utah  County,  Utah,  along- 
side highway  US6  at  an  elevation  of  5400  ft 
in  a  sagebrush-rabbitbmsh  habitat  [Artemisia 
tridentata-Chrysothamnus  nauseosus).  -\s  I 
approached  a  large  moiuid  on  that  wanii, 
sunny  day,  an  area  of  one-half  square  foot 
around  two  enlarged,  south-facing  openings 
was  literally  red  with  a  mixture  of  \\orkers, 
winged  males  and  females.  I  would  have  had 
difficulty  finding  an  open  space  within  tlie 
mass  of  ants  where  I  could  ha\'e  touched  the 


ground  with  a  pencil.  The  winged  forms 
were  relatively  inactive,  none  in  flight,  and 
the  workers  seemed  to  be  grooming  and  at- 
tending them.  As  I  approached  the  moumd  to 
aspirate  a  sample  of  ants,  the  majority  of  the 
winged  females  and  some  males  quicklv  en- 
tered the  nest  openings.  Some  of  the  males, 
however,  remained  immobile  outside  the 
mound  as  though  mesmerized  bv  the  sToom- 
ing  activities  of  the  workers.  The  workers  on 
the  fringe  of  the  mass  immediatelv  began  ag- 
gressive tactics  toward  me  in  much  more  of  a 
frenzied  movement  than  I  had  heretofore  en- 
countered with  workers  when  winged  forms 
were  not  outside  the  nest. 

After  I  had  taken  my  sample.  I  returned  to 
the  car  to  record  the  data.  Five  minutes  later 
I  again  visited  the  mound  to  see  if  the 
winged  forms  had  left  the  burrow.  The  air 
was  filled  with  fl\ing  ants.  At  the  mound  the 
workers  were  no  longer  congregated  around 
the  openings,  but  were  scurrvin^  around, 
near,  and  over  the  mound.  .\  few  winded 
forms  were  crawling  around  on  the  mound.  I 
was  quickly  deluged  with  flNing  ants,  and  a 
sting  on  my  leg.  presimiablv  from  a  winged 
female,  stimulated  m\"  hastv  retreat  to  the 
car.  where  I  quickly  closed  all  windows.  The 
outside  of  the  car  was  soon  covered  with 
winged  ants  that  were  mating.  Females 
seemed  to  be  much  more  abundant  than 
males.  This  was  consistent  \\ith  m\  findings 
wherever  I  had  excavated  mounds  through- 
out the  state  during  Jime  and  July. 

In  mating,  the  male  moimted  the  female 


'Department  of  Zcxilog),  Bricham  Vount;  I'niversit) .  Privo,  I'tah  S46Q2. 


165 


166 


Great  Basin  Naturalist 


Vol.  40,  No.  2 


dorsally,  clasped  her  around  the  thorax  with 
his  legs,  and  bent  his  abdomen  strongly 
downward  to  contact  her  gentitalia.  She  si- 
multaneously bent  her  abdomen  slightly  up- 
wards to  facilitate  contact.  Once  joined,  the 
partners  sometimes  assumed  different  posi- 
tions than  described  above,  frequently  both 
establishing  leg  contact  with  the  substrate  on 
which  they  were  resting,  although  maintain- 
ing abdominal  junction.  Other  crawling  ants 
frequently  came  in  contact  with  a  mating 
pair,  but  the  contact  was  brief.  At  no  time 
did  I  observe  other  males  remaining  with 
mating  pairs  as  described  by  Strandtmann 
(1942)  for  P.  Comanche.  Copulation  lasted  for 
perhaps  20  to  40  seconds,  whereupon  the  two 
sexes  immediately  separated,  with  the  male 
the  first  to  fly  away.  No  case  was  observed 
where  either  the  male  or  female  used  its 
mandibles  to  grasp  or  chew  on  the  other  as 
described  by  Strandtmann  (1942)  for  P.  Com- 
anche, and  by  Michener  (1948)  for  P.  har- 
batus. 

About  12:.30  p.m.,  after  I  had  observed  the 
mating  activities  of  numerous  pairs  and  the 
numbers  of  ants  crawling  on  the  car  and  fly- 
ing in  the  area  had  considerably  diminished,  I 
returned  to  the  mound.  Few  flying  ants  re- 
mained in  the  air  in  its  vicinity,  but  a  few 
winged  forms  were  crawling  around  the 
mound.  Most  of  the  workers  had  reentered 
the  entrances,  and  those  that  remained  out- 
side had  apparently  resumed  their  normal, 
slower  speed  of  routine  activity. 

I  moved  to  another  area  of  several  mounds 
situated  about  25  yards  north  of  the  site  of 
my  previous  observations.  The  ants  of  several 
of  these  mounds  were  in  various  stages  of 
swarming  activities.  By  periodical  rotation 
between  these,  and  with  some  excavations,  I 
was  able  to  summarize  the  overall  activities 
associated  with  swarming  of  this  species. 

When  the  males  and  females  prepare  to 
swarm,  the  workers  enlarge  the  openings 
leading  from   the   mounds,   the  winged  ants 


and  many  workers  leave  the  nest  and  congre- 
gate around  its  openings.  The  winged  males 
and  females  are  attended  by  myriads  of 
workers  who  groom  them  as  their  bodies 
warm  in  the  sun,  and  at  the  same  time  act  in 
a  protective  capacity  in  relation  to  any 
would-be  predators.  Once  the  mating  flight 
begins,  the  workers  disperse  and  crawl 
around  the  mound  for  a  few  minutes,  then  re- 
enter the  burrow.  Normal  worker  activity 
outside  the  mound  is  resumed  within  a  few 
minutes  after  the  majority  of  the  winged 
forms  have  departed.  A  few  winged  forms 
seem  to  linger  around  the  mound,  apparently 
hesitant  to  leave.  Some  also  delay  leaving  the 
burrow  to  assume  their  flight  of  destiny. 
Swarming  males  and  females  seek  some  high 
point  to  mate,  but  many  pairs  mate  on  the 
ground. 

After  mating,  females  migrate  in  all  direc- 
tions to  locate  sites  for  establishing  new  colo- 
nies. Undoubtedly  some  of  them  hitch  rides 
on  moving  conveyances  such  as  cars,  trucks, 
and  trains.  When  I  arrived  at  my  home  in 
Provo,  30  miles  from  and  several  hours  after 
observing  the  swarming  activity,  a  live 
winged  female  dropped  from  my  car  onto  the 
driveway  and  crawled  into  the  vegetation. 
Such  methods  of  conveyance  likely  extend 
the  dispersal  of  females  over  relatively  great 
distances. 

Literature  Cited 

Chapman,  J.  A.  1957.  A  further  consideration  of  summit 
ant  swarms.  Canadian  Ent.  89:389-95. 

Michener,  C.  D.  1942.  The  history  and  behavior  of  a 
colony  of  harvester  ants.  Scientific  Monthly 
55:248-58. 

1948.  Observations  on  the  mating  behavior  of 

harvester  ants.  J.  New  York  Ent.  Soc.  56:239-42. 

Str.\ndtmann,  R.  W.  1942.  On  the  marriage  flight  of  Po- 
go(io»H/n)U'.v  comanclic  Wheeler.  Ann.  Ent.  Soc. 
America  .35:140. 

Wheeler,  W.  M.  1910.  Ants:  Their  structure,  devel- 
opment, and  behavior.  Columbia  Univ.  Press, 
New  York,  663  pp. 


RELATIONSHIP  BETWEEN  ENVIRONMENTAL  AND  VEGETATIONAL 
PARAMETERS  FOR  UNDERSTORY  AND  OPEN-AREA  COMMUNITIES 


William  E.  Evensoii',  Jack  D.  Brothersoir,  and  Richard  B.  Wilcox' 

.XbstR'VCT.—  Ten  individnals  from  each  of  four  tree  species  were  selected,  and  their  associated  understory  and  ad- 
jacent open-area  communities  were  sampled  for  both  environmental  and  vegetational  parameters,  including  light 
intensity,  pH,  litter  depth,  soil  depth,  and  percentages  of  exposed  rock,  litter  cover,  living  cover,  shrubs,  forbs, 
grasses,  and  annuals.  The  four  tree  species  were  ponderosa  pine.  Rocky  Mountain  juniper,  Gambel  oak,  and  snow- 
linish  ceanothus.  The  studv  site  was  in  the  lower  Uinta  Mountains  about  10  miles  east  of  Kamas,  Utah.  Correlations 
among  the  various  biotic  and  abiotic  parameters  were  examined.  The  interplay  of  these  factors  in  differentiating  the 
understory  and  open-area  communities  is  discussed. 


Understanding  the  relationship  of  vegeta- 
tional  patterns  to  environment  is  a  primary 
goal  of  community  ecology.  One  aspect  of 
such  relationships  is  the  effect  of  overstory 
trees  and  shRibs  on  their  associated  under- 
story communities.  In  a  previous  report  (Wil- 
cox, Brotherson,  and  Evenson  1981),  we  ex- 
amined the  influence  of  four  canopy  species 
on  their  associated  imderstory  plant  commu- 
nities in  comparison  to  neighboring  commu- 
nities in  open  areas  outside  the  canopy  in- 
fluence. The  four  canopy  species  were 
ponderosa  pine  {Piniis  ponderosa  Dougl.), 
Rocky  Mountain  juniper  {Junipenis  scopulo- 
nim  Sarg.),  Gambel  oak  {Quercus  gambelii 
Nutt.),  and  snowbrush  ceanothus  (Ceanothus 
vehitinus  Dougl.). 

Many  previous  studies  have  reported  envi- 
ronmental and  vegetational  differences  be- 
tween understories  and  open  areas.  Light  in- 
tensity (del  Moral  1972,  Cline  1966, 
Blackman  1956)  and  spectral  distribution 
(Federer  and  Tanner  1966)  are  known  to 
strongly  differentiate  understory  and  open- 
area  plant  communities.  Soil  moisture  and 
thickness  of  litter  layer  are  also  important 
factors  (Anderson  1969,  McQueen  1973),  as  is 
soil  improvement  due  to  nitrogen  fixation  by 
such  common  understorv  plants  as  bitter- 
bmsh  {Pursliia  tridentata  (Pursh)  DC.)  and 
snowbrush   ceanothus   (Wollam   and   Young- 


berg  1964,  Rusel  and  Evans  1966,  Webster, 
Youngberg,  and  Wollam  1967). 

Because  these  and  other  environmental 
modifications  are  influenced  by  the  canopy 
species,  cover  (Anderson  1969,  McQueen 
1973,  McConnell  and  Smith  1970)  and  diver- 
sity (Auclair  and  Goff  1971)  of  understory 
communities  are  strongly  dependent  on  the 
canopy  tree  or  large  hebaceous  species  with 
which  they  are  associated  (Gordon  1962, 
Smith  and  Cottam  1967). 

The  present  study  examines  detailed  rela- 
tionships between  the  various  environmental 
and  vegetational  parameters  measured  under 
the  canopies  of  fovir  tree  species  and  in 
nearby  open  areas. 

Study  Area 

The  study  site  is  about  10  miles  east  of 
Kamas,  Utah,  along  the  Yellow  Pine  branch 
of  Beaver  Creek  (Fig.  1).  This  area  was  cho- 
sen because  of  the  homogeneity  of  the  under- 
lying parent  material  (an  alluvial  outwash 
gravel  bed)  throughout  the  site,  its  constant 
slope  and  exposure,  and  its  easy  accessibility. 
The  study  site  is  an  area  of  "zone  jumbling" 
(Cottam  1930)  and  contains  plant  representa- 
tives from  all  life  zones  except  lower  sonoran. 
It  is  an  area  of  highlv  mixed  vegetation, 
varying  from  Douglas  fir  {Pseudotsuga  men- 


'Department  of  Physics.  Brigham  Young  I'niversity,  Provo,  Utah  84602. 

'Department  of  Botany  and  Range  Science.  Brigham  Young  University,  Provo,  Utah  84602. 

'Land  Specialist,  State  of  Utah,  Room  440,  Empire  Buildnig,  231  East  400  South,  Salt  Lake  City,  Utah  84111. 


167 


168 


Great  Basin  Naturalist 


Vol.  40,  No.  2 


ziesii  (Mirb.)  Franco.),  white  fir  (Abies  con- 
color  Lindl.),  and  ponderosa  pine  to  clumps 
of  Gambel  oak  and  snowbnish  ceanothus. 
Also  interspersed  throughout  the  area  are  in- 
dividuals of  lodgepole  pine  (Pinus  contorta 
Dougl.  ex  Loud.),  Rocky  Mountain  juniper, 
quaking  aspen  (Populus  tretnuloides  Michx.), 
and  various  other  plant  species.  All  can  be 
found  at  the  same  elevation  and  in  fairly 
close  proximity. 


Because  of  its  apparent  uniformity,  this  site 
is  especially  well  suited  to  measure  the  rela- 
tionship of  environmental  and  vegetational 
parameters  associated  with  understories  of 
different  tree  species  and  nearby  open  areas. 
In  such  an  area,  the  likelihood  of  factors  oth- 
er than  tree  overstory  affecting  such  relation- 
ships in  a  major  way  is  small. 


"^76  00 


y7(.SS 


Sfe 


iSrW 


Yellow  Pine 
Campground 


Fig.  1.  Map  showing  location  of  stiidv  site. 


June  1980 


EVENSON  ET  .\L.:  PlANT  EcOLOCY 


169 


Methods 

Ten  individuals  of  each  of  four  tree  or 
shrub  species  (i.e.,  ponderosa  pine,  Rocky 
Mountain  juniper,  Gambel  oak,  and  snow- 
bnish  ceanothus)  were  chosen  at  random  in 
the  study  area.  Eight  quadrats  (0.25  m^)  were 
placed  aroimd  each  individual  tree  or  .shrub. 
Four  of  these  quadrats  were  placed  inside  the 
cylinder  of  the  canopy,  and  four  were  placed 
outside  the  influence  of  the  canopv.  To  elimi- 
nate bias,  quadrats  were  consistently  placed 
one  at  each  direction  of  the  compass.  Quad- 
rats were  subdivided  into  four  equal  units  for 
species  frequency  measurements.  Sample 
trees  and  quadrat  sites  were  marked  for  relo- 
cation. 

Presence  or  absence  of  individual  plant 
species  in  the  understory  was  determined  for 
all  four  subunits  of  each  quadrat.  All  species 
rooted  in  the  quadrat  were  recorded.  Fre- 
quency of  each  plant  species  was  determined 
by  dividing  the  number  of  quadrat  sub- 
divisions in  which  a  species  occurred  by  the 
number  of  subdivisions  sampled.  Total  living 
plant  cover  and  composition  of  plant  cover 
by  life  form  were  measured  at  each  quadrat 
using  an  ocular  estimate  method  (Daviben- 
mire  1959). 

Light  intensity  was  measured  in  foot-can- 
dles at  each  quadrat,  and  averages  were  com- 


puted for  the  understory  and  open-area  quad- 
rats associated  with  each  tree  species.  All 
readings  were  taken  between  1200  and  1400 
hours  on  cloud-free  days,  the  last  two  days  of 
the  study. 

Soil  pH  was  measured  by  the  colormetric 
method  in  the  field  to  avoid  pH  changes 
which  can  occur  when  soil  is  stored  moist. 

Litter  depth  was  measured  at  the  center  of 
all  quadrats  taken,  and  soil  depth  was  deter- 
mined by  the  average  of  five  penetrometer 
readings  in  each  quadrat  (one  at  each  corner 
and  one  in  the  center).  Correlations  of  all 
variables  with  each  other  were  nm. 


Results  .4nd  Discussion 

Characteristics  of  the  environment  and  the 
vegetation  types  associated  with  understory 
and  open-area  communities  are  summarized 
in  Table  1  for  the  four  canopy  species.  As  ex- 
pected, light  intensity  values  are  consistently 
lower  for  the  imderstory  communities.  Un- 
derstory communities  have  consistently  high- 
er pH  (more  basic),  except  for  ponderosa 
pine  which  shows  an  understory  tending  to 
be  slightly  more  acidic  than  the  open  area. 
Both  litter  depth  and  soil  depth  are  far  great- 
er in  tlie  imderstory  communities  than  out- 
side the  canopies.  Percent  cover  of  litter  is 


Table   1.  Average  measured  values  of  environmental  and  vegetation  parameters  for  understorv  and  open  area 
comnumities  ±  their  standard  deviations. 


Pintis 

Ceanothus 

Juniper  us 

Quercus 

ponderosa 

vchttintts 

scopulortim 

gamhelii 

Under- 

Open 

Under- 

Open 

Under-          Open 

Under- 

Open 

story 

areas 

story 

areas 

story            areas 

story 

areas 

Light  intensity 

(foot  candles) 

117  ±52 

225  ±65 

123  ±29 

.3.39  ±50 

168  ±77      295  ±67 

76  ±.32 

242  ±67 

Soil  pH 

6.3  ±0.2 

6.4  ±0.1 

6.6  ±0.1 

6.4  ±  0. 1 

7.4  ±0.6      6.4  ±0.2 

6.6  ±  0.3 

6.4  ±0.2 

Litter  depth  (cm) 

6.0±2.7 

0.6  ±0.3 

2.1  ±0.7 

0.1±0.1 

2.5  ±0.3      0.3  ±0.2 

2.3  ±  0.6 

0.2  ±0.2 

Soil  depth  (cm) 

10.5  ±2.6 

5.3±1.1 

6.7±1.7 

4.4  ±1.2 

6.1  ±.3.2      4.0  ±1.6 

8.5  ±2.3 

.5.7  ±0.8 

%  Living  cover 

20.2  ±6.0 

25.8  ±6.8 

20.0  ±4.7 

31.5  ±7.6 

28.0  ±19.1  .32.0  ±8.4 

27.5  ±5.2 

23.4  ±5.3 

%  Litter  cover 

83.6  ±6.0 

39.0  ±11.6  45.6±  10.0  9.1  ±7.5 

49.3  ±20.3  13.1  ±9.0 

72.6  ±5.9 

19.3  ±10.4 

%  Exposed  rock 

10.8  ±5.7 

26.3  ±6.4 

24.9  ±6.8 

43.4  ±11.1 

14.8±11.1  .32.4±8.1 

14.6  ±9.9 

28.5  ±4.7 

%  Shrubs 

26.8  ±  1.3.0  27.1  ±  17.0  .37.0  ±  16.8  2.3.9  ±  1.3.2 

48.0±21.9  22.9±17.8  27.7±11.9  18.4±  1,3.6 

%  Perennial  forbs 

5.2  ±7.4 

11.4  ±11.3 

18.9±9.7 

.36.5±  12.9  8.0±  15.3    31.2±  14.9 

15.6  ±9.2 

27.4  ±12.8 

%  Perennial  grasses 

11.5  ±3.4 

4.0  ±.3.1 

13.9  ±8.5 

7.1  ±5.9 

16.9  ±1.3.9  10.4  ±9.0 

22.8  ±10.2 

14.0  ±9.3 

%  .\nnuals 

56.6  ±11.9 

57.5  ±  14.5 

.30.2  ±12.5 

.32.4  ±10.1 

27.1  ±2.3.9  .3.5.6  ±2.5.5 

.3.3.9  ±13.8  40.2  ±13.1 

Average  number  of 

species/tree 

10.5 

11.9 

14.2 

14.9 

10.9              14.3 

12.7 

14.0 

Average  number 

of  species/ 

quadrat /tree 

4.6 

6.2 

7.1 

6.9 

5.4                6.3 

7.4 

8.4 

170 


Great  Basin  Naturalist 


Vol.  40,  No.  2 


much  greater  in  the  understory,  while  per- 
cent of  exposed  rock  is  greater  in  the  open- 
area  communities.  Percent  living  cover  is 
higher  in  the  open-area  communities  except 
for  oak. 

Species  preferences  for  understory  and 
open  areas  were  obtained  by  taking  the  dif- 
ference of  total  subquadrat  occiu-rences  for  a 
species  in  canopy-covered  and  open  stands 
and  normalizing  by  dividing  by  the  total  oc- 
currences of  that  species  in  all  stands.  The  re- 
sulting index  nms  from  -1  to  1.  Those  species 
with  the  highest  positive  values  are  found 
most  often  under  the  canopy  while  those  spe- 
cies with   the   greatest   negative   values   are 


found  most  often  in  open  areas.  Table  2 
shows  species  preference  indices  and  total 
subquadrat  occurrences  for  each  species  ob- 
served in  the  study  (species  are  listed  alpha- 
betically). Frequency  is  obtained  by  dividing 
the  number  of  occurrences  by  160,  the  total 
number  of  subquadrats  sampled  for  each  tree 
species  within  each  type  (canopy  or  open 
area). 

The  species  preference  index  was  broken 
down  into  components  relating  to  each  of  the 
four  tree  species  as  shown  in  Table  3.  Each 
component  was  obtained  by  taking  the  differ- 
ence of  the  subquadrat  occurrences  for  a  spe- 
cies in  canopy-covered  and  open-area  stands 


Table  2.  Number  of  ocxurrences  of  each  species  observed  for  each  of  the  four  canopy  species;  160  subquadrats 
sampled  in  each  category.  Life  form  codes  are  f=forb,  g  =  grass,  c  =  cool-season  or  spring  ephemeral  but  perennial 
(so  cf  =  cool-season  forb),  sh  =  shrub,  e  =  evergreen,  a  =  annual,  t  =  tree. 


Life 

Preference 

Understory 
stands 

Open-area 
stands 

Species 

form 

index 

Ceve 

Pipo 

Jusc 

Quga 

Ceve 

Pipo 

Jusc 

Quga 

Achillea  millefolium 
Agoseris  glauca 
Agropyron  spicatum 
Agropyron  suhsecundum 
Agropyron  tnichycaulum 
Allium  acuuiinatum 
Amelanchier  alni folia 

f 

f 

g 

g 

g 

cf 

sh 

0.4.3 

-0.31 

0.00 

0.09 

0.56 

-0.30 

-0.50 

3 
13 
0 
0 
0 
1 
0 

4 
9 
0 
2 
12 
3 
1 

11 

3 

2 
2 
2 
3 
0 

32 

10 

0 

2 

6 
0 

2 
22 
0 
0 
0 
8 
0 

3 
23 
0 
0 
0 
9 
1 

9 
6 
2 
4 
0 
6 
2 

6 
15 
0 
1 
6 
1 
0 

Antennario  luzuloides 

f 

1.00 

0 

0 

0 

1 

0 

0 

0 

0 

Antennaria  rosea 

f 

-1.00 

0 

0 

0 

0 

0 

0 

1 

0 

Apocynum  androsaemifolium 
Arctostapliylos  uva-ursi 

f 

esh 

-0.78 
1.00 

2 
0 

0 
0 

0 
4 

0 
0 

9 
0 

0 

0 
0 

0 
0 

Artemisia  tridentata 

esh 

-1.00 

0 

0 

0 

0 

0 

0 

1 

0 

Aster  chilensis  var.  adscendens 

f 

-0.28 

6 

0 

0 

3 

10 

0 

0 

6 

Bromus  ciliatus 

g 

0.32 

11 

11 

4 

9 

2 

5 

6 

5 

Bromus  tectorum 

g 

-1.00 

0 

0 

0 

0 

0 

0 

0 

1 

Carex  geyeri 
Carex  rossii 
Ceanothus  velutinus 

eg 
eg 

esh 

0.36 
0.31 
0.09 

6 

23 

0 

9 
0 
0 

35 

1 
6 

43 

18 

0 

1 

14 

0 

15 
0 
0 

22 
5 

6 
1 
0 

Chenopodium  amhrosioides 
Chenopodium  fremontii 
Chrysopsis  tillosa 
Cirsium  undulatum 

a 
a 
f 
f 

1.00 
-0.,32 
-0.79 
-0..33 

0 
9 

8 
1 

0 
1 
0 
0 

0 
0 
4 
0 

3 
5 
0 
0 

0 
20 

33 
0 

0 
0 
17 
0 

0 

3 

42 

2 

0 

6 

10 

0 

CoUinsia  pnrviflora 
Collomia  linearis 
Comandra  umbellata 

a 
a 
cf 

-0.13 

-0.19 

0.07 

7 
20 
11 

.33 
35 

2 

39 

36 

4 

19 

28 

5 

IS 
8 

41 

66 

1 

36 
39 

8 

.33 

37 
2 

Cryptantha  circttmscissa 
Eriogonum  heracleoides 

a 

esh 

-0.42 
0.07 

6 

1 

1 
14 

0 

0 
0 

16 

1 

0 
0 

0 
9 

1 
9 

Erysimum  asperum 
Euphorbia  rohusta 
Galium  horeale 

f 
f 
f 

-1.00 

1.00 

-1.00 

0 
1 
0 

0 
0 
0 

0 
0 
0 

0 
0 
0 

1 

0 
0 

0 
0 
0 

0 
0 
0 

0 
0 

1 

Gayophytum  ramosissimum 
Uydrophyllum  capitatum 
Ipompsis  agoregata 
Juniperus  communis 
Lomatium  grayi 
Mahonia  repens 

a 

cf 

f 

esh 

f 

esh 

-0.91 
-0.02 
-1.00 

1.00 
-0.05 

0.08 

1 

4 
0 
0 

7 
70 

2 
1 
0 
0 
9 
40 

1 

5 
0 
4 
2 
60 

1 

10 
0 
0 
0 

48 

11 

2 

0 

0 

15 

47 

5 
5 
1 
0 
2 
43 

30 
4 
0 
0 
3 

44 

59 
10 
0 
0 
0 
50 

June  1980 


EvENSON    ET  AL.:  PlANT  EcOLOGY 


171 


associated  with  one  of  the  four  canopv  spe- 
cies. This  difference  was  then  normahzed  by 
dividing  by  the  total  occurrence  of  that  spe- 
cies in  all  stands.  The  four  components  thus 
sum  to  the  species  preference  index  discussed 
above  (within  round-off  error).  The  com- 
ponents of  the  species  preference  index  clari- 
fy how  a  species  preference  for  understories 
or  open  areas  is  associated  with  a  particular 
canopy  species.  Some  luiderstorv  species  are 
highly  associated  with  a  particular  canopy 
tree  species,  and  others  are  not.  For  example, 
Carex  geycri's  preference  for  the  understorv 
is  strongly  associated  with  Gambel  oak.  Yet, 
Stellaria  jamesiana  prefers  the  understory 
much  more  evenly  for  three  of  the  four  cano- 
py species. 

Correlation  analysis  was  performed   to 


study  the  relationship  of  the  environmental 
and  vegetational  parameters  which  were 
measured.  Table  4  shows  the  significant  posi- 
tive and  negative  correlations. 

Light  intensity  correlates  significantly  with 
all  variables  except  percent  annuals.  The 
negative  correlation  of  light  intensity  with 
pH,  Itter  depth,  soil  depth,  and  percent  litter 
cover  is  to  be  expected  because  of  the  gener- 
ally higher  values  of  these  parameters  under 
the  canopies.  Similarly,  there  is  more  exposed 
rock  and  living  cover  in  the  open  areas.  The 
correlations  of  light  intensity  with  life  forms 
reflect  the  preference  of  shrul>s  and  grasses 
for  understory  areas  and  forbs  and  annuals  for 
open  areas  (Wilcox,  Brotherson,  and  Evenson 
1981). 

The    significant    correlations   of   pH    with 


Table  2  continued. 


Understory 

Open 

area 

Life 

Preference 

stands 

stands 

Species 

form 

index 

Ceve 

Pipo 

Jusc 

Quga 

Ceve 

Pipo 

Jusc 

Quga 

Melica  bulbosa 

g 

1.00 

0 

2 

0 

0 

0 

0 

0 

0 

Merten.mi  hrcvistijle 

f 

-l.(X) 

0 

0 

0 

0 

2 

0 

0 

0 

Osniorhizci  obtttsa 

f 

-l.(K) 

0 

0 

0 

0 

0 

0 

0 

1 

Pdcliystima  nujisiiiites 

esh 

0.19 

3 

2 

10 

4 

2 

3 

6 

2 

Penstcmon  sp. 

f 

-1.00 

0 

0 

0 

0 

0 

0 

2 

0 

Piiuis  coutorta 

at 

l.(X) 

0 

0 

0 

1 

0 

0 

0 

0 

Pintis  pondcnmi 

at 

0.00 

0 

I 

0 

0 

0 

1 

0 

0 

Poa  curt  a 

g 

-0.36 

0 

0 

0 

- 

0 

0 

3 

12 

Poa  fcndlcriana 

g 

-1.00 

0 

0 

0 

0 

3 

0 

0 

0 

Poa  pratensis 

g 

0.11 

0 

5 

0 

0 

0 

0 

4 

0 

Pohjgonuin  cloui:,hi.sii 

a 

-0.51 

65 

17 

29 

28 

129 

92 

87 

119 

Potcntilla  olanduhmi 

f 

-0. 1 1 

3 

0 

0 

1 

0 

0 

1 

4 

Pntniis  viroiniana 

sh 

0.54 

1 

1 

6 

2 

0 

2 

1 

0 

Purshid  tridcntatci 

sh 

l.(K) 

1 

0 

0 

1 

0 

0 

0 

0 

{hicrru.s  ^ambelii 

t 

l.(K) 

0 

0 

1 

0 

0 

0 

0 

0 

Rosa  ivoodsii 

sh 

0.20 

5 

5 

6 

S 

6 

4 

2 

4 

Seduiii  stenopctaluni 

f 

-0.36 

6 

4 

15 

3 

20 

11 

21 

( 

Scnccio  u  in  tah  ensis 

f 

-0.64 

2 

0 

0 

0 

4 

0 

1 

4 

Silcnc  menziesii 

ef 

0.33 

0 

2 

0 

0 

1 

0 

0 

0 

Solidaoo  tnissouricnsis 

f 

-0.69 

5 

2 

1 

0 

25 

9 

9 

0 

Solida^o  multiradiata 

f 

0.05 

9 

17 

4 

2 

4 

17 

6 

2 

SoUdaop  sparsiflora 

f 

-0.53 

0 

0 

0 

11 

0 

13 

0 

23 

Stellaria  janwsiana 

cf 

0.51 

55 

94 

27 

105 

11 

36 

16 

27 

y,tipa  cohiiubiaiia 

g 

-l.(K) 

0 

0 

0 

0 

0 

0 

0 

5 

Stipa  k'ttcrmanii 

g 

0.09 

5 

1 

5 

1 

5 

3 

2 

0 

Sijmphoricarpos  oreophilus 

sh 

0.20 

8 

2 

12 

5 

4 

3 

9 

2 

Taraxacum  officinale 

f 

0,(K) 

4 

6 

2 

2 

2 

4 

3 

5 

Tlialictrum  fendleri 

f 

l.(K) 

0 

0 

6 

0 

0 

0 

0 

0 

Thlaspi  montanutn 

f 

-0.20 

.53 

16 

28 

40 

48 

55 

25 

76 

Tragopooon  dubius 

a 

0.25 

5 

0 

0 

0 

1 

0 

2 

0 

Viguicra  midtijlora 

f 

0.00 

9 

0 

2 

2 

5 

6 

0 

2 

Viola  nuttallii 

cf 

0.(X) 

0 

0 

0 

1 

0 

0 

0 

1 

Viola  purpurea 

f 

0.52 

11 

0 

0 

5 

1 

2 

0 

2 

Total  number  of  species 

39 

34 

36 

37 

38 

32 

41 

.39 

172                                                    Great  Basin  Naturalist                                    Vol.  40,  No.  2 

other  variables  again  reflect  the  tendency  to-  nated  communities  with  relatively  low  total 

ward  higher  pH  under  the  canopies.   Sim-  living  cover  and  significant  amounts  of  litter 

ilarly,  for  litter  depth,  soil  depth,  percent  ex-  cover. 

posed  rock,  percent  litter  cover,  and  percent  By  contrast,  shrubs  and  forbs  are  positively 

living  cover  the  correlations  generally  reflect  correlated    with    percent    living    cover,    but 

the  relationship  of  these  parameters  to  the  negatively  correlated  with  each  other.  Shrubs 

canopy-covered  or  open-area  condition.  provided  a  significant   proportion  of  living 

Correlations  of  these  parameters  with  per-  cover  in  any  quadrat  in  which  they  occur  just 

cent  annuals  are  not  so  easily   interpreted,  because  of  their  size.  This  fact  accounts  for 

however.  Annuals  tend  shghtly  to  prefer  the  their  positive  correlation  with  living  cover, 

open  areas,  hence  the  negative  correlation  However,  shrubs  tend  to  prefer  understory 

with  pH.  Their  positive  correlation  with  per-  habitats  and  forbs  prefer  the  open  areas.  So 

cent  litter  cover,  however,  is  better  under-  forbs   are   positively   correlated  with   living 

stood  by  observing  the  negative  correlation  cover  due  to  the  greater  cover  outside  the 

with  all  other  life  forms  and  percent  living  canopies,  while  being  negatively  correlated 

cover.  Annuals  tend  to  grow  in  annual-domi-  with  shrubs. 


Table  3.  Preference  index  components  by  tree  species.  Life  form  codes  are  as  in  Table  2. 


Life 
form 

Preference      _ 
index 

Preference  index 

components 

Species 

Ceve 

Pipo 

Jusc 

Quga 

Antennaria  luzuloides 

f 

LOO 

0.00 

0.00 

0.00 

1.00 

Arctostaphylos  uva-ursi 

ash 

LOO 

0.00 

0.00 

1.00 

0.00 

Chenopodium  ambrosioides 

a 

LOO 

0.00 

0.00 

0.00 

LOO 

Euphorbia  robusta 

f 

LOO 

1.00 

0.00 

0.00 

0.00 

Jttniperus  conimiinis 

esh 

LOO 

0.00 

0.00 

1.00 

0.00 

Melica  bulbosa 

g 

LOO 

0.00 

1.00 

0.00 

0.00 

Pinus  contorta 

at 

LOO 

0.00 

0.00 

0.00 

1.00 

Purshia  tridentata 

sh 

1.00 

0.50 

0.00 

0.00 

0.50 

Querciis  gambelii 

t 

1.00 

0.00 

0.00 

1.00 

0.00 

Tlialictrwn  fendleri 

f 

1.00 

0.00 

0.00 

1.00 

0.00 

Agropyron  trachycaitlum 

g 

0.56 

0.00 

0.44 

0.07 

0.04 

Pruntis  virginiana 

sh 

0.54 

0.08 

-0.08 

0.38 

0.15 

Viola  purpurea 

f 

0.52 

0.48 

-0.10 

0.00 

0.14 

Stellaria  jamesiana 

cf 

0.51 

0.12 

0.16 

0.03 

0.21 

Adiillea  millefolium 

f 

0.43 

0.01 

0.01 

0.03 

0.37 

Carex  geyeri 

eg 

0.36 

0.04 

-0.04 

0.09 

0.27 

Silene  menziesii 

af 

0.33 

-0.33 

0.67 

0.00 

0.00 

Bromus  ciliatus 

g 

0.32 

0.17 

0.11 

-0.04 

0.08 

Carex  rossii 

eg 

0.31 

0.14 

0.00 

-0.09 

0.27 

Tragopogon  dubius 

a 

0.25 

0.50 

0.00 

-0.25 

0.00 

Rosa  woodsii 

sh 

0.20 

-0.02 

0.02 

0.10 

0.10 

Symphoricarpos  oreophilus 

sh 

0.20 

0.09 

-0.02 

0.07 

0.07 

Pachystima  myrsinites 

esh 

0.19 

0.03 

-0.03 

0.12 

0.06 

Poa  pratensis 

g 

0.11 

0.00 

0.56 

-0.44 

0.00 

Agropyron  subsecundum 

g 

0.09 

0.00 

0.18 

-0.18 

0.09 

Ceanothus  celutinm 

esh 

0.09 

0.00 

0.00 

0.09 

0.00 

Stipa  lettennanii 

g 

0.09 

0.00 

-0.09 

0.14 

0.05 

Mahonia  repens 

ash 

0.08 

0.06 

-0.01 

0.04 

-0.00 

Comandra  umhellata 

af 

0.07 

0.07 

0.02 

-0.10 

0.07 

Eriogonum  heracleoides 

ash 

0.07 

0.00 

0.34 

-0.05 

-0.22 

Solidago  multiradiata 

f 

0.05 

0.08 

0.00 

-0.03 

0.00 

Agropyron  spicatum 

g 

0.00 

0.00 

0.00 

0.00 

0.00 

Pinus  ponderosa 

at 

0.00 

0.00 

0.00 

0.00 

0.00 

Taraxacum  officinale 

f 

0.00 

0.07 

0.07 

-0.04 

-0.11 

Viguiera  multiflora 

f 

0.00 

0.15 

-0.23 

0.08 

0.00 

June  1980 


EVENSON    ET  AL.:  PlANT  EcOLOGY 


173 


Summary  and  Conclusions 

Canopy  tree  species  clearly  influence  both 
vegetation  and  environment  in  their  under- 
stories  (Wilcox,  Brotherson,  and  Evenson 
1981).  The  effects  of  this  influence  on  plant 
distributions  are  shown  clearly  for  individual 
species  in  Tables  2  and  3. 

The  correlations  of  the  environmental  and 
vegetational  parameters  among  themselves 
can  be  imderstood  on  the  basis  of  a  few  im- 
portant concepts. 

(1)  The  environmental  parameters  (light  in- 
tensity, pH,  litter  depth,  soil  depth,  percent- 
age of  exposed  rock,  and  percentage  of  litter 
cover)  are  directly  influenced  by  the  pres- 
ence or  absence  of  canopy  cover.  All  correla- 
tions among  these  parameters  are  as  expected 
on  that  basis. 


(2)  The  vegetational  correlations  follow 
primarily  from  the  facts  that  there  is  more 
living  cover  in  open  areas  than  in  the  under- 
stories;  shnibs  and  grasses  tend  to  prefer  the 
understories;  and  forbs  and  annuals  tend  to 
prefer  open  areas. 

(3)  Shrubs  follow  the  pattern  inferred  from 
their  tendency  to  prefer  understory  areas  ex- 
cept for  their  positive  correlation  with  the 
percentage  of  living  cover.  This  is  because 
shrubs  themselves  provide  a  large  fraction  of 
the  living  cover  that  is  found  in  understory 
quadrats. 

(4)  Annuals  are  different.  They  apparently 
tend  to  grow  in  annual-dominated  patches 
with  low  total  living  cover  and  relatively 
high  litter  cover. 


Table  3  continued. 


Life 
form 

Preference 
index 

Preference  index 

components 

Species 

Ceve 

Pipo 

Jusc 

Quga 

Viola  ntittallii 

cf 

0.00 

0.00 

0.00 

0.00 

0.00 

Hydrophylltiiii  capitatum 

cf 

-0.02 

0.05 

-0.10 

0.02 

0.(X) 

Lomatiwn  gnnji 

f 

-0.05 

-0.21 

0.18 

-0.03 

0.(X) 

Potentilla  glandulosa 

f 

-0.11 

0..33 

0.00 

-0.11 

-0.33 

CoUinsia  paniflora 

a 

-0.13 

-0.05 

-0.04 

0.01 

-0.06 

Colloniki  linearis 

a 

-0.19 

-0.04 

-0.11 

-0.01 

-0.03 

TliUispi  montanwn 

f 

-0.20 

0.01 

-0.11 

0.01 

-0.11 

Aster  chilensis  tar.  adscendens 

f 

-0.28 

-0.16 

0.00 

0.00 

-0.12 

Allium  acuminatutn 

cf 

-0.30 

-0.19 

-0.16 

-0.08 

0.14 

Agoseris  oUnica 

f 

-0.31 

-0.09 

-0.14 

-0.03 

-0.05 

Chenopodiutn  fremontii 

a 

-0.32 

-0.25 

0.02 

-0.07 

-0.02 

CirsitiDi  iindiilatuui 

f 

-0.33 

0..33 

().(X) 

-0.67 

0.00 

Poo  eurta 

g 

-0.36 

0.00 

0.00 

-0.14 

-0.23 

Sedtim  stenopetahnti 

f 

-0.36 

-0.16 

-0.08 

-0.07 

-0.05 

Cnjptantha  circuinscissa 

a 

-0.42 

-0.42 

0.04 

0.00 

-0.04 

Amelanchier  alnifolia 

sh 

-0.50 

0.00 

0.(X) 

-0.50 

0.00 

Polygonum  dotiglasii 

a 

-0.51 

-0.11 

-0.13 

-0.10 

-0.16 

Solidago  sparsi  flora 

f 

-0.53 

0.00 

-0.28 

0.00 

-0.26 

Senecio  uintahensis 

f 

-0.64 

-0.18 

0.00 

-0.09 

-()..36 

Solidago  m  isso  u  riensis 

f 

-0.69 

-0.39 

-0.14 

-0.16 

0.(X) 

Apocynum  androsaemifolium 

f 

-0.78 

-0.39 

-0..39 

0.00 

0.00 

Chrysopsis  villosa 

f 

-0.79 

-0.22 

-0.15 

-0..33 

-0.09 

Cayophytum  ramosissimum 

a 

-0.91 

-0.09 

-0.03 

-0.26 

-0..53 

Antennaria  rosea 

f 

-1.00 

0.00 

().(X) 

-l.(X) 

0.(X) 

Artemisia  tridentiita 

esh 

-l.(X) 

0.00 

0.(X) 

-l.(X) 

O.tX) 

Bromus  tectorum 

g 

-l.(X) 

0.00 

0.(X) 

().(X) 

-1.00 

Erysimum  aspcrum 

f 

-1.00 

-1.00 

0.00 

0.(X) 

0.(X) 

Galium  horeale 

f 

-1.00 

0.(X) 

0.(X) 

0.(X) 

-l.(X) 

Ipompsis  aggregata 

f 

-l.(K) 

0.00 

-1.00 

0.(X) 

().(X) 

Me rten s ia  b re f  istyla 

f 

-l.(K) 

-1.00 

0.(X) 

0.(X) 

0.00 

Osmorhiza  obtusa 

f 

-l.(X) 

().(X) 

().(X) 

0.00 

-l.(X) 

Penstemon  sp. 

f 

-l.(X) 

0.(X) 

().(X) 

-1.00 

0.(X) 

Poa  fendleriana 

g 

-1.00 

-1.00 

0.00 

0.00 

().(X) 

Stipa  Columbiana 

g 

-1.00 

0.00 

0.00 

0.00 

-l.(X) 

174 


Great  Basin  Naturalist 


Vol.  40,  No.  2 


Table  4.     Significance  levels  for  correlations  of  life  form  types,  cover,  and  measured  environmental  parameters 
for  all  stands  studied. 


Variables 


7 


8 


10 


11 


1.  Light  intensity 

2.  pH 

3.  Litter  depth 

4.  Soil  depth 

5.  %  exposed  rock 

6.  %  litter  cover 

7.  %  living  cover 

8.  %  shrubs 

9.  %forbs 

10.  %  grasses 

11.  %  annuals 


.001    -.001  +.001    -.001  +.1        -.05    +.001    -.001  NS 

NS        NS    -.05          NS        NS  +  .05      -.01          NS  -.05 

+  .(K)1    -.001  +.001        NS  +.1        -.001        NS  NS 

-.001  +.fK)l    -.05          NS    -.01     +.1  NS 

-.(K)l  +.1         -.1       +.001    -.01  NS 

-.001        NS    -.001  +.001  +.05 

+  .01     +.1             NS  -.001 

-.001        NS  -.001 

NS  -.05 

NS  -.05 


-.01 


Literature  Cited 

Anderson,  R.  C,  O.  L.  Loucks,  and  A.  M.  Swain.  1969. 
Herbaceous  response  to  canopy  cover,  light  in- 
tensity and  throughfall  precipitation  in  co- 
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AucLAiR,  .-v.  N.,  and  R.  G.  Goff.  1971.  Diversity  rela- 
tions of  upland  forests  in  the  western  Great 
Lakes  area.  American  Naturalist  105:499-528. 

Blackman,  G.  E.  1956.  Influence  of  light  and  temper- 
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thorpe,  ed.  Growth  of  leaves.  Butterworths  Scien- 
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Cline,  M.  G.  1966.  Effect  of  temperature  and  light  in- 
tensity on  Scropltularia  marihindica.  Ecologv 
47:782-795. 

CoTTA.M,  W.  P.  1930.  Some  unusual  floristic  features  of 
the  Uinta  Mountains,  Utah.  Proceedings  of  the 
Utah  Academy  of  Sciences  7:48-49. 

Daubenmire,  R.  1959.  A  canopy-coverage  method  of 
vegetational  analysis.  Northwest  Science 
.33:4.3-66. 

del  Moral,  R.  1972.  Diversity  patterns  in  forest  vegeta- 
tion of  the  VVenatchee  Mountains,  Washington. 
Torrey  Botany  Club  Bulletin  99:57-64. 

Federer,  C.  a.,  and  C.  B.  Tanner.  1966.  Spectral  distri- 
bution of  light  in  the  forest.  Ecologv  47:55.5-.561. 

Gordon,  D.  T.  1962.  Growth  response  of  east  side  pine 
poles  to  removal  of  low  vegetation.  U.S.  Depart- 


ment of  .'\gricvdture.  Forest  Service,  Pacific 
Southwest  Forest  and  Range  Experiment  Station 
Research  Note  209.  3  pp. 

McCoNNELL,  B.  R.,  A.ND  J.  G.  Smith.  1970.  Response  of 
understory  vegetation  to  ponderosa  pine  thinning 
in  eastern  Washington.  Journal  of  Range  Man- 
agement 23:208-212. 

McQueen,  D.  R.  1973.  Changes  in  understory  vegeta- 
tion and  fine  root  quality  following  thinning  of 
30-year-old  Finns  radiata  in  central  North  Island, 
New  Zealand.  Journal  of  .\pplied  Ecology 
10:13-21. 

RusEL,  \.  R.,  AND  H.  J.  Evans.  1966.  The  nitrogen  fixing 
capacity  of  Ceanothus  vehitinns.  Forest  Science 
12:164-169. 

Smith,  B.  E.,  and  C.  Cottam.  1967.  Spatial  relationships 
of  mesic  forest  herbs  in  southern  Wisconsin. 
Ecology  48:.546-548. 

Webster,  S.  C,  C.  T.  Youngberg,  .\nd  A.  G.  Wollum 
II.  1967.  Fixation  of  nitrogen  by  bitterbrush  {Pur- 
sliia  tridentata  (Pursh)  DC).  Nature 
216(51 1.3):392-93. 

Wilcox,  R.  B.,  J.  D.  Brotherson,  a.nd  W.  E.  Eve.nson. 
1981.  Canopy  influence  on  understory  commu- 
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Wollum,  A.  G.  II,  and  C.  T.  Youngberg.  1964.  The  in- 
fluence of  nitrogen  fixation  by  non-leguminous 
woody  plants  on  the  growth  of  pine  seedlings. 
Journal  of  Forestry  62:216-321. 


SEASONAL  ACTIVITY  PATTERN  OF  COLUMBIAN  GROUND  SQUIRRELS 
IN  THE  IDAHO  PRIMITIVE  AREA 


Charles  L.  Elliott'  and  jerran  T.  Flinders' 

.\bstr.\ct.—  Data  were  gathered  concerning  the  seasonal  activity  pattern  of  a  population  of  Columbian  ground 
scjuirrels  (Spermophihis  colttmbianus)  in  the  Idaho  Primitive  Area.  Adult  females  were  significantly  more  active  in 
|une  of  all  vears  than  were  adult  males.  \  relationship  between  ground  squirrel  activity  and  temperature  is  postulat- 
ed in  which  the  squirrels  alter  their  activity  so  as  to  avoid  high  temperatures  and  possible  heat  stress. 


Members  of  the  genus  Spennophilus  are 
characterized  bv  a  seasonally  short  period  of 
aboveground  activity  and  a  prolonged  period 
of  hibernation.  During  this  time  of  surface 
activity  ground  squirrels  must  establish  terri- 
tories, breed,  reproduce,  and  gain  .sufficient 
weight  to  survive  the  inactive  season.  The  an- 
nual cycle  of  activity  for  various  species  of 
ground  squirrels  has  been  reported  (Skryja 
and  Clark  1970,  Michener  1974,  Loehr  and 
Ris.ser  1977),  but  these  data  as  they  apply  to 
populations  of  Columbian  ground  squirrels 
{Spennophilus  cohimbianiis)  are  limited. 

Methods 

The  study  was  conducted  at  Cold  Mead- 
ows, an  87  ha  meadow  (elev.  2010  m)  located 
in  the  northeastern  portion  of  the  Big  Creek 
Ranger  District,  Idaho  Primitive  Area.  A  de- 
scription of  the  Big  Creek  area  has  appeared 
elsewhere  (Wing  1969,  Hornocker  1970,  Sei- 
densticker  et  al.  1973).  Ground  squirrels  were 
trapped  from  12-19  June,  17-24  July,  and 
14-21  August,  1976-1978.  Field  work  prior 
to  12  June  was  impractical  because  of  bad 
weather  and  the  inaccessibility  of  the  study 
area.  A  90  x  90  m  grid  with  36  trapping  sta- 
tions 15  m  apart  was  establi.shed  on  the  cen- 
tral portion  of  the  meadow.  One  live  trap  (15 
X  15  X  48  cm)  was  placed  at  each  trapping 
station.  Traps  were  baited  with  carrot  and 
checked  every  hour.  Captured  squirrels  were 
marked  using  the  toe  clipping  sequence  of 
Melchior  and  Iwen  (1965),  sexed,  measured. 


weighed,  time-of-capture  recorded,  and  re- 
leased back  onto  the  grid.  Vegetation  was 
collected  using  the  procedure  outlined  by 
Tadmor  et  al.  (1975).  All  plant  samples  were 
weighed  to  the  nearest  gram  in  the  field  and 
then  brought  back  to  the  laboratory,  where 
they  were  oven  dried  at  64  C  for  three  days. 
The  dried  specimens  were  then  weighed  to 
the  nearest  gram  and  percent  moisture  con- 
tent calculated.  Daily  ambient  temperatures 
were  obtained  using  a  Taylor  Maximum-Min- 
imum thermometer. 

Ground  squirrel  activity  in  this  study  was 
equated  with  the  animals  presence  in  the 
traps.  Bias  due  to  'trap-shy'  or  'trap-happy' 
squirrels  may  have  occurred,  but  attempts  to 
conduct  hourly  visual  censuses  proved  unre- 
liable during  the  latter  months  due  to  the  in- 
crease in  vegetation  height. 

Results  and  Discussion 

The  number  of  Columbian  ground  squirrels 
captured,  including  recaptures,  is  depicted  in 
Table  1.  Adult  female  squirrels  were  signifi- 
cantly more  active  in  June  of  each  year  than 
were  adult  males  (Kolmogorov-Smirnov  Two 
Sample  Test,  P  <0.05).  Activity  for  July  and 
August  was  not  significantly  different  be- 
tween sexes  of  adult  or  young  squirrels. 

Males  are  territorial  during  the  breeding 
season  (Steiner  1970a),  exliibiting  extreme  ag- 
gression toward  other  males  and  occasionallv 
raiding  the  nesting  ground  and  colonies  of 
adjacent  males  (Steiner  1970a,  1970b,  1971, 


'Department  of  Botany  and  Range  Science,  Brigham  Young  University,  Provo,  Utah  84602. 


175 


176 


Great  Basin  Naturalist 


Vol.  40,  No.  2 


1972).  This  aggressive  behavior  and  Lam- 
beth's (1977)  findings  that  Columbian  ground 
squirrels  utilize  a  core  area  or  arena  of  activi- 
ty between  21  and  40  m  in  size  may  account 
for  the  greater  number  of  females  captured 
in  June.  Dominant  males  would  have  been 
excluding  other  male  squirrels  from  the 
trapping  grid. 

The  hibernation  entry  sequence  described 
by  Manville  (1959)  and  Michener  (1974)  for 
S.  columbianus  was  not  apparent  for  the 
Cold  Meadows  colony.  This  disparity  may  be 
artificial  because  adult  groimd  squirrels  dur- 
ing August  become  very  lethargic  and  in- 
active and  were  extremely  difficult  to  trap. 

Shaw  (1925)  has  suggested  that  early 
spring  activities  of  Columbian  ground  squir- 
rels are  largely  controlled  by  temperature 
alone,  whereas  estivation  is  induced  by  the 
drying  of  the  vegetation.  Howell  (1938)  felt 
that  "the  date  of  beginning  estivation  was  de- 
termined chiefly  by  the  ripening  of  the  vege- 
tation and  consequent  reduction  of  the  mois- 


ture content  in  their  (ground  squirrels)  food, 
and  in  part  also  by  the  accumulation  of  fat  in 
the  body."  Nansel  and  Knoche  (1972)  also  ob- 
served Columbian  ground  squirrels  and  pos- 
tulated that  hibernation  was  a  response  to 
drought  prior  to  a  decrease  in  temperature. 
Cold  Meadows  plant  moisture  content  de- 
clined as  the  season  progressed  (Table  2),  as 
did  the  squirrels'  aboveground  activity  (Table 

Peak  daily  activity  of  ground  squirrels  at 
Cold  Meadows  appears  to  be  determined  by 
temperature.  Table  3  shows  the  monthly  time 
interval  exhibiting  the  greatest  percentage  of 
adult  ground  squirrel  activity.  The  mean 
maximum  temperature  for  June  1976  (Table 
4)  was  significantly  greater  than  June  1977 
and  June  1978.  The  maximum  percent  of  ac- 
tivity for  June  1976  occurs  later  in  the  day 
than  for  June  1977  or  June  1978.  This  same 
type  of  activity  shift  in  relation  to  significant- 
ly greater  maximum  temperatures  is  evident 
for  July;  July   1978   is  significantly   greater 


Table  1.  Live  trapping  results  by  age  and  sex  for  Columbian  ground  squirrels  at  Cold  Meadows,  Idaho  Primitive 
Area,  1976-1978. 


June 

July 

August 

Adult 

Adult 

Juvenile 

Adult 

Juvenile 

Male 

:        Female 

Male 

Femal 

e 

Male 

Female 

Male 

Female         Male 

Female 

1976 

12 

25 

15 

32 

15 

13 

5 

5 

13 

5 

1977 

15 

39 

7 

12 

11 

8 

3 

2 

2 

4 

1978 

24 

32 

18 

13 

6 

7 

2 

6 

/ 

6 

Table  2.  Mean  percent  moisture  content  ( ±  standard  deviation)  of  plant  species  exhibiting  the  highest  frequency 
of  occurrence  at  Cold  Meadows,  Idaho  Primitive  Area,  1976-1978. 


1976 


1977 


1978 


Species' 


July  August  June 


July 


August  June 


July 


August 


Achillea  millefolium  66  ±1  72  ±4  72  ±1  72  ±5  58  ±1  82  ±3  77  ±2  60  ±4 

Carex  aquatilis  65  ±5  63  ±6  61  ±  7  51  ±  4  30  ±  1  67  ±2  58  ±3  45  ±5 

Fragaria  virginiana  67  ±  8  66 ±  2  65  ±  8  65  ±  7  52  ±  1  76  ±  1  67  ±  2  55  ±  1 

Penstemon  procerus  68  ±6  65  ±2  72  ±3  66  ±5  47  ±1  73  ±  1  69  ±2  63  ±2 

Phleum  alpimim  63  ±5  57  ±7  57  ±1  49  ±8  27  ±1  72  ±2  59  ±4  30  ±1 


l:n  =  10  samples/species/month. 


Table  3.  Peak  activity  time  intervals  for  adult  Columbian  ground  squirrels  at  Cold  Meadows,  Idaho  Primitive 
Area,  1976-1978. 


1976 


1977 


1978 


June  12-19 
July  17-24 
August  14-21 


1700-1800  hours 

13(K)-14(X) 

1600-17(X) 


1400-1500 
1200-1300 
1200-1300 


1400-1500 
1000-1100 
13(K)-1400 


June  1980 


Elliott,  Flinders:  Columbian  Ground  Squirrels 


177 


Table  4.  Monthly  mean  maximum  temperatures 
(±  standard  deviation)  at  Cold  Meadows,  Idaho  Primi- 
tive Area,  1976-1978. 


1976 


1977 


1978 


June  12-19      22±2C'  20±2l  20*2^ 

July  17-24       22±l2  22±li-2  24±l''2 

August  14-21  17  ±2''^  25±1^  17±2'-2 

1;  Means  tested  for  significance  using  unpaired  t-test  at  the  0.05  level. 
2:  Means  tested  for  significance  using  Cochran  and  Cox  test  at  the  0.05 
level  (Ferguson  1959:143). 


than  July  1976  and  July  1977.  August  1977 
mean  daily  maximum  temperature  is  signifi- 
cantly greater  than  August  1976  and  August 
1978;  yet  the  expected  shift  in  peak  activity 
does  not  occur.  This  may  be  due  to  the  small 
activity  sample  size  for  August  1977. 

The  relationship  of  temperature  to  activity 
indicates  that  the  higher  the  average  daily 
maximum  temperature,  the  earlier  or  later  in 
the  day  peak  activity  will  occur.  If  this  hy- 
pothesis is  valid,  then,  for  those  sampling  pe- 
riods where  the  average  maximum  temper- 
atures were  not  significantly  different,  the 
time  interval  of  peak  activity  should  be  sim- 
ilar. This  relationship  is  observed  for  June 
1977  and  June  1978,  and  July  1976  and  July 
1977  (Table  3). 

Betts  (1976)  observed  a  lower  elevation 
(1360  m)  colony  of  S.  columhianus  in  western 
Montana  and  reported  that,  with  the  increase 
in  temperatures  during  lactation  and  post- 
lactation  periods,  there  was  an  increase  in 
morning  and  late  afternoon  activity  and  a  de- 
crease in  midday  activity.  Betts  postulated 
that  temperature  or  solar  radiation  may  limit 
the  amount  of  consecutive  time  Columbian 
ground  squirrels  can  spent  aboveground. 

The  observations  of  Betts  (1976)  and  data 
reported  here  indicate  that  the  scheduling  of 
surface  activity  for  these  squirrels  is  an  ap- 
parent behavioral  response  designed  to  es- 
cape heat  stress. 

We  thank  the  University  of  Idaho  for  per- 
mission to  use  the  facilities  at  the  Taylor 
Ranch  Field  Station,  Idaho  Primitive  Area. 

Literature  Cited 

Betts,  B.  J.  1976.  Behaviour  in  a  population  of  Colum- 
bian ground  scjuirrels,  Spernu)philus  coliuiibianus 
columhianus.  .^nini.  Behav.  24:652-680. 

Ferguson,  C.  \.  1959.  Statistical  analysis  in  psychology 
and  education.  McGraw-Hill  Book  Co.,  Inc.,  New 
York.  .347  pp. 


HoRNocKER,  M.  G.  1970.  An  analysis  of  mountain  lion 
predation  upon  mule  deer  and  elk  in  the  Idaho 
Primitive  Area.  Wildlife  Monogr.  21:l-.39. 

Howell,  \.  H.  1938.  Revision  of  the  North  American 
ground  squirrels.  North  American  Fauna 
56:85-90. 

Lambeth,  R.  E.  1977.  The  Coliunhian  groimd  squirrel  in 
subalpine  forest  openings  in  the  Idaho  Batholith. 
Unpublished  thesis,  Univ.  Idaho,  Moscow.  1 1.3 
pp. 

Loehr,  K.  a.,  and  a.  C.  Risser,  Jr.  1977.  Daily  and  sea- 
sonal activity  patterns  of  the  Belding  ground 
squirrel  in  the  Sierra  Nevada.  J.  Mammal. 
58:445-448. 

Manville,  R.  H.  1959.  The  Columbian  ground  squirrel 
in  northwestern  Montana.  J.  Mammal.  40:26-45. 

Melchior,  H.  R.,  and  F.  A.  Iwen.  1965.  Trapping,  re- 
straining and  marking  .\rctic  ground  squirrels  for 
behavioral  observations.  J.  Wildlife  Manage. 
29:671-679. 

Michener,  D.  R.  1974.  .\nnual  cycle  of  activity  and 
weight  changes  in  Richardson's  ground  squirrel, 
Spermopliilus  richardsonii.  Can.  Field-Nat. 
88:409-413. 

Nansel,  D.,  and  L.  Knoche.  1972.  Blood  changes  in  tor- 
pid and  non-torpid  Columbian  ground  squirrels, 
Spermophilus  cohimbianus.  Comp.  Biochem. 
Physiol.  41A:  175-179. 

Seidensticker,  J.  C  IV,  M.  G.  Hor.nocker,  W.  V. 
Wiles,  a.nd  J.  P.  Messick.  1973.  .Moimtain  lion 
.social  organization  in  the  Idaho  Primitive  Area. 
Wildlife  Monogr.  .35:1-60. 

Shaw,  W.  T.  1925.  Duration  of  the  aestivation  and  hi- 
bernation of  the  Columbian  ground  squirrel. 
Ecology  6:75-81. 

Skryja,  D.  D.,  and  T.  W.  Clark.  1970.  Reproduction, 
seasonal  changes  in  body  weight,  fat  deposition, 
spleen  and  adrenal  gland  weight  of  the  golden- 
mantled  ground  squirrel  Spennophihis  lateralis 
lateralis,  (Sciuridae)  in  the  Laramie  Mountains, 
Wyoming.  Southwestern  Nat.  15:201-208. 

Stelner,  a.  L.  1970a.  Etude  descriptive  de  quelques  ac- 
tivities et  comportements  de  base  de  Spermo- 
philus columhianus  columhianus  (Ord).  I.  Loco- 
motion, soins  du  corps  curiosite  et  alarme, 
reproduction.  Rev.  Comp.  Animal  4:3-21. 

1970b.  Etude  descriptive  de  quelques  activities  et 

comportements  de  base  de  Spennophilus  colum- 
hianus coltimhianus  (Ord).  II.  Vie  de  groups.  Rev. 
Comp.  .Animal  4:23-42. 

1971.  Plav  activitv  of  Columbian  ground  squir- 
rels. Z.  Tierpsychoi.  28:247-261. 

1972.  Mortality  resulting  from  intraspecific  fight- 
ing in  some  ground  squirrel  populations.  J.  .Mam- 
mal. 53:601-603. 

Tadmor,  N.  H.,  a.  Briegher,  I.  Noy-Meir,  R.  W. 
Benjamin,  and  E.  Eyal.  1975.  .^n  evaluation  of 
the  calibrated  weight-estimate  method  for  mea- 
suring production  in  annual  vegetation.  J.  Range 
Manage.  28:65-69. 

Wlng,  L.  D.  1969.  Ecology  and  herbivore  use  of  five 
mountain  meadows  in  the  Idaho  Primitive  ,\rea. 
Unpublished  thesis,  Univ.  Idaho,  Moscow.  215 
pp. 


HABITAT  AND  PLANT  DISTRIBUTIONS  IN  HANGING  GARDENS 
OF  THE  NARROWS,  ZION  NATIONAL  PARK,  UTAH 

George  P.  Malanson' 

,\bstract.—  Hanging  gardens  are  insular  plant  comnuinities  of  the  Colorado  Plateau.  This  study  examines  hanging 
gardens  in  the  Narrows,  Zion  National  Park,  Utah.  The  floristic  similarity  of  gardens  and  the  presence  of  species  in 
classes  of  variables  which  characterize  the  habitat  are  disclosed.  Although  the  gardens  are  isolated  and  dissimilar,  the 
individual  species  are  not  restricted  in  the  range  of  habitat  found.  Floristic  dissimilarity  cannot  be  attributed  to 
differences  in  habitat.  These  results  are  compared  to  studies  of  hanging  gardens  in  eastern  Utah. 


Hanging  gardens  are  plant  communities 
growing  at  seeps  on  the  canyon  walls  of  the 
Colorado  Plateau.  The  hanging  garden  envi- 
ronment is  characterized  by  shallow  soils  at  a 
seep  from  bedrock.  Seeps  occur  where  water 
has  percolated  through  a  porous  formation 
until  meeting  a  less  permeable  layer  of  rock. 
Then  the  water  flows  laterally  until  a  canyon 
intersects  this  plane.  The  narrow  canyons  of- 
ten shade  the  hanging  gardens.  Compared  to 
other  environments  of  the  Colorado  Plateau, 
the  hanging  gardens  are  cool  and  moist. 

The  Narrows  of  the  North  Fork  of  the  Vir- 
gin River  in  Zion  National  Park  is  an  arch- 
etypal hanging  garden  locale.  In  an  8  km  sec- 
tion there  are  about  60  gardens,  varying  in 
size  from  a  few  square  centimeters  to  over 
100  m-.  Most  of  these  are  at  permanent  seeps 
with  small  discharges  of  water.  The  hanging 
gardens  of  the  Narrows  assume  a  variety  of 
shapes,  but  in  general  they  occupy  a  place 
where  erosion  has  modified  the  steepness  of 
the  canyon  wall.  Often  these  places  are  hori- 
zontal bands.  Other  gardens  occur  where  ver- 
tical jointing  has  concentrated  the  seepage. 
Some  gardens  occupy  remnants  of  potholes, 
and  others  are  on  bulges  of  travertine.  A  few 
are  in  alcoves.  Most  hanging  gardens  in  the 
Narrows  are  close  to  the  level  of  the  river, 
where  they  may  be  vulnerable  to  flash  floods. 

This  study  examines  the  relationship  be- 
tween the  plant  species  and  the  habitat  in  the 
hanging  gardens  of  the  Narrows.  The  con- 


cepts of  Ramensky  (1924)  and  Gleason  (1926) 
are  the  basis  for  hypothesizing  that  the  pres- 
ence and  importance  of  species  at  sites  are 
determined  by  their  individual  tolerances  and 
requirements  in  relation  to  the  habitat.  This 
idea  can  be  evaluated  by  examining  the  flo- 
ristic similarity  between  sites  and  the  in- 
cidence of  species  across  a  range  of  variables 
that  characterize  the  biotopes.  The  hypoth- 
esis leads  one  to  expect  a  positive  relation- 
ship between  floristic  similarity  and  sim- 
ilarity of  biotope  (i.e.,  if  sites  represent  a 
single  habitat). 

A  few  authors  have  investigated  the  vege- 
tation of  hanging  gardens.  In  a  general  eco- 
logical stvidy  of  Zion  National  Park,  Wood- 
bury (1933)  outlined  the  stages  of  primary 
succession  that  occur  at  seeps.  Welsh  and 
Toft  (1976)  described  a  variety  of  garden 
types  in  Glen  Canyon,  Utah,  based  on  the 
form  created  by  the  erosion  of  the  rock,  and 
traced  the  geographical  affinities  of  the  spe- 
cies they  found.  They  called  hanging  gardens 
"relictual  refugia"  because  the  gardens  pro- 
vided sites  for  species  from  other  southwest 
locations,  boreal  forests,  and  earlier  epochs. 
Welsh  and  Wood  (1976)  concluded  that 
hanging  gardens  have  a  stable  structure,  attri- 
buting change  in  species  importance  mea- 
-sured  over  a  one-year  interval  to  measure- 
ment error.  Wood  and  Welsh  (1976)  found 
productivity   to   be   relatively   high   for   this 


'Department  of  Geography,  University  of  Utah,  Salt  Lake  City.  Utah  84112.  Pre.scnt  address:  Department  of  Geography,  University  of  California,  Los 
Angeles,  CaWfomia  90024. 


178 


June  1980 


Malanson:  Hanging  Gardens 


179 


type  of  vegetation,  and  presumed  it  to  be 
steady. 

Nebeker  et  al.  (1977)  believed  that  the  flo- 
ristic  dissimihiritv  found  in  hanging  gardens 
of  eastern  Utah  indicated  ''random  assort- 
ments of  individuals  capable  of  exploiting  the 
environments  of  individual  sites."  They  con- 
clude that  over  25  percent  of  the  species 
were  ecological  specialists  of  hanging  garden 
habitats,  within  the  Colorado  Plateau  region. 
A  few  studies  briefly  mention  hanging  gar- 
dens. Clover  and  Jotter  (1944),  Flowers 
(1959),  and  Loope  (1976)  describe  hanging 
gardens  in  general  ecological  studies  of  the 
Colorado  River  area.  Raines  (1976)  noted 
that  hanging  gardens  can  be  critical  habitats 
for  small  mammals  in  drought  years.  All 
these  studies  emphasize  the  importance  of 
habitat  and  support  the  hypothesis  of  this 
study. 

Methods 

I  sampled  29  hanging  gardens  in  an  8  km 
section  of  the  Narrows  and  in  0.75  km  of  a 
tributary  (Orderville  Canyon)  between  June 
and  September  1977.  Sites  were  chosen  infor- 
mally on  the  basis  of  access,  but  a  representa- 
tive range  of  garden  sizes  was  sought.  To  es- 
timate percent  foliar  cover  of  species,  I 
recorded  the  number  of  decimeters  inter- 
cepted by  a  species  along  line  transects 
spaced  at  2  m  intervals  and  perpendicular  to 
the  long  axis  of  each  garden.  In  gardens  nimi- 
bered  1,  2,  and  3,  I  placed  a  2500  cm-  quad- 
rat every  2  m  on  transects  spaced  at  2  m  in- 
tervals and  recorded  the  number  of  100  cm- 
grids  occupied  bv  each  species.  This  method 
proved  to  be  impractical,  although  in  com- 
parison tests  the  results  were  not  significantly 
different. 

From  the  values  of  relative  cover  for  each 
garden,  I  calculated  the  similarity  of  each 
pair  of  hanging  gardens  according  to  Ru- 
zicka's  (1958)  index: 

SI  =  (2A/2B)(100) 

where  A  is  the  smaller  and  B  the  larger  value 
of  a  species  in  two  sites,  considering  all  spe- 
cies present  in  either  site. 

I  measured  11  variables  to  characterize  the 
biotope  of  each  hanging  garden.  Soil  and  wa- 
ter pH  at  each  site  was  measured  color- 


imetrically  (Microessential  Laboratory  0.2 
unit  paper).  I  gathered  grab  samples  of  soil 
from  the  surface  where  there  was  any  depth, 
not  taking  any  litter.  I  took  125  ml  samples 
of  water  from  the  seeps.  In  a  few  cases  sam- 
ples could  not  be  taken  because  of  a  paucity 
of  soil  or  water.  The  USDA  Soils  Testing 
Laboratory  at  Logan,  Utah,  measured  soil 
and  water  conductivity  as  a  surrogate  of  sali- 
nity, and  the  total  phosphorous  content  of 
the  soil.  I  measured  the  slope  of  each  garden 
by  taking  the  height:depth  ratio  and  calcu- 
lating a  percent  slope.  I  sampled  soil  depth 
every  4  m  across  the  center  of  each  garden 
with  a  wire  rod  to  obtain  a  single  average  soil 
depth  measure;  in  smaller  gardens,  one  or 
two  evenly  spaced  measures  were  taken. 

Direct  solar  radiation  was  derived  by  a 
computer  program  of  Williams  et  al.  (1972), 
which  accounts  for  slope,  azimuth,  and  lati- 
tude of  the  site,  and  intervening  topography 
that  shades  the  site.  The  program  computes 
the  calories  per  square  centimeter  for  any 
one  day,  and  I  summed  the  120th,  180th,  and 
240th  days  of  the  year  to  bracket  the  grow- 
ing season.  The  scale  of  resolution  does  not 
account  for  all  possible  variations  in  micro- 
relief.  In  locations  with  particular  sunblock- 
ing  features,  such  as  alcoves,  I  reduced  the 
computed  value  10  percent.  At  one  garden 
where  the  three-day  sum  was  zero,  although 
I  observed  direct  radiation  at  the  site,  I  arbi- 
trarily assigned  a  value  of  50  cal/cm-. 

I  measured  three  spatial  variables:  area, 
isolation,  and  distance  to  the  Gateway  to  the 
Narrows  Trail.  I  derived  the  area  of  each  gar- 
den from  the  grid  formed  by  the  species  in- 
tercept transects.  Isolation  is  defined  here  as 
the  sum  of  the  distances  from  each  garden  to 
its  nearest  three  neighbors.  These  distances 
were  measured  on  a  topographic  map 
(ZNHA  1977).  The  Gateway  to  the  Narrows 
Trail  is  the  scene  of  much  pedestrian  tourist 
traffic,  but  above  the  terminus  human  use  de- 
clines rapidly.  The  distance  to  the  trail,  also 
measured  on  the  topographic  map  (ZNHA 
1977),  may  affect  invasion  by  nonnative  spe- 
cies. 

I  analyzed  the  relationships  between  spe- 
cies presence  and  absence  and  the  11  envi- 
ronmental variables.  These  variables  were 
considered  separately  because  there  was  low 
correlation  between  any  two.  I  followed 


180 


Great  Basin  Naturalist 


Vol.  40,  No.  2 


Strahler's  (1978)  use  of  the  Gh  statistic  to  dis- 
close significant  differences  between  species 
presence  and  absence  in  ordinal  categories  of 
the  environmental  variables.  Although  the 
data  are  ratio  scaled  and  the  categories  are 
ordinal,  the  presence  of  many  zeros  in  the 
species  matrix  and  the  number  of  tie  scores 
among  the  environmental  variables  make 
more  powerful  tests  less  dependable.  Only 
the  13  most  common  species  occur  frequent- 
ly enough  to  be  tested.  I  divided  each  of  the 
11  environmental  variables  into  four  or  five 
ordinal  categories  (Table  1).  The  number  of 
categories  and  the  ranges  are  arbitrary,  ex- 
cept that  each  category  represents  at  least 
one  garden.  I  attempted  to  categorize  the 
variables  so  that  the  results  of  the  Gh  tests 
would  represent  the  relationships  between 
species  presence  and  a  continuous  change  in 
the  variable. 

I  also  calculated  a  species-area  curve  of 
the  gardens  by  a  regression  of  log  number  of 
species  on  log  area.  Following  the  Whitehead 
and  Jones  (1969)  treatment  of  island  floras,  I 
deleted  the  smallest  garden  on  successive  cal- 
culations to  find  the  point  below  which  the 
area  detracted  from  the  fit  of  the  curve. 

Results 

Forty-eight  species  were  coimted  in  the  29 
hanging  gardens.  Nine  taxa  were  identified 
only  to  genus  or  family,  and  four  rare  plants 
could  not  be  identified  (Table  2).  Richness 
ranged  from  2  to  20.  The  frequency  of  occur- 


rence ranged  from  1  to  17.  Although  no  spe- 
cies occurred  in  all  gardens,  the  13  species 
occupying  five  or  more  gardens  are  used  here 
as  diagnostic  species,  because  species  of  mod- 
erate constancy  are  good  indicators  of  varia- 
tion in  the  environment  (Mueller-Dombois 
and  Ellenberg  1974).  The  many  rare  species 
(35  occupying  less  than  five  gardens)  are  sim- 
ply not  common  enough  to  give  useable  cor- 
relations with  garden  environments.  For 
complete  tables  of  data  on  species  cover,  flo- 
ristic  similarity,  and  the  environmental  varia- 
bles, see  Malanson  (1978). 

The  floristic  similarity  of  the  hanging  gar- 
dens is  low,  ranging  from  0  to  77,  and  aver- 
aging 10.23  for  all  gardens.  This  value  of  sim- 
ilarity is  close  to  the  results  of  Nebeker  et  al. 
(1977). 

All  hanging  gardens  provide  a  mesic  to 
hygric  biotope.  For  water  and  soil,  values  of 
pH  ranged  from  5.9  to  7.2  and  6.3  to  7.4,  and 
values  of  salinity  from  0.3  to  2.5  jumhos/cm 
and  230  to  1420  mhos/cm,  respectively.  In 
this  study,  average  soil  phosphorous  content 
ranges  from  2.1  to  84  ppm.  Direct  solar  radi- 
ation varies  from  0  to  683  cal/cm*  for  the 
three-day  sum.  Average  soil  depth  ranges 
from  0.1  to  48.4  cm,  but  exceeds  10  cm  in 
only  three  gardens.  Slopes  average  between 
40  percent  and  vertical.  Area  ranges  from  2 
to  100  m-  among  samples.  Most  values  of  iso- 
lation are  low,  24  gardens  are  less  than  300  m 
from  the  nearest  three  neighbors.  All  but  four 
distances  from  the  Gateway  to  the  Narrows 


Table  1.  Classes  of  the  environmental  variables. 


Classes 

Variables 

1 

2 

3 

4 

5 

Area  (m-) 

0-10 

10-25 

25-50 

50 

Distance  from 

trail  (m) 

50O-ia30 

1631-2760 

2761-3890 

3891-5020 

5021-6150 

Isolation  (m) 

0-80 

81-160 

161-240 

241-320 

,320 

Phosphorous 

(ppm) 

0-5.0 

5.1-15 

16-36 

37-87 

Slope  (percent) 

1-50 

51-125 

126-275 

276-525 

526 

Soil  depth  (cm) 

0-1 

1-2 

2-4 

4-8 

8 

Soil  pH 

6.3-6.4 

6..5-6.6 

6.7-6.8 

6.9-7.4 

Soil  salinity 

(mhos/cm) 

0.1-0.4 

0.5-0.8 

0.9-1.2 

1.3-1.8 

1.9-2.5 

Solar  radiation 

(cal/cm-) 

0-150 

151-300 

.301-4,50 

451-600 

601-750 

Water  pH 

5.9-6.2 

6.3-6.5 

6.6-6.8 

6.9-7.2 

Water  salinity 

(mhos/cm) 

0-300 

301-600 

601-900 

900 

June  1980 


Malanson:  Hanging  Gardens 


181 


Trail  are  clustered  bimodally  between  500 
and  2000  m  and  between  3000  and  5000  ni. 

The  results  of  the  species  presence  tests  in- 
dicate that  there  are  few  relationships  be- 
tween gradients  of  the  environmental  varia- 
bles and  the  presence  and  absence  of 
important  species  in  the  sampled  gardens 
(Table  3).  Of  the  143  tests,  only  15  show  a 
significant    difference    at    p  =  0.05,    and   one 

Table  2.  Hanging  garden  plant  species. 


Species 


Frequency 


Abies  concolor 
Acer  negiindo 
A  did  utti  in  cap  i7/«.s-  veneris 
Adiantuni  pcdatum 
Amaranth tts  c^raecizans 
Amiphalis  margaritaceae 
Apocyniim  cannahinum 
Aquiligia  spp. 
Aralia  racemosa 
A rteniisia  ludovicia na 
Aster  eatonii 
Berheris  rcpens 
Brickelia  grandiflora 
Bromus  ciliattts 
Calaniagrostis  scoptilorum 
Cirsiinn  arizonicuin 
Cystoptcris  fragiUs 
Dodecatheon  pulchellum 
Dryopteris  filix-nias 
Eleocharis  sp. 
Epipactis  gigantea 
Equisetttm  hyemale 
Fraxintis  vehitina 
Galium  aparine 
Hepaticae 
Heuchera  versicolor 
Junctts  sp. 
Lobelia  cardinalis 
Mimiilus  cardinalis 
Mimtdtis  guttatus 
Muhlenbergia  andina 
Middenbergia  mexicana 
Nasturtium  officinale 
Pea  nevadensis 
Rhus  radicans 
Rubus  leucodennis 
Riimex  sp. 
Salix  sp. 

Smilacina  stellata 
Sphagnum  sp. 
Sphagnaceae 
Taraxacum  officinalis 
Thalictrum  fendleri 
Viola  spp. 
unidentified  #  1 
unidentified  #2 
unidentified  #3 
unidentified  #4 


1 
3 

15 
6 
1 
4 
1 

13 

12 
1 
8 
3 
1 
3 
5 
1 

17 
9 
2 
3 
3 
1 
3 
4 

10 
2 
2 
3 

13 
1 
1 
2 
2 
2 
2 
3 
1 
1 
7 

14 
9 
4 
2 
3 
1 
1 
2 
1 


would  expect  7  of  the  tests  to  prove  signifi- 
cant by  chance  alone.  Only  solar  radiation 
consistently  returns  significant  results. 

The  hanging  gardens  are  smaller  isolates 
than  those  described  by  Whitehead  and  Jones 
(1969)  (1.25  to  8.65  ha);  yet  the  changing  spe- 
cies-area curves  are  similar.  The  exclusion  of 
the  four  smallest  gardens  (6  m-)  improves  the 
regression  coefficient  from  0.55  to  0.67,  and 
changes  the  slope  from  0.28  to  0.48. 

Discussion 

The  biotopes  of  the  hanging  gardens  were 
delimited  in  this  study  by  the  environmental 
variables  of  soil  and  water  pH  and  salinity, 
soil  phosphorous,  soil  depth,  slope,  direct  so- 
lar radiation,  area,  isolation,  and  distance  to 
the  Gateway  to  the  Narrows  Trail.  In  gener- 
al, the  biotopes  are  within  the  habitat  of  the 
species,  and  a  particular  species  composition 
is  not  maintained  by  the  environment.  The 
distribution  of  species  among  the  hanging 
gardens  in  the  Narrows  is  not  strongly  affect- 
ed by  their  tolerances  and  requirements. 
High  values  of  solar  radiation  seem  to  mete 
against  the  mosses  and  ferns,  but  these  values 
occur  in  only  13  percent  of  the  gardens  sam- 
pled. The  great  dissimilarity  between  hang- 
ing gardens  cannot  be  attributed  to  dissimilar 
habitats,  although  they  are  insular  commu- 
nities. 

In  this  regard,  the  hanging  gardens  of  the 
Narrows  may  be  very  different  from  those  of 
the  Arches  and  Canyonlands  area.  There  the 
habitat  differences  found  over  a  wide  geo- 
graphic area  are  more  likely  to  be  significant 
in  affecting  species  presence  and  garden  sim- 
ilarity. In  the  Narrows,  the  proximity  of 
many  gardens  and  their  probable  suscepti- 
bility to  flash  floods  prevents  a  strict  com- 
parison with  the  research  in  eastern  Utah. 
The  Narrows  presents  a  case  in  which  we 
must  look  beyond  the  structure  of  the  habitat 
to  find  an  explanation  of  plant  distributions. 
Malanson  and  Kay  (in  preparation)  consider 
disturbance  frequencies  a  likely  alternative. 

Acknowledgments 

I  am  thankful  for  the  encouragement  and 
advice  of  Jeanne  Kay,  Kimball  T.  Harper, 
and  Paul  A.  Kay  on  all  aspects  of  this  re- 


182 


Great  Basin  Naturalist 


Vol.  40.  No.  2 


search.  Kezia  M.  Nielsen,  Garry  F.  Rogers, 
Lois  A.  Amow,  and  Anne  M.  Travis  helped  in 
the  gathering  and  preparation  of  the  data. 
This  research  was  funded  in  part  by  grants 
from  the  University  of  Utah  Student  Re- 
search Grants  in  Geography,  the  Zion  Natu- 
ral Historv  Association,  and  Sigma  Xi,  the 
Scientific  Research  Society,  and  an  equip- 
ment loan  bv  Robert  W.  Austin. 


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McELLER-DoMBOis.  D..  AND  H.  Ellenberg.  1974.  Aims 
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Nebeker,  G.  T.,  K.  T.  1L\rper.  J.  D.  Brotherson,  .\nd  S. 
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Provo. 


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Pages  37-73  excerpted  in  E.  J.  Kormondv,  ed. 
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RuzicKA,  M.  1958.  Anwendung  mathematisch— statistis- 
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Welsh,  S.  L.,  and  C.  .\.  Toft.  1976.  Biotic  conununities 
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Murdock,^S^  L.  Welsh,  and  B.  W.  Wood,  eds. 
Navaho-Kaiparowits  environmental  baseline 
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ronmental Studies.  Brigham  Young  Universitv. 
Provo. 

Welsh,  S.  L.,  .a.nd  B.  W.  Wood.  1976.  Structure  of  a  se- 
lected hanging  garden  in  Glen  Canvon  of  the 
Colorado  River  drainage  in  J.  R.  Murdock.  S.  L. 
Welsh,  and  B.  W.  Wood.  eds.  Navaho-Kaiparo- 
wits environmental  baseline  studies  1971-1974. 
Center  for  Health  and  Environmental  Studies. 
Brigham  Young  University,  Provo. 

\\  hitehead.  D.  R..  and  C.  E.  Jones.  1969.  Small  islands 
and  the  equilibrium  theory  of  insular  biogeo- 
graphy.  Evolution  23:171-179. 

Williams,  L.  D.,  R.  G.  B.arry,  and  J.  T.  .\ndrews.  1972. 
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Health  and  Environmental  Studies,  Brigham 
Young  Universitv.  Provo. 

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Springdale,  Utah. 


Table  3.  Class  of  environmental  variable  in  which  a  species  was  most  significanth  limited  (classes  from  Table  1). 

Variables 


Species 


Soil  Soil  Solar  Water 

Distance      Isolation         Slope  Depth         Salinitv      Radiation        Salinity 


Adiantum  capillus-veneris 
Adiantum  pedatum 
Aster  eatonii 

CaUimagrostis  seoptilorum 
Cijstupteris  fragilis 
Dodeea th eon  p tt  Ich ellu m 
Mimtdus  cardinalis 
Smilacina  stellata 
Sphagnimi  sp. 
Sphagnaceae  sp. 


4-5 


1-2 


I 
4-5 


3-5 


3-5 


5 

4-5 

5 

3 
1&5 


3-4 

1-2 


SHORT-TERM  EFFECTS  OF  LOGGING  ON  RED-BACKED  VOLES  AND  DEER  MICE 

Thomas  M.  Cainpbcll  HI  and  Tim  VV.  Clark- 

Abstract.—  Clearcutting  and  selective  logging  effects  on  red-backed  voles  (Clethrionomtjs  gapperi)  and  deer  mice 
(Peromysciis  maniculatus)  were  studied  (September-November,  1975;  June-October,  1976)  in  Bridger-Teton  Nation- 
al Forest,  Wvoming.  Five  selective  cuts  (total  137  ha)  removed  57  percent  (range  34-74  percent)  of  the  trees.  One 
clearcut  (9.6  ha)  eliminate  84  percent  of  the  trees.  Soils  remained  mesic  in  selective  cuts,  but  became  xeric  in  the 
clearcut.  Snap-trapping  indicated  that  voles  were  most  abundant  on  the  imlogged  and  selectively  cut  mesic  sites  (76 
percent  of  408  captures),  whereas  deer  mice  were  more  common  on  the  xeric  clearcut  (80  percent  of  60  captures). 
Species  composition  remained  unchanged  on  selective  cuts  following  logging  (77  percent  voles  of  256  captures),  but 
changed  from  predominantlv  voles  to  mostly  deer  mice  (80  percent  of  60  captures)  in  the  clearcut.  Intraspecific  age 
and  sex  ratios,  litter  sizes,  and  morphological  measurements  were  compared  between  logged  and  unlogged  areas. 


The  short-term  logging  effects  on  the 
structure  and  dynamics  (i.e.,  habitat,  num- 
bers, and  morphological  and  reproductive 
characteristics)  of  red-backed  voles  {Cleth- 
rionomys  gapperi)  and  deer  mice  {Peromysciis 
maniculatus)  were  examined  on  a  646  ha 
study  area  in  the  Bridger-Teton  National 
Forest,  about  40  km  north  and  8  km  east  of 
Jackson,  Wyoming.  It  is  on  the  backslope  of 
an  escarpment  that  nms  west  from  Mt.  Leidy 
to  a  point  southwest  of  Toppings  Lakes.  This 
backslope  is  a  series  of  benches  with  L5  to  40 
percent  slopes  (Brady  1974)  at  elevations  of 
2300  to  2700  m.  Soils  include  loams  at  the 
surface,  with  siltv  clay  to  clav  loam  subsoils 
(Knight  1973). 

Climate  is  characterized  by  long,  cold  win- 
ters with  deep  snow,  a  short  growing  season 
(average  60  days),  and  a  low  mean  annual 
temperature  of  1  C.  Snowfall  averages  345 
cm  annually  and  can  occur  any  month.  Mean 
annual  precipitation  is  69  cm,  predominantly 
snow  (Department  of  Commerce  1975). 

Jackson  Hole  vegetation  has  been  de- 
scribed by  Read  (1952),  and  Beetle  (1961).  A 
climax  spruce-fir  {Picea  engelmannii-Abies 
lasiocarpa)  forest  covers  the  study  area  ex- 
cept on  recent  bums  (last  80  years)  and  at 
lower  elevations,  where  lodgepole  pine 
{Pinus  contorta)  is  dominant,  and  in  areas  of 
intermixed   coniferous   forest   and   meadows 


where  Douglas  fir  (Pseudotsuga  menziesii) 
also  occurs.  Limber  pine  {Pinus  flexilis)  is 
sparsely  scattered  throughout.  Understory  is 
dominated  by  highbrush  huckleberry 
{Vaccinium  globuhre),  grouse  whortleberry 
(V.  scoparium),  mountain  ash  {Sorbus  scopu- 
lina),  and  sedges  {Carex  spp.). 

Methods 

The  density  of  all  trees  (dbh  >  15  cm), 
saplings  (dbh  7.5-15  cm),  and  seedlings  (dbh 
<  7.5  cm  regardless  of  height)  by  species  was 
measured  on  10  randomly  placed  quadrats 
(0.004  ha  each)  in  each  of  the  six  harvest 
blocks  before  and  after  logging.  Soil  beneath 
the  ground  litter  was  classified  as  mesic  if  it 
felt  damp  or  xeric  if  it  felt  dry. 

Small  mammals  were  snap-trapped  on 
each  harvest  block  just  prior  to  and  immedi- 
ately after  logging  and  on  each  harvested 
block  and  adjacent  undisturbed  sites  for  up  to 
one  vear  thereafter  at  monthly  intervals. 
Each  sample  consisted  of  60  traps  in  three 
lines  with  10  trap  stations  16  m  apart  per  line 
and  2  traps  3  m  apart  per  station.  Traps  were 
baited  with  peanut  butter  and  checked  daily 
for  3  consecutive  days  for  a  total  of  180  trap 
davs  (TD)  per  sample  (one  trap  set  for  24 
hours  equals  one  trap  day).  The  species;  sex; 
age  (juvenile  or  adult);  length  of  body,  tail. 


'Department  of  Fishery-  and  Wildlife  Biology,  Colorado  State  University.  Fort  Collins,  Colorado  80522.  Reprints:  Box  2705,  Jackson.  Wyoming  83001. 
■Department  of  Biology,  Idaho  State  I'niversity.  Pocatello.  Idaho  83209. 


183 


184 


Great  Basin  Naturalist 


Vol.  40,  No.  2 


right  hind  foot,  and  right  ear;  and  reproduc- 
tive status  were  recorded  for  all  individuals 
trapped.  Testes  length  for  males  and  numbers 
of  embryos  or  placental  scars  for  females 
were  also  recorded.  Age  was  based  on  pelage, 
size,  and  reproductive  status. 

Statistical  tests  were  based  on  the  Chi 
square  method  of  analysis  imless  otherwise 
indicated. 

Results  and  Discussion 

Effects  of  Logging  on  Vegetation 

Collectively,  logging  altered  approx- 
imately 24  percent  of  the  study  area.  Table  1 
shows  the  harvest  schedule.  Selective  cutting 
removed  a  mean  of  52,  52,  and  77  percent  of 
the  trees,  saplings,  and  seedlings,  respective- 
ly, and  mesic  ground  conditions  persisted  for 
one  year  on  these  sites  (Table  2).  Clear- 
cutting  removed  88,  70,  and  79  percent  of 
the  trees,  saplings,  and  seedlings,  respective- 


ly, but  ground  conditions  changed  from  mes- 
ic to  xeric  within  9  months  after  harvest.  Al- 
though not  quantitatively  measured,  logging 
and  skidding  drastically  disturbed  understory 
vegetation  and  litter  in  both  clear  and  selec- 
tive cuts. 


Effects  of  Logging  on  Small  Mammals 

Five  species  were  captured:  red-backed 
voles,  deer  mice,  western  jumping  mice 
{Zapiis  princeps),  yellow  pine  chipmunk  (£«- 
tamius  amoenus),  and  masked  shrews  {Sorex 
cinereus).  Ninety-eight  percent  of  478  cap- 
tures were  voles  and  deer  mice;  only  these 
two  species  are  discussed  here. 

Abundance:  Voles  were  significantly 
(P<0.01)  more  abundant  on  mesic  soils  (76 
percent  of  408  captures.  Fig.  1)  throughout 
the  study,  but  deer  mice  were  significantly 
(P<0.01)  more  numerous  on  the  one  xeric 
site  sampled  10  months  after  clearcutting  (80 


Table  1.  Logging  schedule  for  the  six  timber  harvest  blocks.  Toppings  Lakes  Study  Area,  Bridger-Teton  National 
Forest,  Wyoming. 


Size 

Harvest 

Harvest  schedule 

Block 

Interval 

No. 

(ha) 

method 

Cut 

Skidded 

(days) 

1 

19.4 

Selective  cut 

8/19/76 

8/21/76 

2 

2 

9.6 

Clearcut 

9/08/75 

9/10/75 

2 

3 

75.0 

Selective  cut 

9/15/76 

10/31/76 

46 

^ 

8.1 

Selective  cut 

9/10/76 

9/10/76 

0 

5 

9.6 

Selective  cut 

9/09/75 

8/20/76 

315 

6 

36.4 

Selective  cut 

9/13/76 

9/23/76 

10 

^Logging  was  suspended  after  4  ha  were  logged  aiid  it  became  apparent 
that  the  expected  timber  voKimes  were  not  there. 


Table  2.  The  changes  in  tree  overstory  densities  for  clear  and  selective  cuts  on  the  Toppings  Lakes  Study  Area, 
Bridger-Teton  National  Forest,  Wyoming. 


Mean  size 
of  harvest 
block  (\mf 


Mean  number  per  hectare* 


before 
logging 


after 
logging 


Selective  cuts 
Trees 
Saplings 
Seedlings 

Clearcut 
Trees 
Saplings 
Seedlings 


29.7  ±  27.7  SD 


9.6 


1064  ±  233 

866  ±     84 
11027  ±  4124 


1213 

743 

17203 


507  ±  47 

416  ±  196 

2592  ±  545 


150 

230 

3770 


"Means  and  standard  deviations  for  selective  cuts  (N  =  5). 


June  1980 


Campbell,  Clark:  Rodent  Ecology 


185 


percent  of  60  captures).  Koehler  et  al.  (1975) 
found  similar  results  in  Idaho  on  undisturbed 
niesic  sites  and  on  sites  xerified  by  forest  fire. 
Red-backed  voles  apparently  require  a  heavy 
cover  of  vegetation  or  logs  (Gashwiler  1959, 
LaBue  and  Darnell  1959,  Hooven  1969, 
Krefting  and  Ahlgren  1974)  and  reside  pri- 
marily in  cool,  damp  forests  (Townsend  1935, 
Bailey  1936).  The  greater  diversity  of  under- 
story  plants  of  mesic  sites  (Daubenmire  and 
Daubenmire  1968)  apparently  provide  food 
and  cover  for  red-backed  voles,  but  deer 
mice  prefer  xeric  habitats. 

Species  composition  on  logged  and  un- 
logged  mesic  sites  (selective  cut  #5)  did  not 
differ  significantly  (P>0.10)  one  year  after 
logging  (Fig.  2).  Conversely,  the  disturbed 
xeric  site  (clearcut  2)  had  a  highly  significant 
(P<0.01)  change  in  species  composition  fol- 
lowing harvest  (Fig.  2).  Preharvest  and  imme- 
diate postharvest  data  (September  and  Octo- 
ber 1975,  respectively)  showed  a  community 
composed  primarily  of  red-backed  voles  (73 
percent  of  113  captures).  Nine  to  12  months 


after  harvest  (June  to  September,  following 
winter  inaccessibility)  composition  had 
changed  to  80  percent  deer  mice  of  60  ro- 
dent captures.  These  deer  mouse  capture 
rates  indicate  a  larger  population  than  that 
on  the  original,  undisturbed  forest.  This  is  at- 
tributed to  the  xerification  of  this  site  as  a  re- 
sult of  logging.  Similar  increases  of  deer  mice 
were  observed  by  Tevis  (1956),  Gashwiler 
(1959,  1970),  Koehler  et  al.  (1975),  and  Hoov- 
en and  Black  (1976)  in  forests  altered  by  tim- 
ber harvest  and  forest  fires.  Clearcuts  may  be 
more  attractive  to  deer  mice  (Gashwiler 
1959)  because  they  tend  to  move  into  dis- 
turbed, depopulated  areas  (Stickel  1946). 

Age  ratios:  On  newly  logged  selective 
cuts,  juveniles  outnumbered  adults  4.1  :  1.0 
for  red-backed  voles  and  5.0  :  1.0  for  deer 
mice,  but  9  to  12  months  later  juveniles  had 
decreased  significantly  (P<0.01)  to  0.8  :  1.0 
for  voles  and  0.6  :  1.0  for  deer  mice.  Juveniles 
also  outnumbered  adults  on  the  newly  clear- 
cut  site  1.6  :  1.0  for  voles  and  1.8  :  1.0  for 
deer  mice. 


90    -■ 


80 


70 


60 


50    -■ 


40 


30 


20 


10 


RED-BACKED   VOLES 


DEER  MICE 


LOGGED 
(mesic ) 


UN LOGGED 
(mesic) 


LOGGED 
(mesic) 


UNLOGGED 
(xeric) 


SELECTIVE  CUTS 


CLEAR   CUTS 


Fig.  1.   Percent  captures  of  red-backed  voles  and  deer  mice  on  logged  and  unlogged  areas.  Bridger-Teton  National 
Forest,  Wyoming.  Figures  at  the  top  of  the  bars  equal  the  number  of  animals  trapped. 


186 


Great  Basin  Naturalist 


Vol.  40,  No.  2 


Juveniles  also  comprised  75  percent  of  all 
deer  mice  captured  in  a  recent  burn  (Sims 
and  Buckner  1972).  Powell  (1972)  found 
three  times  as  many  red-backed  vole  juve- 
niles in  a  recently  blown  down  forest  than  in 
an  undisturbed  forest.  He  concluded  that 
standing  forests  were  preferred  red-backed 
vole  habitat  and  adults  drove  juveniles  into 
the  less  preferred,  disturbed  habitat  (presum- 
ably via  aggressive  behavior). 

Sex  ratios:  The  overall  sex  ratio  for  red- 


backed  voles  was  1.2M:1.0F,  comparable  to 
1.3M:1.0F  found  in  an  Oregon  study  (Gash- 
wiler  1959).  Adults  had  an  even  sex  ratio,  but 
juvenile  males  significantly  (P<0.05)  out- 
numbered juvenile  females  (1.5M:1.0F).  Vole 
sex  ratios  (adults,  juveniles,  and  total)  did  not 
differ  significantly  (P>0.05)  in  unlogged 
mesic,  logged  mesic,  or  logged  xeric  sites. 

Sex  ratios  for  all  deer  mice  captured  (1.0 
M:1.0  F);  for  adults  (1.6  M:1.0  F);  for  juve- 
niles (0.6  M:1.0  F);  and  for  logged  mesic,  un- 


70 


50   ■- 


40 


30   -- 


20 


10 


y 


70 


40 


30 


20   -- 


10 


Block  2  -  Clearcut 


Sept    Jun   Jul   Aug   Sept 
1975  1976 

Block  5  -  Selective  Cut 


^ 


#     #  Total  Captures 
#—  —  —•  Red-backed  Voles 
# #  Deer  Mice 


-\ 1 1 H 


Block    2    -    Unlogged 


y 


—I 1 1 1 H 

Sept   Jun    Jul   Aug   Sept 
1975  1976 

Block  5  -  Unlogged 


Fig.  2.  Numbers  of  red-backed  voles  and  deer  mice  trapped  on  logged  and  unlogged  sites,  Bridger-Teton  National 
Forest,  Wyoming  (September  1975,  Jvme-September  1976). 


June  1980 


Campbell,  Clark:  Rodent  Ecology 


187 


logged  mesic,  or  logged  xeric  sites  did  not 
differ  significantly  (P>0.05)  from  1.0  M  :  1.0 
F. 

Reproduction:  Reproductive  data  for 
June-September  1976  showed  that  most  re- 
production ceased  after  August  (Tables  3  and 
4).  Males  of  both  species  with  scrotal  testes 
were  captured  through  August  but  not  in 
September.  Packard  (1968)  and  Clark  (1973) 
used  male  rodents  exhibiting  scrotal  testes  as 
an  indication  of  population  breeding.  Fe- 
males of  both  species  carried  embryos  and 
were  lactating  from  June  to  September,  but 
September  embryos  were  near  term  (Table 

4). 

Mean  litter  size  and  SD  based  on  embryo 
and  placental  scar  counts  was  6.0  ±  0.9 
(range  4-9)  for  44  red-backed  voles  and  6.1 
±  0.3  (range  6-7)  for  10  deer  mice.  These 
were  similar  to  other  studies  of  red-backed 
voles  in  Crand  Teton  National  Park  (Clark 
1973)  and  of  deer  mice  in  northwestern 
Wyoming  (Clark  1975,  Long  1964).  Repro- 
ductive timing  or  litter  sizes  for  either  spe- 
cies did  not  differ  between  logged  and  im- 
logged  areas. 


Body  measurements:  Linear  measure- 
ments for  red-backed  voles  and  deer  mice  are 
shown  in  Tables  5  and  6,  respectively.  Adult 
females  of  both  species  were  insignificantly 
larger  than  males.  No  significant  (P>0.05, 
AOV)  intraspecific  age  differences  in  mea- 
surement of  either  sex  were  observed  be- 
tween logged  and  unlogged  areas  9  to  12 
months  after  harvest.  Male  red-backed  voles 
from  this  study  and  nearby  Grand  Teton  Na- 
tional Park  (N'  =  7,  Clark  1973)  did  not  differ 
significantly  (P>0.10,  t-test). 


Conclusions 

Both  selective  cutting  that  removed  about 
57  percent  of  the  trees  and  clearcutting  of 
spruce-fir  forest  resulted  in  an  increase  of 
juvenile  red-backed  voles  and  deer  mice  two 
weeks  to  a  month  after  logging  that  was  not 
apparent  9  to  12  months  later.  Clearcutting 
changed  species  composition  9  to  12  months 
later  from  predominantly  red-backed  voles 
on  unlogged  areas  to  predominantly  deer 
mice.  The  change  was  attributed  to  soil  xeri- 


Table  3.   Male  reproductive  condition  of  red-liacked  voles  and  deer  mice,  Bridger-Teton  National  Forest.  Wyo- 
ming (June-September  1976). 


Number 

of 

Number  of 

red-hacked  voles 

deer  mice 

Condition 

June 

J"iy 

Aug. 

Sept. 

Total 

June 

July 

Aug.       Sept. 

Total 

Scrotal 

9 

7 

12 

0 

28 

4 

9 

1 

0 

14 

Nonscrotal 

0 

0 

11 

9 

20 

0 

1 

13 

6 

20 

Total 

9 

7 

2.3 

9 

48 

4 

10 

14 

6 

34 

Table  4.   Reproductive  condition  of  red-backed  vole  and  deer  mouse  adult  females.  Bridger-Teton  National  For- 
est, Wyoming  (June-September  1976). 


N 

umber 

of 

Number 

of 

red-backed  voles 

deer  mice 

Condition 

June 

July 

Aug. 

Sept. 

Total 

June 

July 

Aug. 

Sept. 

Total 

Nimiber  of  females  examined 

8 

5 

13 

18 

44 

1 

6 

10 

1 

18 

Percent  pregnant 

50 

80 

23 

72 

54 

100 

67 

20 

100 

44 

Niunber  of  embrvos 

Mean 

6.3 

6.8 

6 

5.9 

6.1 

6 

6.5 

6 

6 

6.3 

Range 

6-7 

5-9 

6 

4-9 

4-9 

6 

6-7 

6 

6 

6-7 

Percent  with  placental  scars 

50 

40 

-"- 

28 

48 

0 

.33 

80 

0 

.56 

Number  of  placental  scars 

Mean 

5.5 

6 

6 

.5.6 

5.8 

0 

6 

6 

0 

6 

Range 

5-6 

6 

6 

4-6 

4-6 

0 

6 

6 

0 

0-6 

Number  of  females  lactating 

4 

6 

13 

17 

40 

1 

0 

10 

1 

17 

188 


Great  Basin  Naturalist 


Vol.  40,  No.  2 


fication  on  the  clearcut.  Voles  continued  to 
predominate  on  selectively  cut  sites,  and  soils 
remained  mesic  there.  For  both  species  adults 
consistently  outnumbered  juveniles  in  June, 
July,  and  August  in  both  logged  and  unlogged 
areas,  but  juveniles  outnumbered  adults  in 
areas  newly  logged  in  September.  Sex  ratios, 
timing  of  reproduction,  litter  size,  and  body 
measurements  for  both  species  did  not  differ 
significantly   between   unlogged,    selectively 


cut,  or  clearcut  areas  9  to  12  months  after 
logging. 

Acknowledgments 

This  study  was  supported  by  the  Bridger- 
Teton  National  Forest,  Jackson,  Wyoming. 
Special  thanks  is  extended  to  Mr.  G.  GreuU, 
Wildlife  Biologist,  U.S.  Forest  Service.  Mr. 
W.  Barmore  offered  valuable  criticism  of  the 
manuscript. 


Table  5.  Body  measurements  of  258  red-backed  voles  from  unlogged,  clearcut,  and  selectively  cut  areas,  Bridger- 
Teton  National  Forest,  Wyoming  (June-September  1976). 


Sex  and  age* 


Mean  length,  SD,  and  (range)  in  mm 


Body 


Tail 


Ear 


Hind  Foot 


Control 

M2+  (n=18) 
F2+  (n  =  23) 
Ml-(n  =  40) 
Fl-(n  =  26) 


139.4  ±8.8(1 19-152) 
143.4  ±10.9(118-158) 

115.2  ±12.1(88-137) 
112.2  ±10.5(90-132) 


39.1  ±2.6(34-45) 

40.5  ±4.4(26-47) 
,33.0  ±4.5(26-43) 

.33.2  ±3.7(25-42) 


15.8  ±1.8(1,3-21) 
16.5±  1.1(14-19) 
14.9±  1.8(10-17) 
14.9±  1.7(10-18) 


18.4  ±0.9(17-19) 
18.0  ±1.0(16-19) 
16.9  ±1.2(15-18) 
17.0±  0.7(15-18) 


Clearcut 

M2+  (n  =  3) 
F2+  (n  =  4) 
Ml-(n  =  4) 
Fl-(n  =  l) 


147.7  ±6.4(144-155)  41.0  ±3.5(39-45)  14.3  ±0.6(14-15)  18.7  ±0.6(18-19) 

139.6  ±10.4(125-147)  40.3  ±2.9(37-44)  17.0±  0.8(16-18)  17.8  ±  1.0(17-19) 

113.3  ±13.8(100-130)  33.5  ±6.2(27-40)  13.0±  2.0(12-16)  16.8  ±0.5(16-17) 

117.0-       -  .35.0-       -  17.0-       -  18.0-       - 


Selective  cut 

M2+  (n  =  27) 
F2+  (n  =  17) 
Ml-(n  =  59) 
Fl-(n  =  36) 


138.9±  10.1  (115-160)  40.8 ±3.5(26-48)  15.9±  1.7(12-18)  18.4 ±  1.0(16-19) 

142.7  ±11.2(122-159)  40.6  ±4.7(37-47)  15.9±  1.8(11-18)  17.9±  0.8(16-19) 

121.9  ±8.5(96-1.37)  ,35.6  ±.3.5(26-42)  15.6±  1.6(11-19)  17.3  ±0.7(16-19) 

115.0±  10.7(93-138)  .3,3.5  ±,3.6(25-40)  15.4  ±  1.9(10-18)  17.2  ±0.7(16-19) 


*2+  =  adult;  1-  =  juvenile. 


Table  6.  Body  measurements  of  118  deer  mice  from  unlogged,  clearcut,  and  selectively  cut  areas,  Bridger-Teton 
National  Forest,  Wyoming  (June-September  1976). 


Mean  length,  SD,  and  (range)  in  mm 


Sex  and  age° 


Body 


Tail 


Ear 


Hind  foot 


Control 

M2-I-  (n=ll) 
F2+  (n  =  5) 
Ml-(n  =  8) 
Fl-(n  =  9) 

Clearcut 

M2+  (n  =  12) 
F2+  (n  =  10) 
Ml-(n=9) 
Fl-(n=17) 

Selective  cut 

M2+  (n  =  12) 
F2+  (n  =  4) 
Ml-(n  =  6) 
Fl-(n=15) 

'2+   =  adult;  1-  =  juvenile. 


157.5±  11.1(140-17,5) 
156.6±  10.1(146-170) 
139.4  ±5.0(133-146) 
146.0±6.,3(137-155) 


68.4  ±4.8(62-77) 
69.2±4..3(63-7.3) 
6.3.9  ±2.0(61-66) 
67.0  ±3.4(62-72) 


158.7±  11.1(148-175)  69.8  ±5.0(6,3-77) 

166.3  ±  14..3(144-188)  7.3.4  ±  7.1(62-85) 

146.4  ±7.2(133-156)  67.4  ±3.6(61-74) 

142.5  ±  7.6(130-155)  63.4  ±  5.2(55-72) 


154.1  ±  10.1(141-174)  67.4  ±5.2(61-76) 

160.1  ±  17.8(145-183)  69.8  ±  8.3(62-80) 
146.8  ±6.6(139-157)  66.5  ±3.7(64-74) 

143.2  ±9.6(128-160)  65.0  ±5.0(57-74) 


18.5  ±0.8(1.3-20) 
18.8  ±0.5(1.3-19) 
18.3  ±1.0(17-20) 
18.8  ±0.8(17-20) 


18.0  ±1.7(15-20) 
18.3  ±1.3(17-20) 
18.7  ±1.0(18-20) 
17.6  ±1.8(14-20) 


18.3  ±1.4(15-20) 
18.5±  1.0(17-19) 
18.2  ±0.8(17-19) 
17.9  ±1.5(16-20) 


18.4  ±0.9(19-20) 
19.6  ±0.9(19-21) 
19.1  ±0.6(18-20) 
18.6±  1.0(17-20) 


19.4  ±0.7(19-21) 
19.3  ±0.7(18-20) 
19.1  ±0.9(18-20) 

18.5  ±1.0(17-20) 


19.1  ±0.,5(  18-20) 
19.3  ±1.0(18-20) 

19.2  ±1.0(18-20) 
18.8  ±0.8(17-20) 


June  1980 


Campbell,  Clark:  Rodent  Ecology 


189 


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Daube.nmire,  R.,  and  J.  B.  Daubenmire.  1968.  Forest 
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TERMINAL  BUD  FORMATION  IN  LIMBER  PINE 


Ronald  M.  Lanner'  and  James  A.  Bryan' 


,\bstr.'\ct.—  The  progress  of  bud  development  was  studied  in  limber  pines  growing  in  the  mountains  of  north- 
eastern Utah.  Initiation  of  new  bud  scales  began  in  mid-June,  several  weeks  after  elongation  of  the  current  shoot  had 
begim.  Needle  primordia  first  appeared  in  September  and  continued  to  form  through  the  winter,  until  all  were  pres- 
ent in  May.  This  winter  activity  is  believed  to  be  fostered  by  surface  temperatures  on  the  terminal  buds  considerably 
higher  than  ambient  air  temperatures. 


The  annual  shoot  of  limber  pine  (Pinus 
flexilis  James),  a  five-needled  species  com- 
mon in  the  Rocky  Mountains,  consists  of  a 
monocyclic  spring  shoot  formed  by  the  elon- 
gation of  a  winter  bud.  This  is  the  most  fa- 
miliar shoot  development  pattern  in  northern 
pines  and  has  been  classified  as  the  Resinosa 
pattern  (Lanner  1976).  But  even  among  spe- 
cies of  this  habit,  there  is  diversity  in  the  de- 
velopmental schedule  of  bud  morphogenesis. 
For  example,  in  some  species  formation  of 
the  new  winter  bud  begins  while  the  old  one 
is  still  elongating,  but  in  others  bud  formation 
is  delayed  until  the  cessation  of  current-sea- 
son shoot  growth.  Further,  the  timing  of 
.short-shoot  and  needle  morphogenesis  is  also 
subject  to  variation.  In  this  report  we  de- 
scribe the  annual  cycle  of  development  of 
limber  pine  terminal  buds  to  resolve  the 
questions  of  when  the  short  shoots  and  nee- 
dles are  formed,  both  in  terms  of  calendar 
date  and  in  regard  to  the  growth  stage  of  the 
spring  shoot. 

The  trees  studied  grow  at  an  elevation  of 
2130  m  on  a  steep  southeast  slope  in  Logan 
Canyon,  northeastern  Utah.  They  vary  from 
12  to  27  cm  DBH  and  4  to  7  m  in  height.  At 
each  of  18  collection  dates  during  1978,  2 
terminal  buds  of  vigorous  first-order  branches 
in  the  upper  crown  were  harvested  from 
each  of  at  least  2  of  the  18  study  trees.  Buds 
were  stored  in  formaldehyde-acetic  acid-eth- 
yl alcohol  (FAA),  dissected  with  standard  mi- 
cro-dissecting tools,  and  examined  at  12-lOOx 


with  a  Wild  M-5  stereomicroscope.  Two 
shoots  had  steel  pins  inserted  at  the  base  of 
tlie  bud  as  a  datum  for  observations  of  shoot 
elongation. 

On  each  of  the  harvested  buds  we  dis- 
sected at  least  two  short-shoot  budlets  from 
the  proximal  (basal)  end  of  the  bud,  and  two 
from  the  distal  (apical)  end.  We  counted  the 
budlet  scales  (future  fascicle  sheath  scales) 
and  needle  primordia,  if  any.  The  study  took 
place  during  a  single  calendar  year,  so  we  ac- 
tually observed  the  late  development  and 
elongation  of  the  1978  spring  shoot  and  the 
early  development  of  the  1979  spring  shoot. 
Ideally,  a  study  of  this  kind  should  begin  with 
early  morphogenesis  of  a  bud  and  end  with 
the  maturation  of  the  resulting  shoot. 

Results 

One  of  the  marked  shoots  started  to  elon- 
gate during  the  interval  20-30  May,  and  the 
other  during  the  interval  30  May-5  June. 
These  shoots  completed  their  elongation 
growth  prior  to  8  July  and  13  July,  respec- 
tively. Final  lengths  of  these  shoots  were  30.5 
and  21.5  mm.  Pollen  was  shed  during  the  pe- 
riod 9-13  July. 

Initiation  of  the  primary  bud  scales  (cata- 
phvlls)  of  the  newlv  forming  terminal  bud  be- 
gan between  11  and  18  June.  After  cataphylls 
formed,  meristems  appeared  in  the  axils  of 
most  of  them.  These  axillary  meristems  be- 
came the  apical  meristems  of  the  short-shoot 


'Department  of  Forestry  and  Outdoor  Recreation,  Utah  State  University,  Logan,  Utah  84322. 


190 


June  1980 


Lanner,  Bryan:  Limber  Pine 


191 


budlets  that  would  later  become  needle  fas- 
cicles. Scales  formed  on  these  budlets  are  fu- 
ture fascicle  sheath  scales.  Sheath  scale  ini- 
tiation began  in  proximal  budlets  in  early 
July,  and  in  distal  budlets  in  early  August. 
Scale  production  continued  in  both  types  of 
budlets  luitil  about  mid-Januarv. 

The  earliest  needle  primordia  started  to 
appear  in  proximal  budlets  in  late  August  and 
continued  to  appear  over  a  period  of  three 
months.  Needle  primordia  in  distal  budlets 
did  not  begin  to  form  until  about  mid-No- 
vember, but  continued  appearing  up  to  early 
May,  about  5.5  months  later. 

When  bud  scale  initiation  began,  about 
two-thirds  of  the  elongation  growth  of  the 
marked  shoots  had  been  completed.  Bud 
scale  initiation  probably  continued  through- 
out the  remainder  of  the  elongation  period. 
Sheath  scale  initiation  began  in  proximal 
budlets  about  the  time  shoot  elongation  was 
ceasing,  and  it  began  in  distal  budlets  after 
the  cessation  of  shoot  growth.  When  needle 
primordia  started  to  form,  shoot  growth  had 


been  inactive  for  several  weeks.  The  needle 
primordia  formed  in  the  spring  developed 
prior  to  the  onset  of  shoot  elongation  in  late 
May. 

When  the  first  buds  were  harvested  22  Jan- 
uary 1978,  short-shoot  budlets  from  proximal 
parts  of  the  terminal  bud  contained  10-12 
scales  (x  =  11.2)  and  all  had  their  full  com- 
plement of  5  needle  primordia.  But  budlets 
located  at  the  distal  end  of  those  terminal 
buds  tended  to  have  fewer  scales  (8-11;  x  = 
10.0)  and  averaged  only  1.7  needle  primordia 
(Table  1).  The  difference  in  scale  number  was 
maintained  even  beyond  8  May,  when,  for 
the  first  time,  all  the  distal  budlets  contained 
their  full  complement  of  5  needle  primordia 
(Table  1). 

Discussion  and  Conclusions 

The  initiation  of  bud  scales  was  first  noted 
on  18  June,  when  the  1978  shoot  had  attained 
68  percent  of  its  final  length,  showing  that 
bud  morphogenesis  began  during  the  period 


Table  1.   Progress  of  sheath  scale  and  needle  initiation  in  proximal  and  distal  short  shoot  budlets  of  first-order 
terminal  buds  of  Limber  Pine,  and  of  shoot  elongation. 


Proximal  budlets 

Needle 

Distal  budlets 

Needle 

Length  of 
1978 

number 

rimordium 

primordium 

bud/shoot  as 

Scale 

ninnber 

Scale 

number 

number 

percent  of 
final  length 

1978  Date 

X 

range 

X 

X 

range 

X 

1978  annual 

shoot 

22  Jannary 

11.2 

10-12 

5.0 

10.0 

8-11 

1.7 

31.5 

21  March' 

10.0 

9-12 

5.0 

9.6 

9-10 

2.5 

31.5 

12  April 

10.3 

9-12 

5.0 

9.5 

9-10 

3.3 

31.5 

8  May 

10.0 

9-11 

5.0 

10.6 

10-11 

5.0 

31.5 

20  May 

9.5 

8-11 

5.0 

9.5 

9-10 

5.0 

31.5 

4  Jnne 

10.0 

10 

5.0 

9.5 

9-10 

5.0 

45.0 

1 1  Jnne 

13.0 

13 

5.0 
1979  annual 

11.0 
shoot 

11 

5.0 

55.5 

1 1  Jnne 

0 

0 

0 

0 

0 

0 

55.5 

IS  June 

0 

0 

0 

0 

0 

0 

68.0 

2Jnly 

0 

0 

0 

0 

0 

0 

95.0 

13  July 

2.0 

2 

0 

0 

0 

0 

100 

23  July 

4.5 

3-6 

0 

0 

0 

0 

1(X) 

2  August 

5.0 

5 

0 

0 

0 

0 

1(K) 

16  August 

6.0 

6 

0 

0.75 

0-2 

0 

100 

26  August 

6.0 

.5-7 

0 

2.3 

2-3 

0 

1(X) 

15  September 

8.0 

8 

5.0 

7.3 

6-9 

0 

100 

5  October 

8.3 

8-9 

3.0 

7.3 

6-9 

0 

100 

15  November 

8.8 

8-10 

2.2 

7.0 

6-8 

0 

100 

9  December 

8.0 

8 

5.0 

8.0 

8 

1.8 

100 

192 


Great  Basin  Naturalist 


Vol.  40,  No.  2 


of  shoot  elongation.  In  this  regard  limber 
pine  resembles  P.  strobus  L.  and  P.  lamber- 
tiana  Dougl.  (Lanner  1976).  It  provides  fur- 
ther evidence  that  shoot  elongation  does  not 
inhibit  the  initiation  of  lateral  structures  on 
the  shoot  apical  meristem.  Bud  morphoge- 
nesis in  limber  pine  is  delayed,  however,  in 
comparison  to  that  of  lodgepole  pine  (P.  con- 
torta  Dougl.),  one  of  its  associates  in  this  area 
(Van  Den  Berg  and  Lanner  1971).  In  lodge- 
pole,  initiation  of  the  new  bud  and  elonga- 
tion of  the  shoot  began  almost  simultaneously 
early  in  May. 

A  given  stage  of  development— i.e.,  attain- 
ing a  certain  number  of  sheath  scales  or  a 
certain  number  of  needle  primordia— is 
reached  earlier  in  proximal  than  in  distal 
short  shoots.  Thus,  in  the  1978  winter  buds, 
the  full  complement  of  5  needle  primordia 
was  present  in  proximal  short  shoots  in  Janu- 
ary collections,  but  was  not  found  in  distal 
short  shoots  until  May.  This  is  another  case  of 
the  developmental  gradient  in  short-shoot 
maturation  described  in  the  more  complex 
buds  of  lodgepole  pine  (Van  Den  Berg  and 
Lanner  1971),  the  much  larger  buds  of  slash 
pine,  P.  elliottii  Engelm.  (Lanner  1978),  and 
in  eastern  white  pine  (Owston  1969). 

Perhaps  the  most  unusual  finding  reported 
here  is  that  morphogenetic  activity  contin- 
ued in  wintering  limber  pine  buds.  Buds  col- 
lected early  in  1978  showed  periodic  increas- 
es in  needle  primordium  number  in  distal 
budlets.  Buds  collected  late  in  1978  showed 
consistent  increases  in  needle  primordium 
number  in  both  proximal  and  distal  budlets, 
and  in  sheath  scale  number  in  distal  budlets 
(Table  1).  Yet  temperatures  at  a  nearby  tem- 
porary weather  station  at  a  comparable  ele- 
vation fell  as  low  as  -21  C  in  January  and 
-12  C  in  November  and  December  (pers. 
comm.,  S.  A.  Loomis).  In  lodgepole  pine 
studied  just  a  few  miles  from  this  site,  budlets 
overwintered  with  less  than  their  full  com- 
plements of  sheath  scales  and  needle  pri- 
mordia, but  no  changes  were  noted  during 
the  winter  (Van  Den  Berg  and  Lanner  1971). 
Budlets  of  slash  pine  actively  initiated  scales 
and  needle  primordia  during  the  winter 
months,  but  this  was  in  the  much  milder  cli- 
mate of  Florida  (Lanner  1978). 

Seed  cone  primordia  have  also  been  re- 


ported as  morphogenetically  active  during 
the  winter.  Duff  and  Nolan  (1958)  observed 
changes  in  ovulate  strobili  of  red  pine  (P.  resi- 
nosa  Ait.)  between  October  and  January  in 
the  cold  climate  of  Chalk  River,  Ontario.  Gif- 
ford  and  Mirov  (1960)  also  reported  female 
strobilus  development  in  ponderosa  pine  (P. 
ponderosa  Laws.),  but  in  the  milder  climate 
of  the  Sierra  Nevada  west  slope. 

Such  meristematic  activity  may  be  permit- 
ted by  surface  temperatures  considerably 
higher  than  those  of  the  ambient  atmosphere. 
For  example,  Tranquillini  and  Turner  (1961) 
have  reported  maximum  monthly  temper- 
atures of  Swiss  stone  pine  (P.  cernbra  L.)  nee- 
dles 2.7  and  7.4  C  higher  than  air  temper- 
atures during  November  and  March, 
respectively,  in  the  Austrian  Alps.  In  March, 
needles  reached  a  maximum  of  18.4  C, 
though  the  mean  air  temperature  for  that 
month  was  only  0.3  C.  At  our  study  site,  even 
in  January,  the  coldest  month  of  the  year,  air 
temperature  on  three  occasions  attained  al- 
most 6  C.  Bud  surface  temperatures  may 
have  reached  as  high  as  12  C  or  more,  well 
above  the  apparent  threshold  for  meristemat- 
ic activity. 

Literature  Cited 

Duff,  C.  H.,  and  N.  J.  Nolan.  1958.  Growth  and  mor- 
phogenesis in  the  Canadian  forest  species.  III. 
The  time  scale  of  morphogenesis  at  the  stem 
apex  of  Pintis  resinosa  Ait.  Canadian  J.  Bot. 
36:687-706,  4  pis. 

GiFFORD,  E.  M.,  Jr.,  and  N.  T.  Mirov.  1960.  Initiation 
and  ontogeny  of  the  ovulate  strobilus  in  ponde- 
rosa pine.  Forest  Sci.  6:19-25. 

Lanner,  R.  M.  1976.  Patterns  of  shoot  development  in 
Pinus  and  their  relationship  to  growth  potential. 
Pages  223-243  in  M.  G.  R.  Cannell  and  F.  T. 
Last,  eds.  Tree  physiology  and  yield  improve- 
ment. Academic  Press,  New  York  and  London. 

1978.  Development  of  the  terminal  bud  shoot  of 

slash  pine  saplings.  Forest  Sci.  24:167-179. 

Owston,  P.  W.  1968.  The  shoot  apex  in  eastern  white 
pine:  its  stnicture,  seasonal  development,  and 
variation  within  the  crown.  Canadian  J.  Bot. 
47:1181-1188. 

Tranquillini,  W.,  and  H.  Turner.  1961.  Untersuchu- 
ngen  iiber  die  Pflanzentemperaturen  in  der  sub- 
alpinen  Stufe  mit  besonderer  Beriicksichtung  der 
Nadeltemperaturen  der  Zirbe.  Mitteil.  Forstlich. 
Bundes-Versuchs.  Mariabrunn  59(1):  127-151. 

Van  Den  Berg,  D.  A.,  and  R.  M.  Lanner.  1971.  Bud  de- 
velopment in  lodgepole  pine.  Forest  Sci. 
17:479-486. 


STINGER  UTILIZATION  AND  PREDATION 
IN  THE  SCORPION  PARUROCTONUS  BOREUS 

Bnice  S.  Gushing'  and  Anne  Matherne' 

.\bstr.\ct.—  The  iitihzution  of  the  stinger  and  the  predatory  technique  of  the  scorpion,  Paruroctanus  boretis,  was 
studied  under  laboratory  conditions.  During  the  study,  83  feedings  were  observed.  Age  of  the  scorpions  and  the  per- 
centage of  prey  stung  by  them  were  used  to  classify  the  scorpions  into  groups.  The  scorpions  aged  1.3-61  days  always 
stung  prey,  .\fter  62  days  the  scorpions  began  to  selectively  utilize  the  stinger.  Utilization  declined  until  it  reached 
.30  percent  in  the  adult  stage.  The  stinger  is  apparently  necessary  for  prey  capture  only  in  the  earlv  life  stages. 


The  role  of  the  stinger  in  scorpion  behav- 
ior has  never  really  been  studied  or  estab- 
lished. The  available  information  is  vague 
and  inconsistent.  Alexander  (1959)  reported 
that  different  groups  of  scorpions  utilize  the 
stinger  differentially.  Pocock  (1893)  and 
Stahnke  (1966)  stated  that  scorpions  paralyze 
prey  only  when  it  does  not  submit  to  passive 
consumption.  Finally  Hadley  and  Williams 
(1968)  reported  that  Vepvis  confusus  and 
Paruroctonus  mesaensis  and  P.  baergi  stung 
prey  (Williams  1972).  These  reports  leave  the 
adaptive  significance  of  this  potent  device  in 
doubt.  In  this  study,  we  attempted  to  estab- 
lish the  use  of  the  stinger  through  controlled 
experimentation  and  observation. 

Methods  and  Materials 

Scorpions  were  collected  from  south- 
western Oregon  on  the  Malheur  National 
Wildlife  Refuge  and  adjacent  region.  They 
were  kept  in  10-gallon  terrariums  containing 
three  inches  of  soil  and  flat  rocks  from  the 
natural  habitat.  In  most  ca.ses,  two  scorpions 
were  placed  in  each  tank  to  induce  in- 
traspecific  responses.  A  large  population  of 
grasshoppers  and  a  small  number  of  beetles, 
crickets,  and  termites  were  found  in  the  area, 
and  the.se  were  selected  in  relative  propor- 
tions as  the  prey. 

To  maintain  natural  conditions,  the  terra- 
riums remained  outdoors  except  during  peri- 


ods of  observation.  Observations  were  con- 
ducted at  night  using  red  light.  Red  light 
provided  visibility  for  us,  but  apparently  did 
not  affect  the  scorpions,  which  possess  vision 
in  the  blue  and  ultraviolet  wavelengths 
(Machan  1968). 

Results 

Over  a  two-year  period  several  different 
groups  of  Paruroctonus  boreus  were  ob- 
served. Predation  techniques  were  the  same 
for  scorpions  of  all  ages  and  sizes.  Emergence 
from  cover  occurred  between  2130  and  2300 
hours.  If  emergence  did  not  occur  by  2.300 
the  scorpions  did  not  forage  that  night.  After 
emergence,  contact  with  prey  was  estab- 
lished through  random  encounter  or  active 
stalking.  When  actively  stalking,  the  scor- 
pions traveled  with  the  pedipalps  extended 
forward  and  held  apart  at  a  distance  approx- 
imately equal  to  the  maximum  width  of  the 
abdomen.  The  telson  was  arched  over  the  ab- 
domen with  the  caudal  vesicle  above  the 
midabdomen.  When  a  potential  prey  was  de- 
tected the  scorpions  rushed  it. 

Upon  contact,  the  scorpions  used  their 
pedipalps  to  grasp  the  prey  by  one  or  more 
appendages.  If  stinging  occurred  at  all,  it  oc- 
curred at  this  time.  The  telson  was  arched 
over  the  abdomen  and  at  the  same  time  the 
abdomen  was  quickly  raised.  This  imparted  a 
downward  stabbing   motion   which   allowed 


'Department  of  Wildlife  Biology,  University  of  Montana.  Missoula,  Montana  59812. 
"Department  of  Zoology,  Louisiana  State  University.  Baton  Rouge,  Louisiana  7080.3. 


193 


194 


Great  Basin  Naturalist 


Vol.  40,  No.  2 


the  stinger  to  penetrate  the  prey's  abdomen. 
Resistance  by  the  prey  subsided  within  one 
minute  after  it  was  stung.  Whether  or  not  the 
prey  was  paralyzed,  it  was  held  motionless  in 
the  pedipalps  for  10  to  30  minutes  and  then 
transferred  to  the  chelicerae.  Upon  leaving 
the  site  of  capture  the  prey  was  slung,  ven- 
tral side  up,  onto  the  cephalothorax.The  prey 
was  carried  about  the  terrarium  in  this  posi- 
tion for  up  to  several  hours.  When  walking, 
the  scorpions  waved  their  pedipalps  in  front 
of  their  path  in  a  "blindman"  fashion,  a  slow 
exploratory  touching. 

Except  when  scorpions  were  with  young, 
prey  was  taken  beneath  cover  for  con- 
sumption. Consumption  time  varied  between 
2  and  48  hours.  Feeding  began  at  the  head  of 
the  prey  and  continued  until  the  prey  was 
consumed.  The  hard  exoskeleton  of  beetles 
was  left  as  an  empty  husk.  Several  of  these 
husks  were  found  with  scorpions  in  the  field. 
All  scorpions  used  the  same  feeding  tech- 
niques. However,  the  scorpions  also  under- 
went regular  periods  of  nonfeeding  which 
lasted  up  to  five  months. 

The  stinger  was  removed  from  two  adult 
scorpions.  These  scorpions  fed  six  times  and 
utilized  the  same  techniques  as  unimpaired 
individuals.  However,  they  never  attempted 
to  sting  any  of  the  prey. 

During  intraspecific  aggression  or  canni- 
balism, the  method  of  capture  was  as  de- 
scribed above  with  minor  modifications.  If 
there  was  a  significant  difference  in  size,  the 
smaller  scorpion  attempted  to  avoid  conflict, 
but  the  larger  one  often  pursued.  When  ag- 
gression occurred,  the  scorpions  grasped  each 
other  by  the  pedipalps  and  repeatedly  at- 
tempted to  sting.  A  size  difference  always  re- 
sulted in  the  death  of  the  smaller  scorpion. 
Consumption  proceeded  normally  after  im- 
mobilization. In  two  instances  of  the  scor- 
pions being  the  same  size,  both  animals  were 
killed. 

Immature  scorpions  did  not  capture  prey 
until  they  were  13  days  old.  Prior  to  this  time 
they  consumed  their  casting  left  at  birth  and 
their  first  exuvium.  After  the  juveniles  dis- 
persed at  9  to  11  days,  the  female  began 
feeding  with  an  alteration  in  feeding  tech- 
nique. The  adult  female  consumed  prey  in 
the  open  and  its  young  congregated  about 
the  adult's  cephlathorax.   On  day   14  the 


young  began  to  capture  prey.  Table  1  sum- 
marizes the  percentages  of  prey  stung  by  the 
early  instars  and  all  other  age  groups. 

Discussion  and  Analysis 

No  age  group  beyond  84  days  was  ob- 
served due  to  our  inability  to  keep  juveniles 
alive.  This  resulted  mainly  from  a  high  de- 
gree of  cannibalism  and  mishandling  of  a  few 
remaining  scorpions.  Data  on  cannibalism 
were  excluded  from  Table  1  because  of  the 
bias  it  would  introduce  inasmuch  as  in- 
traspecific aggression  always  elicited  stinger 
utilization.  Two  scorpions  per  tank  and  the 
limited  dispersal  range  of  the  young  led  to  an 
unnatural  increase  in  incidents  of  cannibal- 
ism. Data  on  the  six  feedings  by  the  scorpions 
with  stinger  removed  were  also  excluded  due 
to  their  inability  to  sting. 

During  active  stalking,  the  scorpions 
rushed  prey.  Some  stimulus  must  have  been 
present  which  alerted  the  scorpions.  Paruroc- 
tonus  boreus,  like  other  desert  scorpions,  may 
be  able  to  detect  and  utilize  Rayleigh  waves 
for  prey  location  (Brownell  1977).  A  Rayleigh 
wave  is  a  slow-moving  secondary  vibration 
created  by  movement  and  propagated 
through  sand. 

The  stinger  was  not  essential  for  feeding  by 
the  adult  scorpions.  A  low  percentage  of  prey 
was  paralyzed,  and  stinger-impaired  individ- 
uals were  able  to  feed  without  difficulty. 
Scorpions  are  also  capable  of  surviving  pro- 
longed periods  without  food  (Stahnke  1966). 
Considering  these  factors,  we  suggest  that  the 
amount  of  food  which  would  be  lost  to  an 
adult  incapable  of  stinging  would  not  have  a 
significant  or  deleterious  effect  upon  its  sur- 
vival. 

If  stinging  occurs,  it  is  triggered  by  two 
stimuli.  One,  as  stated  by  Pocock  (1893)  and 


Table  1.  Relationship  between  age  and  stinger  utili- 
zation. 


Age  group 

Number  of 

Number 

Percent 

in  days 

feedings 

stung 

stung 

01-12° 

0 

0 

00.0 

13-61 

40 

40 

100.0 

62-84 

13 

9 

69.2 

Adult 

30 

10 

30.0 

"Fed  on  exuviae  and  adult  pellets. 


June  1980 


CusHiNx;,  Matherne:  Scorpion  Predation 


195 


Stahnke  (1966),  is  an  attempt  by  the  prev  to 
resist  capture.  This  is  not  the  onlv  stiuiuhis. 
Struggling  hard-bodied  or  powerful  prey, 
such  as  grasshoppers,  were  stung.  Termites 
and  other  soft-bodied  prey  were  held  in  the 
pedipalps  until  resistance  subsided.  The  fac- 
tors that  elicited  a  sting  were  resistance  in 
combination  with  the  characteristics  of  the 
prey  species.  This  strongly  suggests  a  form  of 
selective  stinger  utilization. 

Tlie  selection  process  develops  over  time. 
The  first  instars  feed  on  the  exuviae  and  pos- 
sibly on  small  pellets  dropped  by  the  adult 
during  feeding  (Stahnke  1966).  Utilization  of 
pellets  for  food  suggests  the  reason  for  the 
change  in  feeding  technique  by  the  adult  fe- 
male, with  the  young  gathered  about  her 
cephalothorax.  The  next  age  group,  13-61 
days,  paralyzed  all  prey  (Table  1).  The  cause 
of  this  may  be  that  the  pedipalps  were  not 
sufficientlv  developed  at  this  point  to  hold 
prey  against  struggle.  Therefore,  in  order  to 
insure  the  maximimi  number  of  feedings  and 
promote  growth  and  development,  the  juve- 
niles must  sting  prey  at  first  contact.  As  de- 
velopment occurs  the  pedipalps  strengthen, 
and  certain  prey  types,  such  as  small  ter- 
mites, no  longer  must  be  paralyzed.  This 
would  reduce  the  use  of  toxin  and  be  energy 
efficient  by  reducing  the  manufacture  of 
more  toxin. 

Stinger  utilization  drops  from  100  to  30 
percent  in  the  adults  (Table  1).  This  decrease 
began  about  the  second  month  and  continued 
until  the  adult  stage.  The  actual  percentage 
utilization  in  the  group  aged  62-84  days  may 
have  been  biased  in  that  this  group  was  not 
fed  a  representative  class  of  prey,  but  instead 
was  fed  whatever  small  insects  and  arachnids 
happened  to  be  available.  Nevertheless,  this 
group  still  demonstrates  the  beginning  of  the 


process   of  differential   selection   for  stinger 
utilization  in  that  not  all  prey  was  stung. 

In  conclusion,  the  stinger  fimctions  as  a 
necessary  device  for  prey  capture  by  the 
early  instars.  As  physical  development  oc- 
curs, the  pedipalps  can  hold  certain  types  of 
prey  and  there  is  a  reduction  in  the  use  of  the 
stinger.  This  decline  continues  until  the  adult 
stage,  where  only  a  small  percentage  of  prey 
is  stung  and  these  are  not  essential  for  survi- 
val. However,  the  stinger  is  still  utilized  for 
intraspecific  aggression  and  possibly  for  de- 
fense. 

Acknowledgment 

We  thank  Dr.  John  Mates,  University  of 
California,  Davis,  for  his  early  help  in  this 
project,  and  Dr.  Andrew  Sheldon,  Depart- 
ment of  Zoology,  University  of  Montana,  for 
his  critical  review  of  this  manuscript. 


Literature  Cited 

Alex.\nder,  a.  J.  1959.  A  survey  of  the  l)iologv  of  scor- 
pions of  South  Africa.  Afr.  Wildl.  13(2):99-106. 

Brownell,  p.  H.  1977.  Conipressional  and  surface 
waves  in  sand:  used  by  desert  scorpions  to  locate 
prey.  Science  197:479-481. 

RIDLEY,  N.  F.,  AND  S.  C.  WiLLiAMs.  1968.  North  .Ameri- 
can scorpions  in  relation  to  feeding.  Ecology 
49(4):726-734. 

Macha.n,  L.  1968.  Spectral  sensitivity  of  scorpions'  eyes 
and  possible  role  of  shielding  pigment  effect.  J. 
Exp.  Biol.  49(1):95-105. 

PococK,  R.  I.  189.3.  Notes  upon  the  habits  of  some  living 
scorpions.  Nature.  48:1()4-I()V. 

Stahnke,  H.  L.  1966.  Some  aspects  of  scorpion  behavior. 
Southern  Calif.  Acad.  Sci.  Bull.  62(2)65-80. 

W'illia.ms,  S.  C.  1972.  Four  new  scorpion  species  be- 
longing to  the  genus  Paruroctonus.  Occasional 
Papers^Calif.  Acad.  Sci.,  No.  94. 


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TABLE  OF  CONTENTS 

Feeding  ecology  of  Gila  boraxobius  (Osteichthyes:  Cyprinidae)  endemic  to  a  thermal 

lake  in  southeastern  Oregon.  Jack  E.  Williams  and  Cynthia  D.  Williams  101 

First  record  of  the  pallid  hat  (Antrozous  palliclus)  from  Montana.  Jeff  Shryer  and 

Dennis  L.  Flath '. 115 

A  Cliiivcantliiuiti  spider  bite.  Dorald  M.  ,\llred  116 

Identity  of  narrow-leaved  Chitisotliamnits  visciclifloni.s  (Asteraceae).  Loran  C.  .\nder- 


ll'i 


Ribulose  diphosphate  carboxylase  activities  in  cold-resistant  common  mallow,  Malva 
neglecta  Wallr.  and  a  cold-sensitive  tomato,  Lycopersicon  esculentum  L.,  Ace 
55  var.  William  R.  Andersen  and  Jack  D.  Brotherson 121 

Recovery  of  Gambel  oak  after  fire  in  central  Utah.  L.  M.  Kunzler  and  K.  T.  Harper  ..        127 

Relationships  among  total  dissolved  solids,  conductivity,  and  osmosity  for  five  A;- 

temia  habitats  (Anostraca:  Artemiidae).  Nicholas  C.  Collins  and  Gray  Stirling         1.31 

Spawning  of  the  least  chub  (loticJtthijs  phlegt'thuntis).  Thomas  M.  Baiigh  1.39 

Transferrin  polymorphism  in  bighorn  sheep,  Otis  canaclcn.^is,  in  Colorado.  Patrick 

W.  Roberts,  Donald  J.  Nash,  and  Robert  E.  Keiss 141 

The  genus  Eriogoniim  Michx.  (Polygonaceae)  and  Michel  Gandoger.  James  L.  Reveal         143 

Parasites  from  two  species  of  suckers  (Catostomidae)  from  southern  Utah.  J.  Craig 

Brienholt  and  Richard  A.  Heckmann 149 

Soil  water  withdrawal  and  root  distribution  under  grubbed,  sprayed,  and  undis- 
turbed big  sagebni.sh  vegetation.  David  L.  Sturges 157 

Swarming  of  the  western  harvester  ant,   Pogononnjnnex  occiclentalis.    Dorald   M. 

Allred 165 

Relationship  between  environmental  and  vegetational  parameters  for  understory  and 
open-area  communities.  William  E.  Evenson,  Jack  D.  Brotherson,  and  Rich- 
ard B.  Wilcox  167 

Seasonal  activity  pattern  of  Columbian  ground  sqviirrels  in  the  Idaho  primitive  area. 

Charles  L.  Elliott  and  Jerran  T.  Flinders  175 

Habitat  and  plant  distributions  in  hanging  gardens  of  the  narrows,  Zion  National 

Park,  Utah.  George  P.  Malanson  178 

Short-term  effects  of  logging  on  red-backed  voles  and  deer  mice.  Thomas  M.  Camp- 
bell III  and  Tim  W.  Clark  183 

Terminal  bud  formation  in  limber  pine.  Ronald  M.  Lanner  and  James  A.  Bryan 190 

Stinger  utilization  and  predation  in  the  scorpion  Paiuwctoniis  boreits.  Bruce  S.  Gush- 
ing and  -•Vnne  Matherne 193 


HE  GREAT  BASIN  NATURALIST 


lume  40  No.  3 


September  30,  1980 


Brigham  Young  University 

MUS.   COMP.   ZOOl_. 

MAY  14'^ ' 

HARVAl-vO 
UMfVERSI-TY 


GREAT  BASIN  NATURALIST 

Editor.  Stephen  L.  Wood,  Department  of  Zoology,  Brigham  Young  University,  Provo,  Utah 

84602. 
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tural Sciences;  Ernest  L.  Olson,  Director,  Brigham  Young  University  Press,  University 
Editor. 

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ISSN  0017-3614 


The  Great  Basin  Naturalist 


Published  at  Provo,  Utah,  by 
Brigham  Young  University 

ISSN  0017-3614 


Volume  40 


September  30,  1980 


No.  3 


SPATIOTEMPORAL  VARIATION  IN  PHENOLOGY  AND  ABUNDANCE  OF 
FLORAL  RESOURCES  ON  SHORTGRASS  PRAIRIE 

V.  J.  Tepedino''^  and  N.  L.  Stanton^ 

Abstr.\c:t.—  Phenolog\-  and  abinidance  of  the  floral  resources  used  by  bees  and  other  flower-visiting  insects  were 
recorded  weekly  from  permanent  quadrats  for  two  years  on  two  shortgrass  prairie  sites  in  the  Laramie  Basin.  Wyom- 
ing. Each  site  was  composed  of  three  distinct  plant  communities.  Residts  suggest  considerable  spatiotemporal  varia- 
bilit\  in  floral  resources.  Specifically,  most  species  at  both  sites  showed  the  following  temporal  variation:  (1)  bloom 
times  between  one  and  two  weeks  earlier  in  the  second  year;  (2)  differences  of  approximately  1-2  weeks  in  bloom 
span  between  years;  (3)  substantial  differences  in  abundance  of  flowers  between  years.  Species  blooming  during  the 
last  half  of  the  census  period  were  significantly  more  variable  in  flower  abundance  between  years  than  those  bloom- 
ing earlier.  Spatial  variation  was  shown  both  by  differences  between  sites  and  between  plant  communities  within 
sites  in  the  direction  and  magnitude  of  between-year  changes  in  floral  abundance.  For  example,  floral  production  at 
Boulder  Ridge  in  1976  was  much  lower  than  in  1975,  but  at  The  Dirt  Farm  between-year  differences  were  less  pro- 
nounced and  depended  upon  season.  Similarity  measures  and  cluster  analysis  .suggest  differences  in  the  structure  of 
the  bloom  season  at  both  sites  between  years,  and  a  relatively  rapid  turnover  of  floral  composition  within  years  such 
that  bees  face  a  very  different  flora  over  the  latter  part  of  their  flight  seasons  relative  to  that  encountered  initially. 
Evidence  from  other  reports  support  the  hypothesis  of  spatiotemporal  unpredictability  of  floral  resources. 


Spatial  and  temporal  heterogeneity  in  re- 
source supply  is  receiving  increasing  atten- 
tion in  ecological  hypotheses  concerning 
both  species  diversity  and  consumer  foraging 
strategies  (see  reviews  in  Wiens  1976,  Levin 
1976).  An  important  component  of  such  re- 
source heterogeneity  is  its  predictability 
(Wiens  1976).  When  resources  are  unpredic- 
table in  time  or  space,  generalists  are  hypoth- 
esized to  dominate  the  consumer  faima  be- 
cause they  are  less  vulnerable  than  specialists 
to  fluctuations  in  the  supply  of  particular  re- 
sources (Pianka  1966,  1970,  Levin  1968, 
Cody  1974,  Moldenke  1975).  Though  this  hy- 
pothesis is  intuitively  attractive,  existent  evi- 
dence is  equivocal  (Futuvma  1976,  Rabenold 
1978). 

In  addition,  the  resource  predictability  hy- 
pothesis has  been  extended  to  explain  latitu- 
dinal diversity  gradients  by  proposing  that 
high  tropical  diversity  is  the  result  of  clo.se 


species  packing  of  specialists  in  a  predictable 
environment  (Pianka  1966,  1970).  However, 
the  assertion  of  large  differences  in  predict- 
ability between  temperate  and  tropical  re- 
gions has  recentlv  been  challenged  bv  Wolda 
(1978),  who  has  shown  that  precipitation  pat- 
terns appear  no  less  variable  in  the  tropics 
and  that  insect  populations  in  the  seasonal 
tropics  fluctuate  as  much  as  their  temperate 
counterparts. 

The  impression  that  high  .spatiotemporal 
variability  in  resources  is  more  common  in 
certain  regions  is  primarily  intuitive  or  rests 
on  anecdotal  evidence;  convincing  documen- 
tation is  lacking  (Ricklefs  1973).  Clearly,  if 
resource  predictability  and  .spatiotemporal 
heterogeneity  are  to  play  more  than  a  hypo- 
thetical role  in  ecology,  more  quantitative 
field  studies  are  needed  that  measure  changes 
in  resource  levels  and  their  use  across  space 
and  time. 


Department  of  Zoology  and  Physiolog)'.  University  Station  Box  3166,  University  of  Wyoming,  Laramie,  Wyoming  82071. 

Present  address:  USD.VSE.\-Ali-\VR,  Bee  Biology-  and  Systematics  Laboratory,  Utah  Stale  University,  UMC  53,  Logan,  Utah  84322. 


197 


198 


Great  Basin  Naturalist 


Vol.  40,  No.  3 


For  many  consumer  guilds,  it  is  difficult  to 
distinguish  between  what  is  and  what  is  not  a 
resource  (Haigh  and  Maynard  Smith  1972). 
In  contrast,  flowers,  the  trophic  resources  of 
bees  and  other  animals,  are  easy  to  delimit 
and  quantify.  In  this  report  we  present  two 
years'  data  on  variation  in  phenology  and 
abimdance  of  entomophilous  flowers  on  two 
shortgrass  prairie  sites  in  SE  Wyoming.  Fu- 
ture papers  will  relate  these  data  to  the  struc- 
ture of  the  bee  community  (Tepedino  and 
Stanton,  in  preparation). 

The  Great  Plains  are  subject  to  the  wide 
and  unpredictable  fluctuations  in  temper- 
ature and  moisture  availability  which  typify 
interior,  temperate  climates.  Shortgrass 
prairie  experiences  the  most  unpredictable 
fluctuations  in  precipitation  of  all  North 
American  grassland  biomes  (Wiens  1974). 
For  example,  two  of  every  five  years  may  be 
expected  to  deviate  by  a  minimum  of  25  per- 
cent from  mean  yearly  precipitation  and  one 
of  12  deviates  by  at  least  50  percent;  this  var- 
iation is  temporally  impredictable  (Wiens 
1974).  In  addition,  precipitation  frequently 
occurs  in  localized  patches  and  is  spatially 
unpredictable  (Coupland  1958). 

Low  climatic  predictability  prompted  the 
following  hypotheses  for  flower  production: 

1.  Floral  phenology  and  abundance  exhibit 
wide  year-to-year  variations  at  a  given 
site,  both  at  the  level  of  the  species  and, 
more  generally,  for  the  whole  commu- 
nity. 

2.  If  climatic  variation  is  spatially  local- 
ized, then  within-year  differences  be- 
tween sites  in  floral  phenology  and 
abundance  should  be  evident. 

3.  Between-year  variability  is  modified  by 
seasonal  effects.  In  particular,  the 
spring  flora  exhibits  greater  year-to- 
year  variability  than  the  summer  flora 
(Leopold  and  Jones  1947). 

4.  Within-year  predictability,  i.e.,  the 
probability  of  encountering  similar  flor- 
al composition  for  several  consecutive 
weeks,  is  greatest  during  the  summer 
blooming  season. 

Methods  and  Study  Sites 

Study  Sites 

The  study  was  conducted  in  the  southern 
part  of  the  Laramie  Basin,  a  semiarid  inter- 


mountain  valley  in  Albany  County,  Wyom- 
ing. In  general,  soils  are  shallow,  rocky,  and 
poorly  developed.  Precipitation  in  nearby 
Laramie  averages  25.6  cm  per  year,  with  70 
percent  falling  from  April  through  Septem- 
ber. The  growing  season  is  short,  varying  be- 
tween 80  to  100  days,  with  killing  frosts  com- 
mon in  June  and  early  September.  With  few 
exceptions,  the  flora  is  composed  of  perennial 
species. 

The  Dirt  Farm 

Located  approximately  16  km  southeast  of 
Laramie,  The  Dirt  Farm  site  is  1.6  ha  in  area 
at  an  altitude  of  2250  m.  The  vegetation  is 
divided  into  three  contiguous  communities.  A 
cushion  plant  commimity  covers  0.77  ha  and 
is  located  on  windswept  shallow  soils  with 
frequent  bedrock  exposure.  Abundant  species 
include  Phlox  bryoides.  Astragalus  serico- 
leucus,  A.  spatidatiis,  Arenaria  Jiookeri,  and 
Paronychia  sessiliflora,  all  of  the  caespitose, 
herbaceous  growth  form.  Adjoining  the  cush- 
ion plant  community  is  a  level  area  of  0.33 
ha  dominated  by  the  shrub,  Cercocarpus 
montanus,  mountain  mahogany.  Associated 
species  include  Allium  textile,  A.  cernuum, 
and  the  half-shrub  Chrysothamnus  vis- 
cidiflorus.  Soils  are  very  shallow  or  nonex- 
istent here  also,  with  the  roots  of  the  shrubs 
penetrating  cracks  in  the  bedrock.  Extending 
south  from  the  mountain  mahogany  commu- 
nity is  a  0.5  ha  section  of  typical  shortgrass 
prairie.  Soils  are  deeper  here  than  in  the 
other  two  communities. 

Boulder  Ridge 

The  Boulder  Ridge  site  covers  1  ha  and  is 
located  approximately  38  km  southwest  of 
Laramie  (22  km  southwest  of  The  Dirt  Farm) 
at  an  altitude  of  2425  m.  The  vegetation  is 
foothill  scrub  (Porter  1962)  and  is  divided 
into  three  communities.  A  central  section  of 
0.35  ha  is  dominated  by  the  shrub  Cerco- 
carpus montanus  with  a  few  individuals  of 
the  shrubs  Primus  virginiana,  Amelanchier  al- 
nifolia,  and  Ribes  cereum.  Abundant  associ- 
ated herbs  are  Allium  textile,  A.  geyeri,  A. 
cernuum,  Ranunctdus  ranuncuUnus,  and  Ce- 
rastium  arvense.  The  western  part  of  the 
communitv   is  level,  but  the  eastern  end 


September  1980 


Tepedino,  Stanton:  Bee  Ecology 


199 


slopes  at  an  angle  of  approximately  30  de- 
grees. At  slope  bottom  movmtain  mahogany 
gives  Way  to  a  community  of  0.28  ha  domi- 
nated by  sagebrush  {Arteinisia  tridentato). 
The  most  abimdant  associates  are  Castilleja 
flava,  Astmoalus  flexuosiis,  CoUinsia  parii- 
folia,  Orthoccirpus  luteus,  and  Chrysopsis  vil- 
losa.  Bordering  the  study  site  at  its  eastern 
and  western  ends  is  a  heterogeneous  commu- 
nity of  0.37  ha  that  includes  representatives 
of  the  cushion  plant,  shortgrass  prairie,  and 
sagebrush  communities.  Nomenclature  is  that 
of  Harrington  (1954),  Porter  (1965),  and  We- 
ber (1967). 

Methods 

Censusing.—  Floral  phenology  and  abun- 
dance of  species  with  entomophilous  flowers 
were  estimated  for  two  years  at  each  site  by 
weekly  censuses  of  the  number  of  flowers  by 
species  in  permanent  m^  quadrats.  Censusing 
extended  from  the  last  week  in  May  through 
the  last  week  of  August,  except  for  the  Boul- 
der Ridge  site  in  1975,  which  began  one 
week  later  because  of  a  spring  snow  storm. 
Quadrats  were  chosen  by  using  a  stratified 
random  sampling  technique  to  insure  that 
each  community  was  sampled  in  proportion 
to  its  percentage  of  the  entire  study  area.  Ap- 
proximately 1.6  percent  of  the  total  area  of 
each  site  was  sampled.  Individual  flowers 
were  counted  in  all  cases  except  for  most 
Compositae,  where  heads  were  counted,  and 
the  Umbelliferae  and  Polygonaceae,  where 
umbels  were  counted. 


Analysis.—  To  avoid  the  assumptions  of 
normality  and  homoscedasticity,  non- 
parametric  statistics  were  used.  The  Sign 
Test  (Conover  1971)  was  used  to  test  for  be- 
tween-year  differences  in  total  floral  abun- 
dance at  each  site  by  comparing  the  total 
number  of  flowers  in  each  permanent  quad- 
rat for  each  set  of  paired  sampling  dates 
(Table  1).  Comparisons  were  made  for  each 
site  as  a  whole  and  by  vegetation  tvpe. 

To  provide  a  measure  of  the  similarity  be- 
tween sampling  dates  both  within  and  be- 
tween years  for  each  site  we  used  the  Czeka- 
nowski  measure  (also  known  as  the  Brav- 
Curtis  Index:  Goodall  1973)  to  generate  sim- 
ilarity matrices,  which  were  then  subjected 
to  cluster  analysis.  The  Czekanowski  Index  is 
written 

PS  =  22  min  (x.y.)/    2  (x.  +  y.), 

1=1  i=  1 

where  Xi  and  y^  are  the  number  of  flowers  of 
species  i  on  dates  x  and  y. 

The  matrices  were  analyzed  by  single, 
complete,  and  average  linkage  clustering  al- 
gorithms using  the  BMDP  computer  package 
(Dixon  1975)  and  the  best  grouping  method, 
decided  by  calculating  the  cophenetic  corre- 
lation coefficient  of  Sokal  and  Rohlf  (Sneath 
and  Sokal  1973).  The  average  linkage  method 
provided  the  best  results,  and  only  these  are 
reported.  Matrices  for  each  site  for  all  census 
date  comparisons  as  well  as  separate  within- 
site,  within-year  matrices  were  clustered. 
Only  the  four  within-site,  within-year  clusters 
are  reported  here  because  we  consider  them 
most  informative. 


Table  1.  Actual  floral  census  dates  and  their  corresponding  census  code  numbers  for  each  study  site. 


Census 

Dirt  Farm 

Boulder 

Ridge 

Code 

1975 

1976 

197,5 

1976 

1 

26  May 

24  Mav 

No  census 

27  May 

2 

2  June 

31  May 

5  June 

3  June 

3 

9  June 

7  June 

13  June 

10  June 

4 

16  June 

16  June 

20  Jiuie 

18  June 

5 

23  June 

21  June 

27  June 

24  June 

6 

30  June 

28  June 

4  July 

1  July 

7 

7  July 

6  Julv 

11  July 

7  Julv 

8 

14  Julv 

12  Julv 

18  July 

18  Julv 

9 

21  Julv 

21  Julv 

25  July 

25  Julv 

10 

28  July 

28  July 

31  July 

31  July 

11 

6  August 

5  August 

7  August 

8  .\ugiist 

12 

13  August 

11  August 

14  August 

14  August 

13 

19  August 

18  August 

23  August 

19  August 

14 

26  August 

28  August 

29  August 

28  August 

200 


Great  Basin  Naturalist 


Vol.  40,  No.  3 


Results 

Precipitation  and  temperature.—  Precipi- 
tation patterns  over  the  two  years  of  study 
were  quite  different  (Data  from  weather  sta- 
tion at  Laramie  Airport).  Rainfall  in  spring 
and  early  summer  1975  was  much  heavier 
than  normal  (Fig.  la),  and  total  precipitation 
for  the  year  was  6.1  percent  above  normal. 
Conversely,  1976  was  a  dry  year  with  below 
normal  rainfall  for  every  month  from  March 
through  June.  By  the  end  of  June  precipi- 
tation was  32.0  percent  below  normal.  July 
and  August  received  greater  than  average 
rainfall,  and  by  the  end  of  the  study  precipi- 
tation was  only  17.0  percent  below  normal 
for  the  January  through  August  1976  period. 

In  general,  temperatures  were  warmer  in 
1976  (Fig.  lb).  In  particular  1976  was  warm- 
er from  April  through  July,  a  period  which 
was  also  (July  excluded)  much  drier  than  nor- 
mal (Fig.  la).  The  frost-free  period  in  1975 
extended  from  16  June  to  5  September  (79 
days)  and  in  1976  from  25  June  to  termi- 
nation of  the  study  (29  August)  (63  days). 


JFMAMJJASOND 


6- 

I 

a 

E,. 

u 

c 

« 

o"- 

, 

/\ 

— 

,\ 

1 

x^\ 

.«- 

0] 

/  ^' 

, 

J 

/"'/  ''\ 

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Q. 

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K^ 

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/  ' \  / 

I     /\ 

/ 

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■  -i 

1 

"7 

/i  \j 

~^v'V> 

y^ 

/ 

1975 


MAM 

1976 


Fig.  1.  Monthly  precipitation  (a)  and  temperature  (b) 
records  from  the  Laramie  .\irport  (Brees  Field);  a)  solid 
line  =  1975,  1976,  dashed  line  =  normal;  b)  solid  line 
=  1975,  dashed  line  =  1976. 


Species  Composition.—  We  recorded  63 
and  73  entomophilous  plant  species  during 
the  two  years  at  The  Dirt  Farm  and  Boulder 
Ridge,  respectively  (Appendix  A).  Total  flow- 
ers by  species  and  year  are  also  shown  in  Ap- 
pendix A.  The  family  Compositae  was  repre- 
sented by  the  largest  number  of  species  at 
both  sites,  followed  by  the  Cruciferae  at  The 
Dirt  Farm  and  the  Scrophulariaceae  at  Boul- 
der Ridge. 

Dirt  Farm 

Phenology.— 'The  flowering  phenology  of 
selected  species  is  shown  in  Figure  2.  Phe- 
nological  variation  between  years  took  two 
forms:  differences  in  first  bloom  and  in  bloom 
span.  Evidence  for  variability  in  first  bloom 
comes  from  several  sources.  First,  when  total 
floral  abundance  is  graphed  by  date  for  each 
year,  it  is  clear  that  both  June  and  August 
peaks  were  advanced  in  1976  (Fig.  3).  It  is 
worthwhile  to  note,  however,  that  the  ad- 
vancement in  each  peak  is  not  due  to  similar 
responses  in  each  vegetation  type.  The  early 
peak  is  advanced  due  to  responses  of  the 
cushion  plant  and  mountain  mahogany  com- 
munities, and  the  shortgrass  and  mountain 
mahogany  communities  account  for  advances 
in  the  late  peak  (Fig.  3).  Earlier  flowering  in 
1976  was  due  to  a  warmer  spring  and  sum- 
mer relative  to  1975. 

Phenological  advancement  in  1976  is  also 
seen  when  first  bloom  dates  are  compared  by 
species.  Analysis  shows  that  27  of  38  species 
differed  by  at  least  one  week  in  anthesis. 
Eleven  species  were  excluded  because  they 
were  in  bloom  when  censusing  began,  and  14 
others  were  eliminated  because  they  flower- 
ed in  only  one  year.  Of  the  27  species  that 
differed  in  phenology,  24  were  earlier  by  an 
average  of  10  days  in  1976  (X2  =  16.33, 
P<0.001). 

Are  there  seasonal  differences  between 
early-  and  late-blooming  species  in  phenolo- 
gical predictability?  lo  test  this  the  census 
period  was  halved  and  species  grouped  ac- 
cording to  the  half  in  which  they  began 
blooming  (Fig.  3).  Twelve  of  18  first-half  spe- 
cies showed  a  mean  difference  of  one  week  in 
beginning  bloom,  and  15  of  20  second-half 
species  showed  mean  advance  of  12.1  days. 
Species  blooming  during  the  last  half  of  the 


September  1980 


Tepedino,  Stanton:  Bee  Ecology 


201 


1 
2 
3 
4 
5 
6 
7 
8 
9 
10 
11 
12 
13 
14 
15 
16 
17 
18 
19 
CO  20 
021 
U22 
d)  23 
CL24 
00  25 
26 
27 
28 
29 
30 
31 
32 
33 
34 
35 
36 
37 
38 
39 
40 
41 
42 


Census  week 
^456789 


10  11 


12 


13  14 
-J I 


Fig.  2.  Bloom  spans  of  selected  species  at  The  Dirt  Farm  for  1975  (solid)  and  1976  (dashed);  stars  =  did  not  flow- 
.  Clensiis  date  numbers  refer  to  Table  1.  species  numbers  to  Appendix  A. 


202 


Great  Basin  Naturalist 


Vol.  40,  No.  3 


MOUNTAIN  MAHOGANY 


SHORTGRASS 


200r 


120  - 


en 

UJ 
Q 


tr    lOOr 

UJ 
CD 

:^ 

3 


I    2   3   4  5   6   7  8  9  10  II  12  13  14 
JUNE h-  JULY  -\ AUG 


2   3  4  5    6  7  8    9  10  II  12  1314 
JUNE  — +-  JULY  H —  AUG 


CENSUS    WEEK 


Fig.  3.  Total  floral  abundance  per  ni^,  irrespective  of  species,  at  The  Dirt  Farm  for  each  census  date  in  1975  and 
1976.  Census  date  numbers  refer  to  Table  1.  Asterisks  mark  significant  differences  (P<.05)  between  years  for  paired 
dates. 


September  1980 


Tepedino,  Stanton:  Bee  Ecology 


203 


season  showed  significantly  greater  phenolo- 
gical  differences  than  those  blooming  during 
the  first  half  (Mann-Whitney  U-Test, 
P<0.05). 

Between-year  comparisons  of  bloom  spans 
are  a  second  indicator  of  phenological  varia- 
bility. Differences  of  at  least  one  week  in 
bloom  span  were  shown  by  22  of  38  species. 
Average  difference  in  bloom  span  for  the  22 
species  was  two  weeks  with  a  range  of  one  to 
seven  weeks.  Ten  species  had  longer  spans  in 
1975  and  12  had  longer  spans  in  1976.  A 
comparison  by  seasonal  grouping  of  bloom 
span  data  into  early  and  later  blooming  spe- 
cies shows  that  longer  bloom  spans  during 
the  last  half  of  the  season  occurred  mostly  in 
1976,  and  1975  had  more  longer  blooming 
species  during  the  first  half  (X^  =  2.76,  P  = 
0.097). 

Floral  Abundance.—  Differences  in  floral 
abundance  between  years  is  first  shown  by 

DIRT  FARM 


comparing  total  number  of  flowers  by  date 
(Fig.  3).  Total  abundance  comparisons  show 
the  early  peak  to  be  higher  but  the  late  peak 
lower  in  1975.  The  late  peak  difference  is 
due  largely  to  profuse  flowering  of  Eriogo- 
nitm  effusum.  Exclusion  of  this  species  re- 
sults in  much  closer  agreement  of  abundances 
from  late  July  to  mid-August. 

When  total  floral  abundance  is  partitioned 
into  component  communities  it  is  again  evi- 
dent that  there  is  no  typical,  overall  site  re- 
sponse (Fig.  3).  For  example,  the  mountain 
mahogany  community  shows  nine  significant 
between-year  differences  in  abundance,  with 
1976  having  more  flowers  on  eight  dates.  In 
contrast,  between-year  differences  in  the 
cushion  plant  community  show  1975  with 
more  flowers  for  seven  or  nine  significant 
comparisons  and  the  shortgrass  community 
with  more  flowers  in  1975  for  five  of  six 
dates.  A  cold  period  during  the  week  begin- 

BOULDER  RIDGE 


60r 


50  - 


40  - 


iij 

o     30h 

UJ 
Q. 


20  ■ 


10  - 


0 


z 


< 

X 

a: 


< 

X 


o 

o 
llJ 


r 


1.00-       2.00-     >3.00 
1.99  2.99 


< 

X 


o 
o 

UJ 

(A) 


F 


< 

X 


CO 


T 


1.00-       2.00-      >l3.00 
1.99  2.99 


RHL 


Fig.  4.  Frequency  distril)iition  of  the  ratio  of  the  number  of  flowers  in  the  most  abundant  year  to  the  number  of 
flowers  in  the  least  abundant  year  for  each  species  (RHL)  at  both  sites. 


204 


Great  Basin  Naturalist 


Vol.  40,  No.  3 


ning  16  Jiine  1976  reduced  floral  production 
appreciably  on  the  mountain  mahogany  com- 
munity and  probably  affected  the  shortgrass 
community  as  well.  During  this  period  tem- 
peratures were  below  freezing  for  three 
nights  and  snow  and  sleet  fell  twice.  The  ef- 
fect of  such  periods  upon  organisms  in  the 
Rocky  Mountains  has  been  described  by 
Ehrlichetal.  (1972). 

Floral  abundance  was  also  compared  by 
species  between  years.  An  expression  of  dif- 
ferences in  abundance  is  the  ratio  of  total 
number  of  flowers  observed  in  the  year  of 
highest  production  divided  by  total  number 
of  flowers  in  the  year  of  lowest  production 
(RHL).  We  eliminated  from  this  analysis 
those  species  that  either  flowered  in  only  one 
year  or  failed  to  produce  at  least  50  flowers 
in  either  vear.  For  the  remaining  44  species 
mean  RHL  =  5.39  (SD  =  8.56,  range  = 
1.04-48.60),  suggesting  that  the  abundance  of 
flowers  of  most  species  may  show  significant 
bet ween-y ear- variation.  The  data  are  shown 
as  histograms  in  Figure  4.  Of  the  44  species, 
26  were  more  abundant  in  1975  and  18  were 


more  abundant  in  1976.  Thus,  no  year  effect 
was  observed. 

Do  RHL  values  display  a  seasonal  pattern? 
Again,  species  were  grouped  according  to  the 
half  of  the  season  in  which  they  began  flow- 
ering and  the  Mann- Whitney  U-Test  used  to 
test  for  differences  in  RHL  between  groups. 
The  comparisons  show  that  plants  blooming 
during  the  latter  half  of  the  season  were  sig- 
nificantly more  variable  (P  =  0.05),  i.e.,  had 
larger  RHL  values  than  those  blooming  ear- 
lier. No  year  effect  was  evident  since  both 
years  had  an  almost  equal  proportion  of  spe- 
cies with  highest  RHL  values  in  each  half  of 
the  year. 

Similarity  and  Cluster  Analysis.—  We  used 
cluster  analysis  to  elucidate  differences  be- 
tween census  dates  within  and  between  years 
and  to  illustrate  seasonal  groupings.  High 
similarity  values  for  paired  between-year 
sampling  dates  were  expected;  however,  the 
data  do  not  support  this  hypothesis.  Mean 
similarity  for  paired  census  dates  was  only 
0.519  (SD  =  0.196,  range  0.204-0.854). 
Mean  similarity  was  highest  (0.593  [SD    = 


DIRT  FARM 


BOULDER    RIDGE 


0.8  r 


23456789  10  II 
I JUNE 1—  JULY  H 


0.8  r 

0.7 

0.6 

0.5 

0.4 

0.3 

0.2 

0.1 

0.0 


/ 

/ 

/|976 


\ 


J I I L 


J I I I I 


23456789  10 
JUNE \—  JULY  — I 


CENSUS    WEEK 


Fig.  5.  Czekanowski  similarity  measures  between  floral  ahiindance  for  each  census  date  and  the  third  subsequent 
week  at  both  sites.  Census  date  numbers  refer  to  Table  I. 


September  1980 


Tepedino,  Stanton:  Bee  Ecology 


205 


0.149,  range  =  0.265-0.772])  when  each 
1975  date  was  compared  with  the  week  prior 
to  that  date  in  1976.  Thus,  the  similarity  data 
provide  additional  evidence  for  phenological 
advancement  in  1976. 

Within-year  similarity  comparisons  were 
also  quite  variable.  We  reasoned  that,  since 
bees  are  the  predominant  pollinators  on 
shortgrass  prairie  and  because  females  of 
most  species  of  solitary  bees  fly  for  a  mini- 
mum of  four  weeks  (Linsley  1958),  a  conserv- 
ative estimate  of  within-year  resource  varia- 
bility for  a  bee  would  be  the  similarity 
between  the  week  of  emergence  and  three 
weeks  later.  This  measure  is  conservative  be- 
cause we  used  similarity  measures  between 
dates  that  are  one  week  less  than  the  mini- 
mum flight  span.  In  Figure  5  we  graph  the 
results  for  each  year.  Except  for  the  last 
month  of  the  census  period,  values  are  very 
low.  Any  species  emerging  during  the  first 
two-thirds  of  the  blooming  season  would  face 
a  very  different  flora  during  the  latter  part  of 
its  flight  season  relative  to  that  encountered 
initially. 

Cluster  analysis  aids  in  depicting  seasonal 
groupings  and  transitional  periods  within  the 
blooming  season  (Fig.  6).  The  number  of  clus- 


ters formed  at  a  value  of  .50  differs  between 
years  as  does  the  number  of  unclustered 
dates,  suggesting  that  the  "structure"  of  the 
blooming  season  may  differ  from  year  to 
year.  For  example,  five  clusters  plus  one  un- 
clustered date  form  in  1975,  but  four  clusters 
and  four  unclustered  dates  form  in  1976. 

Within  both  years  late  season  dates  cluster 
strongly,  again  indicating  higher  within-year 
predictability  for  late  summer  bees.  Almost 
all  other  clusters  are  composed  of  only  two 
consecutive  census  dates,  indicating  a  high 
rate  of  turnover  in  floral  composition  for  the 
first  two-thirds  of  the  blooming  sason. 

Boulder  Ridge 

Phenohgij.—  Flowering  also  began  earlier 
in  1976  at  Boulder  Ridge  (Fig.  7).^  Of  the  73 
species  censused,  33  were  either  in  bloom 
when  censusing  began  of  flowered  in  only 
one  year  and  were  eliminated  from  this  anal- 
ysis. Of  the  remaining  40  species,  32  showed 
phenological  differences  of  at  least  one  week. 
Twenty-six  of  the  32  species  were  earlier  an 
average  of  10  days  in  1976  (X^  =  12.50, 
P<  0.001).  The  seasonal  differences  in  ad- 
vancement   between    first-    and    second-half 


DIRT   FARM 


0.0  r 
0. 

0.2 
0.3 

>   0.4 

q: 

<    0.5 

1 

'^    0.6 

0.7 

0.8 

0.9 

1.0 


1975 


1976 


a 


I     2    3   4   5    6    7    8    9  10   II   12  13  14 
I JUNE 1—  JULY  —I AUG 


I     2    3    4   5    6    7   8    9   10  II    12  13  14 
I JUNE (—  JULY  — I AUG 


CENSUS    WEEK 


Fig.  6.   Dendrograms  of  floral  similarity  between  census  dates  for  each  year  at  The  Dirt  Farm.  Census  date  niim- 
hers  refer  to  Table  1 . 


206 


Great  Basin  Naturalist 


Vol.  40,  No.  3 


CO 

a; 
u 

(D 
Cl 
00 


Census  week 


1 

2 
3 

4 
5 
6 

7 
8 
9 
10 
11 
12 
13 
14 
15 
16 
17 
18 
19 
20 
21 
22 
23 
24 
25 
26 
27 
28 
29 
30 
31 
32 
33 
34 
35 


3 

I 


4    5    6 


7 

I 


8 


9 

I 


10  11 

_J 1_ 


12  13  14 
_i I I 


Fig.  7.  Bloom  spans  of  selected  species  at  Boulder  Ridge  for  1975  (solid)  and  1976  (dashed);  stars  =  did  not  flower. 
Census  date  numbers  refer  to  Table  1,  species  numbers  to  Appendix  A. 


species  observed  at  The  Dirt  Farm  were  not 
detected  here. 

Most  species  differed  in  bloom  span  be- 
tween years.  After  eliminating  species  that 
began  blooming  prior  to  censusing,  29  of  the 
remaining  40  differed  by  at  least  one  week. 
Average  difference  in  bloom  span  for  the  29 
species  was  17  days  with  a  range  of  one  to 


five  weeks.  Chi-square  analyses  for  sea.sonal 
patterns  were  insignificant.  Year  effects, 
however,  were  present;  longer  bloom  spans 
were  concentrated  in  1975  (20  of  29,  X2  = 
4.17,F<0.05). 

Abundance.—  Differences  in  total  floral 
abundance  between  years  were  remarkable 
(Fig.    8).    Twelve   of   the    13   comparisons 


September  1980 


Tepedino,  Stanton:  Bee  Ecology 


207 


showeci  significantly  greater  abundance  in 
1975.  There  was  a  marked  midyear  peak  in 
1975  that  is  only  suggested  in  1976. 

In  contrast  to  The  Dirt  Farm,  floral  abun- 
dance for  all  vegetation  types  at  Boulder 
Ridge  was  higher  in  1975  (Fig.  8).  Between- 
year  differences  on  the  heterogeneous  area 
were  primarily  responsible  for  the  total  flow- 
er differences.  The  last  10  dates  showed  sig- 
nificantly more  flowers  on  this  section  in 
1975  (Fig.  8).  The  second  peak  in  the  total 
abundance  curve  in  1975  (Fig.  8),  absent  in 


1976,  was  due  mainly  to  the  heterogeneous 
area.  Differences  observed  on  the  other  sec- 
tions showed  floral  abundances  in  1976  to  be 
consistently  below  those  of  1975,  although 
many  of  these  comparisons  were  not  signifi- 
cant. 

The  RHL  ratio  was  used  to  compare  abun- 
dance by  species  between  years.  We  elimi- 
nated all  species  with  less  than  50  flowers  in 
either  years  and  those  that  flowered  in  only 
one  year.  For  the  remaining  38  species,  mean 
RHL     =     5.47    (SD     =     6.56,    range     = 


BOULDER   RIDGE 


c/) 
q: 

UJ 

o 


UJ 
CD 


60 

50 

40 

30 

20 

10 

0 
I     2    3  4    5    6    7  8   9    10  II    12  13  14 

28i-   TOTAL 
24  h 


' 

HETERO- 
GENEOUS 

r- 

,1975 

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A 

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1976 

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1111 

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1     1     1 

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8  - 
4  - 


0 


\^\ 


.1976 


--      \ 


MOUNTAIN   MAHOGANY 


J I L 


J. 


J_J l_l L-J I 


20 

0 

0 

28 

24 

20 

16 

12 

8 

4 

0 


2    3   4   5    6    7    8    9   10  II    12  13  14 
SAGEBRUSH 


2    3   4   5    6   7   8   9   10  II   12  13  14         I    2    3   4   5  6   7    8   9  10  II   12  13  14 

|— JUNE      I         JULY— I AUG  |— JUNE  — | JULY 1 — AUG 

CENSUS    WEEK 

Fig.  8.  Total  abundance  per  m^,  irrespective  of  species,  at  Boulder  Ridge  for  each  census  date  in  1975  and  1976. 
Census  date  numbers  refer  to  Table  1.  Asterisks  mark  significant  differences  (P<.05  between  years  for  paired  dates. 


208 


Great  Basin  Naturalist 


Vol.  40,  No.  3 


1.07-27.50;  Fig.  4).  Again,  there  were  large 
between-year  differences  in  abundance  for 
most  species  on  the  site. 

We  tested  for  seasonal  differences  in  RHL 
values  using  the  same  methods  as  for  The 
Dirt  Farm.  Again,  RHL  values  for  the  latter 
part  of  the  blooming  season  are  significantly 
higher  (P<0.05).  However,  in  contrast  to 
The  Dirt  Farm,  each  year  is  not  equally  rep- 
resented by  species  with  high  RHL  values. 
Twenty-eight  of  the  38  species  were  more 
abundant  in  1975  (X^  =  8.53,  P<  0.005).  This 
phenomenon  is  particularly  striking  for  the 
latter  part  of  the  blooming  season,  when  only 
one  of  16  species  was  more  abundant  in  1976. 

Similarity  and  Cluster  Analysis.—  As  with 
The  Dirt  Farm  data,  the  expected  high  sim- 
ilarity for  paired  between-year  comparisons 
was  not  evident.  Mean  similarity  for  paired 
census  dates  was  only  0.424  (SD  =  0.200, 
range  =  0.184-0.896).  Again,  highest  mean 
similarity  was  between  1975  dates  and  the 
week  prior  to  the  identical  date  in  1976  (x  = 
0.518,  SD  =  0.161,  range  =  0.309-0.869). 

Within-year  similarities  [between  each 
census  date  and  the  census  taken  three  weeks 
later]  were  as  low  as  at  The  Dirt  Farm  (Fig. 
5).  The  graph  for  1975  is  uniformly  low  with- 
out the  appreciable  rise  la^te  in  the  season 


present  in  The  Dirt  Farm  and  Boulder  Ridge 
1976  data. 

Cluster  analysis  again  suggests  "structural" 
differences  between  the  blooming  seasons 
(Fig.  9).  Four  clusters  plus  three  unclustered 
dates  form  in  1975,  and  five  clusters  and 
three  unaffiliated  dates  are  found  in  1976.  As 
at  The  Dirt  Farm,  late  season  dates  cluster 
most  densely  and  all  but  one  other  cluster  is 
composed  of  only  two  consecutive  dates  at 
the  .50  level. 

Discussion 

Plant  species  varied  substantially  between 
years  in  the  onset  and  length  of  the  blooming 
period  and  in  the  number  of  flowers  pro- 
duced. Comparison  of  phenology  and  abun- 
dance, both  between  sites  and  among  vegeta- 
tion types  within  sites,  shows  other  important 
differences.  Though  phenological  advance- 
ment in  1976  was  a  uniform  occurrence  at 
both  sites,  changes  in  the  direction  and  mag- 
nitude of  floral  abundance  were  not.  A  com- 
parison of  total  floral  abundance  between 
The  Dirt  Farm  and  Boulder  Ridge  shows  that 
the  two  sites  behaved  quite  differently.  Floral 
production  at  Boulder  Ridge  in  1976  was 
consistently  well  below  that  of  1975.  In  con- 


0.0  - 
0.1  - 
0.2  - 
0.3  I- 

>-    0.4 

o: 

2    0.5  h 

^    0.6 

0.7 

0.8 

0.9 

1.0 


1975 


BOULDER    RIDGE 


1976 


Q 


2    3  4   5  6    7   8   9   10  II   12  13  14 
I—  JUNE  —I JULY \ —  AUG 


CENSUS    WEEK 


I    2    3   4  5    6    7  8   9   10  II   12  13  14 
I —  JUNE  —\ JULY 1 AUG 


Fij;.  9.  Dciidrograiii.s  of  floral  similarity  between  census  dates  for  each  year  at  Boulder  Ridge.  Census  date  num- 
bers refer  to  Table  1 . 


I 


September  1980 


Tepedino,  Stanton:  Bee  Ecology 


209 


trast,  floral  production  was  significantly  high- 
er at  The  Dirt  Farm  in  1976  over  the  latter 
part  of  the  blooming  season.  Evidently,  the 
spring  and  early  summer  drought  was  either 
more  severe  at  Boulder  Ridge  or  the  plant 
communities  at  Boulder  Ridge  were  more 
susceptible  than  those  at  The  Dirt  Farm. 

Between-site  differences  appear  due  to 
varying  responses  to  weather  by  each  vegeta- 
tion type  within  each  site.  At  Boulder  Ridge 
all  commmiities  displayed  consistently  lower 
floral  productivity  in  1976,  but  at  The  Dirt 
Farm  each  community  responded  indepen- 
dently. Indeed,  at  The  Dirt  Farm  each  spe- 
cies seemed  to  display  an  independent  re- 
sponse as  shown  by  the  lack  of  any  year 
effects  on  the  distribution  of  RHL  values  ei- 
ther in  between-year  comparisons  or  be- 
tween-season  comparisons.  These  observa- 
tions suggest  that  variation  in  floral 
production  is  expressed  as  spatially  localized 
patches  of  high  or  low  abundance  that 
change  from  year  to  year. 

Because  we  collected  data  for  only  two 
years,  it  is  necessary  to  ask  how  representa- 
tive of  routine  variability  these  results  are. 
Schemske  et  al.  (1978),  in  a  study  of  seven 
spring  herbs,  found  the  onset  of  flowering  to 
range,  by  species,  from  8  to  22  days  over 
three  years.  More  importantly,  peak  flower- 
ing did  not  usually  coincide  with  optimal 
pollinator  conditions.  Long-term  bloom  re- 
cords for  several  regions  in  North  America 
are  available  for  analysis.  In  several  cases 
data  are  available  for  period  of  up  to  30 
years  in  the  same  area  (Lindsey  and  Newman 
1956-Indiana;  Smith  1915-Ohio;  Hulbert 
1963-Kansas;  Hodson  1971-Minnesota).  In 
examining  these  data  we  have  used  varia- 
bility in  first  flowering  as  an  indication  of  re- 
source predictability  since  this  phenophase  is 
common  to  all  studies.  In  brief,  we  find  that 
almost  all  variability  in  the  date  of  first 
bloom  is  accounted  for  with  10  years  of  ob- 
servations and  that  the  range  of  first  bloom  is 
between  four  and  five  weeks  for  most  species 
(Tepedino  and  Stanton,  impublished  manu- 
script). Other  support  for  phenological  varia- 
bility exists.  Recently  West  and  Gasto  (1978) 
reported  that  the  onset  of  bloom  of  two  arid 
land  shrubs  in  northwestern  Utah  varied  over 
seven  years  by  44  and  39  days.  Thus,  the  sub- 
stantial phenological  variability  recorded  in 


our  study  over  two  years  is  low  relative  to 
what  can  be  expected  over  a  10-year  period. 

Long-term  studies  of  floral  abundance  are 
few.  Tamm  (1948,  1956,  1972a,  b)  counted 
flowers  of  several  species  in  permanent  quad- 
rats in  forest  and  meadow  in  mid-Sweden  for 
14  to  29  years.  All  species  showed  large  ir- 
regularities in  year-to-year  floral  abundances 
from  no  flowers  in  some  years  to  profuse 
abundance  in  others. 

Short-term  studies  are  more  numerous. 
Ackerman  and  Bamberg  (1974)  reported 
large  variation  in  floral  abundance  over  a 
three-year  period  for  Lijciinn  andersonii  in 
Nevada.  Bykov  (1974),  in  a  general  review  of 
vegetation  dynamics  of  the  arid  Turanian 
Plain,  reported  wide  variation  in  floral  abun- 
dance of  both  ephemerals  and  perennials.  Sa- 
rukhan  (1974)  supplied  floral  abundance  data 
for  three  species  of  Ranunculus  for  two  years 
from  permanent  plots,  with  all  species  pro- 
ducing many  more  flowers  in  the  first  year. 
Holway  and  Ward  (1965),  studying  the  alpine 
plant  community  in  northern  Colorado  over 
two  years,  noted  that  floral  production  in  the 
second  year  was  much  lower.  Davies  (1976) 
used  the  same  five  trees  of  each  of  two  spe- 
cies and  recorded  the  number  of  individuals 
flowering  over  an  eight-year  period  in  west- 
ern Australia.  Combining  data  for  both  spe- 
cies {Acacia  pruinocarpa,  Hakea  lorea;  Davies 
1976;  Table  6)  showed  that  in  three  of  eight 
years  the  number  of  individuals  flowering 
was  40  percent  or  less.  Data  on  fruit  crops 
also  were  presented  for  10  species  of  shrubs 
and  trees  for  10  years.  If  we  can  assume  that 
fruit  crop  bears  at  least  a  partial  relation  to 
floral  production  (Grubb  1977),  floral  pro- 
duction was  irregular  in  9  of  the  10  species. 
Schemske  (1977,  1978)  has  shown  that  the 
number  of  flowers  of  two  woodland  herbs 
censused  in  78  permanent  m^  quadrats  varied 
considerably  between  years.  Moldenke  (1976) 
noted  that  floral  production  varies  widely  be- 
tween years  in  California  grasslands.  Treshow 
(1979),  in  a  six-year  study  of  the  pinyon- 
juniper  community  in  Utah,  has  shown  that 
forb  cover  in  almost  every  year  differed  sig- 
nificantly from  each  other  year. 

Year-to-year  variation  in  floral  abundance 
is  not  restricted  to  "unpredictable"  temper- 
ate zone  communities  (Federov  1966).  Mass 
flowering  via  synchronization  of  all  members 


210 


Great  Basin  Naturalist 


Vol.  40,  No.  3 


of  a  particular  species  or  many  species  in  a 
community  in  the  tropics  at  periodic  inter- 
vals is  well  known  (Whitmore  1975).  Med- 
way  (1972)  and  McClure  (1966)  provided 
data  showing  widespread  irregularity  in  flow- 
ering for  46  species  of  tropical  rain  forest 
trees  in  Malaya.  Although  most  of  the  obser- 
vations were  recorded  on  very  few  individ- 
uals, it  is  enlightening  to  learn  that  the  per- 
cent of  species  flowering  each  year  ranged 
from  44  to  88  over  the  period  from  1963  to 
1968,  with  an  average  of  58  percent.  Of  42 
species  observed  for  the  entire  six-year  peri- 
od, only  11  (26.2  percent)  flowered  every 
year,  and  24  (57.1  percent)  flowered  in  three 
or  fewer  years.  In  a  study  of  flowering 
phenology  in  Ceylon,  Koelmeyer  (1959)  re- 
ported: "There  was  no  regularity  in  the  se- 
quence of  years  of  flowering  and  years  in 
which  there  was  no  flowering  in  the  individ- 
ual trees.  The  result  is  the  absence  of  a  defi- 
nite cycle  of  flowering." 

The  data  seem  clear.  Where  data  on  year- 
to-year  floral  abundances  have  been  record- 
ed, large  variations  in  floral  production  are 
the  rule  rather  than  the  exception. 

Variability  in  floral  resources  may  also  be 
modified  by  seasonal  influences.  First,  some 
parts  of  the  blooming  season  may  exhibit 
more  year-to-year  variability  than  others. 
Leopold  and  Jones  (1947)  hypothesized  that 
early  blooming  species  are  more  "turbulent" 
in  first  bloom  than  those  which  bloom  later 
in  the  year.  We  reexamined  the  phenology 
data  of  Leopold  and  Jones  (1947)  for  Wiscon- 
sin using  multiple  regression  analysis  and 
found  that  their  Sauk  County  data  do  show  a 
significant  inverse  correlation  between  range 
of  flrst  bloom  and  average  first  bloom  date. 
Though  the  Dane  County  data  show  the 
same  pattern,  it  is  not  significant  (Tepedino 
and  Stanton,  unpublished  manuscript). 

The  Wyoming  data  do  not  support  the  hy- 
pothesis of  greater  year-to-year  "turbulence" 
in  the  spring  flora.  At  The  Dirt  Farm,  be- 
tween-year  differences  in  flrst  bloom  were 
signiflcantly  greater  for  plants  that  bloomed 
over  the  second  half  of  the  census  period.  At 
Boulder  Ridge  no  significant  difference  be- 
tween flrst-  and  second-half  plant  species  was 
detectable.  In  addition,  between-year  differ- 
ences in  abundance  (as  judged  by  RHL  ratios) 
were  significantly  greater  for  the  last  half  of 


the  blooming  season  at  both  sites.  Whether 
these  differences  were  due  to  only  two  years' 
data  from  Wyoming  or  to  conditions  that  are 
site  specific  is  not  clear. 

The  second  way  in  which  floral  resources 
may  vary  seasonally  is  in  predictability  of 
subsequent  resource  abundance  and  composi- 
tion. From  this  perspective  the  spring  flora 
is,  indeed,  more  turbulent;  predictability,  as 
judged  by  floral  similarity  values  calculated 
at  three-week  intervals  (Fig.  5),  was  low  rela- 
tive to  average  flight  time  for  bees  until  the 
latter  third  of  the  bloom  season  when  com- 
posites became  dominant.  This  result  may  be 
somewhat  misleading  however,  because  more 
species  begin  bloom  in  spring  than  in  late 
summer  and  low  spring  similarity  values  are 
due  in  large  part  to  species  additions. 

Low  year-to-year  predictability  in  floral 
phenology  and  abundance  must  exert  strong 
selection  on  flower-visiting  insects.  This  is 
particularly  true  for  bees  because  every  stage 
in  their  life  cycle  is  obligately  dependent 
upon  floral  resources  for  food.  When  re- 
sources are  unpredictable  in  time  and/or 
space,  selection  should  favor  generalized  con 
sumers.  Alternatively,  specialization  would 
require  precise  synchronization  between  bee 
emergence  and  anthesis  of  the  host  plant, 
particularly  when  the  host  has  a  brief  bloom 
span.  It  is  unclear  how  such  precise  synchro- 
nization might  be  affected.  In  most  plant  spe- 
cies studied  photoperiodic  stimuli  initiate 
flower  formation,  but  subsequent  devel- 
opment and  anthesis  is  profoundly  modified 
by  diverse  factors  such  as  moisture  and  nutri- 
ent availability  and  temperature  (Evans  1969, 
Schwabe  1972).  Our  knowledge  of  the  stimuli 
used  by  bees  to  cue  emergence  in  a  natural 
setting  is  scanty  (Linsley  1958),  but  in  the 
laboratory  temperature  alone  is  a  reliable 
stimulus  for  several  species  {Megochile  rotiin- 
data  (Fabricius),  Osmia  lignaria  Say,  Hylaetis 
bisintiatiis  Forster,  Nomia  melanderi  Cock- 
erell,  and  several  others;  G.  E.  Bohart,  F.  D. 
Parker,  P.  F.  Torchio,  pers.  comm.,  pers. 
obs.).  Thus,  though  anthesis  is  determined  by 
a  complex  of  factors,  bee  emergence  may  be 
primarily  responsive  to  temperature.  Because 
of  these  differences  in  potential  stimuli  used 
by  the  two  groups,  synchronization  may  be 
rare.  In  this  regard,  Linsley  (1958)  noted  that 


September  1980 


Tepedino,  Stanton:  Bee  Ecology 


211 


studies  of  oligolectic  bees  frequently  reveal      consistent  with  documented  fluctuations  in 


poor  synchronization. 

Even  if  specialized  bees  could  achieve 
close  synchronization  with  host  plant  an- 
thesis,  the  problem  of  year-to-year  variation 
in  resource  quantity  still  remains.  Attempting 
to  track  specific  floral  resources  that  vary 
widely  could  cause  large  fluctuations  in  the 
populations  of  bee  specialists,  thereby  in- 
creasing the  probability  of  local  extinction 
(Tepedino  1979).  The  frequently  expressed 
view  that  most  temperate  bees  are  special- 
ized (van  der  Pijl  1966,  Faegri  and  van  der 
Pijl  1971,  Heinrich  1976,  Raw  1976, 
Heithaus  1979)  will  probably  require  modifi- 
cation because  such  specialization  seems  in- 


floral 


resources. 


Acknowledgments 

We  thank  J.  M.  Loar  and  T.  M.  Root  for 
providing  meticulous  assistance  in  the  field,  J. 
Meyer  for  drawing  the  figures,  and  Dr.  L.  L. 
McDonald  for  advice  on  cluster  analysis.  The 
manuscript  was  improved  by  comments  by 
M.  S.  Boyce,  K.  T.  Harper,  D.  H.  Knight,  P. 
Lincoln,  M.  D.  Marcus,  and  A.  R.  Moldenke. 
The  study  could  not  have  been  conducted 
without  support  from  NSF  Grant  BMS75- 
14044.  "Doc"  and  Peggy  Wollbrinck  of  Lar- 
amie and  the  proprietors  of  The  Dirt  Farm, 
Inc.,  graciously  allowed  use  of  private  land. 


Appendix  A.  The  number  of  flowers  recorded  in  permanent  m^  quadrats  at  The  Dirt  Farm  and  Boulder  Ridge  in 
197.5  and  1976.  Nomenclature:  Harrington  (1954),  Weber  (1967),  Porter  (1965).  Numbers  in  the  far  left  column  refer 
to  Figures  2,  7;  the  first  number  to  Figure  2  (Dirt  Farm),  the  second  to  Figure  7  (Boulder  Ridge).  A  zero  signifies 
nonrepresentation. 


Species 


Dirt  Farm 
1975  1976 


Boulder  Ridge 
1975  1976 


Berberidaceae 

(0,8) 

Berberis  repens  Lindl. 

Boraginaceae 

(19,  0) 

Crijptantha  flovoculata  (A.  Nels 

(23,  0) 

Cijnoglossitm  officinalis  L. 

(14,  0) 

Hackelia  florihwida  (Lehm.) 

Lappulo  redoicskii  (Horneni.) 

(10,  0) 

Lithospermtim  incisum  Lehm. 

(0,5) 

Mertetisia  Iwinilis  Rydb. 

(38,  0) 


(0,  23) 
(24,  19) 

(0,3) 
(22,  20) 


(40,  34) 

(35,  28) 
(39,  0) 
(42,  0) 

(0,  24) 


Cactaceae 

Opiintia  pohjacantha  Haw. 

Capparidaceae 

Cleorne  serrtilata  Pursh 

Caryophyllaceae 

Arenoria  fendlcri  A.  Gray 
Arcnaria  hookcri  Nutt. 
Cerastitim  arvense  L. 
Paronychia  sessiliflora  Nutt. 
Stellaria  media  (L.) 

Compositae 

Acliillea  miUefoliuin  L. 
Antennaria  microphyUa  Rydb. 
Antennaria  rosea  Greene 
Artemisia  frigida  Willd. 
Aster  rtibrotincttis  Blake 
Chnjsopsis  villosa  (Pursh) 
Chrysothamntts  nauseosus  (Pallas) 
Chrysothamntis  viscidiflorus  (Hook.) 
Cirsitim  tmdulatum  (Nutt.) 
Erigeron  canus  A.  Gray 


101 


52 


68 


575 

60 

- 

32 

50 

47 

— 

— 

67 

109 

— 

— 

_ 

— 

5 

2 

133 

16 

9 

9 

12 

5 

16 

42 

_ 

_ 

122 

58 

1165 

628 

1577 

474 

_ 

— 

.332 

50 

3489 

.3000 

718 

40 

— 

— 

9 

— 

382 

19 



_ 

.3.343 

959 



_ 

83 

- 

311 

114 

.393 

- 

,393 

73 

87 

18 

262 

116 

2426 

876 

1882 

73 

- 

- 

1906 

4216 

11 

3 

17 

4 

2 

— 

- 

2 

817 

314 

212 

Appendix  A  continued. 


Great  Basin  Naturalist 


Vol.  40,  No.  3 


(12,  2) 


(41,  35) 

(34,  29) 

(30,  14) 

(6,0) 

(13,  9) 

(0,30) 


(0,  18) 


(32,  0) 


Species 


Erigeron  nematophyUus  Rydb. 
Erigeron  pumilus  Nutt. 
GailUirdia  aristaia  Pursh 
Gutierrezia  sarothrae  (Pursh) 
Haplopappus  nutiaUii  T.  &  G. 
HelkintheUa  uniflom  (Nutt.) 
Hijmenoxijs  acaiiUs  (Pursh) 
Hymenoxijs  torreijana  (Nutt.) 
Senecio  catum  Hook. 
Senecio  integerrimus  Nutt. 
Solidago  spathulata  DC 
Taraxicttm  sp.  Hall 
Townsendid  scricea  Hook. 

Crassulaceae 

S«/i("i  stenopetalum  Pursh 


Cniciferae 

(4,0) 

Arabis  fendleri  (Wats.) 

Arcihis  holbocUn  Hornem. 

(21,  0) 

Dcsnirainia  sophia  L. 

Dmha  nemorosa  L. 

(15,  11) 

b'.rijsimum  capitatum  (Dougl.) 

Halimolohos  virgata  (Nutt.) 

(8,  10) 

LcsqucrcUa  ludoviciana  (Nutt 

(0,7) 

Phiisaria  amtralis  (Payson) 

(17,  0) 

Smjmbrium  ultissimum  L. 

Sunjmhrium  linefolium  Nutt. 

Euphorbiaceae 

Euphorbia  sp.  L. 

Gentianaceae 

Swertia  radiata  (Kellogg) 

Labiatae 

Hedcoma  drtDtitiiondii  Benth. 
Snitelhirid  bhtionii  Porter 


Legiuuinosae 
(25,  0)  Astragcdm  hisulaitus  Hook. 

Astragalus  crassicarpus  Nutt. 
(0,  21)  Astragalus  flexuosus  Dougl. 

(9,  0)  Astragalus  sericolettcus  Gray 

Astragalus  shortianus  Nutt. 
(7,  0)  Astragalus  spatulatiis  Sheld. 

(0,  .32)  Astragalus  striatus  Nutt. 

(0,12)  TItcrmopsis  rhoinhifolia  Nutt. 

Liliaceae 
(37,  31)  Allium  cernuum  Roth 

(0,  16)  Ellium  getjeri  Wats. 

(18,  17)  .A//ii/m  textile  Nels.  &  Macbr. 

Caloehorttis  nuttallii  Torrev 
(0,  1)  Leticocrintim  montanum  Nutt. 

(11,  0)  VAjgadenus  venenosus  Wats. 

Linaceae 
Lidi/Hi  lewisii  Pursh 


Dirt  Farm 

Boulder  Ridge 

1975 

1976 

1975 

1976 

434 

406 

895 

1014 

59 

— 

12 

- 



_ 

7 

- 

2163 

3170 

1512 

735 



45 

— 

— 

45 

35 

50 

3 

62 

81 

82 

24 

342 

248 

— 

— 

499 

701 

21 

53 

10 

11 

— 

19 





227 

100 

_ 

— 

57 

— 

4 

_ 

2 

— 

279 


971 


125 
9 


325 


75 
9 


88 


1768 


366 


535 


89 

166 

— 

— 



— 

13 

17 

128 

225 

— 

- 



35 

581 

207 

— 

122 

- 

160 

12 

— 

- 

- 

1839 

332 

466 

360 

48 

— 

219 

84 

24 

52 

- 

- 

27 

— 

— 

— 

187 


_ 

— 

4.38 

31 

21 

57 

- 

- 

_ 

— 

18 

- 

386 

329 

- 

- 



_ 

41 

— 

- 

- 

84 

43 

1771 

2693 

478 

98 

_ 

_ 

853 

607 

622 

1284 

187 

200 

1 

— 

— 

— 



— 

73 

122 

211 

314 

- 

- 

25 


17 


September  1980 
Appendix  A  continued. 


Tepedino,  Stanton:  Bee  Ecology 


213 


Species 


Malvaceae 

SpluicKiIcea  coccinea  (Pursh) 

Onagraceae 

(31,  0) 

Gaum  coccinea  Nutt. 

Oenothera  cownopifolia  T.  &  G. 

Poleinoniaceae 

Gilia  apicata  Nutt. 

Mirrostcris  htivtilis  (Dougl.) 

(1,0) 

Plilox  hnjoides  Nutt. 

(2,6) 

Phlox  hoodii  Rich. 

Polygonaceae 

Eriogonum  ahitttm  Torr. 

(36,  0) 

Eriogonum  effiistini  Nutt. 

(0,  33) 

Eriogonum  janicsii  Benth. 

(28,  22) 

Eriogonum  umheUatum  Torr. 

Portulacaceae 

Clatjtonia  hinceohita  Pursh 

Primulaceae 

Androsace  septentrionalis  L. 

Ranunculaceae 

(26,  0) 

Delphinium  nehoni  Greene 

(0.  4) 

Ranunctihts  ranunetdinus  (Nutt.) 

Rosaceae 

(0,  15) 

Amelanehier  ahiifolia  Nutt. 

(16.  13) 

Ccrcoearpus  montonus  Raf. 

Potcntilla  coneinna  Richards 

PotcntUla  fi-ssa  Nutt. 

(29,  0) 

PotentiUa  liippiana  Lehm. 

Santalaceae 

Commandra  umbellata  (L.) 

Saxifragaceae 

Ribes  cereum  Dougl. 

Scrophulariaceae 

(0,  25) 

CastUIeja  flava  Watson 

CoUinsia  parviflora  Dougl. 

Oddiocarpus  hiteus  Nutt. 

(20,  0) 

Penstemon  angustifolius  Nutt. 

(27,  0) 

Penstemon  erianthcrus  Pursh 

(33,  27) 

Penstemon  laricifohus  exHifolius  (A.  Neis. 

(0,  26) 

Pemtemon  strictus  Benth. 

Umbelliferae 

(5,  0) 

Harbouria  trachijpleura  (A.  Grav) 

(3,  0) 


Violaceae 

Viola  nuttallii  Pursh 


Dirt  Farm 
1975  1976 


28 
317 


41 


67 


Boulder  Ridge 
1975     1976 


- 

- 

4 

6511 

7526 

— 

91 

42 

12 

192 


27 


85 


28,869 

57,115 

- 

- 

- 

- 

84 

298 

852 

284 

282 

73 

19 


22 


11 

56 

13 

14 

- 

- 

4284 

5509 

_ 



220 

8 

mi 

3752 

128 

219 

- 

- 

9 

1 

- 

— 

6 

2 

862 

375 

20 

3 

243 

5 

14 

98 

- 

- 

3 

7 





912 

.361 

— 

— 

565 

402 

— 

- 

571 

32.3 

67 

^ 

- 

- 

61 

12 

4 

1 

449 

220 

966 

a3 

- 

- 

44 

7 

667 

196 

- 

8 

73 

52 



4 

Species  total 
Totals  (both  years) 


56  5.5 

63 


57 


60. 


214 


Great  Basin  Naturalist 


Vol.  40,  No.  3 


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DOG  OWNERS  AND  HYDATID  DISEASE  IN  SANPETE  COUNTY,  UTAH' 

Peter  M.  Schantz'  and  Ferron  L.  Andersen' 


.\bstract.—  a  questionnaire  survey  was  conducted  in  Sanpete  County,  Utah,  to  determine  the  knowledge  of  dog 
owners  concerning  hydatid  disease  and  an  identification  of  some  basic  sheep  management  practices  there.  The 
households  surveyed  included  21  (Group  I)  that  had  one  or  more  dogs  infected  with  Echinococciis  gmuiilostis  tape- 
worms at  more  than  one  annual  field  clinic,  and  19  others  (Group  II)  that  had  one  or  more  dogs  infected  when  the 
studv  first  began  in  1971-72,  but  had  not  had  any  infected  dogs  identified  at  field  clinics  during  subsequent  years. 
The  results  showed  that  92.5  percent  of  households  knew  the  cause  of  the  disease  and  how  it  is  transmitted,  and  that 
9()  percent  knew  of  someone  who  had  been  operated  on  for  surgical  removal  of  hydatid  cysts.  There  was  no  signifi- 
cant difference  in  the  level  of  knowledge  of  the  disease  between  the  two  groups  of  respondents,  nor  in  their  sheep 
management  practices.  Even  though  the  level  of  infection  of  the  parasite  in  dogs  has  decreased  since  the  project 
started,  certain  sheep  management  practices  persist  among  respondents  in  both  groups  that  allow  for  continued 
transmission  of  the  parasite  in  this  region. 


Hydatid  disease  is  an  infection  of  people, 
sheep,  and  some  other  animals  that  produces 
fluid-filled  (hydatid)  cysts  in  the  liver,  lungs. 


or  other  organs  (Fig.  1).  The  cysts  are  the  lar- 
val (immature)  forms  of  a  tapeworm  parasite, 
Echinococciis  granulosus  (Fig.  2),  which  lives 


Fig.  1.  Fluid-filled  hydatid  cysts  in  the  livers  and  lungs  of  infected  sheep. 


'Supported  in  part  by  U.S.  Public  Health  Grant  AI-10.588. 

'Parasitic  Diseases  Division,  Center  for  Disease  Control,  Atlanta.  Ceorgia  303.33. 

'Department  of  Zoology,  Brigham  Young  University,  Provo,  Utah  84602. 


216 


September  1980 


ScHANTz,  Andersen:  Hydatid  Disease 


217 


^  a. 


•^tiga^-'Ti'^- 


Fisi;.  2.  Tlie  adult  tapeworm.  Echinococni.s  oranitlosus  (approximatelv  5-6  mm  in  leiiiith).  removed  from  the  small 
intestine  ot  an  infeeted  dog. 


as  an  adult  in  the  .small  intestine  of  dogs. 
People  and  sheep  contract  the  hydatid  cysts 
when  they  inadvertently  ingest  the  tapeworm 
eggs  passed  in  the  stools  of  infected  dogs. 
This  may  occur  when  people  handle  dogs 
that  harbor  the  parasite,  and  when  sheep 
graze  on  contaminated  pa.stures.  Dogs  be- 
come infected  with  the  tapeworm  when  they 
ingest  hydatid  cy.sts  in  the  viscera  of  sheep. 
The  parasite  occurs  throughout  the  world 
wherever  dogs,  sheep,  and  other  suitable  ani- 
mal hosts  are  kept  together.  The  common 
practice  among  .sheep  ranchers  of  allowing 
dogs  to  eat  the  uncooked  viscera  of  home- 
killed  sheep  provides  optimum  conditions  for 
continued  transmission. 

In  the  United  States,  transmission  of  Ech- 
inococctis  granulosus  in  the  dog-sheep  cycle 
is  known  to  occur  in  several  western  states, 
including  California  (Araujo  et  al.  1975),  Ari- 
zona and  New  Mexico  (Schantz  1977),  and 
Utah  (Spniance  et  al.  1974).  The  most  serious 
problem  is  in  Utah,  where  nearly  50  human 


cases  have  been  diagnosed  since  1944.  Sever- 
al of  these  cases  were  fatal,  and  most  of  the 
others  have  required  surgical  removal  of  the 
hvdatid  cvsts.  Many  of  the  victims  were  resi- 
dents of  Sanpete  County,  which  is  in  the  cen- 
tral part  of  the  .state. 

Since  1971  hvdatid  disease  has  been  stud- 
ied and  control  measures  initiated  through 
the  combined  efforts  of  Brigham  Young  Uni- 
versity (Provo,  Utah),  the  Utah  State  Depart- 
ments of  Health  and  Agriculture  (Salt  Lake 
Citv,  Utah),  and  the  Center  for  Disease  Con- 
trol (Atlanta,  Georgia).  These  measures  have 
included  (1)  the  development  and  distribu- 
tion of  educational  displays  and  brochures  on 
the  life  cycle  of  the  hydatid  tapeworm,  (2) 
the  development  of  adequate  methods  for 
disposal  of  sheep  carcasses  at  community 
dumping  grounds,  (3)  the  periodic  holding  of 
public  health  clinics  to  detect  new  ca.ses  of 
human  infection,  and  (4)  annual  field  clinics 
to  detect  new  or  persistent  cases  of  infected 
dogs  (Fig.  3).  Following  the  implementation 


218 


Great  Basin  Naturalist 


Vol.  40,  No.  3 


Fig.  3.  Sheep  dogs  from  Sanpete  County  restrained  at 
cocnts  gr«nr//o.v(/.v  tapeworms. 

of  these  control  measures,  the  number  of 
dogs  found  infected  at  the  field  clinics  has 
decreased  from  27  percent  in  1971  (Loveless 
et  al.  1978)  to  14  percent  in  1978  (unpub- 
lished ms.).  Most  sheep  ranchers  have  shown 
a  cooperative  attitude  with  regard  to  proper 
disposal  of  sheep  carcasses  or  viscera.  Certain 
individuals,  however,  have  not  been  success- 
ful in  preventing  reinfection  of  their  dogs  as 
evidenced  by  the  fact  that  some  of  their  dogs 
were  found  repeatedly  infected  on  numerous 
occasions.  We  believed  that  if  the  reasons 
could  be  determined  why  some  dog  owners 
were  imable  or  unwilling  to  comply  with  the 
recommended  preventive  measures,  it  might 
be  possible  to  change  or  modify  the  recom- 
mendations to  obtain  more  cooperation,  and 
ultimately  an  improved  control  program. 

Materials  and  Methods 

A  questionnaire  survey  was  conducted  of 
the  owners  of  dogs  that  had  been  found  to  be 


field  clinic  during  examination  for  detection  of  Echino- 

infected  in  Sanpete  County.  The  survey  in- 
cluded 40  households,  21  of  which  had  one  or 
more  dogs  found  infected  at  more  than  one 
annual  clinic  (Group  I)  and  19  others  that 
had  one  or  more  dogs  infected  only  at  either 
the  first  or  second  annual  clinic  (1971  or 
1972),  but  did  not  have  infected  dogs  at  sub- 
sequent clinics  (Group  II).  During  the  visits, 
questions  were  asked  about  dog-feeding  prac- 
tices, dog  control,  sheep-killing  procedures, 
and  knowledge  of  the  life  cycle  and  control 
of  hydatid  disease. 

Results 

What  emerged  from  our  study  may  be  con- 
sidered a  general  description  of  the  habits 
and  practices  of  dog  owners  that  tend  to 
maintain  the  cycle  of  hydatid  disease  in  San- 
pete County.  Each  household  selected  had 
both  sheep  and  dogs.  The  average  number  of 
dogs  per  household  was  2.5  and  the  average 
flock  size  was  approximately  1000.  We  found 
that  nearly  everyone  was  aware  of  the  dis- 


September  1980 


ScHANTZ,  Andersen:  Hydatid  Disease 


219 


ease.  Persons  interviewed  in  90  percent  of 
the  households  knew  of  someone  who  had 
been  operated  on  for  the  disease.  This  was 
usually  someone  from  the  same  town,  and  in 
10  households  (25.0  percent)  the  victim 
known  was  a  member  of  the  nuclear  or  ex- 
tended family.  Moreover,  persons  inter- 
viewed in  92.5  percent  of  households  knew 
the  cause  of  the  disease  and  how  it  is  trans- 
mitted. Specifically,  they  knew  that  people 
become  infected  with  hydatid  cysts  by  in- 
gesting eggs  passed  in  the  feces  of  infected 
dogs,  and  that  dogs  become  infected  with  the 
hvdatid  worms  by  ingesting  the  cysts  in  the 
lungs  and  livers  of  sheep. 

More  than  four-fifths  of  the  households  in- 
dicated they  sometimes  killed  and  butchered 
sheep  on  their  premises  or  in  the  fields.  De- 
spite their  awareness  and  understanding  of 
how  hvdatid  disease  is  transmitted,  nearly 
two-thirds  admitted  their  dogs  had  access  to 
the  sheep-killing  area,  and  nearly  half  said 
the  dogs  sometimes  ate  part  of  the  sheep  car- 
cass. 

The  main  diet  of  dogs  in  more  than  85  per- 
cent of  households  was  commercial  dog  food, 
and  in  none  was  the  main  diet  reported  as 
sheep  muscle  or  organ  meat.  Nevertheless,  it 
was  clear  that  most  dogs  could  possibly  eat 
sheep  at  least  occasionally,  since  in  two- 
thirds  of  households  dogs  were  allowed  to 
roam  free,  and,  therefore  could  scavenge  on 
sheep  carcasses  at  the  town  dump  or  in  the 
fields.  Less  than  one-third  of  households 
regularly  tied  or  locked  up  their  dogs  when 
the  dogs  were  not  working. 

Persons  interviewed  at  more  than  80  per- 
cent of  households  indicated  they  believed 
that  the  recommended  control  measures  were 
adequate  to  break  the  chain  of  transmission 
and  eliminate  the  infection.  Persons  at  only  6 
(15  percent)  of  households  indicated  they  had 
taken  no  active  measures  to  eliminate  the  in- 
fection. At  the  34  households  that  indicated 
they  had  done  something,  the  most  frequent- 
ly mentioned  steps  taken  were  (1)  periodic 
treating  of  dogs  for  tapeworms,  and  (2)  dis- 
carding of  viscera  from  home-killed  sheep  in 
such  a  way  that  dogs  could  not  get  to  it.  Four 
households  indicated  they  no  longer  had  dogs 
because  of  the  potential  of  contracting  hyda- 
tid disease.  There  was  a  general  consensus 
(82.5  percent)  that  government  authority 


should  not  make  it  illegal  for  dogs  to  eat 
parts  of  the  sheep  carcass. 

When  the  households  were  categorized  ac- 
cording to  whether  their  dogs  had  been 
found  infected  at  only  one  of  the  first  clinics 
or  whether  their  dogs  had  been  found  repeat- 
edly infected,  there  were  no  obvious  differ- 
ences that  would  allow  us  to  conclude  whv 
the  first  group  of  households  was  apparently 
successful  in  preventing  reinfection.  There 
were  no  statistically  significant  differences  in 
the  two  groups  regarding  the  number  of  dogs 
or  sheep  they  owned,  the  frequency  that 
sheep  were  butchered  for  home  consumption, 
the  apparent  access  of  dogs  to  sheep  viscera, 
the  household  members'  knowledge  and  un- 
derstanding of  hydatid  disease,  nor  willing- 
ness to  take  measures  to  prevent  the  infection 
in  the  dogs.  In  fact,  the  responses  to  our  ques- 
tions appeared  to  suggest  that  dog  owners 
with  repeatedly  infected  dogs  were  more 
likely  to  have  tied  their  dogs  up  when  not 
working  and  to  have  taken  other  deliberate 
measures  to  prevent  their  dogs  from  eating 
parts  of  the  sheep  carcass.  This  apparent 
anomaly  is  most  likely  explained  by  the  fact 
that. owners  of  repeatedly  infected  dogs  had 
more  recently  been  made  aware  of  what  they 
should  be  doing  to  prevent  infection  than  the 
other  group  of  dog  owners  whose  dogs  had 
been  given  a  "clean  bill  of  health  "  at  the 
most  recent  dog  clinics. 

In  summary,  we  did  not  learn  from  our 
study  why  some  dog-owning  households  were 
successful  in  preventing  reinfection  of  their 
dogs  and  why  others  were  not.  What  was 
clear,  however,  was  that  numerous  opportu- 
nities still  existed  at  these  households  for  dogs 
to  become  infected  with  hydatid  tapeworms. 
As  a  result  of  health  education  and  other  con- 
trol activities,  virtually  all  the  Sanpete  Coun- 
ty dog  owners  interviewed  in  our  survey 
knew  the  basic  facts  ab^nit  hydatid  disease; 
however,  few  had  actually  taken  all  the  nec- 
essary steps  to  insure  its  elimination.  Evi- 
dence obtained  from  the  survey  suggests  that 
manv  dog  owners  apparently  believe  that  pe- 
riodic treatment  of  dogs  is  sufficient  to  solve 
the  problem;  however,  that  may  be  an  over- 
simplified solution.  To  effectively  break  the 
chain  of  transmission,  all  dogs  must  be  pre- 
vented from  eating  the  viscera  of  infected  an- 
imals. This  means  not  only  that  dog  owners 


220 


Great  Basin  Naturalist 


Vol.  40,  No.  3 


must  refrain  from  feeding  such  organs  to 
their  dogs,  but,  since  dead  sheep  are  fre- 
quently discarded  in  open  pits  and  are  acces- 
sible to  roving  dogs,  dogs  must  be  kept  under 
control  at  all  times.  An  additional  feasible 
control  measure  would  be  the  installation  of 
large  metal  pit  covers  or  sturdy  fences  at  the 
animal  pits  in  order  to  prevent  ready  access 
of  roving  dogs  to  animal  carcasses  discarded 
at  those  sites. 

From  its  inception  in  1971,  the  Hydatid 
Disease  Control  Program  has  been  an  entire- 
ly voluntary  campaign.  Results  of  this  survey 
suggest  that  some  additional  incentives  may 
be  necessar\'  to  insure  that  all  dog  owners 
take  the  necessar\'  steps  to  stop  the  transmis- 
sion of  hydatid  disease. 


Literature  Cited 

Aracjo.  F.  p..  C.  W.  Schwabe.  J.  C.  Sawder,  and  ^\^ 
G.  Davis.  1975.  Hvdatid  disease  transmission  in 
California.  A  study  of  the  Basque  connection. 
Am.  J.  Epidemiol.  102:  291-302. 

ScH.\NTZ.  P.  M.  1977.  Echinococcosis  in  American  In- 
dians living  in  Arizona  and  New  Mexico.  Review 
of  recent  studies.  Am.  J.  Epidemiol.  106: 
.370-.379. 

Spru.ance,  S.  L.,  L.  F.  Klock.  F.  Cha.vg.  T.  Fikushima. 
F.  L.  .\ndersen,  .and  1.  G.  K.\g.an.  1974.  Endemic 
hvdatid  disease  in  Utah.  A  review.  Rocky  Mtn. 
Med.  J.  71:  17-23. 

Loveless,  R.  M.,  F.  L.  .\ndersen.  M.  J.  Rwis.ay,  and  R. 
K.  Hedelius.  1978.  Echinococciis  granulosus  in 
dogs  and  sheep  in  central  Utah.  1971-1976.  Am. 
J.  Vet.  Res.  39:  499-502. 


NEW  GRASS  DISTRIBUTION  RECORDS  FOR  ARIZONA,  NEW  MEXICO,  AND  TEXAS' 

Stephan  L.  Hatch- 

Abstr^ct.—  New  distribution  records  are  given  for  seven  grass  species  now  found  in  Arizona,  New  Mexico,  and 
Texas. 


Recent  collections  have  revealed  new  dis- 
tribution records  for  seven  southwestern  grass 
species.  These  records  are  extensions  to  the 
known  distributions  of  these  species  as  re- 
corded in  general  for  the  United  States  bv 
Hitchcock  (1951),  in  Arizona  and  Te.xas  by 
Gould  1 1951.  1975^.  and  in  the  intermountain 
area  by  Cronquist  et  al.  il977>.  The  checklist 
for  New  Mexico  published  by  Martin  and 
Castetter  (1970)  is  the  basis  of  distributions 
for  that  state.  Voucher  specimens  for  these 
new  records  have  been  distributed  in  the 
Tracy  Herbarium  (^TAES). 

Eremopyron  triticeum  (Gaertn.^  Nevski 

.\rizona:  Coconino  Co.:  Locallv  abundant 
as  an  adventive  on  disturbed  sites.  0.5  km 
north  of  Fredonia,  east  side  of  Highway  89A. 
on  a  hard  gra\  clav  soil  at  1600  m  elevation, 
17  May  1978.  Brown  652  (TAES).  This  in- 
troduced grass  is  a  new  state  record  for  Ari- 
zona. Previous  collections  have  been  report- 
ed from  Oregon,  Idaho,  Nevada,  and  Utah  by 
Cronquist  et  al.  il977'.  Montana  (Hitchcock 
1951k  and  New  Mexico  ^Hatch  1977). 

Afrna  barbota  Brot. 

New  Mexico:  Dona  Ana  Co.:  Rare  adven- 
tive along  Interstate  10.  10  km  south  of  Las 
Cnices,  3  April  1978.  Mocliange  21  (TAES). 
This  is  a  new  state  record  for  New  Mexico. 
Previous  reports  show  a  distribution  from 
Washington  to  Arizona  ^Gould  1951). 

Bothriochloa  ischaemum  yh.)  Keng. 

New  Mexico:  Colfax  Co.:  Locally  abun- 
dant as  an  adventive.  3.5  km  south  of  Raton. 


18  August  1978,  Hatch  4072  (TAES).  Dona 
Ana  Co.:  Locally  abundant.  11  km  south  of 
San  Agustin  Pass  on  the  north  slopes  of  the 
Organ  Mountains.  18  September  1977,  Dick- 
Peddie  55  i^TAES).  Grant  Co.:  .\bundant,  32 
km  north  of  Silver  City  along  Highwav  15,  9 
October  1977.  Hoefler'33  TAES).  Sierra  Co.: 
Five  km  south  of  Williamsburg  along  Inter- 
state 25,  26  August  1977,  Hatch  2604 
I  TAES).  This  is  a  new  state  record  for  New 
Mexico.  Gould  (1975)  reported  this  species  as 
being  introduced  in  Te.xas  as  a  pasture  grass. 

Dactyloctenium  aegyptium  (L.i  Beauv. 

New  Mexico:  Dona  Ana  Co.:  Rare  as  an 
adventive  in  the  New  Mexico  State  Univer- 
sitv  Agronomy  Field  Laboratory  land.  Las 
Cruces,  10  September  1977,  Hatch  2565 
TAES).  This  is  a  new  record  for  New  Mexi- 
co. Gould  ^195L  reported  this  species  from 
Arizona  and  later  ^1975)  from  Te.xas.  Gould 
(1975)  stated  that  this  species  was  well  adapt- 
ed to  sandy  soils  of  the  southern  United 
States. 

Eragrostis  superba  Peyr. 

New  Me.xicg:  Dona  Ana  Co.:  A  relatively 
rare  introduced  grass  along  Interstate  10.  6 
km  south  of  Las  Cruces  in  sandy  soil.  15  Oc- 
tober 1977,  Yehca  3S  (T.\ES).  This  is  a  new 
state  record  for  New  Mexico.  The  plant  has 
been  collected  in  Texas  22  May  1957.  Gould 
7550  (TAES),  but  was  not  included  in  Gould's 
(1975)  Grasses  of  Texas.  This  species  has  been 
collected  in  .\rizona.  3  November  1961.  Mat- 
tox  and  White  sn.  (T.\ES).  It  is  native  to 
South  Africa  and  was  introduced  to  the 
United  States  as  P.I.  185516  (No.  39)  12  De- 
cember 1949. 


Technical  Bulletin  T..\.  15803,  Texas  Agricultural  Eipenment  Stabon.  Teias  A  &  M  L'niversit).  College  Station.  Texas  .  iS*3. 
'Department  of  Range  Science.  Tesas  A  &  Vt  Universitv-.  College  Station.  Teias  77843. 


221 


222 


Great  Basin  Naturalist 


Vol.  40,  No.  3 


Leptoloma  arenicola  Swallen 

New  Mexico:  Lea  Co.:  A  rare-abundant 
perennial  grass  on  sandy  sites,  22  km  east- 
northeast  of  Maljamar  along  Highway  82,  10 
August  1976,  Elhworth  23  (TAES).  This  col- 
lection is  a  new  state  record  for  New  Mexico. 
Hitchcock  (1951)  and  Gould  (1975)  report 
this  taxon  as  occurring  only  in  Kenedy  Co., 
Texas.  This  is  a  rhizoinatous  form  of  the 
widespread  L.  cognatum  (Schult.)  Chase, 
classified  by  Gould  (1975)  as  L.  cognatum 
var.  arenicola  (Swallen)  Gould. 

Texas:  Kent  Co.:  Locally  abundant  in 
stands  of  shinnery  oak  {Querciis  havardii 
Rydb.),  3.0  km  south-southeast  of  Girard,  9 
June  1979,  Slosser  sn.  (TAES).  The  Slosser 
collection  is  a  distribution  extension  within 
Texas  of  several  hundred  miles. 

Neeragrostis  reptans  (Michx.)  Nicora 

New  Mexico:  Sierra  Co.:  Locally  abun- 
dant on  the  mudflats  of  Elephant  Butte  Res- 
ervoir, near  Alamosa,  10  September  1978,  So- 


pyn  sn.  (TAES).  This  is  a  new  record  for  New 
Mexico.  Gould  (1975)  reported  this  species  as 
being  abundant  on  exposed  lake  beds  in  the 
central  United  States  from  Kentucky  and 
South  Dakota  to  Louisiana,  Texas,  and  Flor- 
ida. 

Literature  Cited 

Cronquist,  a.,  B.  H.  Holmgren,  N.  H.  Holmgren,  J.  L. 
Reveal,  and  P.  L.  Holmgren.  1977.  Inter- 
mountain  flora.  Vol.  6.  Columbia  Press.  New 
York.  584  pp. 

Gould,  F.  W.  1951.  Grasses  of  the  southwestern  United 
States.  University  of  Arizona  Press,  Tucson.  352 
pp. 

1975.  The  grasses  of  Texas.  Texas  A  &  M  Univer- 
sity Press,  College  Station.  65.3  pp. 

H.\TCH,  S.  L.  1977.  New  grass  distribution  records  for 
New  Mexico  and  the  United  States.  Great  Basin 
Nat.  37:530-531. 

Hitchcock,  A.  S.  1951.  Manual  of  the  grasses  of  the 
United  States.  USDA  Miscellaneous  Publication 
200,  rev.  by  Agnes  Chase.  1051  pp. 

Martin,  W.  C,  and  E.  F.  Castetter.  1970.  A  checklist 
of  gymnosperms  and  angiosperms  of  New  Mexi- 
co. Published  privately. 


A  COMPARISON  OF  EPIPHYTIC  DIATOM  ASSEMBLAGES  ON 
LIVING  AND  DEAD  STEMS  OF  THE  COMMON  GRASS  PHRAGMITES  AUSTRALIS 

Judith   A.  Grimes'.   Larry  L.   St.  Clair',  and  Samuel  R.  Rushforth' 

Abstract.-  Diatoms  epiphytic  on  Phragmites  austmUs  (Cav.)  Trin.  ex  Steaded  stems  were  collected  from  a  single 
clone  at  the  .southern  end  of  Provo  Bay,  Utah  Lake,  Utah.  Diatom  populations  from  both  living  and  dead  stem  sec- 
tions were  analyzed.  Species  diversity  in  each  .sample  was  high,  indicating  that  the  stems  provide  a  relatively  stable 
habitat  for  diatom  epiphytes.  Of  the  23  genera  found,  only  Gomphonema  and  \avicula  showed  significant  trends 
toward  stem  preference.  The  diatoms  in  this  study  support  the  current  view  that  Utah  Lake  is  a  slightly  saline, 
eutrophic  system. 


The  occurrence  of  diatom  assemblages  as 
epiphytes  on  httoral,  emergent  macrophytes 
is  well  documented  (Godward  1934,  1937, 
Knud.son  1957,  Prow.se  1959).  Likewise,  the 
impact  of  such  epiphytes  on  primary  produc- 
tivity and  community  trophic  structure  has 
been  examined  in  several  estuarine  environ- 
ments (Mclntire  et  al.  1971,  Stowe  et  al. 
1971,  and  Main  et  al.  1974),  but  has  been 
largely  ignored  in  freshwater  systems  (Wetzel 
1964).  Tlie  epiphytic  diatom  communities  at- 
tached to  emergents  inevitably  play  a  role  in 
the  overall  productivity  of  lakes  and  es- 
tuaries. They  also  contribute  to  regulation  of 
the  overall  metabolism  of  such  waters  by  al- 
tering the  amount  and  quality  of  alloch- 
thonous  organics  entering  the  lake  by  acting 
as  physical  and  metabolic  traps  or  filters.  The 
attached  diatom  flora  also  serves  as  an  au- 
tochthonous source  of  particulate  organic 
and  dissolved  organic  matter  that  is  readily 
available  to  pelagic  animals.  The  degree  of 
influence  of  these  epiphytic  organisms  on  the 
productivity  of  standing  waters  has  rarely 
been  determined.  However,  Allen  (1971)  esti- 
mated that  up  to  31.3  percent  of  the  total  lit- 
toral production  could  be  attributed  to 
epiphtyic  algae,  with  up  to  21.4  percent  of 
the  total  lake  production  being  attributable 
to  such  attached  communities.  In  addition,  a 
comparison  between  phytoplankton  and  epi- 
phyte production  demonstrated  that  the  lat- 
ter was  equivalent  to  75  percent  of  the 
phytoplankton  production  over  the  annual 
period  (Allen  1971). 


Even  though  epiphytic  commimities  have 
been  demonstrated  to  be  important,  the  dis- 
tribution patterns  of  such  assemblages  on  the 
basis  of  variation  in  ho.st  species  and  host  sub- 
strate conditions  have  received  little  atten- 
tion. Likewise,  the  complex  physiological 
relationship  between  the  host  macrophyte 
and  the  attached  diatom  species  has  received 
less  attention  than  warranted  (Wetzel  1964, 
1965,  1969b,  Allen  1971,  Hough  et  al.  1975). 
The  impact  of  this  relationship  is  fimdamen- 
tal  to  understanding  the  basic  distribution 
patterns  of  epiphytes  not  only  on  different 
macrophyte  host  species  but  also  on  members 
of  the  same  species  at  different  levels  of  se- 
nescence. 

The  purpose  of  this  study  is  to  illuminate 
distribution  patterns  of  diatom  epiphytes  on 
living  and  dead  specimens  of  a  single  macro- 
phyte host  {Phragmites  australis  (Cav.)  Trin. 
ex  Steaded)  in  Utah  Lake,  Utah.  The  data 
from  this  study  will  be  used  as  a  baseline  for 
extended  research  in  Utah  Lake  on  epiphyte 
distribution  patterns  and  epiphyte  impact  on 
lake  productivity  and  trophic  structure. 

Methods 

Samples  were  collected  20  September 
1978  from  a  single  clone  of  Phragmites  ous- 
trcilis  located  at  the  southern  end  of  the 
mouth  of  Provo  Bay  in  Utah  Lake.  Five  sam- 
ples of  living  and  five  of  dead  Phragmites 
australis  stems  were  collected  as  cut  10  cm 
sections,   measured  from   the  water  level 


'Department  of  Botany  and  Range  Science,  Brigham  Young  University,  Provo,  Utah  84602. 


223 


224 


Great  Basin  Naturalist 


Vol.  40,  No.  3 


down.  Samples  were  prepared  according  to 
standard  acid-oxidation  methods,  and  per- 
manent diatom  slides  were  made  using  Naph- 
rax  diatom  momitant  (St.  Clair  and  Rushforth 
1977).  Slides  were  examined  and  diatom  spe- 
cies were  identified  at  lOOOX  with  a  Ziess  RA 
research  microscope  with  bright  field  and 
Nomarski  interference  phase-contrast  acces- 
sories. 

Quantitative  data  on  the  diatom  assem- 
blages were  recorded  by  counting  250-400 
diatoms  for  each  sample.  Previous  studies 
have  shown  that  a  statistically  valid  count 
can  be  obtained  within  this  range  (Squires 
1977).  Each  slide  was  then  thoroughly 
scanned  to  record  the  rare  species.  The  re- 
sults were  converted  into  percent  relative 
density  values  for  all  species  for  each  site. 
Shannon-Wiener  diversity  indices  were  cal- 
culated for  individual  samples  (Shannon  and 
Wiener  1963). 

The  relative  density  figures  for  each 
sample  were  compared  to  all  other  samples 
and  similarity  indices  were  calculated  (Ru- 
zicka  1958).  These  indices  were  clustered 
(Sneath  and  Sokal  1973)  to  identify  unique 
community  associations  within  and  between 
the  living  and  dead  Phragmites  aiistralis  stem 
sections. 

The  most  prevalent  diatoms  encountered 
in  the  study  and  the  diatoms  that  significant- 
ly differed  between  the  living  and  the  dead 
specimens  of  Phragmites  australis  were  deter- 
mined. 

Results 

Twenty-three  genera  and  114  diatom  spe- 
cies were  found  on  the  10  Phragmites  austral- 
is stem  sections  (Table  1).  The  most  prevalent 
diatoms  throughout  the  study  were  Navicula 
graciloides,  Nitzschia  inconspicua,  and  Nit- 
zschia  filiformis  (Table  2).  Nitzschia  dis- 
sipata,  Stephanodiscus  astrea  var.  minutula, 
and  Nitzschia  palea  were  also  common. 
Among  the  prevalent  species,  Amphora  ve- 
neta  was  found  to  occur  only  on  the  living 
stems,  whereas  Navicula  schroeteri  var.  es- 
camhia  was  essentially  restricted  to  the  dead 
stems. 


Table  1.  Alphabetical  list  of  the  diatom  taxa  found 
on  living  and  dead  Phragmites  australis  stem  sections 
from  Provo  Bay,  Utah  Lake,  and  their  average  relative 
density. 


Species 

Living 

Dead 

Achnanthes  hauckiana" 

.07 

_ 

Achnanthes  lanceolata  var.  dtibia" 

— 

P 

Achnanthes  lewisiana 

.06 

P 

Achnanthes  minutissima 

2.20 

.57 

Achnanthes  sp." 

P 

— 

Amphora  ovaUs 

.31 

P 

Amphora  ovalis  var.  pedicuhis' 

— 

.08 

Amphora  perpusilla 

.76 

1.79 

Amphora  veneta 

8.78 

1.26 

Amphora  sp.°° 

.19 

- 

Anomoeoneis  sphaerophora" ° 

P 

— 

AsterioneUa  formosa 

.09 

.08 

Colonels  fenzlioides'  ° 

.02 

— 

Cocconeis  placenttila  var.  eughjpta' 

— 

P 

Cocconeis  placentula  var.  lineata 

.07 

.07 

Coscinodiscus  sp. 

.06 

.06 

Cyclotella  kiitzinghiana' 

— 

.13 

Cijclotella  meneghiniana 

1.87 

1.80 

Cijmbella  affinis' 

— 

P 

Cymbella  miniita" 

— 

.06 

Cijmbella  muelleri" 

.14 

- 

Cymbella  prostrata 

.07 

.05 

Cymbella  sp.  °  ° 

P 

— 

Diatoma  tentie  var.  elongatum 

.13 

.23 

Diploneis  oblongella' ° 

.02 

— 

Epithemia  adnata  var.  porcellus 

P 

P 

Epithemia  adnata  var.  proboscidea" 

- 

P 

Fragilaria  brevistriata" 

— 

P 

Fragilaria  brevistriata  var.  inflata 

.06 

.15 

Fragilaria  construens' 

— 

P 

Fragilaria  constriiens  var.  binodis 

.06 

P 

Fragilaria  constrtiens  var.  venter 

.65 

.11 

Fragilaria  crotonensis' 

— 

.17 

Fragilaria  lapponica" 

.19 

- 

Fragilaria  pinnata" 

.37 

— 

Fragilaria  pinnata  var.  lancettula 

.07 

.13 

Fragilaria  similis 

.19 

.17 

Fragilaria  vaucheriae 

.90 

.63 

Fragilaria  virescens" 

P 

— 

Gomphonema  affine" 

.13 

- 

Gomphonema  gracile" 

.02 

— 

Gomphonema  intricatum' 

— 

P 

Gomphonema  olivaceiim 

.30 

.05 

Gomphonema  parvulum 

.44 

.21 

Gomphonema  subclavatiim 

var.  commutatum 

.08 

P 

Gomphonema  tenellum" 

.02 

— 

Gomphonema  ventricosum' ° 

.44 

— 

Gomphonema  sp.  °  ° 

.06 

— 

Melosira  granulata 

.55 

.65 

Melosira  granulata  var.  angustissima 

1.58 

1.05 

Melosira  italica 

.21 

P 

Navicula  arvensis' 

_ 

.15 

Navicula  arenaria' 

— 

P 

Navicula  aurora" 

.02 

- 

Navicula  capitata  var.  hungarica' 

— 

.1 

Navicula  cincta' 

— 

.59 

Navicula  circumtexta' 

— 

.15 

Navicula  cryptocephala 

.29 

.35 

September  1980 


Grimes  et  al.:  Diatom  Assemblages 


225 


Table  1  continued. 


Species 

Living 

Dead 

Navicula  cryptocephala  var. 

veneta 

3.45 

4.38 

Navicuhi  cxigua" 

— 

P 

Navicuhi  gmciloides 

12.35 

9.17 

Navicula  Iwiifleri  var.  leptocephala 

.08 

P 

Navicuhi  lanccolata  ° 

— 

.08 

Navicula  tnitiima 

.07 

.84 

Navicula  oblonga 

P 

.08 

Navicula  peregrina' 

- 

.08 

Navicula  pupula 

P 

.08 

Navicula  radiosa  var.  tcnclla 

1.13 

2.42 

Navicula  rh yncocephala  ° 

— 

P 

Navicula  salinarum  ° 

— 

.13 

Navicula  salinarum  var.  intc 

rmedia 

.08 

.05 

Navicula  schroetcri  var.  cscambia 

3.06 

6.23 

Navicula  tcnclloidcs 

.15 

1.27 

Navictila  tripunctata 

P 

.33 

Navicula  tripunctata  var.  sch 

lizonemoides     .44 

1.51 

Navicula  sp.° 

— 

.63 

Nitzschid  acicularis 

.51 

.66 

Nitzschia  amphibia 

.37 

.25 

Nitz-schia  apiculata 

.06 

.18 

Nitzschia  di.ssipata 

6.00 

4.95 

Nitz-'ichia  filiformis 

7.10 

6.83 

Nitzschia  frustulum 

.80 

.53 

Nitzschia  gracilis" 

.23 

— 

Ni  tzsch  ia  han  tzsch  ia  na 

4.94 

4.40 

Nitzschia  holsatica 

1.67 

2.35 

Nitzschia  inconspicua 

12.40 

13.25 

Nitzschia  linearis 

.06 

.08 

Nitzschia  longissima  var.  clostcrium 

.08 

.08 

Nitzschia  ovalis 

.18 

..32 

Nitzschia  palea 

6.90 

13.50 

Nitzschia  paleaceae 

4.63 

3.92 

Nitzschia  punctata' 

— 

P 

Nitzschia  sigmoidca ° 

— 

P 

Nitzschia  stagnorum' " 

P 

— 

Nitzschia  s-p.  1 

P 

.42 

Nitzschia  sp.  2°° 

.21 

— 

Ophephora  martiji' 

- 

P 

Rhoicosphenia  curvata 

3.96 

6.32 

Rhopalodia  gibba° 

- 

.05 

Rhopalodia  gibberuhi  var.  vanheurckii 

.10 

P 

Stcphanudiscus  astrea" 

.06 

- 

Stcphanodiscus  astrea  var.  minutula 

5.15 

4.34 

Stcphanodisctis  niagarae ° 

- 

.08 

Surirella  angustata 

P 

P 

Surirella  ovalis  var.  brightwe 

llii° 

— 

.15 

Surirella  ovata°° 

.08 

— 

Sijnedra  actis 

.08 

1.60 

Sijnedra  dclicatissima  var.  angiistissima 

.15 

.02 

Sijnedra  fasciculata  var.  trunctata' 

- 

.08 

Sijnedra  mazamaensis 

.15 

.08 

Sijnedra  socio 

P 

.08 

Synedra  ulna 

P 

.11 

Synedra  ulna  var.  contractu 

.81 

.02 

°°  Species  unique  to  living  Phragmites  australis  stems. 
'Species  unique  to  dead  Pbragmites  australis  stems. 
P     Species  not  recorded  on  the  transects  taken  for  relative  de 
ures  but  found  on  other  sections  of  the  diatom  slide. 


iity  fig- 


Species  diversity  according  to  number  of 
species  encountered  was  high,  averaging  48 
species  per  sample.  However,  there  were 
generally  2  to  4  dominant  species  ranging  be- 
tween 10-18  percent  relative  density  in  each 
sample,  which  allowed  for  only  moderately 
high  Shannon-Wiener  diversity  values  (Table 
3).  Forty-nine  percent  of  the  diatom  species 
was  found  in  30  percent  of  all  samples,  and 
34  percent  of  the  diatom  species  was  found 
in  50  percent  of  all  samples. 

The  results  of  the  cluster  analysis  (Fig.  1) 
demonstrate  the  high  degree  of  similarity  en- 
countered for  all  ten  .samples.  Even  so,  mar- 
ginal separation  into  .samples  from  living  and 
dead  stems  was  obtained. 

A  similarity  matrix  comparing  all  10  sam- 
ples was  constructed. 

Means  of  similarity  indices  for  living  stems, 
dead  stems,  and  between  living  and  dead 
stems  were  computed.  T-tests  were  per- 
formed and  it  was  determined  that  there  was 
no  significant  difference  in  similarity  within 
or  between  these  samples  sets. 

Discussion 

Each  of  our  10  samples  consistently  con- 
tained approximately  50  identifiable  species. 
In  general,  no  one  species  represented  more 
than  18  percent  of  the  total  population  of 
any  sample.  These  conditions  are  indicative 
of  a  diverse  flora  that  is  further  supported  by 
our  Shannon-Wiener  diversity  values  and  the 
average  number  of  species  per  substrate 
(Table  3).  Such  conditions  indicate  that  the 
epiphyte  flora  in  Utah  Lake  is  more  diverse 
than  we  previously  believed.  T-tests  were 
computed  comparing  the  means  of  the  Shan- 
non-Wiener diversity  indices  of  both  .sub- 
strates as  well  as  the  average  number  of  spe- 
cies from  both  living  and  dead  stems.  No 
significant  differences  between  the  values  in 
either  compari.son  existed. 

A  total  of  23  diatom  genera  were  encoun- 
tered during  this  study.  The  number  of  spe- 
cies included  in  these  genera  was  nearly 
equally  distributed  between  living  and  dead 
stems  (Table  4).  However,  substratum  prefer- 
ences were  noted  in  the  genera  Navicula  and 
Gomphonema  and  in  individual  .species  with- 
in several  other  genera.  Of  the  114  species 
found  in  the  study,  22  were  unique  to  living 
and  29  were  unique  to  dead  stems. 


226 


Great  Basin  Naturalist 


Vol.  40,  No.  3 


T\BLE  2.  Important  species  encountered  on  Phragmites  australis  stem  sections  from  Provo  Bay,  Utah  Lake,  with 
their  percent  relative  densities.  Important  species  are  those  species  with  a  percent  relative  density  greater  than  .3 
percent  in  any  one  sample. 


Livin 

g  stems 

Dead  stems 

Species 

1 

2 

3 

4 

5 

6 

/ 

8 

9 

10 

Achnanthes  minutissima 

3.1 

Amphora  perpusiUa 

3.3 

3.2 

Amphora  ceneta 

17.3 

5.6 

7  2 

11.2 

Cyclotella  meneojiiniaiui 

3.0 

Melosira  granulata 

var.  angustissima 

3.9 

S'avicula  cnjptocephala 

var.  veneta 

3.4 

4.9 

4.2 

3.7 

5.4 

3.3 

6.1 

3.5 

Xaiicula  graciloides 

9.9 

13.2 

13.5 

14.0 

11.2 

5.5 

13.4 

14.9 

8.5 

13.6 

Naiicula  radiosa 

var.  tenelki 

3.3 

3.2 

6.2 

Saviciila  schroeteri 

var.  escambia 

5.7 

6.3 

6.2 

6.2 

10.6 

\aviciila  tripunctata 

var.  scliizonemoides 

.3.3 

\itzschia  dissipata 

6.8 

8.7 

3.6 

7.8 

3.1 

4.1 

4.6 

7.9 

3.6 

4.7 

Xitzschia  filifonnis 

3.1 

12.5 

5.4 

11.1 

3.5 

6.6 

3.2 

7.0 

5.2 

12.2 

Xitzschia  inconspicua 

15.5 

23.4 

9.5 

7.3 

12.2 

25.7 

6.2 

12.1 

10.1 

.Vitast7i ia  hatiztsch ia no 

3.1 

8.7 

4.6 

5.4 

12.3 

3.1 

Xitzschia  holsatica 

3.7 

3.3 

5.4 

.3.9 

Xitzschia  palea 

5.0 

5.7 

5.7 

7.8 

10.4 

17.7 

9.4 

12.4 

16.1 

12.0 

Xitzschia  paleacea 

10.6 

4.5 

6.8 

5.0 

3.3 

7  7 

7.4 

Rlioicosph  en  ia  cu  na  ta 

7.7 

6.2 

6.6 

9.4 

12.2 

Stephanodiscus  astrea 

1 

var.  miniitida 

3.4 

5.3 

5.2 

6.2 

5.5 

4.6 

4.1 

3.2 

43      1 

The  most  important  species  in  each  sample  are  indicated  by  boldface  type. 

Table  3.  Shannon-Wiener  diversity  values  for  the  five  living  and  five  dead  Phragmites  australis  stem  sections  from 
Provo  Ba\.  Utah  Lake. 


Sample  No. 


Livins 


Dead 


Living 


Dead 


2.96 
2.96 
2.86 
2.96 
3.07 

2.962 


3.12 
2.98 
3.09 
2.85 
3.15 

3.038 


48 
50 
33 
44 

47 

44.4 


52 
47 
50 
38 
51 

47.6 


*.\verage  number  of  species/substrate. 

An  analysis  of  the  diatom  types  unique  to 
the  living  steins  reveals  that  most  were  peri- 
phvtic  stalk  formers,  whereas  those  imique  to 
the  dead  stems  were  mostly  periphytic  mo- 
bile forms. 

The  distribution  of  species  of  Gompho- 
nema  and  Navicula  on  Phragmites  stems 
showed  significant  deviation  from  random. 
Thus,  of  a  total  of  nine  Gomphonema  species 
encoimtered  during  this  study,  eight  of  these 
occurred  on  living  stems,  five  of  which  were 
restricted  to  hving  stems.  Conversely,  of  four 
species  that  occurred  on  dead  stems,  only  one 
was  restricted  to  that  habitat.  These  data  sug- 


gest that  the  living  stems  provide  a  more 
suitable  substrate  for  several  Gomphonema 
species.  Such  species  tend  to  be  strictly  epi- 
phytic in  distribution,  usuallv  being  attached 
by  a  gelatinous  jelly  stij)e  (Patrick  and  Rei- 
mer  1966).  Whether  the  preference  of  these 
species  for  living  stems  is  relative  to  the 
availability  of  nutrients  or  the  physical  condi- 
tion of  the  substrate  is  yet  to  be  determined. 

Some  interesting  distribution  patterns  were 
also  observed  in  the  26  species  of  Navicula. 
Twenty-five  of  these  were  found  on  the  dead 
stems,  of  which  11  were  restricted  to  that 
substrate.  On  the  other  hand,  onlv  15  Xav- 


September  1980 


Grimes  et  al.:  Diatom  Assemblages 


227 


20 


30 


PERCENT    SIMILARITY 
40  50 


60 


70 

—I 


1—5    Living 
6—10    Dead 


2 

10 
4 


Fig.  1.  Cluster  dendrogram  showing  similarities  of  diatoms  on  living  and  dead  Phragmites  australis  stem  sections 
from  Provo  Bav,  Utah  Lake. 


icula  species  were  observed  on  the  living 
steins  and  only  one  taxon  was  restricted  to 
living  stems. 

We  believe  the  high  number  of  Navicula 
species  in  our  samples  can  be  accounted  for, 
at  least  in  part,  by  the  fact  that  many  are  op- 
portimistic,  occurring  on  a  wide  variety  of 
substrates.  These  opportunistic  Navicula  spe- 
cies occurred  primarily  on  dead  Phragmites 
stems  except  for  one  or  two  species  that  dom- 
inated both  living  and  dead  stems.  The  rea- 
son for  this  is  open  to  speculation,  but  it  is 
probably  related  to  nutrient  interaction,  the 
physical  condition  of  the  substrate,  or  re- 
duced competition  on  the  dead  stems. 

The  hypothesis  that  condition  of  the  Phrag- 
mites stems  had  no  effect  on  the  presence  or 
absence  of  Gomphonema  and  Naviculu  was 
tested  by  chi-square  analysis  using  a  2  X  2 
contingency  table.  The  results  departed  sig- 


nificantly from  random.  This  supported  the 
concept  that  Gomphonema  and  Navicula 
were  separated  on  the  basis  of  habitat  type. 

Consistent  with  other  Utah  Lake  studies, 
the  diatoms  in  this  study  reflect  the  condition 
of  the  lake  waters.  Most  of  the  prevalent  dia- 
toms were  either  alkaphilous  or  alkabiontic 
forms  and  also  indicators  of  eutrophy.  Addi- 
tionally, many  are  known  to  have  the  ability 
to  withstand  elevated  levels  of  dissolved  salts. 
These  data,  together  with  the  elevated  diver- 
sity found  at  Utah  Lake,  support  the  current 
view  that  Utah  Lake  is  a  saline-eutrophic 
ecosystem. 

We  recognize  the  preliminary  nature  of 
the  present  study.  Even  so,  we  believe  the 
differences  shown  in  communities  on  the  liv- 
ing versus  the  dead  stems  are  significant.  Fu- 
ture studies  are  planned  to  expand  our  data 


228 


Great  Basin  \atur.\list 


Vol.  40,  No.  3 


Table  4.  Alphabetical  list  of  diatom  genera  found  on 
Phragniites  australis  stem  sections  from  Pro%o  Bay.  Utah 
Lake,  and  the  occurrence  of  species  from  those  genera 
on  Hvine  and  dead  substrates. 


Genus 


Li\ing 


Dead       m^in 


Achnanthes 

Amphora 

Ationioeneis 

Asterionella 

Caloneis 

Cocconeis 

Coscinodisctis 

Cyclotella 

Cymbella 

Diatonia 

Diploneis 

Epithemia 

Fragilana 

Goinphonema 

Melosira 

Savicula 

Sitzschia 

Ophepbora 

Rhocosphenia 

Rhoipalodia 

Steplianodiscits 

Surirella 

Synedra 


4 
4 
1 
1 
1 
1 
1 
1 
3 
1 
1 
1 
9 
8 
3 
15 
18 
0 
1 
1 
2 
2 
6 


1 
0 

2 

9 
4 
3 
25 
17 
1 
1 
2 
2 

■7 


base  to  the  other  species  of  emergent  macro- 
ph\1:es  in  Utah  Lake.  Furthermore,  we  plan 
studies  to  answer  the  following  questions:  (1) 
Are  some  epiphytes  host  specific?  (2^  What 
patterns  of  seasonal  succession  are  evident  in 
the  epiphytic  flora?  (3)  \Miat  impact  does  the 
epiphxtic  flora  have  on  productivity  and  tro- 
phic structure  of  the  lake?  These  questions 
take  on  added  significance  for  future  re- 
source management  in  light  of  proposed 
large-scale  changes  in  Utah  Lake,  such  as  the 
dikino;  of  Provo  and  Goshen  bavs. 


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19.37.  .\n  ecological  and  taxonomic  investigation 

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Hough,  R.  A.,  .\nd  R.  G.  Wetzel.  1975.  The  release  of 
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McIntire,  C.  D..  and  W.  S.  Overton.  1971.  Distribu- 
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1979.  Algal  response  to  a  thermal  effluent:  study 
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POISONOUS  PLANTS  OF  UTAH 


Jack  D.  Brotherson.'  Lee  A.  Szyska,'  and  William  E.  Evenson- 

Abstr\ct.—  a  list  of  the  major  livestock-poisoning  plants  has  been  compiled  for  the  state  of  Utah.  Two  hundred 
fifteen  taxa  representing  36  families.  119  genera,  and  209  species  occur  within  the  state.  Fortv-one  percent  are  from 
two  families,  the  Asteraceae  and  the  Fabaceae.  The  remaining  families  of  major  imjx)rtance  are:  Poaceae,  Ranuncu- 
laceae.  Solanaceae.  Chenopodiaceae.  Brassicaceae.  .\scelpiadaceae,  Liliaceae.  and  Euphorbiaceae.  Sixtv-nine  percent 
of  the  genera  occur  with  a  single  species.  Thirty-three  percent  of  the  ta.xa  are  introduced  to  the  state.  .Most  of  the 
plants  are  insect  pollinated;  57  percent  are  herbaceous  perennials. 

Most  livestock  poisoning  occurs  during  the  spring.  This  is  due  both  to  concentration  of  toxins  in  emerging  vegeta- 
tion and  to  the  absence  of  more  suitable  forage  on  late  winter  and  spring  ranges.  Green  herbage  is  poisoning  in  about 
80  percent  of  all  taxa,  seeds  and  fruits  in  about  15  percent,  and  the  remaining  5  percent  have  toxic  compounds 
confined  to  flower  heads,  sap.  tubers,  or  roots.  Disturbed  or  cultivated  habitats  and  poorlv  managed  range  harbor  the 
greatest  diversity  of  poisonous  plants.  Wetlands  contain  fewer  poisonous  taxa  than  do  xeric  or  mesic  areas. 

The  predominating  plant  toxins  are  various  alkaloids  and  glycosides.  Sixteen  percent  of  the  plants  have  un- 
characterized  toxins.  Cattle  and  sheep  are  more  susceptible  to  poisoning  than  are  horses,  swine,  or  poultrw 


Records  document  man's  encounters  with 
poisonous  plants  since  ancient  times.  Thev 
have  played  both  positive  and  negative  roles 
in  human  cult\ire  (Dayton  1948).  This  con- 
spicuous dualitv  of  poisonous  plants  remains 
a  major  concern  for  range  management. 
Kingsbury's  (1964)  manual  on  the  poisonous 
plants  of  the  United  States  and  Canada  was 
designed  to  aid  veterinarians  and  ranchers  in 
recognizing  poisonous  plants  and  the  svmp- 
toms  they  produce  in  poisoned  livestock.  Val- 
entine (1978^  prepared  an  extensive  bibliog- 
raphv  on  the  poisonous  plants  of  American 
rangelands,  and  numerous  works  have  been 
published  dealing  with  local  species  lists  and 
descriptions  (Evers  1972,  Mihalopoulus  1974, 
Schmutz  et  al.  1968.  Stoddard  et  al.  1949. 
USDA  1968). 

The  scope  of  this  paper  is  twofold:  to  pro- 
\"ide  a  list  of  taxa  of  the  major  poisonous 
plants  of  Utah,  and  to  present  some  general 
patterns  observed  among  poisonous  plants 
within  the  state.  It  is  hoped  that  this  anno- 
tated compilation  and  discussion  will  prove 
useful  to  range  managers  and  biologists  alike. 

NL\TERIALS  .\.ND  METHODS 

Data  on  poisonous  taxa  were  gleaned  from 
the  published  literature  and  by  consultation 


with  specialists  in  botany  and  toxicolog}-. 
Much  of  the  descriptive  literature  on  poi- 
sonous plants  is  redundant,  consequently, 
only  the  more  recent  works  are  cited  here. 

Criteria  used  in  compiling  the  list  of  poi- 
sonous plants  were: 

1.  The  ta.xon  had  to  be  sufficientlv  abun- 
dant I  either  native  or  introduced)  in 
natural  ecosystems  to  constitute  a  legiti- 
mate threat  to  livestock  or  wildlife.  For 
example,  some  species  of  the  genus  As- 
tragalus are  known  to  be  toxic  but  are 
not  abundant  enough  within  the  state  to 
be  considered  dangerous  (Williams  and 
Bameby  1977). 

2.  Ornamentals  were  included  only  if  they 
have  escaped  widelv  from  cultivation. 
Such  plants  are  frequent  along  ecotones 
or  in  disturbed  habitats. 

3.  Suspicions  of  toxicity  had  to  be  reason- 
abh  well-founded.  The  genus  Astra- 
galus, for  example,  is  represented  by 
more  than  100  species  in  Utah  (Welsh 
1978),  but  only  those  ta.xa  demonstrably 
to.xic  were  included  in  the  present  list- 
ing. 

Additional  variables  considered  for  each 
taxon  were:  life  history  strategy  (annual. 
biennial,    perennial),   patchiness  of  distribu- 


Department  of  Botany  and  Range  Science.  Brigham  Young  University,  Provo,  l"tah  &4602. 
Department  of  Phvsics  and  .\stronomv.  Brigham  Young  University.  Provo,  Utah  84602. 


229 


230 


Great  Basin  Naturalist 


Vol.  40,  No.  3 


tion,  occurrence  as  a  cultivar  or  as  a  common 
range  plant,  growth  form  (vine,  forb,  grass, 
shrub,  or  tree),  generalized  habitat  require- 
ments including  elevation,  soil  texture  and 
acidity,  and  moisture  preference,  nature  of 
the  toxin  and  its  localization  within  the  plant, 
seasonality  of  poisoning,  animals  affected, 
specific  juvenile  mortality  and  abortifacient 
properties.  Not  all  variables  could  be  docu- 
mented for  each  species. 

Results 

Taxonomy 

Conservatively  estimated,  Utah  has  at  least 
215  major  toxic  taxa  of  plants  representing  36 
families,  119  genera,  and  209  species.  Thirty- 
three  percent  of  these  taxa  are  introduced  to 
the  state.  Two  of  these  36  families,  the  Aste- 
raceae  and  the  Fabaceae  contain  41  percent 
of  the  total  known  taxa  of  poisonous  plants 
(Table  1).  In  decreasing  order  of  floral  promi- 
nence, the  eight  next  important  families  are: 
Poaceae,  Ranunculaceae,  Solanaceae,  Che- 
nopodiaceae,  Brassicaceae,  Asclepiadaceae, 
Liliaceae,  and  Euphorbiaceae.  The  number 
of  toxic  taxa  within  a  family  is  not  related  to 
the  degree  of  toxicity  of  individual  taxa:  two 
of  the  most  deadly  plants,  rosary  pea  or  pre- 
catory bean  {Abms  precatorious)  and  poison 
hemlock  {Conium  maculatum)  belong  to  the 
Fabaceae  and  Apiaceae,  one  major  and  one 
relatively  minor  family  if  numbers  alone  are 
considered. 


Table  1.  Poisonous  plant  taxa  by  families. 


Family 


No.  taxa 


Percent 

of  taxa 

occurring  in 

each  family 


Fabaceae 

50 

23 

Asteraceae 

39 

18 

Poaceae 

19 

9 

Ranunculaceae 

13 

6 

Solanaceae 

11 

5 

Chenopodiaceae 

9 

4 

Brassicaceae 

8 

4 

Ascelepiadaceae 

7 

3 

Liliaceae 

7 

3 

Euphorbiaceae 

6 

3 

26  other  families 

46 

21 

Total 

215 

Sixteen  genera  (13  percent)  occur  in  the 
flora  with  more  than  two  taxa.  Twenty-two 
other  genera  (18  percent)  occur  with  two 
taxa  (including  Cannabis,  which  has  two  sub- 
species of  a  single  species).  Sixty-nine  percent 
of  the  genera  occur  with  a  single  species. 

The  taxonomy  of  poisonous  plants  is  not 
readily  explained.  The  poisonous  flora  of  the 
eastern  half  of  the  United  States  resembles 
that  of  Europe  more  than  it  does  that  of  the 
western  states  (Kingsbury  1961),  and  Day- 
ton's paper  (1948)  on  the  poisonous  plants  of 
the  continental  United  States  gives  a  differ- 
ent listing  of  major  families  than  that  found 
for  the  state  of  Utah  alone. 


Table  2.   Major  toxins  found  in   Utah's  poisonous 
plant  flora. 


Toxin 


Percent  of 

taxa 

in  which 

toxin  occurs 


Alkaloids 

22 

Glycosides 

22 

cyanogenetic  glycosides 

goitrogenic  glycosides 

irritant  oils 

coumarm 

steroids  and  triterpenoids 

Seleniinn 

11 

Nitrates 

5 

Unknown  toxins 

16 

Other  toxins 

19 

amines 

oxalates 

resins  and  resinoids 

photosensitizing  compounds 

nutritional  deficiencies 

Mechanically  injurious 

4 

Plant  Toxins 

Most  poisonous  principles  are  secondary 
by-products  of  the  plants'  metabolism  (Kings- 
bury 1964,  Levin  1976).  In  Utah's  flora,  many 
of  these  toxic  metabolites  are  loosely  classed 
as  alkaloids  and  glycosides  (Table  2).  These 
two  biochemical  groups  are  primarily  arti- 
ficial constructs,  and  each  consists  of  numer- 
ous toxins  (only  a  few  of  which  have  been 
identified)  having  generally  similar  molecular 
structures  or  modes  of  activity. 

When  the  impact  of  poisonous  range 
plants  on  livestock  mortality  is  assessed,  how- 
ever, many  of  the  minor  toxins  become  or 


September  1980 


Brotherson  et  al.:  Poisonous  Plants 


231 


serious  concern  to  ranchers.  Selenium  poison- 
ing by  members  of  the  genus  Astragalus  and 
other  species  is  an  important  source  of  mor- 
tahty,  as  is  oxalate  poisoning  by  halogeton 
Halogeton  glomeratus)  and  fivehook  bassia 
Bassia  hyssopi folia),  and  loss  of  sheep  due  to 
photosensitization  by  plants  such  as  spring 
jarsley  {Cijmoptcrus  icatsonii)  and  St.  Johns- 
.vort  {Hypericum  fowiosuui).  Thus,  the  com- 
nonness  of  a  particular  plant  species  or  toxin 
loes  not  necessarily  imply  high  mortality. 

Seasonality  and  Specificity 

Most  livestock  losses  occur  in  the  early 
pring  (Table  3)  as  animals  are  turned  out 
into  slowly  greening  range  (Evers  1972,  Kee- 
er  1978,  Kingsburv  1964,  Kreger  and  Sharp 
978,  Merrill  and  Schuster  1978).  A  second, 
mailer  peak  in  mortality  follows  in  the  sum- 
ler  as  the  more  palatable  vegetation  withers 
^  the  heat  and  toxins  are  concentrated  in 
ruits  and  seeds  of  poisonous  species. 

Cattle  are  susceptible  to  poisoning  by 
lore  taxa  than  sheep,  with  consequently 
igher  mortality  rates  and  greater  economic 
)ss  (Nielson  1978).  Most  of  the  toxic  taxa 
ommonly  found  on  Utah's  rangeland  will 
'oison  all  livestock  but  others  are  relatively 
pecies  specific  (such  as  Delphinium  poison- 
ig  in  cattle— ranges  infected  with  the  vari- 
us  species  of  this  plant  may  be  safely  grazed 
y  sheep).  Other  poisonous  taxa  may  be 
razed  in  moderate  amounts  without  harm  if 
Iternative  palatable  forage  is  available,  and 
inie  species,  for  example,  halogeton,  can  be 
tilized  by  sheep  if  the  animals  are  in- 
oduced  gradually  to  the  plant,  allowing 
leir  rumen  microflora  to  adapt  to 
etoxifying  large  amounts  of  calcium  oxalate 
ameset  al.  1976). 


Table  3.  Livestock  inortalitv  bv  seasons. 


•ason 

Percent  of 

reported  poisonings 

.ring" 

65 

miner 

17 

ill 

11 

inter 

6 

Mortality  especially  high  in  the  late  winter  and  earlv  spring 


The  Ecology  of  Poisonous  Plants 

The  ecology  of  Utah's  poisonous  flora  is 
highly  variable,  but  our  data  permit  a  few 
general  observations.  Most  of  the  plants  are 
insect  pollinated,  and  57%  are  herbaceous 
perennials.  Green  herbage  is  toxic  in  about 
80%  of  all  taxa,  .seeds  and  fruits  in  about  15%, 
and  about  5%  of  the  taxa  have  toxins  con- 
fined to  flower  head,  sap,  tubers  and  roots. 
The  relative  locations  and  concentrations  of 
toxins  within  a  plant  may  vary  through  the 
growing  season,  and  depend  upon  the  par- 
ticular taxon  being  considered. 

Toxic  species  are  likely  to  be  found  any- 
where in  the  state,  although  disturbed  habi- 
tats and  poorly  managed  rangelands  are  espe- 
cially prone  to  harbor  dangerous  species.  The 
predominance  of  poisonous  plants  in  these 
habitats  reflects  both  a  bias  in  the  reported 
literature  and  the  very  real  dangers  of  over- 
grazing in  the  western  states.  Dry  desert  soils 
have  more  taxa  of  poisonous  plants  than  mes- 
ic  or  hydric  ones;  however,  some  of  the  indi- 
vidually most  deadly  plants  occur  in  wet- 
lands. Data  on  elevation,  soil  texture  and 
acidity  were  insufficiently  complete  to  per- 
mit any  valid  conclusions. 

Discussion  and  Summary 

The  benefit  to  the  plant  of  manufacturing 
and  maintaining  toxic  substances  is  unknown. 
A  few  phytotoxins  such  as  abrin  and  cicuto- 
xin  appear  to  function  specifically  as  verte- 
brate poisons  (Kingsbury  1961,  1964)  and 
may  have  evolved  in  response  to  herbivore 
pressures  (Laycock  1978).  Selenium  accu- 
mulators are  toxic  due  to  characteristics  of 
the  soil,  although  primary  accumulators  may 
actually  require  trace  amounts  of  selenium 
for  proper  growth  (Kingsbury  1964). 

There  does  not  appear  to  be  any  univer- 
sally applicable  method  for  managing  range- 
lands  infested  with  poisonous  plants.  Species 
that  occur  in  dense  clumps  or  thickets  may 
be  individually  irradicated  by  chemical  or 
mechanical  means.  More  commonly,  how- 
ever, vast  areas  will  be  infested,  often  as  a  re- 
sult of  overgrazing.  Wise  management  will 
include  aspects  of  the  following  practices: 

1.  Recognition  of  poisonous  plants  and  an 
accurate  assessment   of  their  potential 


232 


Great  Basin  Naturalist 


Vol.  40,  No.  3 


lethality.  Unless  forced  by  hunger,  live- 
stock generally  will  ignore  poisonous 
plants  in  favor  of  more  palatable  for- 
age. 

2.  Removal  of  susceptible  animals. 

3.  Provision  of  sufficient  alternate  forage 
if  animals  must  be  turned  onto  ranges  in 
early  spring  before  palatable  plants  are 
abundant. 

4.  Scheduling  of  range  use  around  live- 
stock susceptibility  patterns. 

There  are  several  good  reviews  of  manage- 
ment practices  (Evers  1972,  Keeler  1978, 
Knieger  and  Sharp  1978,  Merrill  and  Schus- 
ter 1978),  and  the  range  literature  abounds  in 
articles  dealing  with  specific  plants  and  their 
effects  (see  Valentine  1978  for  a  com- 
prehensive listing).  The  purpose  of  the  pres- 
ent paper  is  not  to  review  management  tech- 
niques in  detail,  but  primarily  to  provide  an 
updated  listing  of  Utah's  dangerous  flora. 
Further  research  needs  to  be  done  not  only  in 
management  but  in  toxicology  and  pharma- 
cology so  that  control  of  poisonous  plants 
will  be  a  matter  of  understanding  instead  of 
irradication  and  vast  areas  of  range  can  again 
be  utilized  and  productive. 


POISONOUS  PLANTS  OF  UTAH 

The  present  list  of  plant  taxa  was  gleaned 
from  published  literature  and  by  consultation 
with  experts  in  botany  and  toxicology  as 
cited  above. 

Certain  plant  characteristics  are  desig- 
nated for  each  taxon  in  parentheses  immedi- 
ately following  the  taxon  name.  The  abbre- 
viations are  as  follows: 


p 

Perennial 

B 

Biennial 

A 

Annual 

N 

Native 

I 

Introduced 

T 

Tree 

S 

Shnib 

F 

Forb 

G 

Crass 

R 

Rush 

The  following  list  of  families,  genera,  spe- 
cies, and  varieties  is  arranged  in  alphabetical 
order  for  ease  of  reference.  Plant  synonymy 
follows  Manual  of  the  Plants  of  Colorado 
(Harrington  1964)  and  Utah  Plants  (Welsh 
and  Moore  1973). 


Amaranth ACE AE 

Amaranthus  retroflexus  L.  (AIF) 

Common  name:  pigweed,  carelessweed, 
redroot,  redroot  amaranth. 

Toxin:  nitrates,  under  conditions  of  over- 
fertilization  with  too  little  water. 

Habitat:  common  garden  and  field  weed; 
waste  places. 

Animals  affected:  livestock. 

Reference:  Brakenridge  1956,  Kingsbury 
1964. 

Anacardiaceae 

Toxicodendron   rydbergii   (Small)   Greene 

(PNS) 

Common  name:  poison  ivy. 
Toxin:  3-n-pentadecylcatechol. 
Habitat:  moist  areas  at  lower  elevations. 
Animals  affected:  humans  (dermatitis). 
Reference:  McNair  1923,  Kingsbury  1964. 

Apiaceae 

Cicuta  douglasii  (DC.)  Coult.  &  Rose  (PNF) 

Common  name:  Douglas  waterhemlock. 

Toxin:  alcohol  (cicutoxin). 

Habitat:  swampy  or  wet  habitats  along 
streams  and  in  marshes. 

Animals  affected:  livestock,  humans. 

Reference.:  Alberts  1931,  Kingsbury  1964. 

Conium  tnaculatum  L.  (BIF) 

Common  name:  hemlock,  poisonous  hem- 
lock, spotted  hemlock,  California  or  Nebras- 
ka fern. 

Toxin:  alkaloids  (conine,  N-methyl  conine, 
conhydrine,  lambda-coniceine,  pseudoconhy- 
drine). 

Habitat:  weed  of  roadsides,  ditches,  edges 
of  cultivated  fields  and  other  waste  areas. 

Animals  affected:  livestock,  humans. 

Reference:  Penny  1953,  Kingsbury  1964. 

Daucus  carota  L.  (BIF) 

Common  name:  wild  ;parrot.  Queen  Ann's 
lace. 

Toxin:  nitrates,  under  conditions  of  over- 
fertilization  with  too  little  water. 

Habitat:  waste  places. 

Animals  affected:  horses,  cattle. 

Reference:  Volker  1950,  Kingsbury  1964. 

Pastinaca  sativa  L.  var.  sylvestris  DC.  (BIF) 

Common  name:  common  parsnip. 
Toxin:  unknown. 


September  1980 


Brotherson  et  al.:  Poisonous  Plants 


233 


Habitat:  widely  naturalized  weed. 
Animals  affected:  humans  (dermatitis). 
Reference:  S.  L.  Welsh  (personal  commu- 
nication, 1980). 

Slum  suave  Walt.  (PNF) 

Common  name:  water  parsnip,  hemlock 
water  parsnip. 

Toxin:  unknown. 

Habitat:  marshy  lands  and  wet  soils. 

Animals  affected:  hogs,  cattle. 

Reference:  Fyles  1920,  Kingsbury  1964. 

Apocynaceae 

Apocynum  androsaemifolium  L.  (PNF) 

Common  name:  spreading  dogbane. 

Toxin:  resins,  glycosides. 

Habitat:  common  weed  of  open  places,  in 
coarse  soils  along  streams,  meadows,  and 
wooded  hillsides. 

Animals  affected:  cats,  dogs. 

Reference:  Moore  1909,  Kingsbury  1964. 

Note:  The  closely  related  species  A.  me- 
dium Greene  and  A.  sibericwn  Jacq.  niav 
show  similar  effects. 

Apocynum  cannabinum  L.  (PNF) 

Common  name:  Indian  hemp,  dogbane, 
hemp  dogbane. 

Toxin:  resins,  glycosides. 

Habitat:  common  weed  of  open  places,  in 
coarse  soils  along  streams. 

Animals  affected:  cats,  dogs. 

Reference:  Finnemore  1909,  Kingsbury 
1964. 

Nerium  oleander  L.  (PIS) 

Common  name:  oleander. 

Toxin:  glycosides. 

Habitat:  cultivated  greenhouse  plant, 
street  plant  in  St.  George. 

Animals  affected:  livestock,  humans. 

Reference:  West  1957,  Kingsbury  1964. 

Asclepiadaceae 

Asclepias  asperula  (Decne)  Woodson  (PNF) 

Common  name:  asper  milkweed,  spider 
antelopehorn. 

Toxin:  resinoids,  glycosides  and  an  alka- 
loid. 

Habitat:  open  dry  soils,  flats,  desert  swales, 
sandy  or  rocky  hillsides  with  pinyon,  juniper 
or  oak. 


Animals  affected:  sheep,  cattle,  goats, 
horses,  poultry. 

Reference:  Huffman  1956,  Kingsbury  1964. 

Asclepias  fascicularis  Decne  ex  DC.  (PNF) 

Common  name:  Mexican  whorled  milk- 
weed. 

Toxin:  resinoids,  glycosides  and  an  alka- 
loid. 

Habitat:  dry  hillsides  and  roadsides;  pas- 
tures, moist  streamsides. 

Animals  affected:  sheep,  cattle,  goats, 
horses,  fowl. 

Reference:  Schmutz  et  al.  1968. 

Asclepias  incarnata  L.  (PNF) 

Common  name:  swamp  milkweed. 

Toxin:  resinoids,  glycosides  and  an  alka- 
loid. 

Habitat:  marshes. 

Animals  affected:  sheep,  cattle,  horses, 
poultry. 

Reference:  Hansen  1924,  Kingsbury  1964. 

Asclepias  labriformis  Jones  (PNF) 

Common  name:  labriform  milkweed. 

Toxin:  resinoids,  glycosides  and  an  alka- 
loid. 

Habitat:  in  sandy  soils  along  old  stream 
beds. 

Animals  affected:  sheep. 

Reference:  Holmgren  1945,  Kingsbury 
1964. 

Asclepias  latifolia  (Torr.)  Raf.  (PNF) 

Common  name:  broadleaf  milkweed. 

Toxin:  resinoids,  glycosides  and  an  alka- 
loid. 

Habitat:  dry  plains  in  sandy  soils. 

Animals  affected:  sheep. 

Reference:  Schmutz  et  al.  1968,  Kingsbury 
1964,  Shrift  1958. 

Asclepias  speciosa  Torr.  (PNF) 

Common  name:  showy  milkweed. 

Toxin:  resinoids,  glycosides,  and  an  alka- 
loid. 

Habitat:  prairies  and  open  areas. 

Animals  affected:  sheep. 

Reference:  Fleming  1920,  Kingsbury  1964. 

Asclepias  subverticillata  (Gray)  Vail  (PNF) 

Common  name:  whorled  milkweed,  west- 
ern whorled  milkweed. 

Toxin:  resinoids,  glycosides  and  an  alka- 
loid. 


234 


Great  Basin  Naturalist 


Vol.  40.  No.  3 


Habitat:  dry  plains  and  foothills;  spreads 
rapidlv  along  waterways  and  irrigation 
canals,  forming  dense  stands;  prefers  sandy 
soils. 

.\niinals  affected:  sheep. 

Reference:  Glover  1917,  Kingsbiir\-  1964. 

.\steraceae 

Achillea  miJlefolium  L.  iPNF) 

Common  name:  \arrow. 

To.xin:  alkaloids  and  glycosides. 

Habitat:  various. 

Animals  affected:  livestock. 

Reference:  Hurst  1942,  Kingsbury  1964. 

Ambrosia  tomentosa  Nutt.  (PNF^ 

Common  name:  white  ragweed,  skeleton 
leaf  bursage 

To.xin:  nitrates,  imder  conditions  of  over- 
fertihzation  with  too  little  water. 

Habitat:  dr\"  plains,  hills,  waste  ground  and 
fields. 

Animals  affected:  livestock. 

Reference:  Huffman  1956,  Kingsbury  1964. 

Anthemis  cotula  L.  (AIF) 

Common  name:  dog  fennel,  mayweed. 
ma\Aveed  camomile. 

Toxin:  acrid  substance  irritating  to  mucous 
membranes. 

Habitat:  weedy  plant  of  disturbed  soils, 
fields  and  waste  places;  common  weed  in 
hay. 

.\nimals  affected:  poultry". 

Reference:  Los  Angeles  Count}-  Livestock 
Department  1938,  Kingsbur>-  1964. 

Artemisia  fiUfoUa  Torr.  (PNS^ 

Common  name:  sand  sagebrush,  old  man 
sagebrush. 

Toxin:  volatile  oils. 

Habitat:  sandy  soils. 

Animals  affected:  horses. 

Reference:  Beath  1953,  Kingsbur>-  1964. 

Artemisia  spinescens  (DC.)  Eaton   PNS) 
Common  name:  bud  sagebrush. 
Toxin:  volatile  oils. 
Habitat:  dry  plains  and  hills. 
Animals  affected:  hvestock. 
Reference:  Sampson  1942,  Kingsbur\-  1964. 

Aster  chilensis  Nees  ssp.  adscendens  (Lindl.) 
Cronq,  PNFi 

Common  name:  pacific  aster. 

Toxin:  secondary-  selenium  accumulator. 


Habitat:  widely  scattered  in  moist  habitats. 
Animals  affected:  livestock. 
Reference:    Trealease   and   Beath    1949. 
Kingsbury  1964. 

Aster  glaucodes  Blake  i^PNF^ 

Common  name:  gray  aster. 

Toxin:  secondary-  selenium  accumulator. 

Habitat:  moimtains. 

Animals  affected:  sheep. 

Reference:  Trelease  and  Beath  1949, 
KingsbuT)"  1964. 

Aster  laevis  L.  (PNF) 

Common  name:  smooth  aster. 

Toxin:  secondarv  selenium  accumulator. 

Habitat:  widely  scattered  in  drv  to  moist 
habitats. 

Animals  affected:  livestock. 

Reference:  Trelease  and  Beath  1949. 
Kingsbur)  1964. 

Aster  occidentalis  (Nutt.)  Torr.  6c  Grav 
iPNF) 

Common  name:  western  aster. 

To.xin:  secondary-  selenium  acctunulator. 

Habitat:  mountain  meadows  at  moderate 
elevations. 

Animals  affected:  livestock. 

Reference:  Trelease  and  Beath  1949, 
Kingsbury  1964. 

Aster  pauciflorus  Nutt.  (PNF) 

Common  name:  fewhead  aster. 

Toxin:  secondary-  selenium  accumulator. 

Habitat:  widespread  in  saline  soils. 

Animals  affected:  livestock. 

Reference:  J.  D.  Brotherson  (personal  com- 
munication. 19801 

Bahia  oppositifolia  (Nutt.)  DC.  iPNF) 

Common  name:  bahia.  plains  bahia. 
Toxin:  cvanogenetic  glvcoside. 
Habitat:  dr)'  soils;  plains  and  hills. 
Animals  affected:  cattle,  sheep. 
Reference:    Deem   et  al.    1939.   Kingsburv 
1964. 

Baileya  midtiradiata  Har\-.  &  Gray  (BNF) 

Common  name:  desert  baileya.  cloth  of 
gold,  desert  marigold. 

Toxin:  unknown. 

Habitat:  sandy  and  gravelly  soils  in  dr\' 
areas. 

Animals  affected:  sheep,  goats. 

Reference:  Mathews  1933,  Kingsburv 
1964. 


September  1980 


Brotherson  et  al.:  Poisonous  Plants 


235 


Baileya  pleniradiata  Har\ .  &  Gray  (ANF) 

Common  name:  desert  marigold  baileya. 

Toxin:  unknown. 

Habitat:  mesas  and  deserts  of  southeastern 
Utah. 

Animals  affected:  sheep,  goats. 

Reference:  Schmutz  et  al.  1968. 

Centaurea  repens  L.  (PIF) 

Common  name:  Russian  knapweed. 

Toxin:  unknown;  produces  nigropallidal 
encephalomalacia. 

Habitat:  fields,  roadsides  and  waste  places. 

^Animals  affected:  horses. 

Reference:  Mecke  1979. 

Centaurea  sohtitialis  L.  (AIF) 

Common  name:  yellow  star  thistle,  vellow 
centaurea. 

Toxin:  imknown;  produces  nigropallidal 
encephalomalacia;  also  mechanically  in- 
jurious. 

Habitat:  waste  places,  fields  and  roadsides. 

Animals  affected:  horses. 

Reference:  Mettler  and  Stem  1963,  Kings- 
bur>-  1964. 

Chrysothamnus    nauseosus    (Pall.)    Britton 

(PXS) 

Common  name:  rubber  rabbitbrush. 

Toxin:  unknown. 

Habitat:  dr\.  open  places  at  moderate  and 
low  elevations. 

Animals  affected:  livestock. 

Reference:  Sampson  1942,  Kingsbury  1964. 

Gritidelia  squarrosa  (Pursh)  Dunal  (BXF) 

Common  name:  gum  weed,  gumplane,  cur- 
lycup  gumweed. 

To.xin:  secondarv  selenium  accumulator. 

Habitat:  dr\  open  places;  prairies,  plains, 
roadsides  and  fields. 

Animals  affected:  livestock. 

Reference:  Trelease  and  DiSomma  1960, 
Kingsbury  1964. 

Heleiiium  autumnaJe  L.  (PXF^ 

Common  name:  sneezeweed.  bitterweed. 
Toxin:  unknown  acrid  substance. 
Habitat:  moist  low  ground  in  lowlands  and 
foothills. 

Animals  affected:  sheep  and  cattle. 
Reference:  Kingsbury  1964. 

Helenium  hoopesii  Gray  (PNF) 

Common  name:  sneezeweed.  orange 
sneezeweed. 


Toxin:  glycoside  (dugaldine). 

Habitat:  high  mountain  slopes  and  vallevs, 
often  forming  dense  stands  in  moist,  sunnv, 
undisturbed  localities. 

Animals  affected:  sheep  and  cattle. 

Reference:  Marsh  et  al.  1921,  Kingsbury 
1964. 

Heleomeris  longifolia  Rob.  &  Greenm.  var. 
annua  (Jones)  Yates  (AXF) 

Common  name:  annual  goldeneye,  resin- 
weed,  talloweed. 

Toxin:  unknown. 

Habitat:  ranges,  hills,  plains,  and  river  bot- 
toms. 

Animals  affected:  cattle. 

Reference:  Schmutz  et  al.  1968,  Kingsbury 
1964. 

Hymenoxys  richardsonii  (Hook.)  Cockerel! 
iPXF 

Common  name:  pingue,  Colorado  rubber- 
weed,  pingue  hymeno-xys,  rubberweed. 

Toxin:  unknown;  may  be  associated  with 
mineral  imbalance. 

Habitat:  dry.  rocky  or  clay  soils  of  plains 
and  mountain  slopes  from  1500  to  12,000 
feet. 

Animals  affected:  sheep,  cattle,  goats. 

Reference:  Aanes  1961,  Kingsbury  1964. 

Oxytenia  acerosa  Xutt.   PXS) 

Common  name:  copperweed;  prickly  oxy- 
tenia. 

Toxin:  unknown. 

Habitat:  alkaline  soils  in  draws  or  stream- 
beds  of  desert  ranges  and  foothills. 

Animals  affected:  cattle,  sheep. 

Reference:  Throp  et  al.  1940,  Kingsburv 
1964. 

Psathyrotes  annua  (Nutt.)  Gray  i.\XF) 

Common  name:  annual  psathyrotes. 

Toxin:  unknown. 

Habitat:  dry,  sandy,  often  alkaline  soils,  es- 
pecially of  creek  beds  and  dr)-  washes. 

.\nimals  affected:  sheep. 

Reference:  Binns  et  al.  1962,  Kingsbur\ 
1964. 

Psilostrophe  sparsiflora  (Gray)  A.  Nels. 
(PXF) 

Common  name:  greenstem  paperflower. 

Toxin:  unknown;  induces  kidney  damage. 

Habitat:  dr\-,  open  range. 


236 


Great  Basin  Naturalist 


Vol.  40,  No.  3 


Animals  affected:  sheep. 

Reference:  Huffman  1956,  Kingsbury  1964. 

Rudbeckia  occidentalis  Nutt.  (PNF) 

Common  name:  western  coneflower,  nig- 
gerheads. 

Toxin:  unknown. 

Habitat:  streambanks  and  woodlands. 

Animals  affected:  generally  unpalatable  to 
livestock;  affects  hogs  and  sheep  in  feeding 
trials. 

Reference:  Pammel  1911,  Kingsbury  1964. 

Senecio  integerrimus  Nutt.  (PNF) 

Common  name:  groundsel,  senecio,  lambs- 
tongue  groimdsel. 

Toxin:  alkaloids. 

Habitat:  dry  or  moist  open  woods  and 
slopes,  from  valleys  to  near  timberline. 

x\nimals  affected:  livestock,  humans  (?). 

Reference:  Clawson  1933,  Kingsbury  1964. 

Senecio  longilobus  Benth.  (PNF) 

Common  name:  wooly  groundsel,  thread- 
leaf  groundsel. 

Toxin:  pyrrolizidine  alkaloids. 

Habitat:  dry  slopes,  mesas  and  dry  washes. 

Animals  affected:  cattle,  horses,  sheep, 
goats. 

Reference:  Clawson  1933,  Kingsbury  1964. 

Senecio  spartioides  Torr.  &  Gray  (PNF) 

Common  name:  broom  groundsel. 

Toxin:  pyrrolizidine  alkaloids. 

Habitat:  valleys,  plains;  open  areas  and 
pine  forests. 

Animals  affected:  cattle,  horses,  sheep, 
goats,  humans(?). 

Reference:  Clawson  1933,  Kingsburv  1964, 
Davis  1958. 

Senecio  vulgaris  L.  (AIF) 

Common  name:  common  groundsel. 

Toxin:  pyrrolizidine  alkaloids. 

Habitat:  weed  of  gardens  and  waste  places. 

Animals  affected:  cattle,  horses,  sheep, 
goats,  humans(?). 

Reference:  Steyn  1934,  Kingsbury  1964. 

Solidago  parryi  (Gray)  Greene  (PNF) 

Common  name:  Parry  goldenweed. 

Toxin:  unknown;  causes  milk-sickness  or 
trembles. 

Habitat:  mountains,  coniferous  forests. 

Animals  affected:  cattle. 


Reference:  Schmutz  et  al.  1968,  Kingsburv 
1964. 

Tanacetum  vulgare  L.  (PIF) 

Common  name:  common  tansv. 
Toxin:  abortifacient. 

Habitat:  weed  along  roadsides,  waste  areas, 
ditchbanks  and  other  moist  areas. 
Animals  affected:  cattle,  humans(?). 
Reference:  Cress  1935,  Kingsbury  1964. 

Tetradymia  canescens  DC.  (PNS) 

Common  name:  spineless  horsebrush,  gray 
horsebrush. 

Toxin:  photosensitizing  compounds. 

Habitat:  dry  desert  and  sagebrush  ranges. 

Animals  affected:  sheep. 

Reference:  Kingsburv  1964,  Schmutz  et  al. 
1968. 

Tetradymia  glabrata  Gray  (PNS). 

Common  name:  littleleaf  horsebrush, 
spring  rabbitbrush,  coaloil  brush. 

Toxin:  photosensitizing  compounds. 

Habitat:  dry  desert  and  sagebrush  ranges. 

Animals  affected:  sheep. 

Reference:  Kingsburv  1964,  Fleming  et  al. 
1922. 

Tetradymia  ntittallii  T.  &  G.  (PNS) 

Common  name:  Nuttall  horsebrush. 
Toxin:  photosensitizing  compounds. 
Habitat:  dry  desert  and  sagebaish  ranges. 
Animals  affected:  sheep. 
Reference:  Kingsbury  1964. 

Tetradymia  spinosa  T.  and  G.  var.  long- 
ispina  Jones  (PNS) 

Common  name:  longspine  horsebrush. 

Toxin:  photosensitizing  compounds. 

Habitat:  dry  desert  and  sagebrush  ranges. 

Animals  affected:  sheep. 

Reference:  S.  L.  Welsh  (personal  commu- 
nication, 1980). 

Tetradymia  spinosa  T.  and  G.  var.  spinosa 

(PNS) 

Common  name:  spinv  horsebrush. 

Toxin:  photosensitizing  compounds. 

Habitat:  dry  desert  and  sagebrush  ranges. 

Animals  affected:  sheep. 

Reference:  S.  L.  Welsh  (personal  commu- 
nication, 1980). 

Xanthium  strumarium  L.  (AIF) 

Common  name:  spiny  clotbur,  spiny  cock- 
lebur. 


September  1980 


Brotherson  et  al.:  Poisonous  Plants 


237 


Toxin:  hydroquinone. 

Habitat:  fields  and  wastelands;  along 
shores  of  ponds,  rivers  and  in  flood  plains. 

.\nimals  affected:  livestock,  fowl,  hogs,  hu- 
mans (dermatitis). 

Reference:  Forrest  1938,  Kuzel  and  Miller 
1950,  Kingsbury  1964. 

Xanthocephalum  microcephahim  (DC.) 
Gray  (PNS) 

Common  name:  broomweed,  perennial 
snakeweed,  slinkweed,  turpentine  weed, 
threadleaf  snakeweed,  matchweed,  resin- 
weed. 

Toxin:  saponin. 

Habitat:  dry  stony  plains,  slopes  and  mesas. 

Animals  affected:  cattle,  sheep,  goats, 
swine,  chicks,  rabbits. 

Reference:  Dollahite  1957,  Kingsburv 
1964. 

Xanthocephalum  sarothrae  (Pursh)  Britt. 
and  Rushy  (A\F) 

Common  name:  broom  snakeweed,  snake- 
weed, matchbnish. 

Toxin:  saponin. 

Habitat:  dry  stony  plains,  slopes  and  mesas. 

Animals  affected:  cattle,  sheep,  goats, 
swine,  chicks,  rabbits. 

Reference:  Dollahite  1962,  Kingsburv 
1964. 

BORAGINACEAE 

Amsinckia  intermedia  Fisch,  &  Mey  (AXF) 

Common  name:  tarweed,  fiddleneck,  fire- 
weed  fiddleneck. 

Toxin:  imknown;  potentially  lethal  nitrate 
levels,  pyrrolizidine  alkaloids  (?). 

Habitat:  dry  open  cultivated  ground  or 
waste  areas. 

Animals  affected:  horses,  hogs,  cattle. 

Reference:  McCulloch  1940,  Kingsburv 
1964. 

Note:  The  closelv  related  species  A.  tesse- 
lata  Gray  and  A.  retrorsa  Suksd.  may  show 
similar  effects. 

Cynoglossiim  officinale  L.  (BIF) 

Common  name:  houndstongue. 

Toxin:  unknown. 

Habitat:  waste  places  of  plains  and  hills. 

Animals  affected:  livestock. 

Reference:  Kingsbury  1964,  S.  L.  Welsh 
(personal  commimication,  1980). 


Brassicace.\e 

Brassica  hirta  Moench.  (AIF) 

Common  name:  white  mustard. 

Toxin:  cyanogenetic  glycoside. 

Habitat:  cultivated  weed,  escaped  to  waste 
areas. 

Animals  affected:  cattle,  sheep. 

Reference:  Eaton  1941,  Kingsbury  1964. 

Brassica  kaber  Wheeler  (.\1F) 

Common  name:  charlock,  wild  mustard. 

Toxin:  cyanogenetic  glycoside. 

Habitat:  common  weed  of  grain  crops  and 
in  waste  areas. 

Animals  affected:  cattle,  hogs,  sheep. 

Reference:  Thomson  and  Sifton  1922, 
Kingsbm-y  1964. 

Descurainia  pinnata  (Walt.)  Britt.  (ANF) 

Common  name:  tansv  mustard,  pinnate 
tansy  mustard. 

Toxin:  unknown. 

Habitat:  heavy  stands  on  dry,  sandy  soils. 

Animals  affected:  cattle. 

Reference:  Hershev  1935,  Kingsburv  1964. 

Erysimum  cheiranthoides  L.  (ANF) 

Common  name:  wormweed  mustard, 
treacle  wallflower. 

Toxin:  cvanogenetic  glycoside. 

Habitat:  weed  of  cultivation,  roadsides, 
meadows;  moist  waste  areas  in  valleys  and 
canyons. 

Animals  affected:  hogs. 

Reference:  Thomson  and  Sifton  1922, 
Kingsbur)  1964. 

Stanleya  integrifolia  James  (PXS) 

Common  name:  wholeleaf  desert  prince's 
plume. 

Toxin:  primarv  selenium  accumulator. 

Habitat:  dry  plains  and  hills. 

Animals  affected:  Not  observed  to  be  eaten 
bv  livestock. 
'  Reference:    Beath   et   al.    1953,   Kingsbur\' 
1964. 

Stanleya  pinnata  (Pursh)  Britt.  (PNS) 

Common  name:  prince's  plume,  desert 
prince's  plume. 

Toxin:  primary  selenium  accumulator. 

Habitat:  desert  soils,  dry  plains  and  mesas. 

Animals  affected:  experimental;  normally 
unpalatable. 

Reference:  Beath  et  al.  1953,  Kingsbury 
1964. 


238 


Great  Basin  Naturalist 


Vol.  40,  No.  3 


Stanletja  viridiflora  Nutt.  (PNF) 

Common  name:  greenflower  prince's 
plume. 

Toxin:  primary  selenium  accumulator. 

Habitat:  dry  plains  and  hills. 

Animals  affected:  Not  observed  to  be  eaten 
by  livestock. 

Reference:  Beath  et  al.  1953,  Kingsbury 
1964. 

Thlaspi  arvense  L.  (AIF) 

Common  name:  fanweed,  field  penny- 
cress. 

Toxin:  cyanogenetic  glycoside. 

Habitat:  Common  weed  of  cultivated  and 
waste  places. 

Animals  affected:  livestock. 

Reference:  Thomson  and  Sifton  1922, 
Kingsbury  1964. 

C.wnabinaceae 

Cannabis  sativa  L.  ssp.  sativa  (AIF) 

Common  name:  marijuana,  hemp. 

Toxin:  narcotic  element  contained  in  tetra- 
hydrocannabinol. 

Habitat:  waste  places. 

Animals  affected:  humans,  livestock. 

Reference:  Steyn  1934,  Kingsbury  1964, 
Small  and  Cronquist  1976,  Welsh  1980. 

Cannabis  sativa  L.  ssp.  indica  (Lam.)  Small 
&  Cronq.  (AIF) 

Common  name:  marijuana,  hemp. 

Toxin:  narcotic  element  contained  in  tetra- 
hydrocannabinol. 

Habitat:  waste  places. 

Animals  affected:  humans,  livestock. 

Reference:  Small  and  Cronquist  1976, 
Welsh  1980. 

Caprifoliaceae 

Sambucus  coerulea  Raf.  (PNS) 

Common  name:  blue  elderberry. 

Toxin:  unknown. 

Habitat:  moist  soils  of  plains  and  hills. 

Animals  affected:  cattle,  children  (?) 

Reference:  Schmutz  et  al.  1968. 

Sambucus  racemosa  L.  (PNS) 
Common  name:  red  elder. 
Toxin:  unknown;  concentrated  in  root. 
Habitat:  moist  forests,  7,500-10,000  ft. 
Animals  affected:  cattle,  children(?) 
Reference:  Schmutz  et  al.  1968. 


Caryophyllaceae 
Saponaria  officinalis  L.  (PIF) 

Common  name:  bouncing  bet,  soapwort. 

Toxin:  saponin. 

Habitat:    fields,    waste    places;    cultivated 
and  escaping. 

Animals  affected:  sheep. 

Reference:  Kingsbury  1964. 

Chenopodiaceae 

Atriplex  gardneri  Moq.  (PNS) 

Common  name:  Nuttall  saltbush,  Gardner 
saltbush. 

Toxin:  secondary  selenium  accumulator. 

Habitat:  saline  plains  and  hillsides. 

Animals  affected:  livestock. 

Reference:  Fleming  1920,  Kingsbury  1964. 

Bassia  hyssopifolia  (Pall.)  Volk  (AIF) 

Common    name:    fivehook   bassia,    smoth- 
erweed. 

Toxin:  oxalates. 
Habitat:  dry,  saline  soils. 
Animals  affected:  sheep. 
Reference:  Pammel  1911. 

Chenopodium  ambrosioides  L.  (AIF) 

Common  name:  wormseed  goosefoot. 
Toxin:  antihelminthic  oil. 
Habitat:  weed  of  waste  places. 
Animals  affected:  geese,  humans. 
Reference:  Bamford  1951,  Kingsbury  1964. 

Chenopodium  album  L.  (AIF) 

Common  name:  lambsquarter. 

Toxin:  nitrates,  under  conditions  of  over- 
fertilization  with  too  little  water. 

Habitat:  weed  of  waste  places. 

Animals  affected:  livestock. 

Reference:  Case  1957,  Kingsbury  1964. 

Chenopodium  glaucum  L.  (AIF) 

Common  name:  oakleaf  goosefoot. 
Toxin:  nitrates,  under  conditions  of  over- 
fertilization  with  too  little  water. 
Habitat:  weed  of  waste  pmces. 
Animals  affected:  livestock. 
Reference:  Case  1957,  Kingsbury  1964. 

Halogeton  glomeratus  (Bieb.)  C.  A.  Mey 

(AIF) 

Common  name:  halogeton,  barilla. 

Toxin:  oxalates. 

Habitat:  dry  saline  plains  and  alkaline 
soils;  roadsides. 


September  1980 


Brotherson  et  al.:  Poisonous  Plants 


239 


Animals  affected:  livestock,  especially 
sheep. 

Reference:  Cook  and  Stoddart  1953,  Kings- 
bury 1964. 

Kochia  scoparia  L.  Schrad,  (AIF) 

Common  name:  summer  cypress,  burning 
bush,  Mexican  fireweed.  Belvedere  summer 
cypress. 

Toxin:  photosensitizing  compounds. 

Habitat:  dry  soils,  roadsides  and  waste 
places. 

Animals  affected:  cattle,  sheep,  horses. 

Reference:  Rottgardt  1944,  Kingsbury 
1964,  Schmutz  et  al.  1968. 

Salsola  iberica  Sennen  &  Pau  (AIF) 

Common  name:  Russian  thistle. 
Toxin:  nitrates  (?),  possible  oxalates  (?).  ' 
Habitat:  dry  soils  of  plains  and  foothills. 
Animals  affected:  livestock. 
Reference:  Huffman  et  al.  1956. 

Sarcobatus  vermiculutus  (Hook.)  Torr.  (PNS) 

Common  name:  greasewood,  black  grease- 
wood. 

Toxin:  oxalates. 

Habitat:  dense  stands  confined  to  alkaline 
flats  or  saline  soils  of  low  and  lower  middle 
elevations. 

Animals  affected:  sheep,  sometimes  cattle. 

Reference:  Kouch  1922,  Kingsbury  1964. 

Cyperaceae 

Scirpus  pungens  Vahl  (PNR) 

Common  name:  bulrush,  three-square, 
American  bulrush. 

Toxin:  unknown;  suspected  of  producing 
pulmonary  emphysema. 

Habitat:  wet  or  moist  ground. 

Animals  affected:  cattle. 

Reference:  Beath  et  al.  1953,  Kingsburv 
1964. 


Equisetaceae 

Equisetum  arvense  L.  (PNF) 

Common  name:  horsetail,  foxtail,  rush, 
marsh  horsetail. 

Toxin:  alkaloids. 

Habitat:   sandy   or  gravelly   soils  along 
streams  and  in  moist  fields  and  meadows. 

Animals  affected:  horses. 

Reference:  Gussow  1912,  Kingsbury  1964. 


Equisetum  laevigatum  A.  Br.  (PNF) 

Common  name:  smooth  horsetail. 

Toxin:  thiaminase. 

Habitat:  marshes,  alluvial  thickets,  sandy 
banks;  weed  of  cultivation. 

Animals  affected:  horses. 

Reference:  Samp.son  and  Malmsten  1942, 
Kingsbury  1964. 

Ericaceae 

Kalmia  microphylla  (Hook.)  Heller  (PNS) 

Common  name:  pale  laurel,  bog  laurel,  al- 
pine kalmia. 

Toxin:  resinoids,  andromedotoxin. 

Habitat:  wet  meadows  and  bogs  of  high 
elevation. 

Animals  affected:  sheep,  calves,  goats. 

Reference:  Kingsbury  1964. 

Ledum  glandulosum  Nutt.  (PNS) 

Common  name:  western  Labrador  tea. 

Toxin:  resinoids,  andromedotoxin. 

Habitat:  wet  meadows  and  bogs  of  high 
elevation. 

Animals  affected:  sheep,  cattle. 

Reference:  Kingsbury  1964. 

Euphorbiaceae 

Croton  longipes  Jones  (PNF) 

Common  name:  croton. 

Toxin:  croton  oil  (caustic). 

Habitat:  roadsides,  fields,  and  dry  stream- 
beds;  artemisia  and  pinyon  belts. 

Animals  affected:  livestock. 

Reference:  Schmutz  et  al.  1968,  S.  L. 
Welsh' (personal  communication,  1980). 

Croton  texensis  (Klotz.)  Muell.  Arg.  ex  DC. 

(ANF) 

Common  name:  Texas  croton. 

Toxin:  croton  oil  (caustic). 

Habitat:  roadsides,  fields,  and  dry  stream- 
beds;  artemisia  and  pinyon  belts. 

Animals  affected:  livestock. 

Reference:  Volker  1950,  Kingsbury  1964, 
Schmutz  et  al.  1968. 

Euphorbia  cyparissias  L.  (PIF) 

Common  name:  cypress  spurge,  graveyard 
weed. 

Toxin:  unknown  acrid  principle. 

Habitat:  cultivated  and  escaping  to  road- 
sides and  waste  places. 


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Vol.  40,  No.  3 


Animals  affected:  cattle. 
Reference:  Muenscher  1935,  Kingsbury 
1964. 

Euphorbia  esula  L.  (PIF) 

Common  name:  leafy  spurge. 

Toxin:  unknown  acrid  principle. 

Habitat:  field  weed  of  roadsides  and  waste 
places. 

Animals  affected:  horses,  sheep. 

Reference:  Johnston  and  Peake  1960, 
Kingsbury  1964. 

Reverchonia  arenaria  Gray  (ANF) 

Common  name:  reverchonia,  sand  rever- 
chonia. 

Toxin:  unknown. 

Habitat:  uncommon;  plains  and  hillsides, 
sandy  areas,  Kane  Co. 

Animals  affected:  sheep. 

Reference:  Schmutz  et  al.  1968,  Kingsbury 
1964. 

Ricinus  communis  L.  (AIF) 
Common  name:  castor  bean. 
Toxin:  ricin  (a  phytotoxin). 
Habitat:  cultivated  as  an  ornamental. 
Animals  affected:  livestock,  humans. 
Reference:  Clarke  1947,  Kingsbury  1964. 

F.\BACEAE 

Acacia  greggii  Gray  (PNT) 

Common  name:  catclaw  acacia. 

Toxin:  cyanogenetic  glycoside. 

Habitat:  plains  and  dry  canyons;  forms 
thickets  along  Beaver  Dam  Wash,  Washing- 
ton Co. 

Animals  affected:  sheep. 

Reference:  Schmutz  et  al.  1968,  Kingsburv 
1964. 

Astragalus  asclepiadoides  Jones  (PNF) 
Common  name:  milkweed  milkvetch. 
Toxin:  primary  selenium  accumulator. 
Habitat:  saline  desert  areas. 
Animals  affected:  livestock. 
Reference:  Welsh  1978. 

Astragalus  hisulcatus  (Hook.)  Gray  (PNF) 

Common  name:  two-grooved  poisonvetch, 
two-grooved  milkvetch. 

Toxin:  primary  selenium  accumulator. 

Habitat:  plains  and  bottom  lands,  sage- 
brush zone. 

Animals  affected:  livestock. 

Reference:  Welsh  1978,  Kingsbury  1964. 


Astragalus  convallarius  Greene  var.  con- 
vallarius  (PNF) 

Common  name:  timber  poisonvetch,  lesser 
rushy  milkvetch. 

Toxin:  produces  locoism. 

Habitat:  dry  hillsides,  desert  shrub  to  lower 
montane  zones. 

Animals  affected:  livestock. 

Reference:  Muenscher  1951,  Kingsbury 
1964. 

Astragalus  drummondii  Dougl.  ex  Hook. 

(PNF) 

Common  name:  Drummond  milkvetch. 

Toxin:  produces  locoism. 

Habitat:  plains  and  hillsides,  brushy  places. 

Animals  affected:  livestock. 

Reference:  Welsh  1978,  Kingsbury  1964. 

Astragalus  eastwoodae  Jones  (PNF) 

Common  name:  Eastwood  poisonvetch, 
Eastwood  milkvetch. 

Toxin:  primary  selenium  accumulator. 

Habitat:  dry  hillsides. 

Animals  affected:  livestock. 

Reference:  Welsh  1978,  Kingsbury  1964. 

Astragalus  flavus  Nutt.  ex  Torr.  &  Gray  var. 
argillosus  (Jones)  Barneby  (PNF) 

Common  name:  yellow  milkvetch. 

Toxin:  primary  selenium  accumulator. 

Habitat:  dry  plains  and  hillsides,  salt  desert 
areas. 

Animals  affected:  livestock. 

Reference:  Welsh  1978. 

Astragalus  flavus  Nutt.  ex  Torr.  &  Gray  var. 
candicans  Gray  (PNF) 

Common  name:  Canada  yellow  milkvetch. 

Toxin:  primary  selenium  accumulator. 

Habitat:  dry  plains  and  hillsides,  shales  and 
clays  of  southern  Utah. 

Animals  affected:  livestock. 

Reference:  Welsh  1978. 

Astragalus  flavus  Nutt.  ex  Torr.  &  Gray  var. 
flavus  (PNF) 

Common  name:  yellow  milkvetch. 

Toxin:  primary  selenium  accumulator. 

Habitat:  dry  plains  and  hillsides,  saline  silts 
and  clays  in  saline  desert  areas,  south-central 
Utah. 

Animals  affected:  livestock. 

Reference:  Welsh  1978,  Kingsbury  1964. 


September  1980 


Brotherson  et  al.:  Poisonous  Plants 


241 


Astragalus  iselyi  Welsh  (PNF) 
Common  name:  Isely  milkvetch. 
Toxin:  primary  selenium  accumulator. 
Habitat:  dry  hillsides,  salt  desert  areas. 
Animals  affected:  livestock. 
Reference:  Welsh  1978. 

Astragalus  lentiginosus  Dougl.  ex  Hook.  var. 
araneosus  (Sheld.)  Barneby  (PNF) 

Common  name:  spider  locoweed,  cobweed 
milkvetch. 

Toxin:  produces  locoism. 

Habitat:  dry  hillsides  in  sagebrush. 

Animals  affected:  livestock. 

Reference:  Welsh  1978,  Kingsbury  1964. 

Astragalus  lentiginosus  Dougl.  ex  Hook.  var. 
palans  (Jones)  Jones  (PNF) 

Common  name:  straggling  milkvetch. 

Toxin:  produces  locoism. 

Habitat:  salt  desert  areas,  dry  hillsides  and 
canyons,  mixed  desert  shrub  communities. 

Animals  affected:  livestock. 

Reference:  Welsh  1978,  S.  L.  Welsh  (per- 
sonal communication,  1980),  Kingsbury  1964. 

Astragalus  lentiginosus  Dougl.  ex  Hook.  var. 
wahweapensis  Welsh  (PNF) 

Common  name:  Wahweap  loco,  Wahweap 
milkvetch. 

Toxin:  produces  locoism. 

Habitat:  dry  hillsides  in  sagebrush,  sandy 
soils,  Kane  Co. 

Animals  affected:  livestock,  esp.  horses. 

Reference:  Welsh  1978,  Kingsbury  1964. 

Astragalus  miser  Dougl.  ex  Hook.  var.  ob- 
longifolius  (Rydb.)  Cronq.  (PNF) 

Common  name:  timber  milkvetch,  Ryd- 
berg  weedv  milkvetch. 

Toxin:  miserotoxin. 

Habitat:  widely  scattered  in  lower  mon- 
tane zones. 

Animals  affected:  livestock. 

Reference:  Williams  1969,  Welsh  1978. 

Astragalus  moencoppensis  Jones  (PNF) 

Common  name:  Moenkopi  poisonvetch, 
Moenkopi  milkvetch. 

Toxin:  primary  selenium  accumulator. 

Habitat:  heavy  soils,  salt  desert  through 
pinyon-juniper  areas. 

Animals  affected:  livestock. 

Reference:  Welsh  1978,  Kingsbury  1964. 


Astragalus  mollissimus  Torr.  (PNF) 

Common  name:  Thompson  woolly  loco- 
weed. 

Toxin:  produces  locoism. 

Habitat:  dry  plains  and  hillsides. 

Animals  affected:  livestock. 

Reference:  Welsh  1978,  Kingsbury  1964. 

Astragalus  pattersonii  Gray  ex  Brand.  (PNF) 

Common  name:  Patterson  locoweed,  Pat- 
terson milkvetch. 

Toxin:  primary  selenium  accumulator. 
Habitat:  dry  plains  and  hillsides. 
Animals  affected:  livestock. 
Reference:  Welsh  1978,  Kingsbury  1964. 

Astragalus  praelongus  Sheldon  (PNF) 

Common  name:  stinking  milkvetch. 

Toxin:  primary  selenium  accumulator. 

Habitat:  dry  plains  and  hillsides,  clay  and 
seleniferous  soils. 

Animals  affected:  livestock. 

Reference:  Welsh  1978. 

Astragalus  preussii  Gray  (PNF) 

Common  name:  Preuss  milkvetch. 

Toxin:  primarv  selenium  accumulator. 

Habitat:  dry  plains  and  hillsides,  seleni- 
ferous clays  and  silts. 

Animals  affected:  livestock. 

Reference:  Welsh  1978,  Kingsbury  1964. 

Astragalus  pubentissimus   Torr.   &    Gray 

(PNF) 

Common  name:  Green  River  locoweed. 
Green  River  milkvetch. 

Toxin:  produces  locoism. 

Habitat:  canyons,  mountainsides. 

Animals  affected:  livestock,  mainly  sheep. 

Reference:  Buck  1961,  Kingsbury  1964. 

Astragalus  racemosus  Pursh  var.  treleasi  Por- 
ter (PNF) 

Common  name:  alkali  milkvetch. 

Toxin:  primary  selenium  accumulator, 
causes  "alkali  disease"  or  "blind  staggers." 

Habitat:  Uinta  and  Duchesne  River  forma- 
tions. 

Animals  affected:  cattle. 

Reference:  Welsh  1978. 

Astragalus  rafaelensis  Jones  (PNF) 
Common  name:  San  Rafael  milkvetch. 
Toxin:  primary  selenium  accumulator. 
Habitat:   seleniferous  clays  and  silts,   salt 

desert  shrub  zones,  Emery  Co. 


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Great  Basin  Naturalist 


Vol.  40,  No.  3 


Animals  affected:  livestock. 
Reference:  Welsh  1978. 

Astragalus  sabuhsus  Jones  (PNF) 

Common  name:  straightstem  poisonvetch, 
Cisco  milkvetch. 

Toxin:  primary  selenium  accumulator. 

Habitat:  salt  desert  shrub  zone  on  shales. 

Animals  affected:  livestock. 

Reference:  Welsh  1978,  Marsh  1935. 

Astragalus  saurinus  Barneby  (PNF) 

Common  name:  dinosaur  milkvetch. 

Toxin:  primary  selenium  accumulator. 

Habitat:  salt  desert  shrub  and  pinyon-juni- 
per  zone,  Uintah  Co. 

Animals  affected:  livestock. 

Reference:  Welsh  1978. 

Astragalus  tetrapterus  Gray  (PNF) 

Common  name:  fourwing  poisonvetch, 
four-wing  milkvetch. 

Toxin:  produces  locoism. 

Habitat:  plains,  dry  hillsides. 

Animals  affected:  cattle,  sheep. 

Reference:  Marsh  1920,  Welsh  1978. 

Astragalus  toanus  Jones  (PNF) 
Common  name:  Toano  milkvetch. 
Toxin:  primary  selenium  accumulator. 
Habitat:  clay  soils,  salt  desert  shrub  zone. 
Animals  affected:  livestock. 
Reference:  Welsh  1978,  Kingsbury  1964. 

Astragalus  woodruffii  Jones  (PNF) 

Common  name:  Woodruff  milkvetch. 

Toxin:  primary  selenium  accumulator. 

Habitat:  desert  shnib  zone  on  sandy  and 
sandy-silt  soils. 

Animals  affected:  livestock. 

Reference:  Welsh  1978. 

Lathyrus  latifolius  L.  (PIF) 

Common  name:  perennial  peavine,  per- 
ennial sweet  pea. 

Toxin:  alpha,  gamma-diaminobutyric  acid. 

Habitat:  rangeland. 

Animals  affected:  rats  (experimental), 
sheep  (lathyrism). 

Reference:  Lewis  1949,  Kingsbury  1964,  S. 
L.  Welsh  (personal  commvmication,  1980). 

Lathyrus  sylvestris  L.  (PNF) 

Common  name:  everlasting  sweetpea,  flat 
pea. 

Toxin:  alpha,  gamma-diaminobutyric  acid. 


Habitat:  rangeland. 

Animals  affected:  sheep,  rats  (experimen- 
tal). 

Reference:  Lewis  1948,  Kingsbury  1964. 

Lupinus  alpestris  A.  Nels.  (PNF) 

Common  name:  mountain  lupine. 

Toxin:  alkaloids. 

Habitat:  mountains. 

Animals  affected:  cattle,  sheep. 

Reference:  Beath  et  al.  1953,  Kingsbury 
1964. 

Lupinus  arbustus  Dougl.  ex  Lindl.  var.  cal- 
caratus  (Kellogg)  Welsh  (PNF) 

Common  name:  Douglas  spurred  lupine, 
spur  lupine. 

Toxin:  alkaloids. 

Habitat:  hillsides,  dry  soils. 

Animals  affected:  cattle,  horses,  sheep. 

Reference:  Clawson  1931,  Kingsbury  1964. 

Lupinus  argenteus  Pursh  (PNF) 

Common  name:  silvery  lupine. 

Toxin:  alkaloids. 

Habitat:  dry  flats  and  slopes  in  woods, 
plains  and  hills. 

Animals  affected:  sheep,  cattle,  horses, 
goats,  hogs,  deer. 

Reference:  Marsh  1916,  Kingsbury  1964. 

Lupinus  caudatus  Kell.  (PNF) 

Common  name:  Kellogg's  spurred  lupine, 
tailcut  lupine. 

Toxin:  alkaloids. 

Habitat:  exposed  hillsides. 

Animals  affected:  cattle. 

Reference:  Animal  Disease  and  Parasite 
Research  Division  USDA  1958,  Kingsbury 
1964. 

Lupinus  leucophyllus  Dougl.  (PNF) 

Common  name:  woolly-leaved  lupine,  vel- 
vet lupine. 

Toxin:  alkaloids. 

Habitat:  dry  soil,  foothills. 

Animals  affected:  sheep,  cattle,  horses, 
goats,  hogs,  deer. 

Reference:  Marsh  1916,  Kingsbury  1964. 

Lupinus  pusillus  Pursh  (ANF) 

Common  name:  low  lupine,  rusty  lupine. 

Toxin:  alkaloids. 

Habitat:  dry  plains,  foothills. 

Animals  affected:  sheep. 

Reference:  Sampson  1942,  Kingsbury  1964. 


September  1980 


Brotherson  et  al.:  Poisonous  Plants 


243 


Lupinus  sericeus  Pursh  (PNF) 

Common  name:  silky  lupine. 

Toxin:  alkaloids. 

Habitat:  dry  hillsides  and  valleys. 

Animals  affected:  sheep,  cattle,  horses, 
goats,  hogs,  deer. 

Reference:  Binns  and  James  1961,  Kings- 
bury 1964. 

Medicago  sativa  L.  (PIF) 

Common  name:  alfalfa,  lucerne. 
Toxin:  saponin. 

Habitat:  cultivated  and  escaping. 
Animals   affected:    cattle,   chicks,   hogs, 
sheep. 

Reference:  Walter  1954,  Kingsbury  1964. 

Melilotus  alba  Desr.  (BIF) 

Common  name:  white  sweetclover. 

Toxin:  dicoumarin. 

Habitat:  waste  places  and  fields,  escaped 
from  cultivation. 

Animals  affected:  cattle. 

Reference:  Roderick  1931,  Kingsbury 
1964. 

Melilotus  officinalis  L.  Lam.  (BIF) 

Common  name:  yellow  sweetclover. 

Toxin:  dicoumarin. 

Habitat:  waste  ground,  fields;  used  for  for- 
age and  fertilizer. 

Animals  affected:  cattle,  sheep,  horses. 

Reference:  Roderick  1931,  Kingsbury 
1964. 

Oxytropis  lambertii  Pursh  (PNF) 

Common  name:  white  loco,  white  paint 
loco,  silky  crazyweed. 

Toxin:  unknown  alkaloids;  produces 
locoism. 

Habitat:  prairies  and  mountains,  usually  in 
drier  areas,  lower  to  middle  elevations. 

Animals  affected:  livestock. 

Reference:  Couch  1929,  Kingsbury  1964. 

Oxytropis  sericea  Nutt.  ex  Torr.  &  Gray 

(PNF) 

Common  name:  white  paint  loco,  silky  cra- 
zyweed. 

Toxin:  alkaloids;  produces  locoism. 

Habitat:  open  gravelly  or  well-drained 
slopes  and  hills  at  lower  to  middle  elevations. 

Animals  affected:  livestock. 

Reference:  Porter  1951,  Kingsbury  1964. 


Visum  sativum  L.  (AIF) 

Common  name:  garden  pea. 
Toxin:  unknown. 
Habitat:  cultivated. 
Animals  affected:  sheep,  cattle. 
Reference:  Whiting  et  al.  1957,  Kingsbury 
1964. 

Poinciana  gilliesii  Hook.  (PIS) 

Common  name:  bird  of  paradise. 

Toxin:  unknown;  green  seed  pods  are  gas- 
trointestinal irritants. 

Habitat:  cultivated  ornamental,  small  pop- 
ulation established  in  Washington  Co. 

Animals  affected:  humans,  livestock. 

Reference:  Cann  and  Verhulst  1958,  Kings- 
bury 1964. 

Prosopis  glandulosa  Torr.  (PNT) 

Common  name:  mesquite. 

Toxin:  unknown;  may  cause  a  nutritional 
deficiency. 

Habitat:  dry  ranges,  washes  and  draws  at 
low  elevations,  especially  along  streams 
where  the  water  table  is  high. 

Animals  affected:  cattle. 

Reference:  Adler  1949,  Kingsbury  1964. 

Robinia  pseudoacacia  L.  (PIT) 

Common  name:  black  locust. 

Toxin:  unknown. 

Habitat:  escaped  from  cultivation;  around 
dwellings  or  along  fencerows. 

Animals  affected:  horses,  cattle,  sheep, 
poultry,  humans. 

Reference:  Power  1901,  Kingsbury  1964. 

Thermopsis  montana  Nutt.  (PNF) 

Common  name:  goldenpea,  mountain  ther- 
mopsis, yellow  pea. 

Toxin:  alkaloids. 

Habitat:  common  in  pastures. 

Affected  animals:  cattle. 

Reference:  Schmutz  et  al.  1968. 

Trifolium  hybridum  L.  (PIF) 

Common  name:  .\lsike  clover. 

Toxin:  photosensitizing  compound. 

Habitat:  cultivated;  escaped  to  roadsides 
and  meadows. 

Animals  affected:  horses,  hogs,  sheep, 
cattle. 

Reference:  Fincher  and  Fuller  1942,  Kings- 
bury 1964. 


244 


Great  Basin  Naturalist 


Vol.  40,  No.  3 


Trifolium  praetense  L.  (PIF) 

Common  name:  red  clover. 
Toxin:  unknown. 

Habitat:  cultivated  and  escaping  along 
roadsides  and  ditches. 

Animals  affected:  cattle,  horses,  sheep. 
Reference:  O'Dell  1959,  Kingsbury  1964. 

Trifolium  repens  L.  (PIF) 

Common  name:  white  clover. 

Toxin:  cyanogenetic. 

Habitat:  cultivated  and  escaping. 

Animals  affected:  newborn  pigs. 

Reference:  Garner  1957,  Kingsbury  1964. 

Vicia  villosa  Roth  (PIF) 

Common  name:  hairy  vetch,  winter  vetch. 
Toxin:  photosensitizing  compound. 
Habitat:  cultivated;  occasionally  escaping. 
Animals  affected:  cattle,  horses. 
Reference:  Claughton  and  Claughton  1954, 
Kingsbury  1964. 


Haemodoraceae 

Iris  missouriensis  Nutt.  (PNF) 

Common  name:  wild  iris,  blue  flag,  fleur- 
de-lis,  western  blue  flag. 

Toxin:  unknown. 

Habitat:  moist  soils  along  stream  banks,  in 
marshes  or  moist  mountain  meadows. 

Animals   affected:    calves,   laboratory   ani- 
mals. 

Reference:   Beath   et   al.    1953,   Kingsbury 
1964. 


Hypericaceae 

Hypericum  formosum  H.B.K.  (PNF) 

Common  name:  southwestern  St.  Johns- 
wort. 

Toxin:  photosensitizing  compounds. 

Habitat:  moist  soils  of  plains  and  hills. 

Animals  affected:  cattle,  sheep,  horses, 
goats. 

Reference:  Harris  1951. 


Fagaceae 

Quercus  gambelii  Nutt.  (PNT) 

Common  name:  Gambel  oak. 

Toxin:  tannins. 

Habitat:  throughout  the  state,  often  form- 
ing dense  thickets. 

Animals  affected:  cattle,  sheep,  and  goats. 

Reference:    Boughton    194.3,    Kingsbury 
1964. 


FUMARIACEAE 

Corydalis  aurea  Willd.  (ANF) 

Common  name:  Golden  corydalis. 

Toxin:  alkaloids. 

Habitat:  woods  and  well-shaded  mountain 
slopes. 

Animals  affected:  sheep,  cattle. 

Reference:  Sperry  1955,  Kingsbury  1964. 

Gentianaceae 

Centaurium  calycosum  (Buckl.)  Fern.  (ANF) 

Common  name:  Buckley  centaury,  moun- 
tain pink,  arizona  centaury. 

Toxin:  unknown. 

Habitat:  moist  soil,  river  valleys. 

Animals  affected:  sheep,  goats. 

Reference:    Dollahite   and   Allen    1962, 
Kingsbury  1964. 


JUNCAGINACEAE 

Triglochin  concinna  Davy  (PNF) 

Common  name:  arrowgrass,  goosegrass, 
sourgrass,  podgrass,  Utah  arrowgrass. 

Toxin:  hydrocyanic  acid. 

Habitat:  salty  marshes  and  ponds. 

Animals  affected:  sheep,  cattle. 

Reference:  Schmutz  et  al.  1968. 

Triglochin  debilis  L.  (PNF) 

Common  name:  arrowgrass,  weak  arrow- 
grass. 

Toxin:  hydrocyanic  acid. 

Habitat:  damp  soils,  marshes  and  sloughs; 
usually  where  the  soil  is  alkaline  or  the  water 
calcareous  or  brackish. 

Animals  affected:  sheep,  cattle. 

Reference:  Schmutz  et  al.  1968,  Kingsbury 
1964. 

Triglochin  maritima  L.  (PNF) 

Common  name:  arrowgrass,  seashore  ar- 
rowgrass, shore  arrowgrass. 

Toxin:  hydrocyanic  acid. 

Habitat:  damp  soils,  marshes  and  sloughs; 
usually  where  the  soil  is  alkaline  or  the  water 
calcareous  or  brackish. 

Animals  affected:  sheep,  cattle. 

Reference:  Beath  et  al.  1933,  Kingsbury 
1964. 


September  1980 


Brotherson  et  al.:  Poisonous  Plants 


245 


Lamiaceae 

Lamium  amplexicaule  L.  (AIF) 

Common  name:  henbit,  dead  nettle. 
Toxin:  unknown. 

Habitat:  occasional  weed  of  fields  and 
waste  places. 

Animals  affected:  sheep,  horses,  cattle. 
Reference:  Hurst  1942,  Kingsbury  1964. 

LiLIACEAE 

Allium  schoenoprasm  L.  (PNF) 

Common  name:  chives. 

Toxin:  unknown. 

Habitat:  cultivated;  river  bars,  lake  shores, 
wet  meadows. 

Animals  affected:  horses. 

Reference:  Kobayashi  1950,  Kingsbury 
1964. 

Asparagus  officinalis  L.  (PIF) 

Common  name:  asparagus. 

Toxin:  unknown. 

Habitat:  cultivated  and  widely  escaped. 

Animals  affected:  cattle  and  dairy  cows. 

Reference:  Los  Angeles  County  Livestock 
Department  1938,  Kingsbury  1964. 

Ornithogalum  umbellatum  L,  (PIF) 

Common  name:  star-of-Bethlehem,  snow- 
drop. 

Toxin:  cholchicine  alkaloid. 

Habitat:  weed  of  grasslands  and  thickets. 

Animals  affected:  sheep,  cattle,  children. 

Reference:  Reynard  and  Norton  1942, 
Kingsbury  1964. 

Veratrum  californicum  Durand  (PNF) 

Common  name:  false  hellebore,  corn-lily, 
skunk  cabbage. 

Toxin:  alkaloids. 

Habitat:  bogs  and  wet  meadows  from  7500 
to  9500  feet. 

Animals  affected:  cattle,  sheep,  fowl,  hu- 
mans. 

Reference:  Schmutz  et  al.  1968,  Kingsbury 
1964. 

Yucca  L.  (PNS) 

Note:  members  of  this  genus  have  been  re- 
ported to  contain  saponins,  salicylic  acid,  the 
alkaloid  imperialin,  and  several  resins.  None 
of  the  species  in  which  these  toxins  have  been 
identified  are  found  in  Utah. 

Reference:  Pammel  1911. 


Zigadenus  elegans  Pursh  (PNF) 

Common  name:  death  camas,  mountain 
death  camas. 

Toxin:  alkaloids. 

Habitat:  prairies,  meadows. 

Animals  affected:  cattle,  horses,  hoes,  fowl 
humans. 

Reference:  Marsh  et  al.  1915,  Kingsbury 
1964. 

Zigadenus  paniculatus  (Nutt.)  Wats.  (PNF) 

Common  name:  death  camas,  foothill 
death  camas,  sandcom. 

Toxin:  alkaloids. 

Habitat:  dry  soils;  hills  and  plains. 

Animals  affected:  cattle,  horses,  hogs,  hu- 
mans. 

Reference:  Fleming  et  al.  1921,  Kingsbury 
1964. 

Zigadenus  venenosus  Wats.  (PNF) 

Common  name:  death  camas,  meadow 
death  camas. 

Toxin:  alkaloids. 

Habitat:  moist,  grassy  meadows. 

Animals  affected:  sheep,  cattle,  horses, 
hogs,  humans. 

Reference:  Cameron  1952,  Kingsbury 
1964. 

PiNACEAE 

Pinus  ponderosa  Doug,  ex  Laws  (PNT) 

Common  name:  western  yellow  pine,  pon- 
derosa pine. 

Toxin:  unknown. 

Habitat:  coniferous  forest  at  moderate  ele- 
vations; dry  hillsides,  plateaus,  slopes,  valleys 
and  mesas. 

Animals  affected:  cattle. 

Reference:  MacDonald  1952,  Kingsbury 
1964. 

POACEAE 

Avena  fatua  L.  (AIG) 

Common  name:  wild  oats. 
Toxin:  mechanically  injurious. 
Habitat:  cultivated  land  and  waste  places. 
Animals  affected:  livestock. 
Reference:  Pammel  1911. 

Avena  sativa  L.  (AIG) 

Common  name:  cultivated  oats. 

Toxin:  nitrates,  photosensitizing  com- 
pounds, grass  tetany. 


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Vol.  40,  No.  3 


Habitat:  open  ground,  grasslands,  waste 
places;  lawns  and  golf  courses. 

Animals  affected:  cattle,  horses,  hogs,  tur- 
keys, goats,  sheep  and  wild  ruminants. 

Reference:  Newsom  et  al.  1937,  Kingsbury 
1964. 

Bromus  rigidus  Roth  (AIG) 

Common  name:  ripgut  brome. 

Toxin:  mechanical  injury  from  mature 
awns. 

Habitat:  common  weed. 

Animals  affected:  cattle  and  sheep. 

Reference:  Range  Plant  Handbook  1937. 

Bromus  rubens  L.  (AIG) 

Common  name:  foxtail  chess,  red  brome. 

Toxin:  mechanical  injury  from  mature 
awns. 

Habitat:  common  weed,  dry  and  saline 
soils. 

Animals  affected:  cattle  and  sheep. 

Reference:  Davis  1952. 

Bromus  tectorum  L.  (AIG) 

Common  name:  cheatgrass,  downy  cheat. 

Toxin:  mechanical  injury  from  mature 
awns;  may  also  be  implicated  in  ergot  poi- 
soning. 

Habitat:  common  weed,  especially  in  dry 
places;  plains  and  foothills. 

Animals  affected:  cattle. 

Reference:  Pammel  1911. 

Cynodon  dactylon  L.  Pers.  (PIG) 

Common  name:  Bermuda  grass. 
Toxin:  photosensitizing  compound. 
Habitat:    open   ground,    grasslands,    waste 
places;  lawns  and  golf  courses. 
Animals  affected:  cattle. 
Reference:  Gibbons  1953,  Kingsbury  1964. 

Eragrostis  cilianensis  (All.)  Link  (AIG) 

Common  name:  lovegrass,  stinkgrass,  stick 
grass. 

Toxin:  unknown. 

Habitat:  cultivated  ground,  gardens  and 
waste  places;  weed  in  fields  and  along  road- 
sides. 

Animals  affected:  horses. 

Reference:  Gates  1930,  Kingsbury  1964. 

Festuca  arundinacea  Schreb.  (PIG) 

Common  name:  fescue,  tall  fescue,  alta  fes- 
cue, goar  fescue. 
Toxin:  alkaloids. 


Habitat:  unimproved  pastures;  wet,  heavy 
soils  of  high  organic  content. 

Animals  affected:  cattle. 

Reference:  Maag  and  Tobiska  1956,  Kings- 
bury 1964. 

Glyceria  striata  Lam.  Hitch.  (PNG) 

Common  name:  fowl  mannagrass. 

Toxin:  cyanogenetic. 

Habitat:  wet  areas. 

Animals  affected:  cattle. 

Reference:  Reynard  and  Norton  1942, 
Kingsbury  1964. 

Hilaria  rigida  (Thurb.)  Benth  ex  Scribn. 

(PNG) 

Common  name:  galleta  grass,  big  galleta, 
dixie  grass. 

Toxin:  unknown. 

Habitat:  dry  lands  and  desert  ranges  to 
4000  feet. 

Animals  affected:  cattle. 

Reference:  Quortrup  and  McFarland  1956, 
Kingsbury  1964. 

Holcus  lanatus  L.  (PIG) 

Common  name:  velvet  grass,  mesquite 
grass,  Yorkshire  velvet  grass. 

Toxin:  unknown. 

Habitat:  open  ground,  meadows  and  moist 
places;  occasionally  cultivated. 

Animals  affected:  livestock. 

Reference:  Couch  1932,  Kingsbury  1964. 

Hordeum  jubatum  L.  (PNG) 

Common  name:  squirreltail  grass,  foxtail 
grass,  wild  barley. 

Toxin:  mechanically  injurious. 

Habitat:  weed  in  open  ground,  meadows, 
prairies,  along  streams,  ditches  and  waste 
places. 

Animals  affected:  sheep,  cattle,  horses. 

Reference:  Fleming  and  Peterson  1919, 
Kingsbury  1964. 

Hordeum  vulgare  L.  (AIG) 

Common  name:  cultivated  barley. 

Toxin:  mechanically  injurious. 

Habitat:  cultivated  for  grain  and  along 
road  shoulders;  sometimes  spontaneous  in 
waste  places  but  not  persistent. 

Animals  affected:  hogs,  dogs,  humans, 
poultry. 

Reference:  Christensen  and  Kernkamp 
1936,  Kingsbury  1964. 


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Brotherson  et  al.:  Poisonous  Plants 


247 


Setaria  lutescens  (Wiegel)  Hubb.  (AIG) 

Common  name:  yellow  bristle  grass,  foxtail 
grass,  pigeon  grass. 

Toxin:  mechanically  injurious. 

Habitat:  common  weed  of  cultivated  and 
waste  areas. 

Animals  affected:  livestock. 

Reference:  Bankowski  et  al.  1956,  Kings- 
bury 1964. 

Sorghum  halpense  L.  Pers.  (PIG) 

Common  name:  Johnson  grass. 

Toxin:  hydrocyanic  acid,  nitrates. 

Habitat:  weed  of  cultivated  fields,  waste 
places  and  along  irrigation  ditches  and 
stream  bottoms. 

Animals  affected:  cattle,  sheep,  horses. 

Reference:  Slade  1903,  Kingsbury  1964. 

Sorghum  vulgare  Pers.  (AIG) 
Common  name:  grain  sorghum. 
Toxin:  cyanogenetic  glycoside. 
Habitat:  cultivated. 
Animals  affected:  cattle,  sheep,  horses. 
Reference:  Slade  1903,  Kingsbury  1964. 

Stipa  commata  Trin.  &  Rupr.  (PNG) 

Common  name:  needle-and-thread  grass. 

Toxin:  mechanically  injurious. 

Habitat:  dry  plains  and  hillsides,  sandy  soil. 

Animals  affected:  livestock. 

Reference:  Pammel  1911. 

Stipa  neomexicana  (Thurb.)  Scribn.  (PNG) 

Common  name:  New  Mexican  feather- 
grass. 

Toxin:  mechanically  injurious. 

Habitat:  common  in  dry  rocky  canyons 
and  mesas. 

Animals  affected:  livestock. 

Reference:  Pammel  1911. 

Zea  mays  L.  (AIG) 

Common  name:  com,  maize. 

Toxin:  nitrates,  under  conditions  of  over- 
fertilization  with  too  little  water. 

Habitat:  cultivated  for  grain,  forage  or  si- 
lage. 

Animals  affected:  livestock,  humans. 

Reference:  Brady  et  al.  1955,  Kingsbury 
1964. 

POLYGONACEAE 

Beta  vulgaris  L.  (AIF) 

Common  name:  beet,  sugar  beet,  fodder 
beet,  mangel-worzel,  mangold. 


Toxin:  oxalates;  nitrates,  under  conditions 
of  overfertilization  with  too  little  water. 

Habitat:  cultivated. 

Animals  affected:  livestock. 

Reference:  Baker  and  Eden  1954,  Kings- 
bury 1964. 

Rheum  rhaponticum  L.  (PIF) 

Common  name:  rhubarb. 
Toxin:  oxalic  acid,  oxalates. 
Habitat:  cultivated  and  persisting. 
Animals  affected:  livestock,  humans. 
Reference:  Hansen  1930,  Kingsbury  1964. 

Rumex  acetosella  L.  (PIF) 

Common  name:  sheep  sorrel,  dock. 

Toxin:  oxalates. 

Habitat:  common  weed  of  acid  or  sterile, 
gravelly  soils  of  pastures  and  meadows;  waste 
places. 

Animals  affected:  sheep. 

Reference:  Connor  and  Adams  1951, 
Kingsbury  1964. 

Rumex  crispus  L.  (PIF) 

Common  name:  curly  dock. 

Toxin:  oxalates. 

Habitat:  moist  fields  and  waste  places. 

Animals  affected:  sheep. 

Reference:  Connor  and  Adams  1951, 
Kingsbury  1964. 

POLYPODIACEAE 

Dryopteris  felix-mas  (L.)  Schrott  (PNF) 

Common  name:  male  fern. 

Toxin:  thiaminase. 

Habitat:  mountains;  damp  soils,  deep 
shaded  ravines,  in  cliffs  or  tallus. 

Animals  affected:  horses. 

Reference:  Harvey  et  al.  1944,  Kingsbury 
1964. 

Pteridium  aquilinum  (L.)  Kuhn  (PNF) 

Common  name:  Bracken  fern. 

Toxin:  thiaminase  and  its  coenzymes. 

Habitat:  upland  pastures,  a.spen  zone. 

Animals  affected:  livestock. 

Reference:  Carpenter  1950,  Kingsbury 
1964. 

PORTULACACEAE 

Portulaca  oleracea  L.  (AIF) 

Common  name:  purslane,  pusley. 
Toxin:  oxalates. 


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Habitat:  common  weed  of  garden  and  cul- 
tivated areas. 

Animals  affected:  sheep. 

Reference:  Mathams  and  Sutherland  1952, 
Kingsbury  1964. 

Ranunculaceae 

Aconitum  columbianum  Nutt.  (PNF) 

Common  name:  western  monkshood. 

Toxin:  alkaloids. 

Habitat:  mountains  from  5000  to  10,000 
feet;  along  streams  and  wet  meadows;  moist 
places  and  thickets. 

Animals  affected:  livestock,  humans. 

Reference:  Stern  1960,  Kingsbury  1964. 

Actaea  arguta  Nutt.  (PNF) 

Common  name:  baneberry,  western  bane- 
berry. 

Toxin:  irritant  oil. 

Habitat:  mountains  in  rich,  moist  soil. 

Animals  affected:  livestock,  humans. 

Reference:  Schmutz  et  al.  1968. 

Caltha  leptosepala  DC.  (PNF) 

Common  name:  elkslip  marshmarigold. 
Toxin:  glycoside  (protoanemonin). 
Habitat:  wet  mountain  soils. 
Animals  affected:  livestock. 
Reference:  Schmutz  et  al.  1968. 

Delphinium  andersonii  Gray  (PNF) 
Common  name:  Anderson  larkspur. 
Toxin:  alkaloids. 

Habitat:  subsaline  soils  of  plains  and  hills. 
Animals  affected:  cattle,  sheep. 
Reference:  Miller  1923,  Kingsbury  1964. 

Delphinium  barbeyi  Huth.  (PNF) 

Common  name:  barbey,  larkspur,  tall 
larkspur. 

Toxin:  alkaloids. 

Habitat:  mountains;  meadows  and  open 
woods,  summer  ranges;  common  under  aspen 
and  along  streams. 

Animals  affected:  cattle,  occasionally 
sheep. 

Reference:  Cook  1951,  Kingsbury  1964. 

Delphinium  nuttallianum  Fritz.  (PNF) 

Common    name:    Nuttall    larkspur,    low 

larkspur.  Nelson  larkspur. 
Toxin:  alkaloids. 
Habitat:    moist    soils,    hills,    foothills,    and 

sagebrush  deserts. 


Animals  affected:  cattle,  sheep. 
Reference:  Ewan  1945,  Kingsbury  1964. 

Delphinium  occidentale  Wats.  (PNF) 

Common  name:  duncecap  larkspur,  tall 
larkspur. 

Toxin:  alkaloids. 

Habitat:  mountain  summer  ranges,  com- 
mon under  aspen  and  along  streams;  moun- 
tain meadows. 

Animals  affected:  cattle,  occasionally 
sheep. 

Reference:  Couch  1936,  Kingsbury  1964. 

Ranunculus  acris  L.  (PIF) 

Common  name:  tall  field  buttercup,  tall 
buttercup. 

Toxin:  protoanemonin. 

Habitat:  common  pasture  weed. 

Animals  affected:  livestock. 

Reference:  Tehon  et  al.  1946,  Kingsbury 
1964. 

Ranunculus  cymbalaria  Pursh  (PNF) 

Common  name:  alkali  buttercup;  trailing 
buttercup. 

Toxin:  glycosides  (protoanemonin). 

Habitat:  muddy  banks  along  brackish 
streams  and  marshes. 

Animals  affected:  livestock. 

Reference:  Fleming  1920,  Kingsbury  1964. 

Ranunculus  flammula  var.  filiformis 
(Michx.)  Hook.  (PNF) 

Common  name:  creeping  spearwort  but- 
tercup. 

Toxin:  glycosides  (protoanemonin). 

Habitat:  marshy  ground  of  lakes,  streams 
and  ditches. 

Animals  affected:  livestock. 

Reference:  Hill  and  van  Heyningen  1951, 
Kingsbury  1964. 

Ranunculus  repens  L.  (PIF) 

Common  name:  creeping  buttercup. 
Toxin:  protoanemonin. 
Habitat:   meadows  and  marshes  at  lower 
elevations;  wet  pastures. 
Animals  affected:  livestock. 
Reference:  Gilkey  1958,  Kingsbury  1964. 

Ranunculus  scleratus  L.  (PNF) 

Common  name:  celeryleaf  crowfoot. 
Toxin:  glycosides  (protoanemonin). 
Habitat:    borders    of    lakes,    ponds    and 

streams. 


September  1980 


Brotherson  et  al.:  Poisonous  Plants 


249 


Animals  affected:  livestock. 

Reference:  Fleming  1920,  Kingsbury  1964. 

Ranunculus  testiculatus  Crantz  (AIF) 

Common  name:  burbuttercup;  testiculate 
buttercup. 

Toxin:  glycosides  (protoanemonin). 

Habitat:  general  in  the  intermountain  re- 
gion. 

Animals  affected:  livestock. 

Reference:  Schmutz  et  al.  1968. 

ROSACEAE 

Cercocarpus  montanus  Raf.  (PNS) 

Common  name:  mountain  mahogany, 
birchleaf  mountain  mahogany,  true  mountain 
mahogany. 

Toxin:  cyanogenetic  glycoside. 

Habitat:  stony  hills  and  slopes. 

Animals  affected:  livestock. 

Reference:  Burke  1960,  Kingsbury  1964. 

Prunus  armeniaca  L.  (PIT) 
Common  name:  apricot. 
Toxin:  cyanide. 

Habitat:  cultivated  and  persisting. 
Animals  affected:  livestock,  humans. 
Reference:  Hurst  1942,  Kingsbury  1964. 

Prunus  persica  Batsch.  (PNT) 

Common  name:  peach. 

Toxin:  cyanide. 

Habitat:  cultivated. 

Animals  affected:  livestock. 

Reference:  Reynard  and  Norton  1942, 
Kingsbury  1964. 

Prunus  virginiana  L.  (PNT) 

Common  name:  choke  cherry. 

Toxin:  cyanogenetic  glycoside. 

Habitat:  common  in  hills,  mountains,  along 
streams,  thickets,  fencerows  and  edges  of 
woods. 

Animals  affected:  sheep,  cattle. 

Reference:  Pijoan  1942,  Kingsbury  1964. 

Santalaceae 

Comandra  umbellata  (L.)  Nutt.  (PNF) 

Common  name:  bastard  toadflax. 

Toxin:  alkaloids,  glycosides,  secondary  se- 
lenium accumulator. 

Habitat:  common  weed,  found  in  various 
habitats. 

Animals  affected:  livestock. 


Reference:  Trelease  and  Beath  1949, 
Kingsbury  1964. 

SOLANACEAE 

Datura  meteloides  Dunal  (ANF) 

Common  name:  datura,  stramonium, 
thornapple,  Jimson  weed,  Jamestown  weed, 
apple  of  Peru,  tolgaudia,  sacred  datura,  In- 
dian apple. 

Toxin:  alkaloids  (atropine,  hyoscvamine, 
hyoscine). 

Habitat:  plains,  dry  hills  and  valleys;  culti- 
vated and  escaping. 

Animals  affected:  humans,  horses,  cattle, 
sheep,  hogs,  mules,  chickens. 

Reference:  Hansen  1924,  Kingsbury  1964. 

Datura  stramonium  L.  (ANF) 

Common  name:  Jimsonweed,  sacred  da- 
tura. 

Toxin:  alkaloids  (atropine,  hyoscyamine, 
hyoscine). 

Habitat:  waste  areas,  rich  soils  of  barn- 
yards, heavily  used  portions  of  pastures. 

Animals  affected:  horses,  cattle,  sheep, 
hogs,  mules,  chickens,  humans. 

Reference:  Hansen  1924,  Kingsbury  1964. 

Hyoscyamus  niger  L.  (BIF) 

Common  name:  black  henbane,  henbane. 

Toxin:  alkaloids  (hyoscyamine,  hyoscine, 
atropine). 

Habitat:  widespread  dry  soils  of  roadsides 
and  waste  areas. 

Animals  affected:  humans,  fowl,  livestock. 

Reference:  Long  1917,  Kingsbury  1964. 

Lycium  halmifolium  Mill.  (PIS) 

Common  name:  matrimony  vine,  tea  vine. 

Toxin:  unknown. 

Habitat:  cultivated  and  escaping  around 
homesites  and  cemetaries. 

Animals  affected:  calves,  sheep. 

Reference:  Hansen  1927,  Kingsbury  1964. 

Nicotiana  attenuata  Torr.  ex  S.  Wats.  (.\NF) 

Common  name:  wild  tobacco,  coyote  to- 
bacco. 

Toxin:  nicotine. 

Habitat:  dry,  sandy  stream  beds  and  flats. 

Animals  affected:  horses,  pigs,  livestock, 
humans. 

Reference:  Marsh  et  al.  1927,  Kingsbury 
1964. 


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Vol.  40,  No.  3 


Nicotiana    trigonophylla    Dunal    ex    DC. 

(ANF) 

Common  name:  wild  tobacco,  desert  to- 
bacco. 

Toxin:  nicotine. 

Habitat:  dry  desert  soils. 

Animals  affected:  horses,  pigs,  livestock, 
humans. 

Reference:  Marsh  et  al.  1927,  Kingsbury 
1964. 

Solarium  dulcamara  L.  (PIF) 

Common  name:  European  bittersweet, 
climbing  nightshade,  bitter  nightshade. 

Toxin:  glycoalkaloids. 

Habitat:  woods,  thickets  and  waste  places; 
cultivated  and  escaping. 

Animals  affected:  cattle,  horses,  sheep,  hu- 
mans. 

Reference:  Craig  and  Kehoe  1925,  Kings- 
bury 1964. 

Solanum  eleagnifolium  Cav.  (PNF) 

Common  name:  silverleaf  nightshade, 
white  horsenettle,  trapillo. 

Toxin:  glycoalkaloids. 

Habitat:  serious  weed  of  prairies,  open 
woods  and  disturbed  soils;  dry  ground;  barn- 
yards. 

Animals  affected:  cattle,  sheep. 

Reference:  Buck  et  al.  1960,  Kingsbury 
1964. 

Solanum  nigrum  L.  (AIF) 

Common  name:  black  nightshade. 

Toxin:  glycoalkaloids. 

Habitat:  common  weed  of  fields  and  waste 
places. 

Animals  affected:  livestock,  humans. 

Reference:  Carey  1955,  Kingsbury  1964. 

Solanum  rostratum  Dunal  (ANF) 

Common  name:  buffalo  bur,  Kansas  thistle, 
Texas  thistle,  buffalobur  nightshade. 

Toxin:  glycoalkaloids. 

Habitat:  plains,  roadsides,  barnyards. 

Animals  affected:  hogs. 

Reference:  Simic  1943,  Kingsbury  1964. 

Solanum  triflorum  Nutt.  (ANF) 

Common  name:  three  flowered  nightshade, 
cutleaf  nightshade. 

Toxin:  glycosides. 

Habitat:  prairies,  fields  and  waste  places; 
weed  of  cultivation  and  disturbed  soils. 


Animals  affected:  horses,  cattle. 
Reference:  Pammel  1921,  Kingsbury  1964. 


Typhaceae 

Typha  latifolia  L.  (PNF) 

Common  name:  cattail. 

Toxin:  unknown. 

Habitat:   common  in  moist  soils,  marshes 
and  ponds. 

Animals  affected:  horses. 

Reference:  Hansen  1930,  Kingsbury  1964. 


Zygophyllaceae 
Tribulus  terrestris  L.  (AIF) 

Common  name:  puncture  vine,  caltrap. 

Toxin:    nitrates,    photosensitizing    com- 
pound. 

Habitat:  dry  soils  of  waste  lands,  roadsides 
and  deserts. 

Animals  affected:  sheep. 

Reference:  Durrell  et  al.  1952,  Kingsbury 
1964. 


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Bankowski,  R.  a.,  R.  W.  Wichmann,  and  E.  E.  Stuart. 
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Beath,  O.  a.,  J.  H.  Draize,  and  H.  F.  Eppson.  1933.  Ar- 
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Beath,  O.  A.,  C.  S.  Gilbert,  H.  F.  Eppson,  and  I. 
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BiNNs,  W.,  AND  L.  F.  James.  1961.  A  congenital  defor- 
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BoucHTON,  I.  B.  1943.  Oak  bmsh  poisoning.  Texas  Vet. 
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\V.  D.  Maclay.  1954.  Alfalfa  saponin.  J.  .\m. 
Chem.  Soc.  76:2271. 

Welsh,  S.  L.  1978.  Utah  flora:  Fahaceae  (Legimiinosae). 
Great  Basin  Nat.  .38(3):225-367. 

1980.  Utah  flora;  miscellaneous  families.  Great 

Basin  Nat.  40:38-58. 

Welsh,  S.  L.  and  G.  Moore.  1973.  Utah  plants- 
Tracheophvta.  Brigham  Yoinig  University  Press. 
Provo.  Utah.  474  pp. 

West,  E.  1957.  Poisonous  plants  around  the  home.  Flor- 
ida Agr.  Expt.  Sta.  Circ.  S-IOO. 

Whiting,  F.,  R.  Connell,  P.  J.  G.  Pllmmer,  and  R.  D. 
Clark.  1957.  Incoordination  {Cerebellar  Ataxia) 
among  lambs  from  ewes  fed  peavine  silage.  Ca- 
nad.  J.  Comp.  Med.  and  Vet.  Sci.  21:77. 

Williams,  M.  C.  and  F.  A.  Norris.  1969.  Distribution 
of  miserotoxin  in  varieties  of  Astragalus  miser. 
Dougl.  ex  Hook.  Weed  Science  17:236-2.38. 

Williams,  M.  C.  .\nd  R.  C.  Barneby.  1977.  The  occur- 
rence of  nitro-toxins  in  North  .\merican  Astra- 
galus (Fabaceae).  Brittonia  29:310-326. 


THE  SUCCESSIONAL  STATUS  OF  CUPRESSUS  ARIZOMCA 

Albert  J.  Parker 

.\bstbact.—  Several  investigations  isize-class  analysis,  age-determination  inquiries,  and  germination  tests"*  suggest 
that  Cupressus  arizonica  of  southeastern  .\rizona  is  a  pioneer  species.  The  tree  requires  disturbance  to  remove  or 
reduce  soil  litter,  which  other\%-ise  inhibits  the  reproduction  of  the  species.  Reduction  of  light  intensity  caused  by 
canopy  closure  appears  to  be  less  important  than  litter  accumulation  in  restricting  C.  arizonica  reproduction.  Fol- 
lowing disturbance,  successful  establishment  of  seedlings  may  occur  over  an  e.xtended  period  ,50  to  100  years  I  as 
Utter  graduallv  accumulates.  The  absence  of  C.  arizonica  seedlings  in  present  populations  suggest  that  fire  suppres- 
sion policies  on  federal  lands  where  C.  arizonica  occurs  have  altered  fire  frequency,  and  consequently  have  fostered  a 
short-term  reduction  in  C.  arizonica  establishment.  Only  in  floodplain  en\ironments.  where  flooding  disturbs  the  soil 
surface,  has  much  reproduction  occurred  in  recent  years.  The  long-term  population  pattern  of  C.  arizonica  appears 
stable,  due  to  the  great  longevity  of  the  species. 


Rough-barked  Arizona  c^'press  {Cupressus 
arizonicu  Greene;  all  taxonomy  after  Kearney 
and  Peebles  I960'  is  a  tree  species  of  local 
occurrence  in  the  mountain  ranges  of  south- 
eastern .\rizona,  southwestern  New  Me.xico, 
western  Texas,  and  northern  Mexico.  Though 
it  has  topically  been  characterized  as  a  mois- 
tiu-e  demanding  species  of  riparian  associ- 
ation iWolf  1948a.  \Miittaker  and  Xiering 
1965.  BrouTi  and  Lowe  1974',  recent  findings 
(Parker  1980 >  demonstrate  that  C.  arizonica 
may  occupy  relatively  xeric  landscape  posi- 
tions—including south-facing  slopes,  ridge 
tops,  and  convex  slope  configurations— at 
higher  elevations  laroimd  2000  ml 

In  the  first  pubUshed  report  mentioning 
Cupressus  arizonica  regeneration.  Posev  and 
Goggans  - 1967'  obser\ed  little  reproduction 
anxAvhere  in  the  Southwest,  although  they 
collected  no  data  to  substantiate  or  in- 
vestigate this  phenomena.  Thev  did  express 
concern  that  the  limited  natural  range  of  the 
species,  coupled  with  its  apparent  inability  to 
reproduce  in  many  natural  circtmistances, 
could  indicate  that  its  existence  was  threat- 
ened. This  paper  investigates  the  population 
d\Tiamics,  reproductive  strategy,  and  role  in 
species  replacement  processes  of  C.  arizonica 
in  the  upland  forests  of  southeastern  Arizona. 

Study  Are.a.s 

Populations  of  Cupressus  arizonica  were 
sampled  in   the  Chiricahua   Moimtains,   the 

Department  of  Geography.  Univeran-  of  Georgia,  .Mhens,  Georgia  30602. 


Blue  Range  north  of  Clifton,  the  Dragoon 
Mountains,  and  the  Santa  Catalina  Moun- 
tains, all  located  in  .\rizona  (Fig.  l\  Cupr- 
essus arizonica  is  most  common  in  the  wood- 
land zone  of  these  mountain  ranges 
(1300-1900  m;  Wliittaker  and  Xiering  1965), 
a  zone  characterized  by  the  frequent  co-do- 
minance of  alligator  juniper  Juniperus  dep- 
peana\  pinyons  [Pinus  cembroides  and  P. 
edulisl  and  a  nmnber  of  oak  species,  includ- 
ing silverleaf  oak  (Quercus  hypoleucoides), 
netleaf  oak  [Q.  rugosa),  and  .\rizona  white 
oak  Q.  arizonica"!.  Cupressus  arizonica  is  pro- 
gressively more  restricted  to  riparian  habitats 
where  the  woodland  zone  grades  into  the 
shrub-dominated  desert  and  semidesert  vege- 
tation types  below.  It  is  also  found  in  stands 
at  higher  elevations  (1600-2300  mi  with 
other  conifers,  notably  Arizona  pine  (Pinus 
ponderosa  var.  arizonica^  Apache  pine  {P. 
engehiannii),  Chihuahua  pine  P.  leiophylla), 
and  Douglas-fir  iPseudotsuga  menziesii); 
these  forested  stands  often  have  closed  ca- 
nopies. 

The  climate  of  the  woodland  zone  is  sub- 
humid  and  mild,  with  about  50  cm  annual 
precipitation  and  a  mean  annual  temperature 
of  12.5  C.  The  wettest  periods  of  the  year  are 
middle  to  late  summer  and  midwinter  (Bry- 
son  and  Lown.-  1955 1. 

The  isolated  mountain  ranges  of  this  region 
of  basin  and  range  topography  are  roughly 
linearlv    aligned,    northward    trending,    and 


254 


September  1980 


P.\rker:  Arizona  Cypress  Succession 


iDO 


ARIZONA 


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MTS. 

M3 


i"''*"/"„ 


aV" 

•■;/'-$, 

f.ii-i/'//^ 

•'/; 

% 

''^*i; 

i 

V"'.. 


-:.  km. 

'-/y,...„.?     ARIZONA 

MEXFCO" 


CARTOGRAPH.C  LASGRATOfiY.  uMVERSiTV  Gf  W.SGG\S;f4  -  V.ADISOM 

Fig.  1.  Study  area  uith  the  species  range  inset.  The  location  of  stands  included  in  this  study  are  noted  by  number. 
Open  circles  in  the  inset  represent  the  three  extensive  areas  of  nonriparian  C.  ari:onica  occuTTence  in  southeastern 
.\rizona:  the  Chiricahua  Mountains,  the  Blue  Range,  and  the  Santa  Catalina  Mountains. 


256 


Great  Basin  Naturalist 


Vol.  40,  No.  3 


separated  by  extensive  sediment-filled  low- 
lands. The  bedrock  core  of  these  ranges  in- 
clude both  igneous  and  sedimentary  rocks 
(Fenneman  1931). 

Methods 

Environmental  and  vegetational  measure- 
ments were  obtained  from  19  internally 
homogeneous  stands  of  C.  arizonica.  Within 
each  stand,  three  4  by  25  m  quadrats  were 
oriented  so  that  their  central  long  axes  were 
normal  to  a  contour  line  bisecting  the  stand, 
with  intervals  of  10  m  separating  each  quad- 
rat. The  following  measurements  were  re- 
corded within  each  quadrat:  the  species  and 
circumference  at  breast  height  (1.37  m)  of 
each  tree,  the  number  of  saplings  of  all  tree 
species,  and  the  number  of  C.  arizonica  seed- 
lings. Any  stem  of  at  least  20  cm  circum- 
ference at  breast  height  was  considered  a 
tree.  A  sapling  was  defined  as  the  stem  of  any 
tree  species  which  exceeded  breast  height, 
but  which  was  less  than  20  cm  in  circum- 
ference at  that  height.  All  stems  of  tree  spe- 
cies less  than  breast  height  were  counted  as 
seedlings.  The  depth  of  the  litter  layer  in 
each  stand  was  measured  at  1  m  intervals 
along  the  central  long  axis  of  each  quadrat.  A 
total  of  60  litter  depth  measurements  was 
made  per  stand. 

In  two  stands,  both  in  the  Chiricahua 
Mountains,  all  C.  arizonica  trees  with  a  cir- 
cumference at  breast  height  exceeding  30  cm 
were  cored  with  an  increment  borer  at  breast 
height.  These  two  stands  were  chosen  for 
their  differing  size  structures.  One  stand  was 
characterized  by  trees  few  in  number  but 
large  in  diameter,  but  the  other  stand  had  a 
great  number  of  individuals,  particularly  in 
the  smaller  size-classes.  These  two  stands 
were  selected  as  representative  of  14  of  the 
19  stands  sampled.  The  other  five  stands  has 
a  size  structure  intermediate  in  character  be- 
tween those  chosen  for  coring.  In  total,  79 
trees  were  cored. 

Cores  were  aged  in  the  laboratory,  al- 
though the  aging  was  complicated  by  two 
factors.  First,  several  of  the  C.  arizonica  trees 
possessed  rotted  xylem  tissue  that  caused 
fragmentation  of  some  cores  and  loss  of  re- 
cord from  the  damaged  segments.  Second,  C. 
arizonica  has  been  shown  to  produce  false  an- 


nual rings  (Bannan  1954).  To  circumvent  the 
first  difficulty,  the  length  of  each  core  was 
measured  and  the  following  formula  was  used 
to  compute  a  "tree  ring"  age  for  each 
sample: 

0.95  (^)(g)  =  t 

where:  t  =  extrapolated  tree  ring  age  (yr) 
c    =    circumference  of  the  tree 

(cm) 

g   =   number  of  annual  rings  per 

unit  length  of  core  (yr*cm-i;  this  value  is 

later  referred  to  as  "mean  time  required 

per  unit  of  radial  growth") 

0.95  =  a  coefficient  to  adjust  for  bark 
thickness 

With  respect  to  the  second  complication. 
Clock  and  Agerter  (1963)  were  able  to  utilize 
microclimatic  records  from  a  plantation  of  C. 
arizonica  near  Lubbock,  Texas  (500  km  from 
the  species'  natural  range)  to  investigate  the 
production  of  false  annual  rings  in  this  spe- 
cies. They  reported  that  multiple  false  rings 
may  be  produced  by  an  individual  in  a  single 
year,  and  that  the  number  of  false  rings  is  not 
consistent  from  tree  to  tree,  or  even  from 
place  to  place  on  the  same  tree.  Because  of 
these  difficulties,  precise  absolute  aging  was 
not  possible,  and  tree  ring  ages  reported  in 
this  study  may  be  as  much  as  two  times 
larger  than  the  actual  tree  age.  Nevertheless, 
the  ring  counts  and  extrapolated  ages  provide 
an  indication  of  the  relative  ages  of  individ- 
uals and  thus  allow  identification  and  inter- 
pretation of  the  approximate  age  structures 
of  the  stands. 

Cones  of  C.  arizonica  were  collected  for 
the  purpose  of  determining  the  mean  number 
of  seeds  per  cone,  as  well  as  to  provide  seeds 
for  use  in  germination  tests.  All  250  cones 
collected  were  from  the  second  year  foliage 
of  a  number  of  C.  arizonica  individuals  lo- 
cated in  the  Blue  Range.  Only  closed  cones 
were  collected.  Fifty  of  tht«e  cones  were  ran- 
domly selected,  and  seeds  from  each  cone 
were  counted.  Following  this,  the  other  cones 
were  emptied  of  seeds,  and  all  seeds  were 
then  sorted  by  size,  the  larger  seeds  being 
kept  for  use  in  germination  tests. 

A  series  of  germination  tests  were  per- 
formed on  lots  of  100  seeds  selected  ran- 
domly from  the  seed  source.  These  tests  in- 


September  1980 


Parker:  Arizona  Cypress  Succession 


257 


vestigated  the  relationship  of  C.  arizonica 
germination  success  to  the  following  factors: 
freezing,  flooding,  fire,  litter  buildup,  and 
light  intensity.  The  physical  conditions  in 
which  the  germination  tests  were  performed 
followed  those  of  Wolf  (1948b)  in  general 
(see  Table  4). 

Results 

Stands  were  segregated  into  three  groups, 
according  to  the  size-class  frequency  distribu- 
tion of  individuals  of  Cupressus  arizonica. 
The  eight  stands  of  the  first  group  display  a 


generally  logarithmatic  decrease  in  the  num- 
ber of  individuals  in  successively  larger  size- 
classes,  with  the  exception  that  the  seedling 
layer  is  virtually  absent  (Fig.  2).  Populations 
with  a  size  stnicture  similar  to  that  of  the 
first  group  are  often  characterized  as  stable 
through  time,  with  losses  of  canopy  individ- 
uals being  balanced  by  replacements  from 
the  sapling  layer  (Daubenmire  1968).  The  six 
stands  of  the  third  group,  in  contrast,  lack  a 
logarithmatic  trend.  Instead,  the  number  of 
individuals  in  successively  larger  size-classes 
is  more  constant  (Fig.  2),  suggesting  that  a 
past  period  of  reproduction  has  ended.  The 


400-1 


350- 


Group  1 


1 — \ — \ — ] — I — r-1 


i—\ — I — \ — I — I  T*  I  ' 

(DraiDirjinLnioinminTTTTTT     ■ 

ID    £    r^ 


CO 


CNJ     CD 


in    to    h- 


(DO)    o>ai^CT)Oio>ojoi    a> 


Group  2 


"I  1 1  ""1 — I  I  I  I  I 


ID  <b  1^   CO   a> 

ininiDuSininioiT) 


*-     CNJ     to     ^     *0     to     ^ 


60- 


40- 


20- 


Group  3 


Fig.  2.  Composite  size-class  structure  histograms  for  stands  of  C.  arizonica.  See  text  for  a  definition  of  each  of  the 
three  groups. 


258 


Great  Basin  Naturalist 


Vol.  40,  No.  3 


second  group  includes  five  stands  inter- 
mediate in  character  between  the  other 
groups. 

Even  in  those  cases  which  appear  to  pos- 
sess a  steady-state  population  above  the  sap- 
ling layer,  the  almost  universal  absence  of  C. 
arizonica  seedlings  in  stand  understories 
(Table  1)  is  conspicuous,  and  agrees  with  the 
observation  of  Posey  and  Goggans  (1967). 
Conditions  in  the  two  stands  which  do  con- 
tain a  relatively  large  number  of  seedlings 
are  significant;  in  both  cases,  disturbance  has 
exposed  bare  mineral  soil,  in  one  case  by  nat- 
ural flooding  and  in  the  other  by  logging  ac- 
tivities that  mechanically  stripped  off  the  lit- 
ter from  a  portion  of  the  forest  floor. 

The  behavior  of  other  tree  species  in  C. 
anzontca -dominated  commimities  fall  pri- 
marily into  two  types,  according  to  stand 
size-class  information  (Table  2).  A  first  group 
consists  of  intolerant  pioneer  species,  which 
reproduce  episodically  in  association  with 
disturbance  events,  and  probably  depend  on 
wide  dispersal  of  seeds  to  maintain  represen- 
tation on  a  given  site.  These  species  display 
an  even  aged  structure  in  most  C.  arizonica 
stands.  The  common  upland  conifers  of  the 
region,  such  as  Apache  pine,  Arizona  pine, 
and  Douglas-fir  fit  this  behavior  pattern.  A 
second  group  consists  of  tolerant  species  with 
stable  populations  that  reproduce  either  veg- 
etatively  of  by  widespread  dispersal  from  ad- 
jacent locales.  These  display  continues  repro- 


Table  1.     The  seedling  class. 

No.  seedlings         Mean 


Recently  flooded  stand 
Recently  logged  stand 
All  other  stands  (17) 

Total 


20 
36 
17 

73 


1.0 

3.7 


duction  in  C.  arizonica-dominated  stands, 
with  a  maximum  number  of  stems  occurring 
in  recent  cohorts.  This  group  includes  the 
typical  pinyon/oak  woodland  dominants- 
Mexican  pinyon,  silverleaf  oak,  netleaf  oak, 
and  Arizona  white  oak. 

Core  analysis  indicated  that  the  mean  tree 
ring  age  of  C.  arizonica  in  the  stand  with  few 
but  large  individuals  was  317.1  years,  with 
ages  ranging  from  197.6  to  456.7  years  (Table 
3).  The  majority  of  tree  ring  ages  were  be- 
tween 250  and  375  years.  No  seedlings,  sap- 
lings, or  trees  of  less  than  30  cm  circum- 
ference at  breast  height  were  found  in  the 
stand.  Thus,  the  tree  ring  age  of  the  youngest 
C.  arizonica  individual  was  almost  200  years. 
The  mean  tree  ring  age  of  C.  arizonica  trees 
in  the  stand  with  the  greater  density  of  small- 
er trees  was  116.9  years,  with  an  age  range 
from  62.4  to  178.1  years.  However,  there 
were  also  a  large  number  of  saplings  and 
trees  too  small  to  core,  and  these  size-classes 
probably  extended  the  age  range  down  to 
about  25  years.  The  conspicuous  absence  of 
C.  arizonica  seedlings  in  the  imderstory  of 
this  stand  (Table  3)  suggested  that  reproduc- 
tion has  been  inhibited  during  about  the  last 
quarter  century. 

To  determine  the  degree  to  which  under- 
story  individuals  of  C.  arizonica  were  sup- 
pressed by  the  overstory,  a  linear  correlation 
and  regression  analysis  was  performed  on  the 
mean  time  required  for  unit  growth  (yr*cm-i) 
for  each  tree  against  tree  radius.  A  negative 
correlation  coefficient  would  be  predicted  in 
conditions  of  understory  suppression,  because 
small  trees  would  require  longer  periods  of 
time  to  produce  an  amount  of  radial  growth 
equal  to  that  produced  on  the  larger,  more 
rapidly  growing  trees  in  the  canopy  in  a 
shorter  period  of  time.  The  results  suggest 
that    suppression    of    understory    individuals 


Table  2.  Composite  size-class  structnres  for  upland  conifers  and  pinyon-oak  species.  The  figures  heading  each 
size-class  category  are  the  smallest  possible  circumference  in  that  class,  and  size-classes  include  all  stems  up  to  the 
next  larger  value.  Upland  conifers  include  Arizona  pine,  ponderosa  pine,  Apache  pine,  and  Douglas  fir.  Pinyons  in- 
clude Mexican  pinyon  and  two-leafed  pinyon.  Oaks  include  silverleaf  oak,  netleaf  oak,  and  Arizona  white  oak. 


sapl 

20 

30 

40 

Size-class 
50 

(cm 
60 

circumference) 
70         80 

90 

100 

110 

120 

Upland  conifers 

3 

4 

4 

7 

4 

3 

8 

4 

0 

4 

5 

9 

Pinyons 
Oaks 

63 
111 

27 
27 

18 
18 

13 
24 

4 
18 

7 
8 

2 
4 

2 
3 

1 
0 

1 
1 

0 
0 

0 
1 

September  1980 


Parker:  Arizona  Cypress  Succession 


259 


Table  3.     Characteristics  of  cored  stands. 

First 
stand 


Second 
stand 


Tree  density  (ha"^) 
Total  basal  area 

(dm2-ha-i) 
Mean  basal  area  of  C. 

arizonica  trees 

(dm^'tree"^ 
Number  of  C.  arizonica 

cored 
'Mean  time  required 

per  unit  growth  of  C. 

arizonica  (yr-cnr') 
Mean  extrapolated  tree 

age  of  C.  arizonica  (yr) 
Standard  deviation  of 

extrapolated  tree  age  (yr) 
Oldest  C  arizcmica 

extrapolated  age  (yr) 
Yoimgest  C.  arizonica 

extrapolated  age  (yr) 
Number  of  C.  arizonica 

saplings  and  trees  too 

small  to  core  (20-30 

cm) 
Number  of  C.  arizonica 

seedlings 
Correlation  coefficient 

for  suppression  test 


1333 

3733 

11901 

5854 

9.51 

1.49 

21 

58 

24.2 

16.6 

317.1 

116.9 

63.0 

26.4 

456.7 

178.1 

197.6 

62.4 

0 

81 

0 

0 

-0.833 

-0.673 

'Calculated  by  dividing  the  number  of  annual  rings  per  core  by  the 
length  of  that  core,  it  expresses  the  number  of  years  required  for  an  individ- 
ual to  add  one  cm  of  radial  growth  and  is  the  reciprocal  of  the  growth  rate. 


The  great  number  of  seeds  produced  does 
not  necessarily  insure  abundant  reproduction. 
Sudworth  (1915)  noted  that  seeding  of  C. 
arizonica  was  best  on  moist,  bare  mineral 
soils.  Field  observations  made  during  the 
course  of  data  collection  confirm  this  sugges- 
tion, in  that  C.  arizonica  reproduction  was 
generally  restricted  to  areas  within  2  of  the 
19  stands,  and  then  only  on  exposed  mineral 
soil  associated  with  recent  floods  or  logging. 
Moreover,  the  weighted  average  of  litter 
depth  where  C.  arizonica  seedlings  were  en- 
countered (1.74  cm)  was  only  half  the  mean 
depth  of  litter  for  all  stands  in  this  study  (3.42 
cm),  indicating  that  successful  reproduction 
is  associated  with  reduced  litter  depth. 

Germination  tests  also  indicated  that  a  lit- 
ter layer  sharply  reduced  the  germination 
and  survival  of  seedlings.  Onlv  3.7  percent  of 
the  seeds  germinated  on  the  litter-covered 
substrate,  in  contrast  to  10.0  percent  on  min- 
eral soil  controls  (Table  4).  Of  the  other  fac- 
tors tested,  freezing  of  seeds  prior  to  germi- 
nation, reduced  light  intensity,  and 
immersion  of  seeds  in  water  did  not  alter  ger- 
mination success.  Simulated  exposure  of  seed- 
bearing  cones  to  canopy  fire  conditions, 
while  reducing  germination  success  (6.0  per- 


was  evident  in  both  stands  (Table  3).  The  de- 
gree of  suppression  was  greater  in  the  older 
stand,  as  a  consequence  of  the  longer  period 
of  time  over  which  competition  was  acting. 
These  results  indicate  that  small  C.  arizonica 
trees  may  persist  beneath  a  canopy,  and  sug- 
gest that  the  species  is  shade  tolerant. 

Evidence  from  core  aging  also  .suggests 
that  C.  arizonica  possesses  potentially  great 
longevity.  Based  on  the  rates  of  growth  de- 
termined in  this  study  (Table  3),  and  account- 
ing for  multiple  annual  rings,  some  of  the 
larger  individuals  encountered  in  canyon  bot- 
toms (exceeding  1  m  dbh)  may  be  300  to  500 
years  of  age. 

Not  only  does  the  species  achieve  old  ages, 
but  also  it  produces  great  numbers  of  seed. 
The  mean  number  of  seeds  per  cone  was 
104.2,  with  a  standard  deviation  of  19.8.  It 
was  estimated  that  healthy  trees  of  moderate 
size  (50  cm  circumference)  may  produce 
from  103  to  10*  cones  per  year,  therefore  re- 
sulting in  an  annual  seed  crop  per  tree  of 
from  10^  to  10'^  seeds. 


Table  4.  Germination  test  results.  The  soil  in  each 
pot  was  composed  of  a  mixture  of  50  percent  sand  and 
.50  percent  silt  loam.  The  surface  was  without  litter  cov- 
er. Pots  were  supplied  with  adequate  water  regularly 
and  were  grown  under  high  levels  of  simlight.  The  pots 
were  15  cm  in  diameter  and  allowed  free  drainage  be- 
low. The  air  temperature  was  between  20  and  24  C 
throughout  the  duration  of  the  tests. 


Composite 

Number  of 

germination 

Treatment 

test  pots 

rates  (%) 

Litter  cover  over 

mineral  substrate 

6 

3.7 

Seeds  exposed  to 

freezing  before 

planting 

4 

8.8 

Seeds  immersed  in 

agitated  water 

2 

10.0 

"Seeds  exposed  to 

canopy  fire  conditions 

1 

6.0 

°°  Seeds  exposed  to 

groimd  fire  conditions 

1 

0.0 

Seeds  grown  under 

reduced  light  levels 

2 

8.5 

°°  "Controls 

10 

10.0 

•—Cones  filled  with  seeds  were  exposed  to  80  C  for  eight  minutes. 
■-Cones  filled  with  seeds  were  exposed  to  315  C  for  eight  minutes. 
*— Control  conditions. 


260 


Great  Basin  Naturalist 


Vol.  40,  No.  3 


cent),  did  not  destroy  all  seeds.  In  contrast, 
exposure  to  simulated  ground  fire  conditions, 
which  are  much  hotter,  killed  all  seeds  within 
the  cones.  Thus,  C.  arizonica  seeds  enclosed 
in  cones  on  branches  may  remain  viable  after 
fires,  and  have  the  capability  to  germinate  on 
the  mineral  seedbeds  exposed  by  burning. 

Discussion 

The  absence  of  C.  arizonica  reproduction 
under  adults  of  the  same  species  in  natural 
stands  is  striking.  Two  interpretations  of  this 
observed  absence  of  young  individuals  are 
possible.  First,  successful  reproduction  may 
be  temporally  sporadic  and  linked  to  occa- 
sional optimal  climatic  conditions  that  foster 
waves  of  C.  arizonica  reproduction.  If  such 
optimal  conditions  have  not  occurred  during 
the  last  20  or  30  years,  the  paucity  of  C. 
arizonica  seedlings  would  be  explained.  How- 
ever, successful  reproduction  of  C.  arizonica 
was  encountered  in  two  recently  disturbed 
habitats,  and  this  indicates  that  recent  climat- 
ic conditions  are  not  inimical  to  the  tree's  re- 
production. The  environmental  conditions  as- 
sociated with  these  cases  of  successful 
reproduction  do  support,  however,  a  second 
interpretation  of  the  absence  of  C.  arizonica 
seedlings,  namely,  that  disturbance  is  neces- 
sary to  create  conditions  favorable  for  rege- 
neration, and  that  factors  associated  with  a 
closed  forest  inhibit  seeding  by  the  tree. 

Stand  size-class  analyses  would  seem  to 
both  siipport  and  contradict  this  character- 
ization of  C.  arizonica  as  a  species  requiring 
disturbance  for  reproduction.  Stands  of  the 
first  group  suggest  that  the  species  is  tolerant, 
able  to  maintain  itself  under  a  closed  canopy 
through  time.  Stands  of  the  second  and  third 
size-class  groups  imply  that  the  species  is  in- 
tolerant, with  a  period  of  establishment,  fol- 
lowed by  the  cessation  of  C.  arizonica  repro- 
duction. Therefore,  the  latter  groups  support 
the  hypothesis  that  the  species  is  opportunis- 
tic, capable  of  invading  open  habitats,  but 
not  able  to  compete  effectively  with  other 
species  as  successional  processes  proceed. 

Inspection  of  core  analyses  resolve  this  ap- 
parent paradox.  The  first  stand  cored,  repre- 
senting those  stands  that  are  dominated  by 
large  individuals  and  which  suggest  that  the 
species  is  a  pioneer,  possesses  no  individuals 
with  a  tree  ring  age  of  less  than  200  years. 


The  trees  from  this  stand  exhibit  a  tendency 
to  clump  in  the  250-  to  375-year  tree  ring 
age  range.  Lack  of  successful  C.  arizonica 
regeneration  over  an  extended  period,  com- 
bined with  a  tendency  toward  clumping  of 
ages,  suggests  that,  following  a  lengthy  peri- 
od of  C.  arizonica  establishment,  reproduc- 
tion is  virtually  eliminated.  The  second  stand 
cored,  representing  those  stands  with  a  mixed 
size  structure  and  suggesting  that  the  species 
is  a  climax-type,  possesses  trees  ranging  in 
tree  ring  age  from  25  to  150  years.  This  stand 
possesses,  however,  a  conspicuous  absence  of 
reproduction  during  the  last  quarter  century. 
These  data,  in  the  same  manner  as  those  from 
the  first  stand,  suggest  that  following  a 
lengthy  period  of  colonization  and  reproduc- 
tion of  C.  arizonica,  perhaps  50  to  100  years 
in  length,  reproduction  subsequently  ceases. 
The  population  structures  of  these  two 
stands,  therefore,  differ  only  in  the  elapsed 
times  since  disturbance,  rather  than  in  some 
fundamental  difference  in  the  ecological  be 
havior  of  the  species.  In  both  cases,  C.  arizo- 
nica acts  as  an  intolerant  species,  requiring 
disturbance  to  create  invadable  habitats.  In 
the  denser,  more  youthful  C.  arizonica  popu- 
lation of  the  second  stand,  disturbance  was 
more  recent  than  in  the  older  population  of 
C.  arizonica  in  the  first  stand.  The  initial  in- 
terpretation of  stand  structures  of  the  first 
type— that  the  species  is  a  climax-type,  with  a 
stable,  self-perpetuating  population— is 
shown  to  be  incorrect.  Stands  of  the  first 
group  attain  this  pseudoclimax  type  structure 
only  temporarily  as  a  consequence  of  the  ex- 
tended period  following  disturbance  when  C. 
arizonica  reproduction  is  possible.  Thus,  the 
three  groups  identified  by  differing  size 
structures  may  be  interpreted  as  stages  of  a 
temporal  continuum,  with  each  stage  reflect- 
ing a  longer  period  of  development  since  a 
disturbance  event. 

Other  results  yield  evidence  favoring  the 
interpretation  of  C.  arizonica  as  an  intolerant 
species.  The  longevity  of  C.  arizonica  is  char- 
acteristic of  many  pioneer  species,  as  it  in- 
creases the  allowable  length  of  time  between 
periodic  disturbances,  thus  enhancing  the 
probability  that  perturbation  will  occur  on  or 
near  the  site  while  mature  seed  trees  are 
available  for  colonization.  Seed  counts  and 
estimated  cone  crop  sizes  indicate   that  C. 


September  1980 


Parker:  Arizona  Cypress  Succession 


261 


arizonico  is  a  prolific  seeder,  also  a  character- 
istic of  the  reproductive  strategy  of  a  pioneer 
species,  because  it  increases  the  probability 
that  a  germule  will  be  dispersed  to  an  open 
habitat.  The  inverse  relationship  of  litter 
depth  to  C.  arizonica  seedling  number  in- 
dicates that  partial  or  complete  removal  of 
litter  on  the  forest  floor  is  necessary  to  pro- 
mote reproduction.  The  inhibition  of  litter  on 
germination  in  test  pots  further  corroborates 
the  negative  effects  of  litter  accumulation  on 
this  tree's  regeneration.  This  need  for  mineral 
seedbeds,  created  by  natural  disturbance  (i.e., 
fire  or  flooding)  or  by  certain  human  activi- 
ties (i.e.,  logging)  is  typical  of  pioneer  ele- 
ments. Finally,  the  ability  of  seeds  to  remain 
viable  following  a  canopy  fire  is  an  opportu- 
nistic character  often  favored  by  intolerant 
species  that  must  colonize  open  habitats  fol- 
lowing such  fires. 

In  contrast  to  the  characteristics  suggesting 
that  C.  arizonica  is  a  pioneer  species,  the 
tree's  abilities  to  germinate  in  low  light  con- 
ditions and  to  tolerate  shaded  sites  by  sup- 
pression of  growth  are  characteristics  more 
commonly  associated  with  climax-type  than 
with  pioneer  species.  These  responses  to  light 
are  puzzling  and  deserve  closer  inspection  in 
the  future.  In  general,  however,  it  is  con- 
cluded that  C.  arizonica  is  a  pioneer  species, 
requiring  periodic  disturbance  to  open  new 
sites  for  colonization. 

A  unique  characteristic  of  C.  arizonica 
stand  dynamics  is  the  long  period,  as  long  as 
a  century,  during  which  colonization  and  re- 
production remain  possible  after  the  disturb- 
ance event.  Jenny  et  al.  (1949)  have  demon- 
strated that,  in  ponderosa  pine  forests  at  1220 
to  2220  m  elevation  in  California,  100  to  200 
years  are  required  to  reach  a  near-equilib- 
rium steady-state  condition  of  litter  thickness, 
in  which  mineralization  of  organic  matter 
balances  the  addition  of  litter  produced  by 
the  vegetation.  Such  a  long  period  of  litter 
accumulation  is  characteristic  of  moist,  cool 
montane  conifer  forest  ecosystems,  and  fits 
well  with  the  observed  time  scale  of  C.  arizo- 
nica establishment  on  a  site.  This  peculiarly 
long  period  of  colonization  appears  to  be  re- 
lated to  the  shade-tolerant  character  of  the 
species.  Most  pioneer  species,  being  intoler- 
ant, rapidly  cease  reproduction  as  light  levels 
decrease  under  the  species'  own  canopy.  The 


ability  of  C.  arizonica  to  reproduce  in  shade 
allows  it  to  continue  reproducing  for  a  much 
longer  period  than  most  colonizing  species. 
This  period  is  later  terminated  by  the  more 
gradual  process  of  litter  accumulation. 

Thus,  C.  arizonica  may  be  considered  to  be 
a  pioneer  species  that  is  intolerant  of  litter 
accumulation  under  a  closed  canopv,  even 
though  it  is  tolerant  of  low  light  levels  fol- 
lowing colonization  of  disturbed  sites.  Addi- 
tionally, its  habit  of  seed  persistence  in  the 
cone  is  critical  to  its  maintenance  on  most 
sites.  It  may  not  depend,  then,  on  seed  dis- 
persed from  distant  sources  to  colonize  a  giv- 
en disturbed  area. 

Using  the  Noble  and  Slatyer  (1977)  model 
of  successional  processes  (from  Cattelino  et 
al.  1979),  this  study  suggests  three  tvpes  of 
species  that  exhibit  fundamentally  different 
responses  to  disturbance  events.  First,  the  up- 
land conifers  are  DI  species,  characterized  bv 
widespread  dispersal  capabilities  (D)  and  in- 
tolerance to  a  closed  canopy  (I).  These  trees 
depend  on  colonizing  disturbed  sites  with 
seed  from  an  off-site  source.  Second,  the  pin- 
yon  oak  group  are  DT/VT  species,  which 
persist  on  a  disturbed  site  by  colonization 
from  an  off-site  source  (D— pinyon)  or  by 
vegetative  reproduction  (V— oaks).  These  spe- 
cies maintain  themselves  by  virtue  of  their 
tolerance  of  closed  canopy  conditions  (T). 
Cupresstis  arizonica,  the  third  type,  is  a  CI 
(DI)  species.  As  described  above,  it  is  intoler- 
ant (I)  of  closed  forest  conditions  and  gener- 
ally is  maintained  locally  by  seed  persistence 
in  cones  from  the  canopy  (C),  which  shelter 
viable  seeds  through  disturbance  events.  Be- 
cause of  its  limited  range,  C.  arizonica  is  less 
commonly  maintained  by  dispersal  from  adja- 
cent poulations  (D),  except  perhaps  in  ripa- 
rian stands,  where  surface  water  wash  may 
supply  seeds  to  wash  environments  from  sur- 
rounding upland  slopes. 

These  three  types  of  species,  in  com- 
bination with  differential  longevity  patterns, 
yield  three  possible  pathways  of  community 
development  (Fig.  3).  The  initial  composition 
of  each  case  is  represented  by  at  least  one 
stand  sampled  in  this  study.  The  first  and  sec- 
ond cases  diagram  multiple  successional 
pathways  in  communities  which  possess  rep- 
resentatives of  the  three  behavioral  types.  In 
the  first  case,  the  model  assumes  a  greater 


262 


Great  Basin  Naturalist 


Vol.  40,  No.  3 


Case  1 


<b 

/ 

UC,  CA,  PO 

CA,  PO 

PO 

-J 

1 

I  x<a 


z' 


V 


-^_L 


UCPO 


PO 


—^ — TTL "ii^- 


X  >b 


Case  2 


'  \a<x<b 


UC,  CA,  PO 


UCPO 


b<x<c 


/ 


-^_lJ 


UC,  PO 


PO 


i 

I x>_c I 


Case  3 


N 


\  a<x<b 


J- 


CA,  PO 


-    PO 

T 

L x_l§ ! 


L 


Fig.  3.  Potential  multiple  pathways  in  a  successional  development  model.  Solid  lines  indicate  developmental 
changes  in  the  absence  of  disturbance.  Dashed  lines  represent  disturbance  events.  Boxes  define  the  constraints  on  C. 
arizonica  populations,  if  they  are  to  avoid  local  extinction.  Symbols: 
UC     Upland  conifers 
CA     Cupressiis  arizcmica 
PO     Pinyon-oak  species 
X     period  of  time  since  last  catastrophic  disturbance  event 
a     time  required  by  C.  arizonica  to  reach  maturity  and  produce  seeds 
b     longevity  of  C.  arizonica 
c     longevity  of  upland  conifers 


September  1980 


Parker:  Arizona  Cypress  Succession 


263 


life  span  for  C.  arizonica  than  for  the  upland 
conifers,  but  in  the  second  case  the  model  as- 
sumes that  upland  conifers  live  longer  than 
C.  arizonica.  In  both  cases,  upland  conifers 
are  maintained  indefinitely  in  the  landscape 
by  their  wide  dispersal  of  seeds,  and  pinyons 
and  oaks  are  maintained  by  their  tolerance  to 
a  closed  forest  canopy.  Cupressus  arizonica, 
however,  may  become  locally  extinct,  depen- 
ding on  the  timing  of  disturbance  events.  If 
disturbance  recurs  more  frequently  than  the 
time  required  for  C.  arizonica  maturity,  the 
species  will  not  be  maintained  in  the  commu- 
nity because  the  canopy  seed  reserve  is  de- 
stroyed. Such  local  extinction  assumes  that  no 
introductions  occur  by  dispersal  from  non- 
local populations,  and  that  the  disturbance 
was  large  enough  in  areal  extent  to  kill  the 
entire  local  population  of  C.  arizonica.  A  sec- 
ond mechanism  for  local  extinction  in  each 
case  would  be  for  disturbance  to  be  so  in- 
frequent that  the  local  population  dies  out 
before  reinitiation  of  a  successional  cycle. 
The  third  case  in  the  model  represents  a  sim- 
plified commimity  in  which  the  upland  con- 
ifer element  is  locally  absent,  whether  by 
habitat  restriction  or  chance. 

It  is  apparent  from  this  model  of  succes- 
sion that,  assuming  habitat  factors  remain  un- 
changed, local  extinction  of  C.  arizonica 
would  occur  only  with  excessively  frequent 
or  infrequent  catastrophic  fire  events.  Low 
net  productivity  and  fuel  loadings  do  not 
make  frequent  catastrophic  fire  a  highly 
probable  event  in  montane  forests  of  south- 
eastern Arizona.  Suppression  of  fires  by  hu- 
mans may  temporarily  reduce  C.  arizonica 
reproduction,  but,  unless  the  program  is 
maintained  effectively  for  several  himdred 
years,  it  is  unlikely  to  result  in  extinction  of 
local  populations. 

The  situation  in  riparian  settings  is  even 
more  favorable  for  C.  arizonica  perpetuation. 
In  these  habitats,  population  maintenance  is 
effected  by  dispersal  from  adjacent  upland 
populations,  as  well  as  by  local  preservation 
of  seeds  in  the  canopy.  Extinction  on  such 
sites  would  require  elimination,  not  only  of 
the  local  population,  but  also  of  other  C. 
arizonica  populations  higher  in  the  water- 
shed. Additionally,  flooding  assumes  an  im- 
portant role  as  a  disturbance  mechanism  in 
these  stands.  This  is  particularly  significant 


with  respect  to  C.  arizonica  regeneration,  be- 
cause, while  it  may  not  increase  surface  light 
intensities,  it  will  remove  litter  and  expose 
bare  mineral  soil,  which  triggers  reproduc- 
tion, apparently  irrespective  of  light  condi- 
tions at  the  forest  floor.  (Certainly,  light  in- 
tensities influence  growth  rates  and  vigor, 
but  not  necessarily  establishment  potential.) 

In  effect,  C.  arizonica  appears  to  be  a 
stable,  terminal  element  of  the  restricted 
habitats  in  which  it  presently  occurs.  Its  life 
history  characteristics  (including  its  great  po- 
tential longevity),  population  maintenance 
mechanisms,  and  colonization  patterns  com- 
bine to  make  it  a  persistent  species,  threat- 
ened in  a  temporal  sense  only  by  the  remote 
probability  of  either  very  frequent  or  in- 
frequent catastrophic  disturbance  events. 
Any  short-term  efforts  designed  to  stimulate 
C.  arizonica  reproduction  should  be  ad- 
dressed to  the  elimination  of  fire  suppression 
policies  in  regions  of  its  occurrence.  Long- 
term  preservation  of  this  locally  dispersed, 
intolerant  species  requires  that  open  habitats 
continue  to  be  created  by  catastrophic  per 
turbations  with  a  frequency  of  recurrence 
greater  than  the  age  of  first  viable  seed  pro- 
duction, and  less  than  the  life  span  of  the  in- 
dividuals in  the  population. 


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A  SELF-POLLINATION  EXPERIMENT  IN  PINUS  EDULIS 

Ronald  M.  Lanner' 

Abstract.-  Controlled  pollinations  were  performed  on  fonr  pinyons  (Pinus  cdtilis  Engelm.)  to  compare  the  results 
of  selfing  and  outcrossing.  Final  cone  size  was  the  same  under  both  treatments.  There  was  no  significant  difference 
in  number  of  seeds  per  cone.  Filled-seed  yields  averaged  14.4  percent  in  selfings  and  90.5  percent  in  outcrossings. 
Relative  self-fertility  averaged  about  15  percent,  a  level  comparable  with  that  of  other  pine  species  studied. 


The  breeding  system  of  pinyon  {Pinus 
edulis  Engelm.)  has  received  Httle  study.  In 
several  particulars  pinyon  resembles  other 
members  of  Pinus.  For  example,  it  is  general- 
ly monoecious  (Lanner  1975)  and  the  sexes 
are  usually  segregated  with  the  upper  crown 
tending  to  be  more  female  than  male  and 
vice  versa,  though  there  may  be  broad  over- 
lap. Pollen  shed  and  conelet  receptivity  are 
limited  to  a  short  period  in  the  spring  (Lan- 
ner 1970).  Meiosis  leading  to  microspore  for- 
mation is  temperature  sensitive  during  meta- 
phase  and  anaphase,  and  temperatures  below 
4  C  may  result  in  inviable  polyploid  pollen 
grains  (Chira  1967).  Pinyon  crosses  readily 
with  singleleaf  pinyon  {P.  monophylla  Torr. 
&  Frem.)  and  natural  hybrids  and  in- 
trogressants  are  widely  distributed  where 
their  ranges  overlap  (Lanner  1974,  1975). 

Though  pines  are  generally  considered  to 
be  outcrossing  species,  many  studies  have 
shown  that  self-fertilization  is  usually  at  least 
marginally  successful  in  producing  sound 
seeds.  This  paper  reports  the  results  of  a  con- 
trolled pollination  experiment  in  which  self- 
ing was  compared  to  outcrossing. 

Methods  and  Materials 

Isolation  bags  of  nonwoven  cloth  with  cel- 
luloid windows  were  placed  over  female 
branches  of  four  pinyon  pines  of  unknown 
seed  source  on  the  campus  of  Utah  State  Uni- 
versity 4  June  1976.  Pollen  extracted  the  pre- 
vious year  and  stored  in  a  freezer  was  ap- 
plied with  a  hypodermic  syringe  8  June  1976, 
shortly    after    conelets    emerged    from    their 

'Department  of  Forestry  and  Outdoor  Recreation,  Utah  State  University',  Logan,  Utah  84322. 


covers  of  bud  scales.  Table  1  summarizes  the 
crosses  made.  Pollination  treatments  were 
self-pollen  and  outcross  pollen.  Most  of  the 
outcross  pollen  was  a  bulked  mix  from  all  the 
seed  trees  except  tree  3,  plus  an  additional 
tree.  Tree  3  was  outcrossed  with  trees  9  and 
28.  Isolation  bags  were  removed  22  June 

1976.  Cones  were  inventoried  13  July  1976 
and  in  June  1977;  they  were  harve.sted  25 
August  1977  prior  to  cone  dehiscence.  Seeds 
were  extracted  in  the  laboratory  as  cones  de- 
hisced at  room  temperature.  Seeds  were  tal- 
lied as  filled  or  empty  on  the  basis  of  weight 
and  seed  coat  color:  empty  seeds  are  easilv 
detected  in  handling  and  by  their  light-col- 
ored seed  coats. 

Results 

1.  Cone  survival—  Within  2  weeks  of  pol- 
lination 12  of  the  31  .selfed  cones  and  2  of  the 
35  outcrossed  cones  had  aborted.  By  15  June 

1977,  a  year  after  pollination,  one  more  self- 
ed cone  and  4  more  outcrossed  cones  had 
aborted  and  been  shed.  No  further  losses  oc- 

Table  1.  Distribution  of  pollination  treatments 
among  trees. 


Pollination  treatment 

Tree 

Selfed 

Outcrossed 

Number  of  conelets 

.3 

— 

16 

4 

7 

5 

.36 

12 

8 

58 

12 

6 

Total 

31 

35 

265 


266 


Great  Basin  Naturalist 


Vol.  40,  No.  3 


curred  prior  to  maturity.  Final  survival  rates 
were  57  percent  for  selfed  cones  and  83  per- 
cent for  outcrossed  cones. 

2.  Cone  size—  Lengths  of  14  mature  selfed 
cones  ranged  from  3.3-3.9  cm  (x  =  3.6  cm). 
Lengths  of  16  outcrossed  cones  showed  iden- 
tical ranges  and  mean. 

3.  Seed  yield  — Yields  of  seeds  per  cone 
are  summarized  in  Table  2. 

In  all  three  trees  that  were  both  selfed  and 
outcrossed  the  yield  of  seeds  per  cone  was 
slightly  higher  under  selfing,  but  the  differ- 
ences were  nonsignificant  in  all  cases. 

Yield  of  filled  seed  and  relative  self- 
fertility  are  summarized  in  Table  3.  Filled 
seed  percent  averaged  14.4  percent  in  self- 
ings  and  90.5  percent  in  outcrossings.  Rela- 
tive self-fertility,  a  statistic  designed  to  neu- 
tralize variation  not  associated  with  embryo 
genotype  but  which  can  influence  seed  yield 
(Sorensen  1970),  ranged  from  about  10  to  21 
percent  and  averaged  about  15  percent. 

Discussion  and  Conclusions 

It  is  unclear  why  survival  of  selfed  cones 
was  exceeded  by  that  of  the  outcrossed  cones. 
All  losses  of  selfed  cones  occurred  by  13  July 
1976,  just  five  weeks  after  pollination.  Fer- 

Table  2.  Number  of  seeds  per  cone  after  selfini;  and 
outcrossing. 


Pollination  treatment 

Tree 

Selfed 

Outcrossed 

3 

_ 

13.9 

4 

5.8 

4.8 

.36 

10.4 

9.4 

58 

5.2 

3.7 

Total 

7.76 

10.52 

Table  3.  Yield  of  filled  seed  by  pollination  treatment 
and  relative  self-fertility  of  seed  trees. 


Pollination  treatment 

and  yield  of  filled  seed 

Relative 

Tree 

Selfed 

Outcrossed 

self-fertility' 

Percent 

4 

8.7 

89.5 

9.72 

36 

15.7 

74.2 

21.2 

.58 

15.4 

lOO.O 

15.4 

MEAN 

14.4 

90.5 

15.4 

'Relative  self-fertility  (Sorenson 

1970)  is  computed  as: 

No.  of  filled  seed/total  number  of  seed  in  selfings 

No.  of  filled  seed/total  number  of  seed  in  crossings 


tilization  does  not  occur  in  pines  until  about 
one  year  after  pollination  and  cones  are 
made  up  entirely  of  female  parent  tissue,  so 
early  cone  abortion  is  not  related  to  such 
genetic  causes  as  homozygosity  of  recessive 
genes.  The  most  likely  explanation  of  a  lower 
average  level  of  cone  set  among  selfed  cones 
is  a  lower  level  of  pollination.  Pollen  of  indi- 
vidual trees  was  in  shorter  supply  than 
bulked  pollen,  and  may  have  been  used  too 
sparingly  in  a  few  cases,  re.sulting  in  close  to 
zero  pollination  of  ovules.  Sorenson  (1970) 
has  reported  75  percent  cone  set  in  out- 
crossed  cones  of  ponderosa  pine  {P.  ponderosa 
Laws.)  and  70  percent  in  .selfed  cones,  but  the 
difference  was  not  statistically  significant. 

Reduction  in  filled  seed  per  cone  by  selfing 
as  compared  to  outcrossing  is  a  common  find- 
ing among  pines.  Soreason  (1970)  found  that 
filled  seed  yields  in  ponderosa  pines  average 
23.7  percent  for  .self  and  66.5  percent  for  out- 
crosses;  in  Scotch  pine  (P.  sijlvestris  L.)  self- 
ing produced  13.4  percent  filled  .seed  and 
outcrossing  71.4  percent  (Johnsson  1976). 
Bramlett  and  Pepper  (1975)  reported  average 
filled  seed  yields  of  16.4  percent  and  90.7 
percent  for  selfed  and  outcrossed  cones  of 
Virginia  pine  (P.  virginiana  Mill.).  Squillace 
and  Kraus  (1962)  found  that  selfing  slash  pine 
{P.  elliottii  Engelm.)  resulted  in  9  sound  seeds 
per  cone,  and  outcrossing  produced  34  seeds 
per  cone.  Snyder  (1968),  also  working  with 
sla.sh  pine,  generalized  that  selfing  yields 
about  15  percent  as  many  seeds  as  wind  polli- 
nation. In  this  experiment,  the  filled-seed 
yield  of  selfed  pinyon  was  about  15  percent 
that  of  the  outcrosses. 

The  reduced  .sound-.seed  yield  in  selfed 
pines  and  other  conifers  is  due  to  the  en- 
hanced homozygosity  of  embryonic  lethal  al- 
leles. The  results  of  the  crosses  reported  here 
suggest  that  the  number  of  lethals  in  pinyon 
is  comparable  to  that  in  other  species  of 
Pinus. 


Acknowledgments 

This  work  was  funded  by  the  Mclntire- 
Stennis  program  and  is  published  as  Utah  Ag- 
ricultural Experiment  Station  Journal  Paper 
2549.  Mark  Buckbee  and  Michael  Jenkins  a.s- 
sisted  in  the  field  and  laboratory  work. 


September  1980 


Lanner:  Pinyon  Pollination 


267 


Literature  Cited 

Bramlett,  D.  L.,  and  \V.  D.  Pepper.  1974.  Seed  yield 
from  a  diallel  cios.s  in  Virginia  pine.  Pages  49-55 
in  Seed  Yield  from  Southern  Pine  Seed  Orchards 
Colloq.  Proc.,  Macon,  Ceorgia. 

CniRA,  E.  1967.  Pollen  grains  of  Pinus  edulis  with  more 
than  the  haploid  number  of  chromosomes.  Silvae 
Genet.  16:14-18. 

JonNssoN,  H.  1976.  Contributions  to  the  genetics  of 
empty  grains  in  the  seed  of  pine  (Pintts  silvestris). 
Silvae  Genet.  25:10-14. 

La.nner,  R.  M.  1970.  Origin  of  the  sunmier  shoot  of  pin- 
von  pines.  Canadian  ).  Bot.  .38:1759-1765. 

1974.  Natural  hybridization  between  Pinus  edulis 

and  Pinus  monophylla  in  the  American  South- 
west. Silvae  Genet.  2.3:108-116. 


1975.  Pinon  pines  and  junipers  of  the  south- 
western woodlands.  Pages  1-17  in  G.  F.  Gifford 
and  F.  E.  Busby,  eds.,  Proc.  Symp.  on  Pinyon- 
Juniper  Ecosystems.  Utah  State  University,  Lo- 
gan. 

Snyder,  E.  B.  1968.  Seed  yield  and  nurserv  performance 
of  self-pollinated  slash  pines.  Forest  Sci. 
14:68-74. 

SoRENSEN,  F.  C.  1970.  Self-fertility  of  a  central  Oregon 
source  of  ponderosa  pine.  USD.\  Forest  Service 
Res.  Pap.  PNW-109,  9  pp. 

SyuiLLACE,  A.  E.,  and  J.  F.  Krals.  1962.  Effects  of  in- 
breeding on  seed  yield,  germination,  rate  of  ger- 
mination, and  seedling  growth  in  slash  pine. 
Pages  59-63  in  South.  Forest.  Tree  Imp.  Comm. 
and  Soc.  Amer.  Foresters  Tree  Imp.  Comm.  For. 
Genet.  Workshop  Proc. 


COMPARATIVE  FLORAL  BIOLOGY  OF  PENSTEMON  EATONII  AND 
PENSTEMON  CYANANTHUS  IN  CENTRAL  UTAH:  A  PRELIMINARY  STUDY 

Lucinda  Bateman' 

Abstract.—  A  comparison  of  the  floral  visitors  of  two  closely  related  plant  species,  Penstemon  ctjanonthus  and  P. 
eatonii  suggests  that  flower  shape  and  color  may  affect  the  number  and  type  of  pollinators,  and  the  ability  of  the 
plant  to  set  fniit.  Penstemon  cijananthus,  which  is  most  attractive  to  hymenopteran  visitors,  has  a  blue  flower,  large 
in  diameter,  that  is  positioned  as  a  convenient  "landing  pad."  Although  many  types  of  insects  visit  the  flower,  the 
transport  of  pollen  directly  to  flowers  of  another  individual  of  the  same  species  is  somewhat  inefficient,  since  fruiting 
success  is  low  (66.7  percent).  The  tubular  red  flowers  of  P.  eatonii  are  narrow  and  droop  downward  from  the  stem. 
The  nectar  is  accessible  to  a  specific  and  well-adapted  visitor,  the  hummingbird.  This  less  promiscuous,  bird-polli- 
nated species  sets  fruit  more  successfully  (82.4  percent)  than  P.  cijananthus. 


Observations  of  animal  visitors  to  flowers 
suggest  that  the  broad  range  of  phenology, 
size,  structure,  color,  and  odor  evident 
among  flowers  of  any  complex  plant  commu- 
nity is  related  to  the  size,  morphology,  be- 
havior, and  sensory  acuity  of  the  animals  vis- 
iting the  flowers.  It  has  been  observed,  for 
instance,  that  nocturnal  blooming  flowers  are 
specially  adapted  to  night-flying  insects  or 
bats  (Faegri  and  Pijl  1971).  Bees  appear  to  be 
more  influenced  by  flower  shape  than  color. 
Bees  also  have  appendages  specialized  for 
collection  and  transport  of  pollen,  since  pol- 
len is  an  important  food  item  for  their  off- 
spring. Accordingly,  flowers  visited  by  bees 
may  be  white,  blue,  or  yellow,  but  commonly 
offer  a  generous  reward  of  both  nectar  and 
pollen  (Proctor  and  Yeo  1972,  Raven,  Evert, 
and  Curtis  1976).  Flowers  whose  most  fre- 
quent visitors  are  nonhovering  individuals 
such  as  bees  are  usually  so  structured  as  to 
provide  a  "landing  pad"  near  the  flower's  re- 
productive parts  and  nectar  or  pollen  "re- 
ward" (Free  1970). 

In  view  of  the  fact  that  few  insects  are  be- 
lieved able  to  distinguish  red  (Raven,  Evert, 
and  Curtis  1976),  it  seems  signiflcant  that  red 
flowers  worldwide  are  regularly  visited  by 
birds,  known  to  be  more  stimulated  by  that 
color  than  any  other  (Faegri  and  Pijl  1971).  It 
is  an  interesting  and  probably  not  imrelated 
fact  that  red  flowers,  unlike  most  flowers  of 
other  colors,  are  essentially  odorless  (Grant 


1966).  Significantly,  insects  have  keen  olfac- 
tory senses,  but  those  faculties  in  birds  tend 
to  be  poorly  developed  (Proctor  and  Yeo 
1972,  Faegri  and  Pijl  1971).  The  corolla  of 
many  flowers  visited  by  birds  is  typically 
tubular  and  narrow  and  without  a  landing 
platform,  excluding  all  but  the  smallest  insect 
intruders  (Raven,  Evert  and  Curtis  1976). 
Clearly,  flowers  that  are  tubular,  red,  and 
odorless  should  offer  minimal  attraction  to 
insects,  specifically  nonhovering  insects,  but 
should  be  highly  attractive  to  birds  (Boyd  and 
Brown  1978). 

Phenological,  structural,  color,  and  odor 
differences  among  the  flowers  of  any  particu- 
lar complex  plant  community  undoubtedly 
increase  the  degree  of  fidelity  between  par- 
ticular flower  types  and  specific  insects. 
Since  flowering  periods  of  different  species  in 
a  common  plant  community  often  overlap, 
flower-pollinator  fidelity  should  enhance  re- 
productive success  of  plant  species  having 
such  flowering  overlap.  Plants  able  to  selec- 
tively entice  pollinators  should  be  more  suc- 
cessful in  the  distribution  of  their  pollen.  The 
animal  visitor  should  simply  find  it  more 
profitable  to  visit  nonpromisCiious  flowers, 
since  there  is  a  greater  probability  that  such 
flowers  will  yield  a  reward  on  any  given  visit. 
Promiscuous  pollinators  would  be  particu- 
larly detrimental  to  the  reproductive  success 
of  rare  to  moderately  common  plants  that  are 
obligate  outcrossers,  and  that  flower  simulta- 


'Department  of  Botany  and  Range  Science,  Brigham  Young  University,  Prove,  Utah  84601.  Present  address:  1212  Ash  Avenue,  Provo.  Utah  84601. 


268 


September  1980 


Bateman:  Comparative  Floral  Biology 


269 


neously  with  a  variety  of  other  species,  since 
pollen  of  a  given  species  could  be  expected 
to  be  largely  dislodged  from  the  body  of  the 
promiscuous  pollinator  before  it  encountered 
another  individual  of  that  plant  species  (Le- 
vin and  Anderson  1970). 

Evidence  suggests  that,  by  chance  muta- 
tion, plants  gradually  develop  characteristics 
attractive  to  the  most  consistent  pollinators 
in  the  commimity.  Surviving  individuals  of 
the  species  become  specialized  for  visits  from 
the  more  efficient  pollinators.  Tantalizing 
odors  beckon  hungry  insects,  and  brightly 
colored  corollas  are  a  signal  to  the  pollinators 
of  the  presence  of  a  nectar  reward.  These 
forms  of  advertisement  attract  pollinators, 
and  thereby  accomplish  a  more  efficient 
spread  of  pollen. 

Statement  of  the  Problem 

Few  studies  have  been  made  of  the  com- 
parative floral  biology  of  two  species  of  the 
same  genus  growing  in  a  common  environ- 
ment and  flowering  simultaneously.  The  pur- 
pose of  this  study  is  to  compare  the  floral 
morphology,  insect  visitors,  and  fruit  set  of 
two  closely  related  species,  Pensternon  eaton- 
ii  Gray  and  Penstemon  cijananthus  Hook.  At 
the  site  studied,  these  species  grow  in  close 
proximity,  although  individuals  of  P.  cya- 
nanthiis  are  approximately  twice  as  numer- 
ous. I  have  tested  the  following  hypotheses: 
(a)  Corolla  size  and  color  affect  the  number 
and  types  of  pollinators,  (b)  P.  eatonii,  with  a 
red,  narrow  corolla  tube,  will  attract  fewer 
insects  and  will  be  visited  by  hummingbirds. 
(c)  P.  cijananthus,  with  a  broader,  blue  co- 
rolla, will  be  more  promiscuous,  attracting  a 
variety  of  insects,  (d)  The  less  promiscuous, 
bird-pollinated  species  will  set  fruit  more 
successfully. 

Materials  and  Methods 

Penstemon  eatonii  plants  produce  from 
5-10  inflorescences,  each  approximately  50 
cm  in  length.  Narrow  tubular  corollas  hang 
downward  along  the  upright  stem.  The  five- 
lobed  corolla  is  red.  Four  fertile  stamens  lie 
within  the  corolla  tube,  and  one  sterile  sta- 
men protrudes  beyond  the  corolla  orifice 
(Fig.  lA). 


Fig.  lA.  Penstemon  cifanantlms  flower  shape  and  ar- 
rangement of  reproductive  organs  (above);  IB.  Penste- 
mon eatonii  flower  shape  and  arrangement  of  reproduc- 
tive organs. 

Penstemon  cyananthus  generally  produces 
four  to  five  times  as  many  inflorescenses  as  P. 
eatonii,  each  approximately  45  cm  in  length. 
Each  blue,  five-lobed  corolla  is  displayed  at 
an  ascending  angle  from  the  stem  and  has 
fused  petals  and  five  stamens,  four  fertile  and 
one  sterile.  Two  stamens  are  as  long  as  the 
petals,  but  the  other  two  are  only  two-thirds 
that  length  (Fig.  IB). 

The  data  were  collected  29  June,  and  2 
July  1979  on  the  west  face  of  Mount  Tim- 
panogos,  in  Battlecreek  Canyon,  near  Pleas- 
ant Grove,  Utah.  A  small  stream  flows 
through  the  relatively  dry  study  site.  Some  of 
the  major  contributors  to  the  vegetation  of 
the  area  are:  sagebrush  {Artemisia  tridentata 
Nutt.),  rabbitbrush  {Chrysothamnus  nau- 
seosus  [Pall.]  Britt.),  poison  ivy  (Tox- 
icodendron radicans  L.),  scrub  oak  [Quercus 
gambelii  Nutt.),  big  tooth  maple  {Acer  gradi- 
dentatum  Nutt.),  chokecherry  {Prunus  virgi- 


270 


Great  Basin  Naturalist 


Vol.  40,  No.  3 


niana  L.),  squawbush  (Rhus  trilobata  Nutt.), 
and  various  grasses. 

Observations  of  the  plants  were  made  daily 
in  the  first  half  of  each  hour  from  0730  to 
1800  hours.  On  29  June  P.  cyananthus  was 
observed  during  the  first  hour,  P.  eatonii  dur- 
ing the  second,  and  so  on  throughout  the  day. 
On  2  July  the  observation  order  was  reversed. 
The  two  plants  observed  during  these  time 
periods  were  within  four  feet  of  each  other. 

Fruit  set  data  were  collected  14  July  1979, 
about  0.4  km  down  the  canyon  from  the  first 
observation  site.  Individuals  of  each  species 
were  randomly  selected  and  checked  for 
height,  spent  flowers,  developing  fruit,  open 
flowers,  and  buds.  Ten  stems  of  each  of  8  dif- 
ferent P.  cyananthus  plants  were  analyzed. 
Because  of  fewer  stems  per  plant,  13  P.  ea- 
tonii individuals  with  up  to  10  stems  per 
plant  were  also  tallied.  Of  the  total  flowers 
that  had  been  produced  per  inflorescence, 
both  species  showed  at  least  90  percent  spent 
flowers,  the  remainder  of  the  number  con- 
sisting of  open  flowers  and  buds. 

Average  fruit  set  was  calculated  using  the 
formula: 

No.  filled  fruits 


No.  spent  flowers 

Plant  nomenclature  follows  Welsh  and 
Moore  (1973).  Insect  family  names  are  taken 
from  Borror  and  White  (1970).  Bird  identi- 
fication is  from  Bobbins,  Brium,  and  Zim 
(1966). 


S    30- 


7  AM    8  9         10         11         12  1  2  3  4  5  6  PM 


Fig.  2.  Temperature  readings  in  degrees  Celsius  for 
29  June  (dotted  line)  and  2  July  (solid  line). 


Results 

Insect  activity  was  minimal  during  the 
early  morning  hours,  but  as  air  temperature 
rose,  greater  numbers  and  more  types  of  in- 
sects appeared.  Temperatures  for  the  two 
days  of  observation  were  similar  (Fig.  2). 

Penstemon  cyananthus  attracted  a  greater 
number  of  total  visitors  (153  in  two  days), 
and  also  more  insect  families  (9)  than  P.  ea- 
tonii (Table  1).  Not  all  insect  visitors  came  to 
the  plant  in  search  of  pollen  or  nectar.  Some 
coleopterans  landed  on  the  showy  petals  as  if 
to  rest,  making  no  attempt  to  enter  the  flow- 
er. Still  other  types  of  insects  crawled  among 
the  stems  and  leaves.  Neither  of  these  types 
of  visitors  were  recorded  as  pollinators.  The 
most  numerous  and  determined  visitors  were 
hymenopterans  in  search  of  pollen  (Table  1). 
Hymenopterans  accounted  for  85  percent  of 
the  visitors  and  over  89  percent  of  the  flow- 
ers visited.  Thirteen  percent  of  the  visitors  to 
P.  cyananthus  were  lepidopterans  and  the  re- 
maining 2  percent  were  dipterans. 

Penstemon  eatonii  attracted  fewer  insect 
visitors  (23)  or  15  percent  as  many  as  P.  cya- 
nanthus. The  visitors  that  were  observed  be- 


30 
28 

•s 

/     ^ 

26 

/ 

"^ 

24 

/ 

\       .'^ 

22 

I                                                                           1 

\  ,'    » 

1 

0                 \ 

20 

1 

\ 

18 

1 
1 

\ 
\ 

16 

^ 

\ 

14 

• 

\ 

r 

\ 

12 

* 
* 

\ 

10 

/ 

1 
1 

8 

/ 

I 

X                 . 

• 

6 

^     Av 

4 
2 

^y^^^X.^^^^ 

0 

••  ^    tL^^^>»^  /                 ^"^ 

T 

Fig.  3.  Hourly  distribution  of  individual  flower  vis- 
itors to  Penstemon  cijaniinthtis  (dotted  line)  and  P.  ea- 
tonii (solid  line)  throughout  the  observation  period. 


Fig.  4.  Hourlv  distribution  of  insect  families  that  vis- 
ited Penstemon  ri/anantlius  (dotted  line)  and  P.  eatonii 
(solid  line)  throughout  the  observation  period. 


September  1980 


Bateman:  Comparative  Floral  Biology 


271 


longed  to  three  hymenopteran  families.  Dur- 
ing tlie  heat  of  the  day,  when  the  greater 
numbers  of  insects  were  actively  foraging,  4 
or  5  attempted  to  enter  the  narrow  corolla 
tube.  Few  were  successful  in  their  efforts. 
Once  during  the  observation  period,  a  broad- 
tailed  hummingbird  visited  the  plant  at  0930, 
before  the  temperature  had  risen  above  24  C 
(75  F).  The  bird  moved  systematically  down 
the  canyon,  stopping  at  every  P.  eatonii  indi- 
vidual within  10  or  20  m  of  either  side  of  the 
path.  Upon  reaching  the  observation  plant. 


the  bird  sampled  five  or  six  flowers  on  four  of 
the  six  stalks,  pausing  less  than  one  second  at 
each  flower.  It  hovered  in  the  air  .slightly  be- 
low each  flower  (Figs.  3  and  4). 

Penstemon  cyananthu.s  averaged  66.7  per- 
cent fruit  set  per  plant,  and  P.  eatonii  showed 
a  much  higher  82.4  percent  (Table  2). 

Discussion 

The  results  of  this  study  clearly  indicate 
that  the  two  penstemons  considered  have  de- 


Table  1.  Activity  and  presence  of  each  family  throvigho\it  the  observation  period.  The  first  (upper)  n\imber  in- 
dicates the  number  of  individual  visitors;  the  second  (lower)  indicates  the  total  number  of  flowers  visited. 


Hour  observed 

Family 

8 

9 

10         11         12           1 

2 

3 

4 

5 

6 

Total 

Penstenion  cyanantlnis 

.\pidae' 
Chrysididae' 

1 
2 

1 
3 

3 
10 

3 
12 

1 
3 

3 

8 

6 
22 

Halictidae' 

Hesperiidae- 

Megachilidae' 

'Nymphalidae 

Pieridae- 

Syrphidae' 

Vespidae' 

Total 


1 

3 

1 

6 

2 

4 

2 

7 

3 

9 

10 

25 

2 
11 

6 
17 


10 

27 


14 
36 


5 

4 

3 

2 

2 

1 

21 

6 

4 

5 

6 

15 

2 

3 

2 

1 

1 

29 

13 

25 

/ 

16 

10 

10 

18 

2 

75 

6 
4 

39 

26 

15 

47 

5 

183 
6 

6 

2 
2 

1 
1 

1 

1 

11 

1 
1 

3 

3 

8 

2 

8 

2 

3 

25 

22 

6 

26 

9 

15 

89 

r; 

28 

26 

21 

24 

7 

153 

s 

(.3 

66 

46 

61 

23 

370 

Penstenwn  eatonii 


Formicidae' 

Halictidae' 

Megachilidae 

'Trochilidae' 

Total 


1 
15 

2 
16 


2 

9 

5 

17 

1 
15 

2 

1 

2.3 

5 

2 

54 

Key  to  the  orders:  1,  Hymenoptera;  2,  Lepidoptera^  3.  Diptera;  4,  Apodiformes  {'Selasphorus  platycercus). 


272 


Great  Basin  Naturalist 


Vol.  40,  No.  3 


Table  2.  Percent  fruiting  success  per  plant  of  Penste- 
mon  cyananthus  and  P.  eatonii  (Number  of  fruit/spent 
flowers/plant).  (Difference  significant  at  the  .05  level) 


Plant 

P.  cyananthus 

P.  eatonii 

% 

% 

1 

84.7 

45.5 

2 

47.5 

93.3 

3 

59.8 

73.1 

4 

47.6 

68.5 

5 

74.8 

89.7 

6 

75.5 

90.0 

7 

69.7 

77.4 

8 

79.3 

9 

90.8 

10 

86.1 

11 

95.4 

12 

82.4 

13 

100.0 

Average  % 

66.7 

82.4 

veloped  different  ways  of  attracting  pollina- 
tors. The  narrow  red  corolla  tube  of  P.  eaton- 
ii physically  excludes  all  but  a  few  small  in- 
.sect  visitors,  and  the  absence  of  odor  appears 
to  minimize  attractiveness  to  insect  visitors. 
The  absence  of  any  sort  of  a  landing  pad  hin- 
ders the  ability  of  nonhovering  visitors  to 
successfully  work  the  flowers.  The  only  ob- 
vious diurnal  pollinator,  a  hummingbird,  is 
le.ss  frequent  but  more  systematic  and  specif- 
ic. That  the  job  of  pollination  is  done  more 
efficiently  by  such  a  specific  pollinator  is  sug- 
gested by  the  higher  fruit  set. 

The  more  promiscuous  P.  cyananthus  also 
enjoys  a  fairly  high  fruit  set.  It  does  this, 
however,  with  larger,  more  accessible  blos- 
soms and  with  no  assurance  that  its  pollina- 
tors will  be  species  specific.  The  flowers  are, 
of  necessity,  displayed  so  as  to  form  a  con- 
venient landing  pad  for  approaching  hyme- 
nopterans. 


Limitations 

Although  the  initial  implications  are  clear, 
these  data  constitute  only  preliminary  results. 
Data  were  collected  during  daylight  hours 
late  in  the  flowering  season,  when  flowers  of 
both  species  contained  little  or  no  nectar. 
Only  one  hummingbird  was  observed  directly 
during  the  study,  but  my  presence  may  have 
frightened  usual  avian  visitors  away.  No  at- 
tempt was  made  to  observe  early  evening  or 
nocturnal  pollinators.  The  degree  to  which 
either  species  is  capable  of  self-pollination  is 
unknown. 

Literature  Cited 

Bond,  H.  W.,  and  W.  Brown.  1978.  The  exploitation  of 
floral  nectar  in  Eucalyptus  incrassata  by  honey- 
eaters  and  honeybees.  Unpublished  manuscript. 

BoRROR,  D.  L.,  and  R.  E.  white.  1970.  A  field  guide  to 
the  insects.  Houghton  Mifflin  Co.,  Boston. 

Faegri,K.,  and  L.  van  der  Pijl.  1971.  The  principles  of 
pollination  ecology.  2d  ed.  Pergamon  Press,  Ox- 
ford, England. 

Free,  J.  B.  1970.  Effect  of  flower  shapes  and  nectar 
guides  on  the  behavior  of  foraging  honeybees.  Be- 
havior 37:269-285. 

Grant,  K.  A.  1966.  A  hypothesis  concerning  the  preva- 
lence of  red  coloration  in  Californian  humming- 
bird flowers.  American  Nat.  100:85-98. 

Levin,  D.  A.  1969.  The  effect  of  corolla  color  and  out- 
line on  interspecific  pollen  flow  in  Phlox.  Evolu- 
tion 23:444-445. 

Proctor,  M.,  and  P.  Yeo.  1972.  The  pollination  of  flow- 
ers. Taplinger  Publishing,  New  York. 

Raven,  P.,  R.  F.  Evert,  and  H.  Curtis.  1976.  Biology  of 
plants.  2d  ed.  Worth  Publishers,  New  York. 

Bobbins,  C.  S.,  B.  Bruun,  and  H.  Zim.  1966.  Birds  of 
North  America.  Western  Publishing  Co.,  New 
York. 

Welsh,  S.  L.,  and  G.  Moore.  1973.  Utah  plants: 
Tracheophyta.  Brighain  Young  University  Press, 
Provo,  Utah. 


DIFFERENTIAL  HABITAT  UTILIZATION  BY  THE  SEXES  OF  MULE  DEER 

Michael  M.  King'  and  H.  Duane  Smith' 

Abstract.-  Habitat  segregation  trends  have  been  observed  and  published  for  the  sexes  of  mule  deer  {Odocoilem 
hcmionus)  based  on  elevation  and  slope  exposure.  Despite  these  brief  descriptions,  quantitative  studies  on  habitat 
segregation  by  the  sexes  of  mule  deer  are  lacking.  Results  of  research  conducted  in  central  Utah  indicated  no  signifi- 
cant difference  in  elevation  positions  used  by  males,  but  did  show  significant  difference  in  utilization  of  studv  sites 
based  on  slope  exposure,  relative  percentage  forb  cover,  and  relative  percentage  hiding  cover.  Males  were  most  com- 
mon at  sites  characterized  by  low  forb  abundance  and  hiding  cover,  and  on  south-facing  exposures.  Females  were 
most  common  at  sites  characterized  by  high  forb  abundance  and  hiding  cover,  and  on  north-facing  exposures.  Pos- 
sible advantages  of  habitat  separation  to  both  sexes  and  management  implications  are  discu.ssed. 


Mule  deer  {Odocoileus  hemionus),  like 
many  other  ungulates,  seem  to  exhibit  habitat 
partitioning  between  sexes  (Darling  1937, 
Estes  1974,  Geist  1974,  1977,  Gest  and  Pe- 
tocz  1977,  Hirth  1977,  Leuthold  1978). 
DeVos  et  al.  (1967)  indicated  that  male  and 
female  mule  deer  are  separated  throughout 
the  year  with  the  exception  of  the  breeding 
season.  Dasmann  and  Taber  (1956)  found  that 
males  occupied  more  open  south-facing 
slopes  and  females  occupied  densely  vegeta- 
ted north-facing  slopes.  Several  workers  have 
suggested  that  males  prefer  higher  altitudes 
and  ridge  tops  more  than  do  females  (Cowan 
1956,  Miller  1970).  The  same  trend  was  ob- 
served in  a  Nevada  mule  deer  herd  by  Robin- 
ette  et  al.  (1977)  where  subalpine  and  alpine 
conditions  were  prevalent.  Males  were  found 
predominantly  above  3000  m  elevation, 
whereas  females  were  more  often  below  2500 
m  elevation.  Although  habitat  separation  by 
male  and  female  mule  deer  has  been  report- 
ed, little  attempt  has  been  made  to  quantify 
differential  habitat  use  or  to  describe  site  dif- 
ferences other  than  to  suggest  slope  exposure 
and  elevational  differences. 

The  objectives  of  this  study  were:  (1)  to  de- 
termine quantitatively  if  male  and  female 
mule  deer  differentially  utilize  habitat,  (2)  to 
suggest  possible  advantages  to  habitat  separa- 
tion by  sexes  of  mule  deer,  and  (3)  to  identify 
critical  management  problems  related  to  dif- 
ferential resource  utilization  between  male 
and  female  mule  deer. 


Study  Area 

The  study  was  conducted  on  the  Bighorn 
Ranch,  a  privately  owned  ranch  in  the  Nebo 
Range  of  the  Wasatch  Mountains,  Utah.  The 
study  area  was  approximately  1130  ha  in 
size,  ranging  from  2200  to  2500  m  elevation. 
Human  access  is  restricted,  thus  providing  a 
relatively  undisturbed  area  for  observation  of 
mule  deer  behavior,  distribution,  and  habitat 
utilization.  Ridge  tops,  south-facing  slopes, 
and  other  well-drained  areas  were  dominated 
by  Gambel  oak  {Quercus  gamhelii)  and  big 
sagebrush  {Artemisia  tridentata)  communities 
with  little  herbaceous  growth.  Drainage  bot- 
toms, north-facing  slopes,  and  well-watered 
areas  were  dominated  by  quaking  aspen  {Pop- 
ulus  tremuloides).  Rocky  Mountain  maple 
(Acer  glabrum),  and  chokecherry  {Prttnus  vir 
giniana)  communities,  with  numerous  forb 
species  in  the  understory. 

Methods 

Preliminary  observations  made  in  Septem- 
ber 1977  to  determine  deer  distribution  in 
the  various  watersheds  of  the  ranch  indicated 
habitat  segregation  between  male  and  female 
mule  deer.  Based  on  that  survey,  the  follow- 
ing spring  nine  study  sites  (Fig.  1)  were  se- 
lected where  deer  numbers  were  relatively 
high.  Other  areas  had  equally  as  many  deer, 
but  excessive  area  or  distance  from  access 
roads  made  observation  unrealistic. 


'Department  of  Zoology,  Brigham  Young  University.  Provo,  Utah  84602. 


273 


274 


Great  Basin  Naturalist 


Vol.  40,  No.  3 


THE    BIGHORN    RANCH 


O  Study  Sites 


—  Ranch  Border 


Nephi    and 
Fountain  Green 


Highway  89 


N 


Highway  89 


Fountain  Green 


Fij;.  1,   Map  ot  the  Bit^honi  Ranch  showing  the  approximate  location  of  study  sites. 


September  1980 


King,  Smith:  Deer  Habital  Utilization 


275 


Each  study  site  was  observed  weekly  from 
1  June  1978  to  1  September  1978.  Observa- 
tions were  made  from  established  observation 
points  or  by  vehicle  from  sunrise  until  late 
morning  or  from  early  afternoon  until  dark. 
Ob.servation  time  for  each  site  was  alternated 
weekly  between  morning  and  evening  so  ap- 
proximately equal  observation  time  was 
spent  at  each  site  during  each  time  period.  A 
variable  15-45X  spotting  scope  and  12X  bi- 
noculars were  used  for  daytime  observation, 
and  a  200,000  candlepower  spotlight  oper- 
ated through  the  electrical  system  of  the  ve- 
hicle, along  with  spotting  scope  and  binocu- 
lars, was  used  for  observation  at  night.  Total 
observation  time  for  the  study  exceeded  900 
hours. 

Observed  deer  were  recorded  according  to 
sex,  slope  position  (Fig.  2),  and  slope  expo- 
sure. A  2X4  contingency  analysis  (Zar  1974) 
was  performed  to  determine  significant  dif- 
ferences in  utilization  of  slope  positions  by 
males  and  females.  Total  numbers  of  males 
and  females  recorded  at  north-  and  south-fac- 
ing exposures  were  also  subjected  to  contin- 
gency analysis  to  determine  slope  exposure 
usage  differences.  To  characterize  study  sites 


two  critical  parameters,  relative  abundance 
of  forb  and  hiding  cover,  were  examined  at 
each  site.  Forbs  were  defined  as  succulent, 
low-growing,  nonwoody  vegetation,  and  hid- 
ing cover  as  vegetation  more  than  2  m  in 
height.  Both  estimates  were  determined  by  a 
line-point  transect  method  for  determining 
relative  abundance  of  vegetation  (Kershaw 
1973). 

Simple  correlation  procedures  relating  rel- 
ative abundance  of  forb  and  hiding  cover 
with  the  corresponding  male/female  ratio  for 
each  site  were  used  to  determine  if  utiliza- 
tion of  sites  by  males  and  females  differed 
significantly  based  on  forb  abundance  and 
hiding  cover  (Zar  1974).  The  maximum  prob- 
ability accepted  for  statistical  significance 
was  0.05;  probabilities  less  than  0.01  were 
considered  highly  significant. 

Results  and  Discussion 

Analysis  of  slope  positions  used  by  males 
and  females  (Table  1,  Fig.  3)  showed  no  sig- 
nificant difference  in  slope  position  utiliza- 
tion by  sex  (P  =  0.06).  The  data,  though  not 
significant  at  the  0.05  level,  approach  signifi- 


SLOPE     POSITIONS 


^^^^jM-i^-jt-^is^aN. 


Fig.  2.  Diagram  of  the  four  slope  positions  at  each  site;  slope  position  1  =  canyon  bottom  to  '4  slope,  slope  position 
2=1/2  slope  to  '/2  slope,  slope  position  3  =  '/2  slope  to  %  slope,  and  slope  position  4  =  \  slope  to  ridgetop. 


276 


Great  Basin  Naturalist 


Vol.  40,  No.  3 


Table  1.  Total  number  of  male  and  female  mule  deer  at  each  slope  position  for  all  study  sites.  2X4  contingency 
analysis  indicates  a  nonsignificant  difference  in  utilization  of  slope  position  by  male  and  female  mule  deer  (X-  =  7.42, 
df  =  3,  P  =  0.06).  Numbers  in  parentheses  are  expected  values. 


Slope  position 

Sex 

1 

2 

3 

4 

Total 

Males 

74 
(86.9) 

63 

(56.4) 

42 

(39.9) 

14 

(9.8) 

193 

Females 

157 

(144.1) 

87 
(93.4) 

64 
(66.1) 

12 

(16.2) 

320 

Total 

231 

150 

106 

26 

513 

cance  and  indicate  support  for  elevational 
segregation  observed  in  other  areas.  Further 
categorical  analysis  to  determine  usage  of  in- 
dividual slope  positions  by  males  and  females 
shows  that  considerable  differences  exist  in 
utilization  of  slope  positions  1  and  4  by  males 
and  females  (Fienberg  1977),  with  more  fe- 


males than  males  at  slope  position  1,  and 
more  males  than  females  at  slope  position  4 
(position  1  male  =  -.208,  position  1  females 
=  +  .208;  position  4  males  =  -I-  .245,  posi- 
tion 4  females  =  -.245;  positive  values  in- 
dicate most  usage  at  slope  position).  Analysis 
of  slope  exposure  use  by  males  and  females 


UJ 

< 


O 
111 

< 


100    -I 


75    - 


^         50    - 


+      FEMALES 
•      MALES 


Ik 
O 


O 

o 

u 

85 


25 


T 1 r 

1  2  3 

SLOPE    POSITION 


Fig.  3.  Percentage  of  total  males  and  total  females  observed  at  each  slope  position. 


September  1980 


King,  Smith:  Deer  Habital  Utilization 


277 


showed  a  significant  difference  (P< 0.005), 
with  males  most  often  at  south-facing  slopes 
and  females  most  often  at  north-facing  slopes 
(Table  2,  Fig.  4). 

Calculated  male /female  ratios  for  each  site 
correlated  with  corresponding  relative  per- 
centages of  forb  and  hiding  cover  (Table  3, 
Fig.  5)  showed  a  highly  significant  negative 
correlation  between  forb  abundance  and 
male/female  ratios  (r  =  -.89,  df  =  7, 
P< 0.005)  and  a  significant  negative  correla- 
tion between  male/female  ratios  and  hiding 
cover  (r=  -.69,  df  =  7,  P<0.05).  Therefore, 
as  forb  and  hiding  cover  increased,  the 
male/female  ratios  decreased,  indicating  that 
females  select  areas  characterized  by  rela- 
tively high  forb  and  hiding  cover  densities, 
but  males  select  areas  characterized  by  low 
forb  and  hiding  cover  densites. 


Table  2.  Total  number  of  male  and  female  mule  deer 
at  each  slope  exposure  for  all  study  sites.  2X2  contin- 
gency analysis  indicates  a  significant  difference  in  utili- 
zation of  slope  exposure  bv  male  and  female  mule  deer 
(X^  =  29.3,  df  =  1,  P<  0.005).  Numbers  in  parentheses  are 
expected  values. 


Slope  exposure 

Sex 

North-facing 

South-faciiiu 

T..i,,l 

Males 

64 

(93.7) 

129 

(99.3) 

193 

Females 

185 

(155.3) 

135 

(164.7) 

320 

Total 

249 

264 

513 

To  comprehend  implications  of  this  pat- 
tern of  spatial  separation,  po.ssible  advantages 
accrued  by  males  and  females  in  their  spring- 
summer  habitats  should  be  examined.  During 
the  spring-summer  season  perhaps  the  most 
critical  events  to  females  are  production  and 


100  -1 


3 

o 

Q. 
X 


Q. 

o 


< 


< 

ui 

< 


z 
< 

UJ 

< 


75  - 


50  - 


67 


33 


25  - 


NORTH-FACING 
EXPOSURE 

SOUTH-FACING 
EXPOSURE 


MALES 


FEMALES 


Fig.  4.  Percentage  of  total  males  and  total  females  observed  at  each  slope  exposure. 


278 


Great  Basin  Naturalist 


Vol.  40,  No.  3 


§ 

U 

flO 

O 

Ik 


> 
< 

Ul 


> 
O 

o 

z 

o 

z 

Ul 

> 


50      - 


40      - 


30     - 


20      - 


10      - 


r=  -.89 
P<0.005 


50      - 


40       - 


=         30       - 


20       - 


10       - 


»'=-.69 
P<0.05 


T" 

1.0 


2.0 


1^ 

3.0 


"T" 

4.0 


male/female     ratios 


Fig.  5.  Correlation  of  relative  percentage  fort)  cover  and  relative  percentage  hiding  cover  with  corresponding 
male/female  ratio  for  each  study  site. 


September  1980 


King,  Smith:  Deer  Habital  Utilization 


279 


Table  3.  Relative  percentage  forb  cover,  relative  percentage  hiding  cover,  total  number  of  male  and  female  mule 
deer,  and  male/female  ratios  for  each  study  site. 


Site  No. 


Relative 
%  forb  cover 


Relative 
hiding  cover 


Males 


Females 


M/F  ratios 


40.50 

19.00 

7 

63 

0.11 

32.25 

14.00 

6 

39 

0.15 

41.25 

19.(X) 

12 

61 

0.20 

46.50 

22.(X) 

2 

25 

0.08 

37.75 

16.00 

7 

46 

0.16 

16.50 

15.00 

52 

29 

1.80 

19.00 

13.00 

32 

23 

1.40 

20.75 

8.00 

38 

15 

2.53 

19.75 

17.00 

37 

19 

1.95 

rearing  of  offspring.  This  implies  that  consid- 
erable energy  is  apportioned  to  gestation, 
parturition,  and  lactation  beyond  normal 
body  maintenance  requirements  (Nelson 
1975,  Stebbins  1977).  If  these  requirements 
are  not  met  through  diet  resources,  body  re- 
serves are  utilized,  thus  reducing  offspring 
vigor  and  survival.  Since  energy  demands  for 
offspring  production  exceed  normal  energy 
requirements,  it  is  important  that  females  oc- 
cupy areas  where  nutritious,  high-quality  for- 
age is  readily  available.  Adequate  nutrition 
insures  successful  offspring  production  and 
facilitates  proper  lactation.  Research  in- 
dicates that,  during  the  time  period  critical 
to  fawn  production,  high  moisture  content, 
ease  of  digestion,  and  increased  nutrition 
content  (Smith  1952,  Short  1966,  Short  and 
Reagor  1970,  Boeker  et  al.  1972)  make  the 
forbs  preferred  diet  items  of  mule  deer 
(Smith  1952,  Morris  and  Schwartz  1957,  Lo- 
vass  1958,  Anderson  et  al.  1965,  Crouch 
1966,  Dasmann  et  al.  1967,  Nelson  1975).  It 
has  also  been  shown  that  deer  herds  having  a 
variety  of  succulent  forage  in  their  diets  have 
greater  herd  productivity  and  vigor  than 
those  that  utilize  a  greater  percentage  of 
woody  vegetation  (Biswell  1961,  Julander  et 
al.  1961,  Boeker  et  al.  1972,  Nelson  1975,  Pe- 
derson  and  Harper  1978).  We  assume,  there- 
fore, that  forb  abundance  is  related  to  site 
quality  and  that  areas  of  high  forb  abundance 
are  considered  high  quality,  whereas  areas  of 
low  forb  abundance  are  low  quality.  Deer 
that  select  forb-rich  areas  would  have  survi- 
val advantages  because  of  availability  of 
choice  forage.  It  follows  that  females  should 
select  high-quality  sites  to  assure  adequate 
nutrition  and  energy  for  fawn  production  and 


survival.  However,  it  seems  strange  that 
males  would  occupy  areas  of  low  forb  abun- 
dance when  selection  of  high-forb  areas 
would  more  readily  insure  adequate  energy 
for  increasing  body  size,  accumulating  fat  re- 
serves, and  developing  antlers.  We  suggest  as 
a  partial  explanation  that  it  is  more  advanta- 
geous for  the  sexes  to  be  separated  to  reduce 
chances  of  energy-expensive  agonistic  expres- 
sion between  males  and  females  (Geist  and 
Petocz  1977,  McCullough  1979).  This  allows 
energy  allocation  to  gestation,  parturition, 
and  lactation  rather  than  to  stresses  of  haras.s- 
ment.  Females  on  ranges  uncontested  by 
males  should  leave  more  offspring,  and  those 
behavioral  traits  responsible  for  habitat  parti- 
tioning should  be  selected  to  increase  the 
population.  Males  that  did  not  compete  for 
resources  necessary  for  fawn  production, 
though  occupying  lower-quality  sites,  should 
likewise  leave  more  offspring  to  succeeding 
generations  than  males  whose  behavior  bring 
them  into  competition  with  their  offspring 
(Wilson  1975,  Geist  and  Petocz  1977, 
McCullough  1979). 

Differential  habitat  use  patterns  could  also 
confer  advantages  to  males  and  females  by 
increasing  odds  for  predator  avoidance.  Ecol- 
ogists  working  on  vmgulates  have  suggested  a 
theory  of  predator  avoidance  based  on  the 
relative  degree  of  habitat  openness  and  group 
size.  Ungulates  that  inhabit  densely  vegetated 
areas  usually  occur  as  solitary  animals  or  in 
small  groups  that  use  hiding  as  a  mechanism 
for  predator  avoidance,  whereas  ungulates 
that  occupy  open  areas  are  primarily  herding 
animals  that  rely  on  the  use  of  collective 
senses  and  high  mobility  for  predator  escape 


280 


Great  Basin  Naturalist 


Vol.  40,  No.  3 


(Dasmann  and  Taber  1956,  Kitchen  1974, 
Hirth  1977). 

Life  history  studies  of  mule  deer  (Linsdale 
and  Tomich  1953,  Robinette  et  al.  1977), 
along  with  personal  observations,  give  sup- 
portive evidence  to  this  hypothesis.  Males 
and  females  form  different-sized  groups  in 
their  preferred  habitats.  During  spring  and 
summer  months  females  seek  isolation  in 
areas  where  hiding  cover  is  relatively  abun- 
dant (Fig.  5)  and  tolerate  few  deer  other  than 
their  offspring  of  the  year.  This  partial. soli- 
tary existence  has  advantages  from  a  predator 
avoidance  standpoint  in  that  females  can  se- 
cret themselves  and  their  offspring  in  dense 
vegetation  during  periods  of  high  vulnera- 
bility to  potential  danger.  After  the  young 
are  bom,  maternal  duties  restrict  the  mobil- 
ity and  escape  efficiency  of  females.  It 
would,  therefore,  be  advantageous  for  fe- 
males with  fawns  to  avoid  open  habitats 
where  predators  can  detect  and  capture  them 
or  their  fawns  more  easily. 

In  contrast,  males  on  the  Bighorn  Ranch 
during  the  same  time  period  were  often  ob- 
served to  form  fraternal  groups  in  areas  with 
relatively  low  abundance  of  hiding  cover 
(Fig.  5).  Male  groups  that  inhabit  open  areas 
of  high  visibility  can  collectively  monitor 
their  surroundings  and  take  advantage  of  rap- 
id flight  when  escape  is  necessary.  This  is 
better  than  hiding  in  dense  cover,  where  ef- 
fectiveness of  concealment  would  be  reduced 
by  large  groups  and  would  increase  chances 
of  detection  by  predators. 

Furtlier  supportive  evidence  for  the  pre- 
dator avoidance  theory  is  provided  by  the 
differential  use  of  slope  positions  1  and  4  by 
males  and  females.  More  females  than  males 
occupied  slope  position  1,  the  lower  position 
of  the  slope  where  the  greatest  abundance  of 
hiding  cover  was  located.  This  suggests  the 
importance  of  cover  to  females.  More  males 
than  females  used  slope  position  4,  the  open 
ridges,  suggesting  preference  by  males  for  the 
areas  of  high  visibility. 

In  this  study  habitat  separation  by  males 
and  females  is  primarily  based  on  slope  expo- 
sure, forb  abundance,  and  hiding  cover,  with 
some   evidence,   though   not   significant,   for 


elevational  segregation.  We  do  not  detract 
from  the  importance  of  elevational  segrega- 
tion as  it  has  been  observed  frequently  in 
other  areas,  but  suggest  to  wildlife  managers 
that  there  are  several  habitat  separation  pos- 
sibilities, depending  on  characteristics  of 
mule  deer  range  in  a  given  area.  We  encour- 
age that  further  research  delineating  segrega- 
tion characteristics,  advantages,  and  mecha- 
nisms be  initiated  so  that  management 
implications  can  be  evaluated. 

Knowledge  of  habitat  separation  between 
the  sexes  of  mule  deer  will  have  considerable 
influence  on  several  critical  management 
problems.  Three  important  problems  as  we 
view  them  are  now  discussed.  First,  as  deer 
populations  are  being  censused  in  various 
areas  and  habitats,  a  prime  concern  is  the  de- 
termination of  an  accurate  sex  ratio.  If  man- 
agers are  not  aware  of  site-specific  habitat 
separation  by  male  and  female  deer,  biases 
favoring  one  sex  over  the  other  will  arise  in 
calculated  ratios  depending  on  the  area  sam- 
pled. Failure  to  determine  accurate  sex  ratios 
will  allow  faulty  plans  to  be  devised  and  im- 
plemented. Second,  through  recognition  of 
specific  habitat  requirements  of  the  sexes,  it 
is  possible  that  habitat  can  be  manipulated 
through  appropriate  techniques  to  create 
conditions  favorable  to  either  sex.  This  will 
allow  sex  ratio  manipulation  depending  on 
management  needs.  Third,  critical  areas  to 
females  and  offspring  as  well  as  males  must 
be  protected  from  detrimental  commercial, 
industrial,  and  recreational  development.  De- 
struction of  important  fawning  areas  through 
development  will  force  females  to  occupy 
suboptimal  habitats  and  result  in  reduced 
fawn  production  and  survival.  Development 
in  areas  occupied  predominantly  by  males 
will  restrict  fall  hunting  and  ultimately  re- 
duce herd  productivity  if  adequate 
male/female  ratios  are  not  maintained. 

An  understanding  of  how  male  and  female 
mule  deer  partition  the  habitant  and  how  hab- 
itats preferred  by  females  differ  from  those 
most  frequented  by  males  will  undoubtedly 
improve  abilities  to  effectively  manage  mule 
deer  habitat.  Proper  use  of  knowledge  re- 
garding differential  habitat  and  resource  uti- 
lization by  the  sexes  of  mule  deer  can  in- 
crease the  efficiency  with  which  agencies 
manage  the  deer  resource. 


September  1980 


King,  Smith:  Deer  Habital  Utilization 


281 


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TEMPORAL  ACTIVITY  PATTERNS  OF  A  DIPODOMYS  ORDII  POPULATION 

Clive  D.  Jorgensen,'  H.  Diiane  Smith,',  and  James  R.  Garcia' 

Abstract.—  Temporal  activity  patterns  for  Dipodomijs  ordii  were  generally  bimodal  during  the  summer,  with  the 
highest  peak  occurring  during  early  predawn  hours  when  conditions  were  optimum  for  water  conservation.  Removal 
of  dominant  members  in  the  population  resulted  in  a  substantial  shift  in  the  activity  pattern  to  increased  activity 
during  the  evening  hours. 


Ecologists  studying  small  mammals  must 
contimially  attempt  to  resolve  the  problems 
of  inadequate  methods  to  study  daily  activity 
patterns  in  natural  environments,  especially 
when  studying  secretive  and/ or  nocturnal 
species  that  require  trapping  procedures. 
Usefulness  of  temporal  activity  data  has  been 
demonstrated  in  simulation  trapping  studies 
(Burnham  and  Overton  1969,  Manly  1970, 
Jorgensen  et  al.  1972),  population  estimator 
studies  (Scott  et  al.  1978),  and  energetics 
studies  (Kenagy  1973),  among  others.  Al- 
though methods  for  obtaining  data  under 
field  conditions  have  not  been  well  devel- 
oped, some  have  been  reported  (Jorgensen 
and  Hayward  1965,  Eider  1968,  Marten 
1973).  Generally,  their  results  are  deficient  in 
one  or  more  of  the  following:  (1)  numbers  of 
recorded  activity  events  per  day,  (2)  timing 
of  the  observed  or  measured  activity  event, 
(3)  ability  to  assign  an  activity  event  to  a  spe- 
cific individual,  and  (4)  correlation  between 
the  measured  activity  under  laboratory  con- 
ditions with  analogous  activity  in  natural  en- 
vironments. 

Harling  (1971)  attempted  to  relieve  some 
of  the  difficulty  in  recording  activity  by  de- 
veloping a  trap  that  could  be  continually 
monitored  during  a  trapping  period.  His 
traps  were  electrically  wired  to  a  central 
communications  console.  Using  walkie-talkie 
communications  between  someone  on  the 
grid  and  another  at  the  console,  Harling 
(1971)  was  able  to  obtain  the  precise  time 
when  an  animal  was  caught  as  well  as  when 
it  was  released. 

We  extended  Harling's  (1971)  methods  to 
include  an  entire  grid  of  traps  comparably 
wired  (Garcia  et  al.  1974)  to  study  a  popu- 

'Department  of  Zoology.  Brigham  Voting  University,  Provo,  Utah  84602. 


lation  of  Dipodomys  ordii  under  field  condi- 
tions. Our  objectives  were  to  determine  the 
optimum  foraging  times  for  D.  ordii  in  the 
salt  desert  shrub  community  of  west-central 
Utah,  and  illustrate  activity  patterns  that 
could  assist  in  interpreting  trapping  data 
from  other  studies  that  include  this  species. 

Study  Site  and  Methods 

Data  were  collected  from  two  sites  at  the 
Desert  Range  Experiment  Station,  Millard 
Co.,  Utah,  from  21  August  to  3  September 
1971  and  25  June  to  27  August  1973.  Al- 
though D.  ordii  was  the  species  trapped  most 
frequently  during  this  study,  Perognathus 
longimernbris  was  common  and  Peromyscus 
maniculatus  and  Onychomys  leucogaster 
were  trapped  occasionally.  Both  sites  were 
sandy  and  dominated  by  Oryzopsis  hyme- 
noides,  Chrysothamnus  nauseosus,  and  Sal- 
sola  kali,  although  Ambrosia  acanthicarpa. 
Astragalus  spp.,  Atriplex  canescens,  Gilia 
hiitchinsifolia,  and  Hilaria  jamesii  also  were 
present. 

The  trap  design  and  surveillance  methods 
were  described  in  detail  by  Garcia  et  al. 
(1974).  A  10  X  10  (100  traps)  grid  was  wired 
to  a  central  communications  console,  where 
one  researcher  recorded  traps  as  they  were 
"set  off."  He  then  informed  an  assistant 
working  on  the  grid  where  trapped  animals 
were.  Animal  data  were  radioed  from  the  as- 
sistant to  the  researcher  at  the  console  by 
walkie-talkie.  Data  were  then  recorded  and 
the  trap  reset.  Animals  seldom  were  detained 
in  traps  for  more  than  a  few  minutes.  Data 
collected  in  our  study  included:  species,  rela- 
tive age  (juvenile,  subadult,  adult),  sex,  repro- 


282 


September  1980         Jorgensen  et  al.:  Dipodomys  Activity  Patterns 


283 


ductive  condition,  and  identification  mark 
(toe  clip).  Supportive  data,  used  as  the  inde- 
pendent variables  in  our  analyses  and  collect- 
ed each  time  an  animal  was  captured,  were: 
ambient  temperature,  soil  temperature,  wind 
speed,  cloud  cover  and  lunar  events  (sub- 
jectively assessed),  light  intensity,  barometric 
pressure,  and  relative  humidity.  Absolute  hu- 
midities (gm/m^)  were  determined  using  the 
methods  described  by  Piatt  and  Griffiths 
(1964),  i.e.,  X  =  217(RH)  (^'J/IOOT,  where  T 
is  degrees  Kelvin  and  e^  is  vapor  pressures  in 
air  saturated  with  water. 

The  period  between  sundown  and  sunrise 
was  divided  into  20  subperiods,  which  were 
used  as  the  time  units  while  plotting  activity. 
Since  activity  periods  changed  slightly  as  day 
lengths  changed,  it  was  necessary  to  establish 
a  standard  set  of  subperiods  before  data  for 
different  periods  could  be  pooled.  Stepwise 
regression  methods  were  used  to  assess  the  ef- 
fects of  independent  variables  on  activity 
rates  among  the  pooled  data  for  all  activity 
subperiods.    Differences    in    activity    among 


sexes,  ages,  and  reproductive  condition  class- 
es were  determined  using  Chi-square  tests  of 
independence. 

An  opportunity  to  assess  possible  effects  of 
socially  dominant  individuals  in  the  D.  ordii 
population  was  noted  after  three  weeks  of  ac- 
tivity data  had  been  gathered  in  1973.  An  an- 
imal was  considered  dominant  if  it  was  the 
only  adult  captured  within  the  area  pre- 
scribed when  its  capture  points  had  been 
connected,  or  if  it  was  the  only  adult  repeat- 
edly caught  in  a  specific  trap.  Eleven  domi- 
nant individuals  among  the  42  individuals  re- 
corded on  the  grid  were  removed  from  the 
poulation  and  activity  data  gathered  for  an 
additional  five  days.  Temporal  activitv  pat- 
terns of  the  populations  before  and  after  the 
selected  removals  were  compared. 

Results  and  Discussion 

Activity  for  the  intact  population  of  D.  or- 
dii was  essentially  bimodal,  with  the  greatest 
activity  occurring  during  the  predawn  hours 


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2.0- 


1.0  - 


Dipodomys  ordii  (Dominants  included) 
Dipodomys  ordii  (Dominants  removed  ) 


I  I  I  ■  ■ 

7  9  11  13  15 

Activity  Subperiods 


17 


19 


21 


Fig.  1.  Activity  patterns  for  Dipodomys  ordii  pooled  among  20  subperiods  over  a  24-hoiir  activity  period. 


284 


Great  Basin  Naturalist 


Vol.  40,  No.  3 


(Fig.  1).  These  observations  agree  with  Jor- 
gensen  and  Hayward  (1965),  although  our 
early  evening  peak  is  not  as  distinct.  Since 
their  data  were  gathered  over  the  entire  year 
from  trapping  rates  per  hour,  we  question 
whether  their  results  are  directly  comparable 
with  ours.  Although  we  have  no  season-spe- 
cific data  to  demonstrate  seasonal  variations 
in  activity  patterns,  any  variation  would  pro- 
duce composite  patterns  difficult  to  interpret 
and  compare. 

Precipitation  (11.4  cm)  was  unusually  low 
in  1971  and  rather  high  (20.5  cm)  in  1973. 

I 


0 
Q. 

a 
o 


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o 
E 
'E 
< 


These  differences  in  moisture  were  accom- 
panied by  comparable  differences  in  vegeta- 
tive production— low  in  1971  and  high  in 
1973  (Jorgensen,  unpublished  data).  Chi- 
square  analyses  resulted  in  only  6  of  a  pos- 
sible 20  tests  being  significant  (p  =  .95).  All 
the  6  cohorts  of  animal  classes  that  were  sig- 
nificant included  adults  (Table  1).  The  ober- 
vations  suggest  that  adults  are  more  sensitive 
to  seasonal  changes  in  precipitation  and  veg- 
etative production  than  immature  classes 
(Fig.  2).  The  apparent  difference  was  a 
change  from  reduced  activity  of  fewer  adults 


Sunset 


Time 


Sunrise 


Fig.  2.   1971  and  1973  activity  patterns  for  adult  Dipodonujs  orclii  pooled  among  20  subperiods  over  a  24-hour 
activity  period. 


September  1980         Jorgensen  et  al.:  Dipodomys  Activity  Patterns 


285 


Table  1.  Significantly  independent  (p  =  .05)  values  of 
Chi-square  tests  for  Dipodontys  ordii  activity  patterns 
(1971  and  1973). 


Variable  contrasts 

d.f. 

X2 

1971  vs.  1973  (all  classes  combined) 

19 

131.37 

1971  vs.  1973  (males) 

19 

95.66 

1971  vs.  1973  (females) 

19 

.51.41 

1971  vs.  1973  (sexuallv  active  males) 

18 

32.48 

1971  vs.  1973  (sexuallv  inactive  males) 

18 

68.29 

1971  vs.  1973  (all  adults) 

19 

117.41 

around  a  relatively  con.stant  level  in  1971  to 
a  higher  level  of  activity  for  more  animals  in 
the  predawn  hours  of  1973,  a  pattern  that 
persisted  when  all  1973  observations  were 
pooled  (Fig.  1). 

The  activity  pattern  with  all  animals  still 
in  the  population  (Fig.  1)  was  then  examined 
to  determine  the  effects  independent  vari- 
ables may  have  had  on  it.  Schmidt-Nielson 
(1964)  reported  that  Dipodomys  merriami 
was  most  active  at  low  ambient  temper- 
atures, high  relative  humidity,  and  high  abso- 
lute humidity.  In  1973,  we  found  that  activi- 
ty increased  as  temperature  decrea.sed  and 
relative  humidity  increased,  a  condition  that, 
along  with  an  increase  in  absolute  humidity, 
occurred  most  frequently  in  the  predawn 
hours.  Dipodomys  ordii  followed  the  same  ac- 
tivity pattern  reported  for  D.  merriami,  a  be- 
havior reported  to  maximize  the  conservation 
of  water  (Schmidt-Nielson  1964)  or  accom- 
modate temporal  competition  with  Dipo- 
domys microps  (Kenagy  1973),  where  their 
distributions  overlap.  This,  coupled  with  in- 
creased activity  as  wind  speeds  decrease  be- 
low 3.2  km/hr  and  after  periods  of  rain,  sug- 
gests that  changes  in  activity  may  be  related 
to  water  conservation. 

An  additional  40  animals  were  marked  and 
their  activity  monitored  for  five  days  after  11 
dominant  animals  had  been  removed  from 
the  grid.  All  except  8  (most  of  which  were 
trapped  near  the  grid  border)  of  these  40  new 
animals  were  juveniles  and  subadults.  The 
pooled  activity  pattern  shifted  to  develop  a 
peak  in  the  early  evening  hours  and  generally 
declin  ed  thereafter  (Fig.  1).  Early  evening  ac- 
tivity exposed  the  animals  to  less  than  optim- 
al conditions  for  water  conservation,  but 
seeds  were  more  abundant  on  the  soil  surface 
because  of  natural  seed-drop  and  accumula- 
tion  during   the   day.   Seeds  of  Oryzopsis 


hymenoides  were  dropped  to  the  ground  in 
rather  large  numbers  while  these  data  were 
being  gathered.  The  change  in  activity  may 
reflect  the  release  from  domination,  early  at- 
tempts to  reestablish  social  dominance,  or 
perhaps  inexperience  among  the  numerous 
immature  members  of  the  population. 

The  optimum  period  of  activity  above- 
ground  during  summer  months  was  during 
predawn  hours  when  conditions  for  water 
conservation  were  enhanced.  Activity  pat- 
terns of  the  intact  population  of  D.  ordii  were 
highest  during  this  predawn  period  of  time. 
When  the  .social  order  was  disrupted  in  1971 
due  to  low  rainfall  and  low  vegetative  pro- 
duction and  in  1973  when  the  dominant  indi- 
viduals were  removed  from  the  population, 
activity  was  highest  during  less  optimum 
conditions  for  water  conservation,  but  more 
nearly  optimal  for  seed  availabilitv. 

Terrell  Johnson  (1979)  found  that  Per- 
omyscus  maniculatus  is  more  efficient  in  har- 
vesting seeds  buried  up  to  0.25  in.  in  sand 
when  the  moisture  content  of  the  sand  is  in- 
creased. This  ob.servation  lends  credence  to 
the  harvesting  strategies  demonstrated  by  the 
activity  pattern  of  the  intact  D.  ordii  popu- 
lation, since  their  peak  activity  occurred 
when  moisture  was  highest.  Paradoxically,  O. 
hymenoides  seeds  were  most  abundant  in  the 
early  evening  after  thev  had  dropped  from 
the  plants  during  the  day.  Rodents  apparent- 
ly selected  harvesting  times  that  either  max- 
imized seed  availability  or  water  con- 
servation. From  our  data,  it  appears  that 
immature  D.  ordii  were  most  active  when 
seeds  were  most  abundant  and  adults  tended 
to  optimize  water  conservation. 

Since  seed-drop  is  ephemeral  and  occurs 
when  voung  animals  are  most  abundant  in 
the  population,  strategies  of  D.  ordii  for  sur- 
vival until  home  ranges  can  be  established 
.seem  enhanced  by  early  temporal  activity. 
Annual  replacement  of  older,  often  non- 
reproductive  adults  that  have  established  and 
control  home  ranges  within  the  population  is 
important  to  population  survival,  since  dry 
years  with  little  or  no  reproduction  are  not 
uncommon.  Replacement  of  the  breeding 
population  would  facilitate  survival  of  adults 
during  nonproductive  years,  until  reproduc- 
tion is  again  feasible.  Replacement  is  likely 


286 


Great  Basin  Naturalist 


Vol.  40,  No.  3 


only  if  the  younger  animals  obtain  a  com- 
petitive advantage  from  some  source  other 
than  size.  This  might  be  provided  by  the 
"flush"  of  energy  available  to  preadults  that 
feed  during  the  evening  hours  when  harvest 
of  high  quality  energy  is  optimized  during 
years  with  high  seed  production.  This  strate- 
gy encourages  the  infusion  of  young  animals 
into  the  population  and  allows  genetic  fix- 
ation of  the  activity  patterns  that  optimize 
the  likelihood  of  survival  for  D.  ordii  until 
the  next  breeding  opportunity. 

Literature  Cited 

BiDER,  J.  B  1968.  Animal  activity  in  uncontrolled  terres- 
trial communities  as  determined  by  a  sand  tran- 
sect technique.  Ecol.  Monogr.  38:269-308. 

BuRNHAM,  K.  P.,  .AND  W.  S.  OvERTON.  1969.  A  simulation 
studv  of  live  trapping  and  estimation  of  popu- 
lation size.  Tech.  Kept.  No.  14,  Oregon  State 
University,  Corvallis,  152  pp. 

G.\RciA,  J.  R.,  H.  D.  Smith,  and  C.  D.  Jorgensen.  1974. 
A  capture-release  method  for  determining  small 
mammal  activity.  Proc.  Utah  .\cad.  Sci.,  Arts  and 
Letters  51:1-11. 


Harling,  J.  1971.  A  technique  for  precisely  timing  cap- 
tures of  Perornysctts  manicuhitus.  Canadian  J. 
Zool.  48:1275-1277. 

Johnson,  T.  K.  1979.  Ability  of  desert  rodents  to  find 
buried  seeds  in  desert  range  communities.  Un- 
published thesis.  Brigham  Young  University,  Pro- 
vo,  Utah.  23  pp. 

Jorgensen,  C.  D.,  and  C.  L.  Hayward.  1965.  Mammals 
of  the  Nevada  Test  Site.  Brigham  Young  Univer- 
sity Sci.  Bull.,  Biol.  Ser.  6(3):  1-81. 

Jorgensen,  C.  D.,  D.  T.  Scott,  and  H.  D.  Smith.  1972. 
Small  mammal  trapping  simulator.  Proc.  1972 
Summer  Computer  Simulation  Conf.:  1154-1168. 

Ken  AG  Y,  G.  J.  1973.  Daily  and  seasonal  patterns  of  activ- 
ity and  energetics  in  a  heteromvid  rodent  com- 
munity. Ecology  54:1201-1219. 

Manly,  B.  F.  J.  1970.  A  simulation  study  of  animal  pop- 
ulation estimation,  using  the  capture-recapture 
method.  Jour.  Appl.  Ecol.  71:13-39. 

Marten,  G.  G.  1973.  Time  patterns  of  Perornysctts  activ- 
ity and  their  correlation  with  weather.  J.  Mam- 
mal. 54:169-188. 

Platt,  R.  B.,  and  J.  R.  Griffiths.  1964.  Environmental 
measurement  and  interpretation.  Reinhold  Publ. 
Co.,  New  York.  235  pp. 

ScHMiDT-NiELSON,  K.  1964.  Descrt  animals:  phys- 
iological problems  of  heat  and  water.  Clarendon 
Press,  Oxford,  England.  277  pp. 

Scott,  D.  T.,  C.  D.  Jorgensen,  and  H.  D.  Smith.  1978. 
Comparison  of  live  and  removal  methods  to  esti- 
mate small  mammal  densities.  Acta  Theriol. 
23:173-193. 


NEW  RECORDS  OF  WESTERN  TRICHOPTERA  WITH  NOTES  ON  THEIR  BIOLOGY' 

Bernard  G.  Swegniau"  and  Leonard  C.  Ferrington,  Jr.- 

Abstract.-  Western  records  for  27  species  of  Trichoptera  are  given;  a  majority  of  the  records  are  from  the  Bear- 
tooth  Mountains  of  northwestern  Wyoming.  In  addition,  examples  of  variation  in  the  male  genitiiia  of  Limnephilus 
coloradensis  (Banks)  are  figured  and  the  female  is  described.  Some  comments  regarding  the  larvae  of  Allomyia 
(Inuinia)  are  pre,sented. 


Twenty-two  species  of  Trichoptera  are  re- 
ported from  Wyoming  and  Montana,  includ- 
ing 20  from  the  immediate  vicinity  of  the 
Beartooth  Mountains,  where  the  University 
of  Pittsburgh's  Pymatuning  Laboratory  of 
Ecology  has  offered  a  summer  field  course  in 
Alpine  Ecology.  In  addition,  7  species  were 
collected  near  a  pond  at  the  Winding  River 
Campground  near  Rocky  Mountain  National 
Park,  Grand  County,  Colorado.  Within  the 
Beartooth  Mountains,  immatures  of  many 
common  limnephilid  genera  were  collected, 
but  the  species  remain  unknown  to  us.  Some 
of  the  more  common  genera  taken  include: 
Discoinoecus,  Hesperophylax,  Homophijlax, 
Lenarchus,  Limnephilus,  and  Psychoghjpho. 

Considerable  variability  was  observed  in 
the  male  genitalia  of  Limnephilus  colorad- 
ensis (Banks).  Examples  of  this  variation  are 
illustrated.  Further,  in  copula  specimens  of 
this  species  were  collected,  thereby  making 
possible  the  identification  of  the  female  that 
is  described  and  figured. 

Collecting  Sites 

The  following  sites,  with  the  exception  of 
the  Grow  Ventre  and  Winding  River  loca- 
tions, are  within  the  Beartooth  Mountains  of 
the  Absaroka  range  of  northwestern  (Park 
Co.)  Wyoming.  The  Star  Lake  site  lies  in 
Park  Co.,  Montana. 

Inlet  Run.—  Elevation  approximated  3140 
m,  109°  29'  W,  44°  58'  N.  The  sample  site  is 
on  the  more  easterly  of  two  first  order,  snow 
melt  streams  that  flow  into  Frozen  Lake.  A 
majority  of  the  specimens  were  collected 
during  a  series  of  diel  drift  studies;  however, 

'Support  for  this  paper  has  been  provided  by  the  Pymatuning  Laboratory  of  Ecology. 
■University  of  Pittsburgh  and  Pymatuning  Laboratory  of  Ecology.  Pittsburgh.  Pennsylvania  15260. 


occasional  specimens  were  collected  by 
sweeping  low  vegetation  or  were  picked 
from  rocks. 

Frozen  Lake.—  Elevation  3070  m;  same  lo- 
cation as  above.  Specimens  were  collected  by 
sweeping  vegetation  or  were  collected  from 
rocks  in  1  m  or  less  of  water. 

Chain  Lakes.-  Elevation  2880  m,  109°  31' 
W,  44°  55'  N.  The  sample  .site  was  at  the 
southeast  shore  of  lower  Chain  Lake  near  the 
point  where  the  stream  draining  Fantan  Lake 
enters.  An  extensive  alpine  meadow  sur- 
rounds the  lake. 

Sawtooth  Lake.-  Elevation  2835  m,  109° 
28'  W,  44°  54'  N.  The  sampling  site  was  on 
an  unnamed  second  order  stream  that  drains 
the  western  slope  of  Sawtooth  Mountain. 
This  stream  drains  open  alpine  meadow  and 
flows  into  the  ea.stern  edge  of  Sawtooth  Lake. 
The  specimens  were  collected  from  exposed 
and  submerged  rocks. 

Beartooth  Butte.—  Elevation  approximatelv 
2910  m,  109°  37'  W,  45°  57'  \.  The  sample 
site  is  a  large  spring  that  originates  on  the 
south  face  of  Beartooth  Butte  at  the  base  of 
an  open  talus  slope.  The  stream  flows 
through  open  meadow  and  enters  Beartooth 
Lake  on  its  western  shore  at  approximately 
2710  m  elevation. 

Moose  Bog.-  Elevation  2740  m,  109°  37' 
W,  44°  56'  N.  This  bog  is  approximately  30 
m  south  of  U.S.  Route  212,  just  ea.st  of  the 
point  where  the  gravel  road  to  Clay  Butte 
lookout  station  begins.  The  specimens  were 
collected  by  sweeping  the  bog  vegetation  on 
the  Sphagnum  mat  near  the  largest  area  of 
open  water. 


287 


288 


Great  Basin  Naturalist 


Vol.  40,  No.  3 


Ghost  Creek.-  Elevation  2650  m,  109°  37' 
W,  44°  56'  N.  The  sample  site  was  near  the 
point  where  the  gravel  road  to  the  University 
of  Pittsburgh's  research  trailer  crosses  the 
stream.  The  stream  is  second  order  and  drains 
intermittent  patches  of  alpine  meadows  and 
coniferous  forest.  Most  specimens  were  col- 
lected during  diel  drift  studies;  however, 
some  specimens  were  taken  by  sweeping  veg- 
etation. 

Clark's  Fork  of  Yellowstone  River  —  Eleva- 
tion 2060  m,  209°  41'  W,  44°  51'  N.  The  riv- 
er at  this  point  is  fifth  order  and  flows  south- 
east, draining  mixed  coniferous  forest  and 
pasture  lands. 

Star  Lafce.- Elevation  2940  m,  109°  55' 
W,  45°  6'  N.  The  sample  site  was  located  in 
the  stream  just  east  of  the  outlet  from  Star 
Lake.  The  stream  width  is  approximately  2  m 
at  this  point.  This  site  is  located  in  Montana. 

Gros  Ventre.- Elevation  1980  m,  110°  40' 
W,  43°  36'  N.  The  sample  site  was  located  in 
Gros  Ventre  Campground  approximately  15 
miles  northeast  of  Jackson  Hole,  Wyoming. 
The  specimens  were  picked  from  the  win- 
dows of  the  campground  restrooms. 

Winding  River.-  Elevation  2590  m,  105° 
53'W,  40°  16'  N.  This  sample  site  was  lo- 
cated just  west  of  the  entrance  to  Rocky 
Mountain  National  Park,  Colorado,  in  the 
Winding  River  Campground.  Specimens 
were  collected  by  sweeping  the  emergent 
vegetation  of  a  small  pond  and  were  picked 
from  the  windows  of  the  campground  rest- 
rooms. 

Species  Collected 

Rhyacophilidae 

Since  less  than  10  percent  of  the  western 
Rhyacophilia  immatures  are  known  (Flint,  in 
litt.),  the  following  identifications  are  only 
tentative.  All  identifications  are  based  on  our 
use  of  Smith  (1968). 

Rhyacophila  acropedes  Banks.—  Wyoming 
(Park  Co.):  Ghost  Creek,  14  August  1974,  10 
larvae,  and  15  August  1974,  7  larvae,  collect- 
ed L.  Ferrington.  15  August  1974,  1  larva, 
collected  G.  Goetz. 

Rhyacophila  hyalinata  Banks.—  Wyoming 
(Park"  Co.):  Ghost  Creek,  19  August  1977,  3 
larvae,  collected  D.  Ferrington.  The  ventral 
surface   of  the   head   in   these   specimens   is 


darkened,  but  on  the  basis  of  distribution  are 
assigned  to  hyalinata  rather  than  vocala 
(Smith  1968). 

Rhyacophila  tucula  Ross.—  Wyoming 
(Park  Co.):  Ghost  Creek,  14  August  1974,  1 
larva,  26  August  1978,  11  larvae,  19  August 

1977,  3  prepupae.  Sawtooth  Lake  vicinity,  23 
July  1975,  1  larva;  Inlet  Run,  21  July  1978,  1 
larva.  Montana  (Park  Co.):  Outlet  of  Star 
Lake,  25  July  1975,  4  larvae.  All  by  D.  and  L. 
Ferrington. 

Rhyacophila  vacciia  Milne.—  Wyoming 
(Park  Co.):  Sawtooth  Lake  vicinity,  23  July 
1975,  2  larvae,  D.  Ferrington. 

Rhyacophila  vagrita  Milne.—  Wyoming 
(Park  Co.):  Ghost  Creek,  14  August  1974,  1 
larva,  L.  Ferrington. 

Rhyacophila  verulla  Milne.—  Wyoming 
(Park  Co.):  Ghost  Creek,  26  August  1978,  2 
larvae,  19  August  1977,  1  larva;  Sawtooth 
Lake  vicinity,  23  July  1975,  4  larvae;  Inlet 
Run,  21  July  1978,  1  larva.  Montana  (Park 
Co.):  Outlet  of  Star  Lake,  25  July  1975,  1  lar- 
va. All  by  D.  and  L.  Ferrington. 

Glossosomatidae 

Glossoma  vebna  Ross.—  Colorado  (Grand 
Co.):  Winding  River  Campground,  2  August 

1978,  1  male,  collected  D.  Ferrington. 

Hydropsychidae 

Arctopsyche  grandis  Banks.—  Wyoming 
(Park  Co.):  Clark's  Fork  Yellowstone  River, 
23  July  1978,  22  larvae,  6  pupae  (1  pharate 
male),  6  males,  D.  Ferrington. 

Hydropsyche  oslari  Banks.—  Wyoming  (Te- 
ton Co.):  Gros  Ventre  Campground,  29  July 
1978,  4  males,  26  females,  D.  Ferrington. 

Hydroptilidae 

Stactobiella  delira  Ross.—  Colorado  (Grand 
Co.):  Winding  River  Campground,  2  August 
1978,  2  males,  1  female,  D.  Ferrington. 

Limnephlidae 

Anabolia  bimaculata  Walker.—  Colorado 
(Grand  Co.):  Winding  River  Campground,  2 
August  1978,  1  male,  D.  Ferrington. 


September  1980 


SwEGMAN,  Ferbington:  Western  Trichoptera 


289 


Chyranda  centralis  Banks.—  Wyoming 
(Park  Co.):  Ghost  Creek,  27  July  1978,  1  fe- 
male, collected  L.  Ferrington,  identified  A. 
Nimmo. 

Dicosmoecus  gilvipes  Hagen.—  Wyoming 
(Park  Co.):  Spring  at  base  of  Beartooth  Butte, 
15  August  1974,  1  female,  G.  Goetz.  Uniden- 
tified larvae  of  Dicosmoecus  have  also  been 
collected  in  large  numbers  from  Inlet  Run. 

Ecclisomijia  conspersa  Banks.—  Wyoming 
(Park  Co.):  Inlet  Run,  22  July  1978,  1  male, 
L.  Ferrington. 

Ecclisomijia  maculosa  Banks.—  Wyoming 
(Park  Co.):  Spring  at  base  of  Beartooth  Butte, 
15  August  1974,  4  pupae  (2  pharate  males,  2 
females),  G.  Goetz;  Ghost  Creek,  26  July 
1978,  3  males,  27  July  1978,  3  males,  4  fe- 
males, L.  Ferrington.  Frozen  Lake  vicinitv, 
22  July  1978,  43  larvae,  D.  Ferrington.  The 
larval  records  are  a  tentative  assignment 
based  on  proximity  of  sites  where  adults  were 
taken.  However,  the  single  record  of  E.  con- 
spersa indicates  two  species  occur  in  this 
area. 

Allomyia  bifosa  (Ross.)—  Wyoming  (Park 
Co.):  Inlet  Run,  21-22  July  1978,  13  males,  2 
females,  69  pupae  (51  pharate  males,  18  pha- 
rate females).  In  addition  to  these  adult-pupal 
records,  larval  records  of  Allomyia  include; 
Ghost  Creek,  14  August  1974,  1  larva,  G. 
Goetz,  14  August  1974,  8  larvae,  and  15 


August  1974,  1  larva,  L.  Ferrington,  18  Au- 
gust 1977,  2  larvae,  R.  Seward,  19  August 
1977,  5  larvae,  and  26  August  1978,  1  larva, 
L.  Ferrington;  Inlet  Run,  10-11  August  1977, 
89  larvae,  17-18  August  1977,  31  larvae,  21 
July  1978,  4  larvae,  all  by  L.  Ferrington.  Two 
types  of  larvae  were  present  in  these  collec- 
tions. One  type  is  without  the  flattened  head 
and  pronounced  carina,  as  in  Allomyia 
(Imania)  scotti  Wiggins  (Wiggins  1977,  Fig. 
10.28).  The  second  type  has  this  character 
and  also  possesses  single  gills  dorsallv  and 
ventrally  on  segments  2,  3,  4,  and  5.  Wiggins 
(pers.  comm.)  suggests  that  two  species  are 
involved.  He  has  associated  gilled  larvae  with 
Allomyia  tripunctata  Banks,  which  is  known 
from  Wyoming.  Adults  of  A.  tripunctata, 
however,  have  yet  to  be  collected  in  the 
Beartooth  Mountains.  These  gilled  larvae 
were  compared  to  gilled  larvae  of  Allomyia 
collected  17  July  1975  at  St.  Vrain  Creek 
above  Peaceful  Valley,  Boulder  Countv,  Col- 
orado by  Dr.  J.  V.  Ward.  The  specimens  from 
the  two  localities  appear  to  be  distinct,  sug- 
gesting that  at  least  two  species  of  Allomyia 
in  this  area  possess  gills. 

Limnephilus  coloradensis  (Banks).— Wyom- 
ing (Park  Co.):  Ghost  Creek,  19  August  1977, 
1  male,  1  female,  and  1  pair  in  copula,  D. 
Ferrington,  Moose  Bog,  9  August  1979,  10 
males,       1       female,       L.       Brooks,       26 


Fig,  1.  Variation  in  the  parameres  (Lateral  appendages  sensu  Ross  1938)  of  Limnephilus  coloradensis  (Banks).  A 
typical  symmetrical  stnicture,  B-E.  Variation  in  parameres  showing  auxiliary  spines. 


290 


Great  Basin  Naturalist 


Vol.  40,  No.  3 


males,  39  females,  C.  Sirianni.  Considerable 
variation  in  the  male  genitalia  occm-s  in  this 
species  (Figs.  lA-E).  The  simplest  type  and 
probably  the  most  common  form  (Fig.  lA)  is 
somewhat  different  from  the  lectotype  fig- 
ured by  Ross  (Ross  1938,  Fig.  76).  Many  spec- 
imens also  possess  auxiliary  spines  (Fig. 
IB-E)  not  unlike  Limnephilus  kennicotti 
Banks.  However,  all  types  will  key  easily  to 
L.  coloradensis  in  Ross  and  Merkley  (1952). 
In  copula  specimens  have  made  possible  the 
correct  association  of  the  female.  The  origi- 
nal description  of  the  male  is  given  by  Banks 
(1899). 

Description  of  female:  Forewing  length 
7-8  mm.  Hindwing  6-7  mm.  Hindwings 
clear.  Forewings  with  brown  irregular  patch- 
es at  stigma,  distally  between  R4  and  Mg  and 
in  costal  area  between  Mg  and  Cui^.  Smaller 
patches  in  discoidal  and  thyridial  areas,  and 
along  A,.  Female  genitalia  as  in  Figures 
2A-B.  Tergite  9  small.  Sternum  9  ventrally 
divided  into  two  distinct  somewhat  circular 
lobes,  roundly  triangular  when  viewed  later- 
ally. Segment  10  reduced  to  a  ventral  flap. 
Appendages  of  segment  10  greatly  enlarged, 
produced  to  points.  Supragenital  plate  rec- 
tangular. Median  lobe  of  vulval  scale 
rounded  apically,  lateral  lobes  quadrate,  pro- 
duced laterad. 

In  general,  the  female  bears  some  resem- 
blance to  Limnephilus  kennicotti  Banks.  Both 


species  were  placed  in  the  fenestratus  group 
by  Schmid  (1955).  In  L.  kennicotti  the  dorsal 
body  of  segment  9  is  indistinguishable  from 
segment  10  (see  Nimmo  1971,  Figs.  419,  420). 
In  L.  coloradensis  the  dorsal  lobe  of  segment 
9  is  distinct.  In  both  species  the  appendages 
of  segment  10  are  greatly  enlarged  and  pro- 
duced to  points  distally.  These  appendages 
show  considerable  variation  in  L.  colorad- 
ensis. In  both  species  the  supragenital  plate  is 
rectangular.  The  lateral  lobes  of  the  vulval 
scale  is  more  quadrate  in  L.  coloradensis. 

Limnephilus  hageni  Banks.—  Colorado 
(Grand  Co.):  Winding  River  Campground,  2 
August  1978,  2  males,  D.  Ferrington.  Wyom- 
ing (Park  Co.):  Moose  Bog,  18  August  1977,  4 
females,  collected  L.  Ferrington,  females 
identified  A.  Nimmo. 

Limnephilus  indivisus  Walker.—  Wyoming 
(Park  Co.):  Moose  Bog,  9  August  1979,  1  fe- 
male, C.  Sirianni. 

Limnephilus  janus  Ross.—  Colorado  (Grand 
Co.):  Winding  River  Campground,  2  August 
1978,  10  males,  8  females,  D.  and  L.  Ferring- 
ton. 

Limnephilus  picturatus  McLachlan.— 
Wyoming  (Park  Co.):  Moose  Bog,  6  August 
1977,  25  males,  22  females,  18  August  1977, 
4  females,  1  female,  L.  and  D.  Ferrington,  9 
August  1979,  33  males,  24  females,  L.  Brooks, 
53  males,  66  females,  C.  Sirianni.  Chain 
Lakes,  12  August  1977,  2  males,  6  females,  D. 


app 


Fig.  2.  Female  genitalia:  A,  ventral  view;  B,  lateral  view,  segments  8  to  10.  app.  =  appendages  of  segment  10. 


September  1980 


SwEGMAN,  Ferrington:  Western  Trichoptera 


291 


Ferrington,  6  Augvist  1979,  57  males,  24  fe- 
males, L.  Brooks. 

Limnephilus  secludens  Banks.—  Colorado 
(Grand  Co.):  Winding  River  Campground,  2 
August  1978,  1  female,  collected  D.  Ferring- 
ton, identified  A.  Nimmo. 

Neothremma  alicia  Banks.—  Wyoming 
(Park  Co.):  Ghost  Creek,  15  August  1974,  33 
larvae,  6  pupae  (1  pharate  male,  5  pharate 
females),  L.  Ferrington;  Sawtooth  Lake,  23 
July  1978,  4  larvae,  1  pharate  male  pupa,  D. 
Ferrington. 

Oligophlebodes  zelti  Nimmo.—  Wyoming 
(Park  Co.):  Ghost  Creek,  27  July  1978,  3 
males,  3  females,  collected  L.  Ferrington. 
Some  variation  exists  between  these  and  Al- 
berta specimens  (Nimmo,  pers.  Comm.).  This 
species  was  previously  known  only  from  Al- 
berta (Nimmo  1971). 

Psychoglypha  aff.  subborealis  Banks.— 
Wyoming  (Park  Co.):  Beartooth  Lake,  17 
August  1977,  5  larvae,  R.  Seward.  These  lar- 
vae possess  banded  legs,  as  in  P.  subborealis 
(Wiggins  1977). 

Lepidostomatidae 

Lepidostoma  phwiale  Milne.—  Wyoming 
(Teton  Co.):  Gross  Ventre  Campground,  29 
July  1978,  8  males,  30  females,  D.  and  L. 
Ferrington. 

Acknowledgments 

We  thank  Deborah  Ferrington  and  Lisa 
Brooks  for  help  in  the  field.  Dr.  Glenn  Wig- 
gins for  his  helpful  comments  regarding  the 


AUomyia  larvae,  and  Dr.  J.  V.  Ward  for  sup- 
plying Alhmyia  larvae  from  Colorado.  Spe- 
cial thanks  are  also  extended  to  Dr.  A.  Nim- 
mo for  his  identifications  and  helpful 
comments.  Dr.  Oliver  S.  Flint,  Jr.  for  obtain- 
ing permission  for  us  to  examine  material  at 
the  U.S.  National  Museum  of  Natural  History 
and  for  checking  the  identifications  of  Hydro- 
psyche  oslari  and  Lepidostoma  pluviale,  and 
to  Dr.  F.  Schmid  for  his  examination  of  L. 
coloradensis.  Drs.  Flint  and  Schmid  also  pro- 
vided critical  comments  of  this  manuscript. 
Dr.  Richard  T.  Hartman  provided  consid- 
erable material  support  through  the  Pyma- 
tuning  Laboratory  of  Ecology. 

Literature  Cited 

Banks,  N.  1899.  Descriptions  of  new  North  .American 
neuropteroid  insecis.  Trans.  .\m.  Ent.  See. 
25:199-218. 

Nimmo,  A.  P.  1971.  The  adult  Rhyacophihdae  and  Lim- 
nephilidae  (Trichoptera)  of  .\lberta  and  eastern 
British  Columbia  and  their  post-glacial  origin. 
Quest.  Entomologicae.  7:234  pp. 

Ross,  H.  H.  1938.  Lectotypes  of  North  American  caddis 
flies  in  the  Museum  of  Comparative  Zoology. 
Psyche  45:1-61. 

Ross,  H.  H.,  AND  D.  R.  Merkley.  1952.  An  annotated 
key  to  the  nearctic  males  of  Limnephilus  (Tri- 
choptera:Limnephilidae).  .Am.  Midi.  .Nat. 
47:435-455. 

Schmid,  F.  1955.  Contributions  a  i'etude  des  Limnophi- 
lidae  (River  drainage  of  Idaho  with  special  refer- 
ence to  the  larvae.  .Ann.  Ent.  Soc.  .Am. 
61:655-674. 

Wiggins,  G.  1977.  Larvae  of  the  North  .American  cad- 
disfly  genera  (Trichoptera).  University  of  Toronto 
Press,  Toronto. 


OBSERVATIONS  OX  SEASONAL  VARIATION 
LN  DESERT  ARTHROPODS  IN  CENTR-\L  NEVADA 

Robert  D.  Pietruszka- 

AByrRACT-  "       "      -i  Malaise  trap  collections  from  terrestrial  arthropod  populations  in  central  Nevada  were 
aQal^•zed  for  .;  periods  during  the  I97S  growing  season.  Mites   Acarina   and  ants   Formicidae   were  the 

taxa  representee  p%  tne  largest  numbers  of  indi%iduals  in  pitfall  trap  collections  throughout  the  season:  Malaise  col- 
lections were  composed  mainlv  of  aerial  taxa  largeh  Diptera  and  H%Tnenoptera  .  Peak  arthropod  abundance  was 
recorded  during  mid-June.  Collection  diversities  for  both  trapping  methods  were  generally  low  due  to  the  abun- 
dance of  a  few  ta-xa.  E\idence  for  spatial  heterogeneity  in  arthropod  populations  was  meager:  temporal  hetero- 
geneitv  in  these  populations,  however,  was  more  apparent. 


Within  shrub  desert  communities  in- 
vertebrate animals  constitute  a  major  part  of 
the  biotic  matrix  \Fautin  19-t6\  forming  an 
important  food  resource  for  many  consimier 
species.  Spatial  and  temporal  variation  in 
such  a  resource  base  can  affect  the  foraging 
patterns  of  individual  consumers  iGill  and 
Wolf  1977.  Mac-\rthiu-  and  Pianka  1966  >  as 
well  as  their  intra-  and  interspecific  ecologi- 
cal relationships  .  Wiens  1976'. 

Commonly,  investigations  of  arthropod 
populations  are  limited  to  faunistic  invento- 
ries e.g..  Allred  et  al.  1965.  Beck  and  Allred 
1968'.  or  to  broad  scale  comparisons  e.s;..  Al- 
hed  1973.  .\lh-ed  and  Gertsch  1976,  Gertsch 
and  .\llred  1965 1.  The  present  report  repre- 
sents a  preliminar\"  analysis  of  one  part  of  an 
ongoing  investigation  of  lizard  ecology"  in  the 
Great  Basin  Desert.  Here  I  will  deal  with  the 
arthropod  food  base  at  a  single  location,  fo- 
cusing upon  the  relative  success  of  two  sam- 
pling schemes,  seasonal  changes  in  arthropod 
abundance  and  diversity",  and  the  apparent 
degree  of  local  spatial  and  temporal  hetero- 
geneity' in  arthropod  populations. 

Study  Site  and  Methods 

The  research  site  is  in  Fairsiew  \'allev,  Ne- 
vada, a  relatively  flat  basin  ranging  in  eleva- 
tion from  1370  to  1500  m.  Vegetation  within 
the  valley  is  dominated  by  Atriplex  confer- 
tifolia,  Sarcobatus  baileyi,  and  Oryzopsis 
hymenoides  and  generally  t\pifies  the  shad- 
sc-ale  zone  common  to  much  of  interior  Ne- 
vada (Billings  1949 1.  Average  monthlv  tem- 


peratures  for  the  site  varv"  from  about  0  C  to 
just  over  23  C;  average  monthly  precipitation 
varies  from  about  .5  cm  to  just  over  1.5  cm: 
mean  growing  season  is  142  days. 

Data  were  analyzed  from  an  insect  sam- 
pling plot  estabhshed  appro.ximately  9.6  km 
N  of  Frenchman,  Churchill  County,  where 
49  pitfall  traps  were  arrayed  along  cardinal 
compass  directions  at  intervals  of  5  m  Fig. 
1l  Traps  were  randomly  located  with  respect 
to  the  vegetation;  the  total  linear  distance  of 
each  line  was  120  m.  Each  pitfall  trap  mea- 
sures 98  mm  in  diameter  by  144  mm  in 
depth:  a  funnel  insert  prevents  escape  of  ar- 
thropods once  captured.  A  Malaise  trap 
Townes  1972)  was  placed  at  appro.ximately 
the  center  of  the  two  lines  iFig.  L. 

Malaise  and  pitfall  traps  were  opened  for  a 
period  of  48  hours,  followed  by  a  closed  peri- 
od of  generally  equal  duration.  This  schedule 
was  maintained  from  13  May  to  24  August 
1978  and  yielded  a  new  sample  at  roughly 
four-day  intervals.  Each  trap  contained  a 
standardized  amount  of  5  percent  formalde- 
hyde solution.  The  captures  of  each  "arm"  of 
the  pitfall  traplines  for  each  trapping  period 
were  combined.  Thus,  each  of  the  N-,  S-,  and 
W-arm  samples  contained  the  contents  of  12 
traps,  and  the  E-arm  sample  contained  those 
of  13  traps.  Contents  of  NIalaise  trap  samples 
were  maintained  separately. 
To  examine  the  major  seasonal  patterns  in 
the  prospective  arthropod  food  base,  samples 
were  analvzed  for  four  trapping  periods  des- 
ignated  13   May.   12  June.   14  July,  and   12 


292 


September  19S0  Pietruszka:  Season.\i,  V.\riation  of  Arthropods 


293 


Fig.  1.  Pitfall  and  Malaise  trap  placements.  Dots  ref>- 
resent  pitfall  traps;  M  represents  Malaise  trap. 

Au2;iist.  These  spanned  the  major  part  of  the 
surface-active  season  for  lizards  in  1978. 
Arachnids  caught  were  identified  to  the  or- 
dinal level:  insects  caught  were  identified  to 
family  or  superfamily  where  practicable,  us- 
ing Borror.  DeLong,  and  Triplehom  J976\ 
Borror  and  \Miite  (1970),  and  Chu  (1949*. 
Samples  from  each  arm  of  the  pitfall  trap- 
ping grid  were  analyzed  separately,  as  were 
Malaise  samples.  Total  counts  of  identified 
groups  were  then  determined  for  each 
sample.  Arthropod  diversity  within  a  sample 
was  assessed  using  B  =  1  — pf.  where  p, 
equals  the  proportion  of  individuals  in  cate- 
gory i  ^MacArthur  1972).  The  relative  de- 
grees of  spatial  and  temporal  heterogeneity 
were  assessed  using  a  similarity  inde.x,  S, 
where 

S  =  l-4(Z|p^-p,.,|)     , 

and  p^^,  and  p,,  are  the  proportions  of  sam- 
ples X  and  y  in  category  i  ^Schoener  1970). 

Results 

Trapping  success.—  For  the  four  trapping 
periods  analyzed,  the  two  methods  employed 
amassed  a  total  catch  of  7176  arthropods. 
The  vast  majoritv  of  these.  6117.  were  col- 
lected along  pitfall  traplines:  1059  arthro- 
pods were  collected  bv  Malaise  trapping.  On 


a  per-trapping  period  basis.  Malaise  trapping 
yielded  an  average  of  265  captures.  This  is  a 
substantially  lower  capture  rate  (up  to  50 
percent  lower  than  when  these  traps  are 
used  in  forested  habitats  Matthews  and  Mat- 
thews 1971).  Pitfall  traps  also  \ielded  a  high- 
er number  of  captiu-es  per  trapping  period, 
averaging  382  per  trapline  arm,  or  just  over 
1500  captures  per  trapping  grid.  However, 
capture  rates  for  the  two  methods  are  not  di- 
rectly comparable  due  to  the  greater  "at 
risk"  area  for  pitfall  traps. 

The  composition  of  collections  obtained  bv 
the  t^vo  methods  also  differed  substantially. 
Pitfall  collections  were  dominated  bv  mites 
(Acarina>,  which  comprised  almost  half  the 
total  collection.  These  were  followed,  in  or- 
der of  numerical  importance,  by  H\Tnenoi>- 
tera  (the  vast  majority-  of  which  were  ants), 
Coleoptera.  and  Diptera.  In  all,  largely  or 
completely  terrestrial  forms  comprised  ap- 
proximately 88  percent  of  the  arthropods  col- 
lected. Malaise  trap  collections,  by  c-ontrast 
were  dominated  by  Diptera.  which  formed 
over  70  percent  of  the  total  collection. 
H\"menoptera,  Homoptera.  Lepidoptera.  and 
Coleoptera  combined  to  form  just  over  one- 
quarter  of  the  total  collection  Table  1). 
These  data  are  consistent  with  those  from 
Malaise  traps  used  in  forested  areas  in  which 
Diptera,  H\"menoptera.  Hemiptera  including 
Homoptera'.  and  Lepidoptera  constitute  at 
least  90  percent  of  each  collection  (Matthews 
and  Matthews  1971). 

Seusorwl  clianges  in  abundance  and  diver- 
sity.— Arthropod  abundance  appeared  to 
peak  during  mid-June,  approximately  1.5 
months  after  the  last  of  the  spring  rains. 
Numbers  of  arthropods  declined  rapidly 
thereafter  to  moderate  levels.  This  seasonal 
trend  is  closely  reflected  in  collections  from 
pitfall  traps  but  not  from  Malaise  trapping 
(Fio.  2.\\  The  low  number  o*  captures  on  12 
June  mav  reflect  an  actual  decrease  in  aerial 
insects,  but  it  is  more  Ukely  that  this  is  a  re- 
flection of  local  changes  in  wind  conditions, 
to  which  this  technique  is  highly  susceptible 
^Matthews  and  Matthews  1971).  This  inter- 
pretation is  strengthened  by  the  observation 
that  aerial  insects  occurred  in  approximately 
equal  numbers  in  the  12  June  and  14  July 
Malaise  samples. 


294 


Great  Basin  Naturalist 


Vol.  40,  No.  3 


Table  1.  Siiminary  of  arthropods  collected  in  pitfall  and  Malaise  trap  samples. 


Pitfall  trap 


Malaise  trap 


Acarina 

Araneida 

Scorpion  ida 

Solpiigida 

Coleoptera 

Carabidae 


Collembola 
Diptera 


Hemiptera 


Homoptera 


Taxa 


Number  of 
specimens 


Relative 
abundance 


Number  of 
specimens 


Relative 
abundance 


3051 


0.4988 


Anthribidae 
Buprestidae 

Curculionidae 

Dascillidae 

Histeridae 

Leiodidae 

Melyridae 

Nitidulidae 

Pedilidae 

Staphylinidae 

Tenebrionidae 

Sminthuridae 
Poduridae 

Anthomyiidae 

Bibionidae 

Bombyliidae 

Calliphoridae 

Cecidomyiidae 

Chironomidae 

Conopidae 

Dolichopodidae 

Empididae 

Lauxaniidae 

Muscidae 

Mycetophilidae 

Pipunculidae 

Psychodidae 

Ptychopteridae 

Sarcophagidae 

Sciaridae 

Simuliidae 

Syrphidae 

Tachinidae 

Therevidae 

Tipulidae 

Xylophagidae 

Acalypterate  muscoids 

Diptera  larvae 

Lygaeidae 

Miridae 

Nabidae 

Pentatomidae 

Tingidae 

Aphididae 
Cercopidae 
Cicadellidae 
Coccoidea 
Nymph /larvae 


104 

0.0170 

16 

0.0026 

2 

0.0003 

30 

0.0049 

17 

0.0028 

5 

0.0008 

2 

0.0003 

5 

0.0008 

2 

0.0003 

244 

0.0399 

1 

0.0002 

1 

0.0002 

3 

0.0005 

243 

0.0397 

38 

0.0062 

7 

0.0011 

24 

0.0039 

1 

0.0002 

3 

0.0005 

1 

0.0002 

37 

0.0060 

18 

0.0029 

1 

0.0002 

8 

0.0013 

4 

0.0007 

0.0002 


1 

0.0002 

3 

0.0005 

45 

0.0074 

1 

0.0002 

18 

0.0029 

1 

0.0002 

42 

0.0069 

2 

0.0003 

84 

0.0137 

18 

0.0029 

2 

0.0003 

2 

0.0003 

1 

0.0002 

10 

0.0016 

5 

0.0008 

102 

0.0167 

2 

0.0003 

1 

0.0002 

4 

29 


3 

5 

197 
5 
1 


4 

11 

3 

9 


41 

203 

21 

1 

1 

3 

248 


1 
60 
11 


0.0009 
0.0057 


0.0019 
0.0009 

0.0019 
0.0009 


0.0038 
0.0274 


0.0028 

0.0047 

0.1860 
0.0047 
0.0009 


0.0038 
0.0104 
0.0028 
0.0085 

0.0009 


0.0387 
0.1917 
0.0198 
0.0009 
0.0009 
0.0028 
0.2342 


0.0028 
0.0038 


0.0009 
0.0567 
0.0104 


September  1980 

Pietruszka:  Seasonal  Variat 

ION  OF 

Arthropods 

295 

Table  1  continued. 

Pitfall  trap 

Malaise 

trap 

Hymenoptera 

Andrenidae 

6 

0.0010 

Apidae 

21 

0.00.34 

1 

0.0009 

Braconidae 

13 

0.0123 

Chalcidoidea 

51 

0.0083 

87 

0.0822 

Chrysididae 

6 

0.0010 

Dryinidae 

2 

0.0003 

Fonnicidae 

1577 

0.2578 

13 

0.012.3 

Halictidae 

66 

0.0108 

5 

0.0047 

Ichneumon  idae 

1 

0.0002 

1 

0.0009 

Mutillidae 

5 

0.0008 

Pom  pil  idae 

4 

0.0007 

2 

0.(K)19 

Sphecidae 

46 

0.0075 

3 

0.0028 

Lepidoptera 

Cosmopterygidae 

19 

0.0031 

38 

0.0359 

Lycaenidae 

1 

0.0002 

Pyralidae 

5 

0.0008 

4 

0.00,38 

Unidentified  larvae 

12 

0.0020 

Neuroptera 

Chrysopidae 

1 

0.0009 

Coniopterygidae 

3 

0.0005 

1 

0.0009 

Henierobiidae 

18 

0.0029 

1 

0.0009 

Myrmeleontidae 

17 

0.0028 

Unidentified  larvae 

3 

0.0005 

Orthoptera 

Acrididae 

2 

0.0003 

Blattidae 

2 

0.0003 

Gryllacrididae 

18 

0.0029 

1 

0.0009 

Mantidae 

1 

0.(XK)2 

Thysanoptera 

Heterothripidae 

4 

0.(XK)7 

6 

0.0057 

Phlaeothripidae 

7 

0.0011 

Thripidae 

9 

0.0015 

Unidentified  larva 

(campodeiform) 

1 

0.(K)02 

Isopoda 

1 

0.0002 

Among  the  major  arthropod  groups  occur- 
ring in  seasonal  samples,  mites  most  closely 
follow  the  general  trend.  It  is  quite  likely,  in 
fact,  that  mite  populations  are  the  major  fac- 
tor underlying  the  observed  seasonal  pattern. 
The  much  greater  abundance  of  mites  tends 
to  mask  other  groups,  such  as  the  Hymenop- 
tera, Diptera,  and  Coleoptera,  which  tend  to 
remain  at  low  to  moderate  levels  of  abun- 
dance throughout  the  season  (Fig.  2B). 

Diversity  values  based  on  pitfall  trapping 
and  Malaise  trapping,  respectively,  were  sub- 
stantially different  from  one  another  during 
all  trapping  periods  of  the  1978  season  (Fig. 
3).  Among  pitfall  samples  arthropod  diversity 
is  generally  low  due  to  the  high  abundance  of 
both  mites  and  ants  (see  Table  1).  The  trend 
toward  increa.sing  diversity  reflects  the  rela- 
tive decrease  in  mite  abundance  in  late  .sea- 
son samples.  Among  Malaise  samples  diver- 
sity values  reflect,  in  part,  the  lower  total 
catch  afforded  by  this  method.  Trapping 


dates  with  the  highest  diversity  values,  12 
June  and  12  August,  had  catches  that  were 
approximately  28  and  8  percent,  respective- 
ly, of  the  catches  for  the  remaining  trapping 
periods.  These  samples  contained  fewer  taxa 
more  equitably  represented,  yielding  greater 
apparent  diversity.  It  is  likely  that  the  values 
for  12  June  and  12  August  Malaise  samples 
are  inordinately  high  due  to  the  sensitivities 
of  the  technique  mentioned  earlier.  Never- 
theless, that  there  should  be  relatively  greater 
diversity  of  aerial  insects  seems  reasonable, 
if  for  no  other  reason  than  their  greater 
mobility. 

Spatial  and  temporal  heterogeneity  of  ar- 
thropods— As  mentioned  above,  spatial  and 
temporal  variation  in  arthropod  abundance 
may  affect  not  only  characteristics  of  individ- 
ual consumer  behavior,  but  also  the  ecologi- 
cal relationships  within  and  between  species. 
As  an  approach  to  spatial  variation  on  a  rela- 
tivelv  small  scale  (minimum  area  effect  of  the 


296 


Great  Basin  Naturalist 


Vol.  40,  No.  3 


8-1 


Hymenoptera 


13 
May 


12 
June 


14 

July 


12 
August 


Fig.  2.  Seasonal  changes  in  arthropod  abundance:  a, 
as  reflected  by  pitfall  and  Malaise  trap  samples;  b,  sea- 
sonal changes  in  the  major  taxa  of  these  samples. 

trapping  grid  is  probably  on  the  order  of  1.4 
ha),  I  calculated  similarity  values  for  all  pos- 
sible combinations  of  grid  arms  for  each 
trapping  period.  An  average  similarity  value 
(S)  was  then  obtained  as  a  measure  of  the 
overall  spatial  heterogeneity  over  the  trap- 
ping grid.  Immediately  apparent  from  this 
analysis  is  the  high  degree  of  similarity  (low 
heterogeneity)  between  grid  arms  at  all 
trapping  periods  (range  =  .730-.893,  Fig.  4). 
Yet,  there  does  appear  to  be  a  trend  toward 
increasing  arthropod  patchiness  with  decreas- 
ing abundance  levels.  The  trend  is  not  statis- 
tically significant,  however,  based  upon  these 
data. 


13 
May 


12 
June 


14 

July 


12 
August 


Fig.  3.  Seasonal  changes  in  arthropod  diversity  as  re- 
flected in  pitfall  and  Malaise  trap  samples.  Pitfall  diver- 
sities represent  the  average  diversity  for  the  four  grid 
arms  at  each  sampling  date. 

Though  evidence  for  spatial  variation  in 
arthropod  numbers  during  1978  is  meager, 
temporal  variation  is  much  more  apparent. 
Average  similarity  values,  based  upon  all  pos- 
sible comparisons  of  each  grid  arm  over  all 
trapping  dates,  were  substantially  lower  than 
for  spatial  variation:  N-arm  =  .621;  E-arm 
=  .648;  W-arm  =  .672;  S-arm  =  .692. 
These  data  indicate  a  substantial  change  in 
the  arthropod  fauna  throughout  the  active 
season  over  a  relatively  small  area.  Com- 
parisons of  Malaise  trap  collections  support 
this  interpretation  (S  =  .384),  but,  as  men- 
tioned above,  the  collections  for  two  of  the 
dates  may  be  suspect. 

Discussion 

Desert  habitats  are  characterized  by  both 
cyclic  and  unpredictable  climatic  changes  on 
micro-  as  well  as  macrogeographic  scales 
(Cloudsley-Thompson  1968,  Logan  1968).  As 
a  result,  these  habitats  are  typified  by  periods 
of  pulsed  production.  It  is  particularly  note- 
worthy that  in  the  first  four  months  of  1978 
Fairview  Valley  received  255  percent  greater 
than  normal  rainfall;  average  temperatures 
between  April  and  August  were  below  nor- 
mal (U.S.  Weather  Bureau  data).  Such  a  com- 
bination of  climatic  events  may  have  provid- 
ed for  a  longer  than  normal  production  pulse, 
resulting  in  a  marked  increase  in  arthropod 
abundance  throughout  the  season.  The  nu- 
merical dominance  by  mites  and  ants  of  col- 


September  1980  Pietruszka:  Seasonal  Variation  of  Arthropods 


297 


12  Aug 


14  July 


13  May 


0.5  15  25 

Arthropods/ Trapping    Period    (  x  10^) 

Fig.  4.  Spatial  heterogeneity  of  arthropods  as  reflect- 
ed in  the  average  similarity,  S,  between  pitfall  trap  grid 
arms  at  each  sample  date. 

lections  spanning  the  entire  season  seems  to 
argue  for  this  possibiHty.  Moreover,  it  has 
been  suggested  that  at  extremely  high  popu- 
lation levels  habitat  patchiness  will  be  re- 
duced, and  localized  areas  may  even  become 
unifonn  in  their  species  distributions  (Wiens 
1976).  Indeed,  this  seems  to  be  what  occurred 
in  1978.  If  this  hypothesis  is  correct,  then  ar- 
thropod patchiness  would  be  predicted  to  be 
more  apparent  during  dry  years  when  abun- 
dance levels  are  low.  The  data  to  test  this 
prediction  have  been  gathered  (for  1979,  a 
substantially  drier  year)  but  have  not  yet 
been  analyzed. 

Finally,  it  is  clear  from  these  analyses  that 
to  effectively  monitor  arthropod  populations 
no  single  methodology  is  sufficient.  The  com- 
binati  ai  of  pitfall  and  Malaise  trapping  pro- 
vides a  reasonable  balance  of  terrestrial  and 
aerial  forms  from  desert  habitats.  Never- 
theless, specific  situations  and  goals  will  ulti- 
mately determine  the  techniques  to  be  used. 

Literature  Cited 

Allred,  D.  M.  1973.  Scorpions  of  the  National  Testing 
Station,  Idaho.  Great  Ba.sin  Nat.  33:251-254. 

Allred,  D.  M.,  and  VV.  J.  Gertsch.  1976.  Spiders  and 
scorpions  from  northern  Arizona  and  southern 
Utah.  J.  Arachnol.  3:87-99. 


Allred,  D.  M.,  D.  E.  Johnson,  and  D.  E.  Beck.  1965.  A 
list  of  some  beeflies  of  the  Nevada  Test  Site. 
Great  Ba.sin  Nat.  25:5-11. 

Beck,  D.  E.,  and  D.  M.  Allred.  1968.  Faunistic  inven- 
tory-BYU  ecological  studies  at  the  Nevada  Test 
Site.  Great  Basin  Nat.  28:132-141. 

Billings,  W.  D.  1949.  The  shadscale  vegetation  zone  of 
Nevada  and  eastern  Galifornia  in  relation  to  cli- 
mate and  soils.  .\mer.  .Midi.  Nat.  42:87-109. 

BoRROR,  D.  J.,  D.  .M.  DeLonc,  and  C.  A.  Triplehgrn. 
1976.  .\n  introduction  to  the  study  of  insects.  4th 
ed.  Holt,  Rinehart  and  Winston,  New  York.  852 
pp. 

BoRROR,  D.  J.,  and  R.  E.  White.  1970.  A  field  guide  to 
the  insects  of  .\merica  north  of  Mexico.  Hough- 
ton Mifflin  Go.,  Bo.ston.  404  pp. 

Ghu,  H.  F.  1949.  How  to  know  the  immature  insects. 
Wm.  G.  Brown  Go.  Publishers,  Dubuque,  Iowa. 
234  pp. 

Gloudsley-Thompson,  J.  L.  1968.  The  Merkhivat  Je- 
bels:  a  desert  community.  Pages  1-20  in  G.  VV. 
Brown,  ed..  Desert  biology.  .Academic  Press,  .New 
York. 

Fautin,  R.  W.  1946.  Biotic  communities  of  the  Northern 
Desert  Shrub  Biome  in  western  Utah.  Ecol.  Mon- 
ogr.  16:251-310. 

Gertsch,  W.  J.,  and  D.  M.  .-Vllred.  1965.  Scorpions  of 
the  Nevada  Test  Site.  Brigham  Young  Universitv 
Sci.  Bull.,  Biol.  Ser.  6(4):  1-15. 

Gill,  F.  B.,  a.nd  L.  L.  Wolf.  1977.  Nonrandom  foraging 
by  sunbirds  in  a  patchy  environment.  Ecolog\' 
58:1284-1296. 

Logan,  R.  F.  1968.  Gauses,  climates,  and  distribution  of 
deserts.  Pages  21-50  in  G.  W.  Brown,  ed..  Desert 
biology.  .Academic  Press,  New  York. 

Mac.\rthur,  R.  H.  1972.  Geographical  ecolog)'.  Harper 
and  Row,  New  York. 

Mac.\rthur,  R.  H.,  and  E.  R.  Pianka.  1966.  Optimal 
use  of  a  patchy  environment,  .^mer.  Nat. 
100:60.3-609. 

Matthews,  R.  W.,  and  J.  R.  Matthews.  1971.  The  .Ma- 
laise trap:  its  utility  and  potential  for  sampling 
insect  populations.  Michigan  Entomologist. 
4:117-122. 

Schoener,  T.  W.  1970.  Nonsynchronous  spatial  overlap 
of  lizards  in  patchy  habitats.  Ecology  51:408-418. 

Townes,  H.  1972.  A  light-weight  Malaise  trap.  Ent. 
News  83:2.39-247. 

Wiens,  J.  A.  1976.  Population  responses  to  patchy  envi- 
ronments. .\nn.  Rev.  Ecol.  Svst.  7:81-120. 


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TABLE  OF  CONTENTS 

Spatiotemporal  variation  in  phenolog)'  and  abundance  of  floral  resources  on  short- 

CTrass  prairie.  V.  J.  Tepedino  and  N.  L.  Stanton  197 

Dog  owners  and  hvdatid  disease  in  Sanpete  County,  Utah.  Peter  M.  Schantz  and 

Ferron  L.  Andersen  216 

New  grass  distribution  records  for  Arizona,  New  Mexico,  and  Texas.  Stephan  L. 

Hatch  221 

A  comparison  of  epiphvtic  diatom  assemblages  on  living  and  dead  stems  of  the  com- 
mon grass  Phragmites  atistralis.  Judith  A.  Grimes,  Larry  L.  St.  Clair,  and  Sam- 
uel R.  Rushforth 22.3 

Poisonous  plants  of  Utah.  Jack  D.  Brotherson,  Lee  A.  Szyska,  and  William  E.  Even- 
son  229 

The  successional  status  oi  Cupressus  arizvnica.  Albert  J.  Parker  254 

A  self-pollination  experiment  in  Pintis  edulis.  Ronald  M.  Lanner  265 

Comparative  floral  biology  of  Penstemon  eatonii  and  Penstemon  cyananthus  in  cen- 
tral Utah:  a  preliminar\^  study.  Lucinda  Bateman  268 

Differential  habitat  utilization  by  the  sexes  of  mule  deer.  Michael  M.  King  and  H. 

Duane  Smith  2/ -3 

Temporal  activitv  patterns  of  a  Dipodomys  ordii  population.  Clive  D.  Jorgensen,  H. 

Duane  Smith,  and  James  R.  Garcia 282 

New  records  of  western  Trichoptera  with  notes  on  their  biolog).  Bernard  G.  Sweg- 

man  and  Leonard  C.  Ferrington,  Jr 287 

Observations  on  seasonal  variation  in  desert  arthropods  in  central  Nevada.  Robert  D. 

Pietniszka 292 


HE  GREAT  BASIN  NATURALIST 


^  lume  40  No.  4 


December  31,  1980 


Brigham  Young  University 

MUS.  COMP.  ZOO' 
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GREAT  BASIN  NATURALIST 

Editor.  Stephen  L.  Wood,  Department  of  Zoology,  Brigham  Young  University,  Provo,  Utah 

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tural Sciences;  Ernest  L.  Olson,  Director,  Brigham  Young  University  Press,  University 
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5-81  650  50104 


ISSN  0017-3614 


The  Great  Basin  Naturalist 


Published  at  Provo,  Utah,  by 
Brigham  Young  University 

ISSN  0017-3614 


Volume  40 


December  31,  1980 


IMPACT  OF  THE  1975  WALLSBURG  FIRE  ON 
ANTELOPE  BITTERBRUSH  {PURSHIA  TRIDENTATA) 

Fred  J.  W'agstaff 

.\bstract.—  Antelope  bitterbrush  {Pur.sliia  tiidentcita)  is  a  preferred  browse  species  that  is  susceptible  to  decreases 
ill  population  density  due  to  fire.  The  reduction  in  density  of  this  species  due  to  fire  was  determined  bv  sampling 
areas  within  and  adjacent  to  the  burn.  The  1975  burn  caused  a  significant  reduction  in  the  population  density  of 
bitterbmsh.  It  \yas  also  determined  that  rate  of  growth  was  lower  for  plants  within  the  burn. 


In  the  summer  of  1975  a  fire  burned  sever- 
al himdred  acres  of  mule  deer  winter  range 
in  Wasatch  County,  Utah.  The  burned  area 
includes  the  area  from  the  junction  of  the 
Wallsburg  road  southeast  to  the  crest  of  the 
west  Daniels  Canyon  ridge  and  to  the  north- 
east along  Highway  40  to  near  the  Midway 
Junction.  Deer  Creek  Reservoir  is  just  across 
the  highway  to  the  northwest  of  the  burned 
area. 

The  study  plots  are  near  the  northeast  cor- 
ner of  the  burn  in  an  area  where  a  population 
density  of  bitterbrush  was  great  enough  to 
permit  quantitative  analysis  of  the  response 
of  this  species  to  fire.  Burned  and  imburned 
areas  were  studied  along  with  some  islands 
that  escaped  burning.  Bitterbrush  {Purshia 
tridentata)  was  selected  as  an  indicator  spe- 
cies because  of  its  status  as  a  preferred 
brow.se  plant  on  mule  deer  winter  ranges. 

Any  factor  that  causes  significant  changes 
in  the  structure  of  the  plant  communities  on 
winter  ranges  is  of  concern.  One  of  the  most 
significant  agents  known  is  fire.  Fire  has  oc- 
curred naturally  since  time  began  and  is  a 
major  factor  in  determining  the  .structure  of 
many  plant  communities.  In  other  commu- 
nities, man-caused  fire  has  introduced  an 


agent  of  change  that  has  modified  vegetation 
over  large  areas. 

In  a  situation  where  prefire  structure  is  so 
important,  the  impact  of  fire  on  structure  of 
the  plant  community  should  be  known.  Will 
plants  be  killed?  How  long  will  the  impacts 
last?  Will  value  of  the  area  as  mule  deer  win- 
ter range  be  completely  lost?  Can  deer  move 
to  another  winter  range?  These  and  other 
questions  occur  and  need  to  be  answered  if 
the  impacts  of  fire  are  to  be  understood. 

The  area  chosen  for  study  has  been  burned 
several  times  in  the  last  one  hundred  years. 
Fire  has  occurred  at  irregular  times  and  over 
different  portions  of  the  area.  This  has  led  to 
a  mosaic  of  vegetation  types  and  age  struc- 
tures. The  1975  fire  was  much  larger  than 
most  of  the  past  fires  and  affected  a  signifi- 
cant portion  of  mule  deer  winter  range  in  the 
area.  With  passage  of  four  years,  the  in- 
ception of  the  postfire  plant  succession 
should  be  identifiable. 

Three  major  hypotheses  were  fornuilated 
to  determine  some  of  the  relative  impacts  of 
the  1975  fire.  First,  the  density  of  bitterbmsh 
had  been  significantly  reduced  by  the  fire. 
Secondly,  use  by  mule  deer  is  less  in  the 
burned  area.  Third,  the  shift  in  deer  u.se  has 


'U.S.  Forest  Service  Interniountain  Forest  and  Range  Experiment  Station  Sl.nib  Sciences  Laboratory,  735  North  500  East.  Provo.  Utah  84601. 


299 


3()0 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


had  a  detrimental  effect  on  surviving  bitter- 
bnish  plants. 

Literature  Review 

Literature  pertaining  to  bitterbrush  is  ex- 
tensive, with  over  200  references  identified. 
The  following  references  support  the  results 
of  the  study. 

FiirsJiid  tridcntata  (antelope  bitterbrush)  is 
highly  desirable  as  browse  on  deer  winter 
range  (Bissell  et  al.  1955,  Giunta  et  al.  1978, 
Hoskins  and  Dalke  1955,  Julander  1952, 
Leach  1956,  Longhurst  et  al.  1952,  Mace 
1957,  Reynolds  1960,  Smith  et  al.  1954, 
Smith  1952).  Since  bitterbmsh  is  so  highly 
preferred,  it  can  be  used  as  an  indicator  spe- 
cies for  use  on  an  area  bv  game  animals  and 
game  winter  range  conditions. 

There  have  been  numerous  articles  written 
about  the  impacts  of  fire  on  bitterbrush 
(Blaisdell  1950,  1953,  Blaisdell  and  Mueggler 
1956,  Countryman  and  Cornelius  1957,  Fer- 
guson and  Basile  1966,  Komarek  1965,  Miller 
1963,  and  Pechanec  et  al.  1954).  They  have 
determined  that  browse  production  of  bitter- 
brush plants  that  have  been  burned  has 
lagged  behind  miburned  control  plants  for 
several  years.  Blaisdell  (1950)  also  showed 
that  relative  densities  of  bitterbrush  in 
burned  and  imbiuned  areas  differed  signifi- 
cantly. Nord  (1965)  developed  data  that  dem- 
onstrates the  existence  of  fire-resistant  eco- 
types  where  most  plants  in  the  population 
resprout  after  fire.  Blaisdell  (1953)  and  others 
have  shown  that  variables  of  fire  intensitv, 
fuel  loading,  and  soil  moisture  affect  re- 
sprouting.  Even  nonsprouting  types  will  have 
some  survivors,  particularly  where  the  fire 
does  not  burn  intensely. 


Results 

The  following  data  were  collected  from  six 
100  ft-  quadrats  in  the  burned  and  adjacent 
unburned  area.  Two  of  the  quadrats  were  at 
the  lower  edge  of  the  burn  in  the  sagebrush- 
bitterbrush  tvpe  and  four  near  the  middle  of 
the  burn  in  the  oak-sagebrush  tvpe.  The  rela- 
tive density  of  live  bitterbrush  plants  is 
shown  in  Table  1. 

There  is  a  striking  difference  between  the 
nimiber  of  living  plants  in  the  burned  and  un- 
burned areas.  None  of  the  burned  plots  had 
anv  surviving  old  plants  because  the  fire  was 
intense  enough  to  kill  the  tops  of  all  bitter- 
brush plants.  It  appears  the  fire  mav  have 
been  hotter  at  the  lower  part  of  the  burn  be- 
cause there  were  no  relic  bitterbnish  plants. 
In  the  midslope  plots,  relics  were  found  for 
most  shrubs  of  various  species. 

All  the  young  bitterbrush  plants  in  the 
burned  area  were  from  resprouting  crowns. 
There  were  voung  plants  in  all  the  unburned 
plots,  indicating  the  species  is  successfully  re- 
producing in  the  study  area. 

In  Table  2,  the  data  collected  on  the 
current-year  twig  growth  are  presented. 
Twenty  twigs  per  plant  on  20  plants  (200 
twigs  in  burned  areas  and  200  in  unbiu'iied) 
were  measured  to  determine  if  there  was  anv 
difference  in  current-year  growth.  The  differ- 
ence in  twig  growth  was  found  to  be  statistic- 
ally significant  at  the  90  percent  level.  In 
other  words,  the  burned  plants  were  growing 
at  a  slower  rate. 

For  each  of  the  six  plots,  mule  deer  fecal 
pellet  groups  were  counted.  All  pellet  groups 
were  counted  without  regard  to  pellet  age. 
Clearly,  the  unl)urned  areas  have  received 
heavier  use  than  the  adjacent  burned  areas,  as 
shown  in  Table  3. 


Table  L  Relative  density  of  bitterbrush  plants  on  six  plots. 


Burned 

Slope 
location 

Bitte 

rbrush 

plants 

Quadrat 

New 

Old 

Dead 

Total 

1 

Yes 

Mid 

2- 

0 

22' 

24 

2 

Yes 

Mid 

f-i- 

0 

22' 

28 

3 

No 

Mid 

(i 

15 

1 

22 

4 

No 

Mid 

9 

15 

1 

24 

5 

Yes 

Lower 

2- 

0 

0 

2 

6 

No 

Lower 

3 

10 

0 

13 

Includes  relics  probably  killed  by  fire. 
Includes  resprouting  after  fire. 


December  1980 


Wagstaff:  Impact  of  Fire  ox  Bitterbrush 


301 


Takle   2.    1979   lcn<4tli    (if  tuitrs  on   20  liitterhrush 
plants. 


Table  3.  Number  of  mule  deer  fecal  pellet  groups  for 
each  of  six  plots. 


Unhiirned  area  plants 
Aveniije 
Plant  twig  length 

mmiber  (inches) 


Burned  area  plants 
Average 
Plant  twig  length 

number  (inches) 


9.5 

1 

9.3 

2 

9.4 

3 

7.05 

4 

10.15 

5 

10.05 

6 

9.8 

7 

6.55 

8 

8.45 

9 

9.9 

10 

9.02 

y 

Discu: 

SSION 

9.2 

7.75 

9.55 

7.6 

6.95 

7.45 

8.45 

9.1 

6.8 

7.15 

8.0 


There  are  several  obvious  differences  be- 
tween the  burned  and  unburned  plant  popu- 
lations. It  is  evident  the  fire  had  a  detrimen- 
tal impact  on  the  density  of  bitterbrush 
plants  and  production  of  this  component  of 
the  plant  community.  A  relatively  low  per- 
centage of  the  bitterbrush  plants  were  able  to 
resprout  and,  therefore,  most  burned  area 
plants  were  eliminated  from  the  community. 

Sagebnish  was  also  largely  absent  in  the 
burned  areas,  but  in  adjacent  unburned  areas 
it  was  a  significant  part  of  the  community. 
The  marked  change  in  the  community  spe- 
cies composition  caused  by  the  fire  is  still 
very  much  in  evidence.  Also  the  size  of  the 
browse  plants  has  been  altered  to  the  point 
that  much  of  the  burned  area  vegetation 
would  be  totally  covered  by  several  inches  of 
snow.  This  physical  barrier  would  limit  use  of 
the  burned  area  even  though  considerable 
forage  may  occur  there. 

Another  obvious  factor  is  the  difference  in 
occurrence  of  fecal  pellet  groups  between 
burned  and  adjacent  unburned  areas.  Since 
the  measurements  were  taken  in  adjacent 
areas,  the  reason  for  the  significant  difference 
seems  easiest  to  explain  on  the  basis  of  physi- 
cal availability  of  browse  diuing  the  winter. 
If  there  are  no  significant  differences  in  nu- 
tritive value  or  availability,  one  would  expect 
essentially  equal  use  near  the  boundary  of  the 
two  areas.  The  difference  in  use  must  occur 
because  of  an  absolute  difference  in  the 
amount  of  browse  available,  which  was  prob- 
ably compounded  by  snow  coverage. 


Plot 


Slope 
position 


Number  of 
pellet 
groups 


Mid 

Burned 

8 

Mid 

Burned 

5 

Mid 

I'nburned 

28 

Mid 

Unburned 

.36 

Low 

Burned 

4 

Low 

Unburned 

16 

All  the  bitterbrush  plants  that  were  exam- 
ined during  this  study  exhibited  substantial 
twig  growth  and  appeared  to  be  vigorous  and 
healthy.  Growth  form  of  the  old  plants  in  the 
unburned  areas  showed  a  fairlx'  open  fcjriu 
that  indicates  little  use  by  sheep  and/or  deer. 
These  plants  did  not  exhibit  the  clubhead 
form  indicative  of  sustained  heavy  use;  nei- 
ther was  there  evidence  of  browsing  on  large- 
diameter  twigs. 

Growth  of  the  Ijitterbrush  plants  in  un- 
burned areas  was  greater,  as  has  been  docu- 
mented in  other  areas  by  Blaisdell  and 
Mueggler  (1956).  This  difference  in  the 
growth  rate  is  expected  to  continue  for  sever- 
al years.  In  terms  of  total  production  of  hit- 
terbnish  forage,  the  burned  area  has  lapsed 
considerably  since  the  fire  and  will  most 
likely  continue  to  do  .so  for  a  long  time.  The 
density  of  plants  has  been  reduced,  as  well  as 
the  size  of  plants.  There  are  fewer  plants  of 
smaller  size  in  the  burned  areas  than  were 
there  prior  to  the  fire.  Since  no  evidence  of 
new  seedlings  could  be  foinid,  it  is  reasonable 
to  conclude  the  burned  area  production  will 
lag  for  nianv  years. 

Deer  herd  populations  are  often  directly 
tied  to  winter  range  availability  and  condi- 
tion because  it  is  the  element  most  often  in 
shortest  supplv.  It  is  not  known  whether  this 
is  the  case  with  the  mule  deer  using  the  study 
area.  There  is  need  for  concern  because  deer 
numbers  are  increasing  and  winter  range  in 
the  area  is  decreasing  due  to  changing  land 
ase.  Both  of  these  trends  cannot  continue  in- 
definitelv  without  deer  niunhers  reaching  the 
limit  of  available  winter  range. 

CONCLL  SIO.NS 

It  is  easv  to  conclude  that  the  197.5  fire 
was  detrimental  to  bitterbnish,  but  it  is  not 


302 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


easy  to  conclude  that  the  numbers  of  deer 
have  been  reduced  because  of  it.  This  is  due 
to  populations  of  deer  being  within  the  ca- 
pacity of  the  remaining  winter  range.  The 
area  is  neither  producing  the  winter  forage 
for  deer  that  it  was  prior  to  the  burn  nor  are 
deer  numbers  anywhere  near  historic  high 
levels. 

If  this  area  is  indeed  a  critical  winter  range 
when  population  numbers  are  larger  and 
other  factors  holding  populations  down  are 
temporary,  additional  concerns  arise. 
Thought  should  be  given  to  introducing  a 
fire-tolerant  ecotype  of  bitterbnish  with  the 
hope  of  hybridizing  this  trait  into  the  com- 
munity, and  effective  means  of  reducing  fire 
occurrence  and  spread  should  be  developed. 

Literature  Cited 

BissELL,  H.  D.,  B.  Harris,  H.  Strong,  and  F.  James. 
1955.  The  digestibility  of  certain  natural  and  arti- 
ficial foods  eaten  by  deer  in  California.  Calif. 
Fish  and  Game  41(1)1.57-78. 

Blaisdell,  J.  p.  1950.  Effects  of  controlled  burning  on 
bitterbni.sh  on  the  upper  Snake  River  plains.  U.S. 
For.  Serv.,  Int.  For.  and  Range  Exp.  Sta.  Res. 
Pap.  20,  .3  pp. 

19.53.  Ecological  effects  of  planned  burning  of 

sagebnish-grass  range  on  the  upper  Snake  River 
plains.  U.S.D.A.  Tech.  Bull.  1075,  .39  pp.,  illus. 

Blaisdell,  J.  P.,  and  W.  F.  Mueggler.  1956.  Sprouting 
of  bitterbnish  (Purshia  tridentata)  following  burn- 
ing or  top  removal.  Ecology  37:.365-370,  illus. 

Countryman,  C.  M.,  and  D.  R.  Cornelius.  1957.  Some 
effects  of  fire  on  a  perennial  range  type.  J.  Range 
Manage.  19:39-41,  illus. 


Ferguson,  R.  B.,  and  J.  V.  Basile.  1966.  Topping  stiniu 
lates  bitterbrush  twig  growth.  J.  Wildl.  .Manage. 
.30(4):  8.39-841. 

Giunta,  B.  C,  R.  Steve.ns,  K.  R.  Jorge.nsen,  and  A.  P. 
Plummer.  1978.  Antelope  bitterbrush— an  impor- 
tant wildland  shrub.  Utah  State  Div.  Wildl.  Re- 
sources Publ.  78-12. 

HosKiNS,  L.  W.,  AND  p.  D.  Dalke.  1955.  Winter  browse 
on  the  Pocatello  big  game  range  in  southeastern 
Idaho.  J.  Wildl.  Manage.  19:21.5-225. 

JuLA.NDER,  O.  1952.  Forage  habits  of  mule  deer  during 
the  late  fall  as  measured  by  stomach  content 
analyses.  U.S.  For.  Serv.  Int.  For.  and  Range  Exp. 
Sta.  Res.  Note-2.  5  pp. 

KoMAREK,  R.  1965.  Fire  and  changing  wildlife  habitat. 
Proc.  Tall  Timbers  Fire  Ecol.  Conf.  2:35-43. 

Leach,  H.  R.  1956.  Food  habits  of  the  Great  Basin  deer 
herds  of  California.  Calif.  Fish  and  Game 
43(4):243-308,  illus. 

LoNGHURST,  W.  M.,  A.  S.  Leopold,  and  R.  F.  Das.ma.nn. 
1952.  A  survey  of  California  deer  herds,  their 
ranges  and  management  problems.  Calif.  Dept. 
Fish  and  Game  Bull.  6.  1.36  pp.,  illus. 

Mace,  R.  LI.  1957.  Oregon's  mule  deer.  Oregon  State 
Game  Comm.  Wildlife  Bull.  No.  3.  25  pp. 

Miller,  H.  A.  1963.  Use  of  fire  in  wildlife  management. 
Proc.  Tall  Timbers  Fire  Ecol.  Conf.  2:19-30. 

NoRD,  E.  C.  1965.  Autecology  of  bitterbnish  in  Califor- 
nia. Ecol.  Monogr.  .35:307-.3.34. 

Pechanec,  J.  F.,  G.  Stewart,  and  J.  P.  Blaisdell.  1954. 
Sagebrush  burning— good  and  bad.  U.S.D..\. 
Farmer's  Bull.  1948.  .34  pp. 

Reynolds,  T.  A.,  Jr.  1960.  The  mule  deer— its  history, 
life  history,  and  management  in  Utah.  Utah  State 
Dept.  Fish  and  Game  Inform.  Bull.  60-4.  .32  pp. 

Smith,  \.  D.,  and  R.  Hubbard.  1954.  Preference  ratings 
for  winter  deer  forages  from  northern  Utah 
ranges  based  on  browsing  time  and  forage  con- 
sumed. J.  Range  Jv-lanage.  7(6):262-265. 

SxuTii,  J.  G.  1952.  Food  habits  of  mule  deer  in  Utah.  J. 
Wildl.  Manage.  16(2):  148-155,  illus. 


TERRESTRIAL  VERTEBRATE  FAUNA  OF  THE  KAIPAROWITS  BASIN 

N.  Diiane  Atwood',  Clyde  L.  Pritchctt',  Richard  D.  Porter',  and  Benjamin  W.  Wood' 

.\bstr^ct.-  This  report  inehides  data  collected  during  an  investigation  by  Brighani  Young  University  personnel 
from  1971  to  1976,  as  well  as  a  literature  review.  The  fauna  of  the  Kaiparowits  Basin  is  represented  by  7  species  of 
mnphihians  (1  salamander,  5  toads,  and  1  tree  frog),  29  species  of  reptiles  (1  turtle,  16  lizards,  and  12  snakes),  183 
species  of  birds  (plus  2  hypothetical),  and  74  species  of  mammals.  Geographic  distribution  of  the  various  species 
within  the  basin  are  discussed.  Birds  are  categorized  according  to  their  population  and  seasonal  status.  .Avian  habitat 
relationships  are  discussed,  and  extensions  of  range  are  reported  for  5  species  of  birds.  Three  threatened  or  endan- 
gered avian  species  occur  in  the  basin.  Four  avian  species  seem  to  have  declined  significantly  in  numbers  in  recent 
\ears. 


The  early  activities  and  exploration  of 
trappers,  missionaries,  and  government  sur- 
vey workers  provided  little  information  to 
om-  knowledge  of  fauna  in  the  Kaiparowits 
Basin.  Most  of  these  early  expeditions  skirted 
around  the  basin  on  all  sides  or  were  con- 
fined to  the  depths  of  the  Colorado  River 
Canyon.  The  Domingues-Velez  de  Escalante 
party  traveled  along  the  north  side  of  the 
Colorado  River  between  Lee's  Ferry  and  the 
well-known  Crossing-of-the-Fathers.  During 
this  portion  of  their  journey,  it  became  neces- 
sary for  the  party  to  eat  their  horses  (Auer- 
bach  1943).  No  mention  is  made  in  Father 
Escalante's  journal  of  any  fauna  observed. 
Records  of  Powell's  expeditions  of  1869-1870 
and  1871-1872  indicate  observations  of  wild- 
life were  limited  to  the  larger  game  animals, 
probably  those  that  could  be  used  for  food. 

In  1892  the  American  Museum  of  Natural 
History  sent  an  expedition  into  the  San  Juan 
region  just  east  of  the  Kaiparowits  Basin  (Al- 
len 1893).  Wetherill,  Flattum,  and  Sterns 
(1961)  made  a  trip  by  boat  up  the  Colorado 
River  from  Lee's  Ferry  to  Rainbow  Bridge. 
Both  expeditions  recorded  the  animals  ob- 
served. 

The  number  of  scientific  investigations  af- 
ter those  of  Powell  into  the  Glen  Canyon 
were  few  indeed  (Crampton  1959).  The  Na- 
tional Park  Service  recognized  this  fact  and, 
in  measure,  filled  some  of  the  gaps  by  send- 
ing out  an  expedition  that  descended  by  boat 


the  San  Juan  and  Colorado  Rivers  to  Rock 
Creek.  From  this  point,  they  spent  about  two 
weeks  on  the  southwest  end  of  the  Kaiparo- 
wits Plateau.  The  results  of  this  work  were 
reported  by  Hall  (1934).  Others  such  as 
Gregory  (1917,  1938,  1945,  1947,  1948)  and 
Gregory  and  Moore  (1931)  have  made  sub- 
stantial contributions  to  our  knowledge  of  the 
Colorado  River  drainage,  particularly  with 
respect  to  the  geology.  Observations  and 
comments  were  also  recorded  regarding  the 
biota. 

Members  of  the  Department  of  Zoology  at 
Brigham  Young  University'  (BYU)  initiated  a 
.series  of  biological  investigations  of  the  up- 
per Colorado  River  Basin;  of  these,  the  fol- 
lowing were  within  the  Kaiparowits  Basin:  In 
July  1927,  a  group  visited  Lee's  Ferry  and 
Bryce  Canyon,  and  in  1936.  a  party  of  four 
biologists  spent  810  man  hours  in  the  Esca- 
lante River  drainage.  In  the  fall  of  1937,  V. 
M.  Tanner  and  C.  L.  Hayward  studied  in  the 
Paria  Valley.  A  party  headed  by  D.  E.  Beck 
entered  the  region  in  the  fall  of  1938  and 
1939,  exploring  the  course  of  the  Escalante 
River.  Beck  also  .spent  the  last  part  of  May 
and  earlv  June  of  1940  in  the  area  east  of 
Willow  Tank  Spring  down  to  the  Escalante 
River.  In  1946,  1952,  and  1953,  other  BYU 
expeditions  entered  the  area  in  both  Kane 
and  Garfield  Counties.  The  Navajo  Mountain 
and  Wahweap  Drainage  were  visited  in  1955 
and  1958  (Hayward  et  al.  1958). 


'U.S.  Forest  Service,  Provo,  Utah  84601. 

■Department  of  Zoology,  Brigham  Young  I'niversity.  Prove,  Utah  84602. 

U.S.  Fish  and  Wildlife'Service,  Provo,  Utah  8460r(.325  North  .300  West.  Mapleton,  Utah  84663). 
'Department  of  Botany  and  Range  Science.  Brigham  Young  University,  Provo,  Utah  84602. 


303 


304 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


11  ,;  .A  30/  ^^J' V    7       Aa;^'^    ;|smok*    NfV^j    -i  ,•     ,    Ky'x         \        ^^'vA 


1/  S  ,v     ^-x^;^  ;, 


NIPPIE       BENCH      "^^^ 


0' 


...^ 


Fig.  1.   Map  of  Brigham  Young  University  Xavajo-Kaiparowits  study  sites,  Kane  and  San  Juan  Counties,  Utah,  and 
Coconino  Count\.  Arizona. 


Benson  (1935)  published  an  important  pa- 
per on  the  fauna  and  flora  collected  during 
an  expedition  led  by  Alexander,  Kellogg,  and 
Benson  in  the  Navajo  Mountain  region  in 
1933.  They  had  six  stations  as  follows:  one 
five  miles  .south  of  the  mountain,  10-12  June; 
one  at  War  God  Spring  on  top  of  the  moun- 
tain at  8400  feet,  13-20  June;  one  at  Bridge 
Canyon,  21-24  June,  two  on  the  mesa  south 
of  the  mountain;  and  the  other  near  Navajo 
Mountain  Trading  Post,  25-26  June.  In  addi- 
tion,  naturalists  assigned   to    Bryce   Canvon 


National  Park  have,  since  1932,  made  contri- 
butions toward  our  knowledge  of  the  fauna 
and  flora  of  the  region. 

Woodbury  and  Russell  (1945),  in  their 
comprehensive  report  on  the  birds  of  the 
Navajo  countrv,  presented  data  on  specimens 
collected  and  observations  made  in  Glen 
Canvon,  on  Navajo  Mountain,  and  on  Kai- 
parowits  Plateau.  Behle  and  three  associates 
made  a  boat  trip  down  the  Colorado  River 
13-17  April  1947.  They  made  observations  at 
various  places  between  Hite  and  Lee's  Ferry, 


December  1980 


Atwood  et  al.:  Kaiparowits  Vertebrates 


305 


including  the  mouth  of  the  Escalante  River, 
river  mile  88;  Hidden  Passage,  river  mile  76; 
Aztec  Creek,  river  mile  68.5;  Crossing-of-the- 
Fathers;  near  Creek,  river  mile  40.5;  and 
Lee's  Ferry.  Behle  and  associates  also  collect- 
ed birds  at  or  near  the  confluence  of  Calf 
Creek  and  the  Escalante  River,  11-14  June 
1953,  and  7-9  May  1954  (Behle  et  al.  1958; 
Behle  1960).  In  1958,  Behle  and  Higgins 
(1959)  made  some  observations  at  Hole-in- 
the-Rock  (20  October),  and  the  confluences 
of  Kane  Creek  and  the  Escalante  River  with 
the  Colorado  River  (19  October);  birds  were 
observed  by  Harold  Higgins  and  Gerald 
Smith  on  a  river  trip  extending  from  1  Julv  to 
9  August  1958  from  Hite  to  Lee's  Ferry 
(Behle  and  Higgins  1959).  We  have  not  given 
the  precise  dates  in  the  species  accounts  giv- 
en below  of  birds  collected  or  seen  bv  Behle 
and  his  associates  on  June  1953  and  Mav 
1954  trips,  or  for  Benson's  (1935)  trips  in 
1933,  becau.se  of  the  short  duration  of  their 
investigations.  If  desired,  such  can  be  obtain- 
ed from  the  literature.  The  dates  of  observa- 
tions from  our  investigations,  however,  are 
reported  herein. 

In  1957,  the  National  Park  Service  in- 
itiated a  salvage  program  for  the  Glen  Can- 
\"on  area  preparatory  to  the  constniction  of 
Glen  Canyon  Dam.  These  data  were  pub- 
lished by  the  University  of  Utah  and  the  Mu- 
seum of  Northern  Arizona  in  their  respective 
journals.  Earlier  (as  a  result  of  a  reconnais- 
sance trip,  8-14  August  1957,  and  as  a  part  of 
tlie  same  project),  Woodbury  et  al.  (1959) 
had  prepared  an  annotated  checklist  of  the 
birds  of  the  proposed  Glen  Canvon  Reservoir 
area  from  various  sources,  including  pub- 
li.shed  and  unpublished  manuscript  records. 
In  addition,  Behle  and  Higgins  (1959)  pub- 
lished a  number  of  previouslv  impubli.shed 
bird  observations  from  Woodburv  and  Rus- 
sell's field  notes  from  the  Rainbow  Bridge- 
Monument  Valley  expedition. 

In  June  1971,  Brigham  Young  University 
and  Northern  Arizona  University  initiated 
the  Environmental  Impact  Studies  for  the 
Navajo  and  later  in  1972  for  the  proposed 
Kaiparowits  Generating  Stations.  Extensive 
collections  and  observations  of  the  terrestrial 
vertebrate  fauna  and  flora  have  been  made 
by  these  groups. 


This  report  on  the  terrestrial  vertebrate 
fauna  is  based  on  species  reported  in  the  lit- 
erature and  collections  or  field  observations 
made  by  BYU  personnel  from  1971  to  the  fall 
of  1976.  The  various  taxa  in  the  annotated 
lists  are  arranged  phylogenetically  following 
the  order  given  in  Schmidt  (1953)  for  the  am- 
phibians and  reptiles;  the  .\merican 
Ornithologists  Union  (AOU)  Check  List,  Fifth 
Edition  (1957)  and  subsequent  supplements 
for  the  bird.s;  and  Hall  and  Kelson  (1959)  for 
the  mammals. 

Most  of  the  studies  dealing  with  the  distri- 
bution and  ecology  of  amphibians  and  rep- 
tiles within  the  Kaiparowits  Basin  have  l^een 
restricted  to  areas  along  the  Colorado  River. 
The  stretch  of  river  generally  known  as  Glen 
Canyon  Gorge  has  been  the  area  most  com- 
monly studied.  The  most  comprehensive  list 
(28  taxa)  of  amphibians  and  reptiles  for  this 
area  was  by  W.  W.  Tanner  (1958a).  Wood- 
bury et  al.  (1959)  published  a  similar  list,  but 
it  contained  only  19  ta.xa.  Both  of  these  stud- 
ies were  designed  to  obtain  data  prior  to  the 
constrution  of  Glen  Canyon  Dam  and  sub- 
sequent development  of  Lake  Powell.  .-Xs  part 
of  the  Navajo-Kaiparowits  environmental 
baseline  studies.  Toft  (1972)  prepared  a  field 
key  ba.sed  on  the  above  literature  and  field 
observations  and  collections  made  during  the 
summer  of  1971  and  1972.  .\dditional  data 
have  been  added  from  subsecjuent  studies 
made  during  the  period  of  1973-1975.  These 
studies  represent  to  date  the  most  extensive 
field  studies  into  the  interior  of  Kaiparowits 
Basin. 


Amphibians 
Ambystomidae  (Salamanders) 

Ambystoma  trigrimim  nehulosum  Hallow- 
ell.  Utah  Tiger  Salamander.  V.  NL  Tanner 
(1930)  and  Weight  ^1932)  Bryce  Canvon.  W. 
W.  Tanner  (1975). 

Pelobatidae  (Spadefoot  Toads) 

Scaphiopus  hammondi  Baird.  Hammonds 
Spadefoot  Toad.  V.  M.  Tanner  (1930)  and 
Weight  (19.321  Bryce  Canvon. 

Scaphiopus  intermontanus  Cope.  Great 
Basin  Spadefoot  Toad.  V.  M.  Tanner  (1940a) 


306 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


Tablf.  1.  Vegetation  types  and  locations  of  perniaiient  sites  for  Biighaiii  Young  University  Navajo-Kaiparowits 
study. 


Site 
No. 


Vegetation  type 


Oryzopsis-Stipa-Ephedra 
Vanclevea 


Location 


At  the  base  of  Cedar  Mt.  on  loo.se, 
deep  sand,  3.2  mi  W  of  GCC 
through  the  highway  fence.  T  4.3  S 
R  2E  S\\'  Part  of  the  NW  quarter 
of  Sec.  21. 


Junipenis-Bouteloua 


On  Cedar  Mt.  4.1  mi  on  from  Site 
1.  .\pprox.  20  vds  N  of  fence  on  left 
of  road.  Exclosure  100  yd  N  of 
fence  and  W  30  yd  R  44  S  T  2E 
NE  corner  of  Sec.  3. 


Hilaria-Ephedra 


Cedar  Mt.  .\pprox.  1.3  mi  on  from 
Site  2.  Road  is  headed  W  and  ca  50 
vd  before  Pole  Line  on  left.  R  44 
S  T  2E  Center  of  Sec.  10,  ca  100 
vd  from  state  line.  Kane  Co.,  UT. 


Coleogyne 


Cedar  Mt.  0.8  mi  on  from  Site  3 
after  turning  left  under  Pole  Line. 
R  6E  T  42.N  SW  part  of  the  NE 
quarter  of  Sec.  5. 


.•\triplex  corrugata- A  triplex 
confertifolia 


.\pprox.  7.0  mi  East  GCC  near 
U.S.  W.  B.  and  Dames  and  Moore 
weather  station  on  Dakota 
formation  on  right,  ca  75  yd  from 
W.  B.  station. 


Pinyon-Juniper- Artemisia 


Smokij  Mt.  14.2  mi  from  Last 
Chance  Jet.  Turn  right  .4  mi  Site 
on  right  of  road.  On  coal  mine  road 
turn  right  at  top  of  Smoky  (survey 
marker)  0.1  mi  then  left  .4  mi  T 
41S  R  4E  NE  part  of  the  SE  quarter 
of  Sec.  8. 


10. 


Gravia-Coleogvne-Hilaria 


Gravia-Hilaria 


Atriplcx-Kochia- 
Artemisia  spinescens 


Orvzopsis 


Smoky  Mt.  12.3  mi  from  Last 
Chance  Jet.  on  top.  E  of  road  near 
dead  juniper.  T  41S  R  8E  NE  part 
of  the  SE  quarter  of  Sec.  21. 

Smokif  Mt.  9.1  mi  from  Last 
Chance  Jet.  (on  left  of  road)  or  5.2 
mi  from  coal  mine  road  jet.  on 
right  side  of  road.  T  41S  R  9E  SW 
part  of  the  SE  quarter  of  Sec.  .34. 

Base  of  Sntokii  Mt.  2  mi  on  from 
Ahlstrom  Point  Jet.  on  left.  Turn  off 
road  in  bottom  of  small  wash  to 
dead  end  at  ca  .2  mi  Site  20  yd  N. 
T  42S  R  5E  NW  part  of  the  NW 
quarter  of  Sec.  .30. 

Alil.strotn  Point  Road  4.7  mi  from 
Ahlstrom  Point  Jet.  T  43S  R  5E  SE 
part  of  the  NW  quarter  of  Sec.  8. 


December  1980 

Tal)Ie  1  tontiiuied. 


Atwood  et  al.:  Kaiparowits  V'ertebrates 


307 


Vegetation  type 


Coleogvne 


Coleotrvne 


Grayia-Ephedra-Orvzopsis 
Boiiteloua-Hilaria 


Grayia-Coleogyne-Bouteloiia 


Popiilus-Tainaiix-L\  ciimi 


Tainari.\-Chr\'sothainn\is 


Spoiol) 


(Joleosivne 


Location 


Ahl.strom  Point  Road  5.0  mi  or  .3 
mi  from  Site  10.  T  4;3,S  R  5E  .\E 
part  of  the  SE  (|iiartL-r  of  Sec.  8. 

\il)))lc  Bench  In  first  well- 
developed  Goleogyne  ca  .3.5  mi 
from  jet.  at  top  after  leaving  Tibbet 
Spring.  On  left  of  road,  there  i,s  a 
.small  ridge  on  right.  T  42  S  R  .3E 
NE  part  of  the  S\\'  qnarter  of  Sec. 
9. 

\ipple  Bench  .\pprox.  .6  mi  on 
fiom  Site  12.  Tnrn  np  wash  to  the 
right  ca  50  ft  into  Sec.  17  from 
marker,  which  is  .1  mi  off  road  up 
the  woods.  T  42  S  R  3E  \E  part 
of  the  NE  qnarter  of  Sec.  17. 

\ip})le  Bench  .Appro.x.  1.1  mi  S  of 
SE  corner  17  on  W  of  road.  2.1  mi 
S  Site  13  T  42  S  R  3E  SE  part  of 
the  SW  ([iiarter  of  Sec.  21. 

Lmt  ('liance  Creek  31.8  mi  from 
C.CXJ  tnrn  \ip  creek  bottom  ca  1  mi 
T  415  R  5E  SE  part  of  the  SE 
quarter  of  Sec.  4. 

.Salt  Wash  at  Middle  Branch  Creek 
Across  the  creek  and  ca  40  vd 
downstream  from  washed  out  stock 
watering  pond.  T  41S  R  4E  SE  part 
of  the  SE  quarter  of  Sec.  .3.5. 

Staleline  7.6  mi  from  (iC^C  on  Utah 
side  of  border  turn  left  toward 
Lone  Rock  .Marina  from  U.S.  89. 
.\pprox.  .2  mi  on  right  of  road. 

Statcline  Same  as  above  except  on 
left  of  road  75  \cl  and  on  a  slight 
rise. 


Coleogyne 


Winihnill  9.3  mi  S  of  Faue  on  89  on 
left  of  highwav  on  pullout  bv 
reflector  post.  T  .39.\  R  8E  SE  part 
of  the  XW  (luarter  of  Sec.  10. 


Bouteloua-Hilaria- 
Muhlenber<;ia 


Pinvon-Juniper 
Coleog)aie 


M  mi  post  .5.35  ca  12  mi  S  of  Page. 
50  vd  \  of  post  and  20  \d  E  of 
road  on  pullout.  T  .39N'R  8E  SW 
part  of  the  .\E  Cjuarter  of  Sec.  20. 

Echo  Cliffs  19.3  mi  S  of  Page  turn 
left  and  on  ridge  ca  '4  mi. 

Xavajo  Plant  Site  (undecided) 


308 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


Table  1  continued. 


Site 
No. 


Vegetation  type 


Location 


23. 


24. 


25. 
26. 


27. 


28. 


29. 


30. 


31. 


32. 


33. 


3.3a. 


33h. 


33c. 


Coleogyne-Ephedra-Grayia 

Hilaria-Ephedra-Grayia 

Atriplex  confertifolia 
Atriplex  corrugata 

Pinyon-Juniper 

Artemisia  tiidentata 

Pinyon-Jiiniper 
Grass-Ephedra 

Coleogvne 
Orvzopsis 


Moist  hanging  gardens 
with  thin-leaved  genera  of 
trees  and  shrubs  various 
genera  of  herbaceous  plants. 

Cercis,  Cladium,  Rhamnus 


Ostrva,  Cirsiium.  Rubus 


Quercus.  Gcltis.  Populus, 
Baccharis,  grass/forlj 


1.2  mi  N  Tibbet  Spring  to  jet.  and 
thence  1.6  mi  E  on  Cathy's  Flat 
Road  50  yd  N  of  road.  T  41S  R  3E 
NE  part  of  the  SE  quarter  of  Sec. 
33. 

2.1  mi  E  Site  2.3.  .\ppro.\.  .2  mi 
beyond  end  of  road.  T  42S  R  3E  SE 
part  of  the  NE  quarter  of  Sec.  3. 

To  be  selected  near  Cath\"s  Flat. 

1(X)  yd  W  of  small  twin  flat-tops  at 
Last  Chance  Summit  on  S  of  road 
ca  13  mi  E  of  Warm  Creek  Jet.  T 
42S  R  .5E  east  central  part  of  the 
SE  quarter  of  Sec.  25. 

The  Pine  ,\pprox.  3  mi  from  cow 
camp  at  head  of  Wesses  Canyon.  T 
40  S  R  2E  part  of  the  SE  quarter  of 
Sec.  2. 

1-1  mi  W  Drip  Jet.  40  yd  SW  from 
Dead  Juniper  on  S  of  road.  T  40S  R 
3E  NE  part  of  the  SW  quarter  of 
Sec.  7. 

Farthest  distance  out  on  Drip 
Point.  T  40S  R  3E  SE  part  of  the 
NE  quarter  of  Sec.  21. 

T  41S  R  1 W  NE  part  of  the  SW 
quarter  of  Sec.  27.  Brigham  Plains 
Bench,  ca  5  mi  N  of  U.S.  Highway 
89,  E  of  Paria  River. 

Grand  Bench  ca  50  mi  E  of  Glen 
Canyon  City.  T  42S  R  6E. 

Grand  Bench  ca  54  mi  E  of  Glen 
Canvon  Citv.  T  42S  R  6E. 


In  Driftwood  Canvon  on  the  .N  side 
of  Lake  Powell  ca'l  mi  NW  of 
Rainbow  Bridge  Canyon.  T  43S  R 
8E. 

Ribbon  Canyon,  between  San  Juan 
Drainage  and  Hole-in-the-Rock  on 
the  E  side  of  Canyon.  T  41S  R  lOE. 

Reflection  Canyon  (Cottonwood 
Gulch),  between  San  Juan  drainage 
and  Hole-in-the-Rock  on  the  W 
side  of  canvon.  T  42S  R  9E. 


December  1980                Atwood  et  al.:  Kaiparowits  Vertebrates 

309 

Table  1  continiifd. 

Site 

No.                                                               Vegetation  type                                                             Location 

.33d.                              Quercus,  Cercis,  Cirsiuin                                                 1  pii  N  of  confluence  of 

Forh/grass                                                                             Colorado/San  Jnan  Rivers  on  E 

side  of  canyon.  T  42S  R  9E  ,5.5  mi  E 
of  Glen  Canyon  Citv  in  Last 
Chance  Creek.  T  41S  R  6E. 

.34. 


Taniarix-dcsert  siiiuh 


K  42S  H9E.55ini  Euftilen 
Canyon  C^itv  in  Last  C^hance 
Creek.  T  4IS  R  6E. 


Willow  Tank  Spring  and  confluence  Calf 
Creek/Escalante  River;  Hayvvard  et  al. 
(1958)  Navajo  Mountain;  Russell  and  Thomp- 
son (1964)  Bryce  Canyon;  BYU  (1972) 
Grosvenor  Arch. 

Bufonidae  (Toads) 

Bufo  cognatus  Sav.  Great  Plains  Toad. 
BYU  (1971)  Site  15. 

Biifo  punctattis  Baird  &  Girard.  Red- 
spotted  Toad.  V.  M.  Tanner  (1940a)  Willow 
Tank  Spring;  Woodbury  et  al.  (1959)  Bridge 
Canyon,  Rock  Creek,  Hidden  Pa.ssage,  Aztec 
Canvon,  Rainbow  Bridge,  and  Warm  Creek; 
BYU  (1971-1973)  Warm  Creek,  Escalante 
drainage,  Wahweap  Creek,  Driftwood  Can- 
yon, Reflection  Canyon,  and  Three  Garden. 

Bufo  woodhoiisei  Girard.  Woodhouses 
Toad.  V.  M.  Tanner  (1940a)  Tropic,  Esca- 
lante, and  Escalante  River;   Hay  ward  et  al. 

(1958)  Navajo   Mountain;   Woodbury   et   al. 

(1959)  Rock  Creek  and  Kane  Creek;  Russell 
and  Thompson  (1964)  Bryce  Canvon;  BYU 
(1972-1973)  Cottonwood  'Wash  Spring,  Re- 
flection Canyon,  and  Tibbet  Spring. 

Hylidae  (Treefrogs) 

Hyla  arenicolor  Cope.  Canyon  Tree  Frog. 
V.  M.  Tanner  (1940a)  confluence  of  the  Esca- 
lante/Colorado  River;  Hayward  et  al.  (1958) 
Paria  Valley;  Woodbury  et  al.  (1959)  Bridge 
Canyon  and  Rock  Creek;  BYU  (1972-1973) 
Driftwood  Canyon  and  Three  Garden. 

Ranidae  (True  Frogs) 

Rana  pipens  brachycephala  Cope.  West- 
em  Leopard  Frog.  V.  M.  Tanner  (1940a)  con- 
fluence   of   the    Escalante/Colorado    Rivers; 


Woodbury  et  al.  (1959)  Bridge  Canyon,  Hole- 
in-the-Rock,  Hidden  Pas.sage,  Rock  Creek, 
West  Canyon,  Padre  Creek,  and  Warm 
Creek;  Russell  and  Thompson  (1964)  Br\ce 
Canyon;  BYU  (1972)  Escalante  River,  Reflec- 
tion Canyon,  and  Ribbon  Canvon. 

Reptiles 

Emydidae  (Water  and  Bo.x  Turtles) 

Chrysemys  picta  belli  Gray.  Western 
Painted  Turtle.  Woodbury  et  al.  (1959)  Rock 
Creek,  Labvrinth  Canvon,  and  three  miles 
above  Face  Canyon;  Miller  (1966)  Hole-in- 
the-Rock. 

Iguanidae  (Iguanid  Lizards) 

Crotaphytus  collaris  subssp.  Western  C^ol- 
lard  Lizard.  Records  of  C.  c.  haileyi  Stejnegar 
are:  V.  M.  Tanner  (1940a)  Paria  River  and 
Henrieville;  Woodbury  et  al.  (1959)  Rainbow 
Bridge  and  Rock  Creek;  W.  W.  Tanner 
(1958a)  collected  specimens  between  Lee's 
Ferry  and  Tuba  City  and  indicated,  "inter- 
gradations  of  .subspecies  C.  c.  haileyi  and  C. 
c.  auriceps  occur  in  the  region  south  of  the 
San  Juan  River  and  west  of  its  confluence 
with  Colorado  River."  BYU  (1971-1974)  Sites 
4  and  11,  .south  ba.se  of  Navajo  Mountain. 
Glen  Canyon,  and  Nipple  Bench.  C.  r.  bicin- 
tores  Smith  &  W.  W.  Tanner  was  named 
from  specimens  collected  at  Crossing-of-lhe- 
Fathers.  An  additional  collection  of  this  ani- 
mal was  made  by  BYU  (1971)  east  of  Smoky 
Mountain. 

Crotaphytus  wislizenii  punctatus  Baird  & 
Girard.  Long-nosed  Leopard  Lizard.  \'.  .M. 
Tanner  (1940a)  Willow  Tank  Spring;  Hay- 
ward  et  al.  (1958)  Navajo  Mountain;  Wood- 


310 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


bury  et  al.  (1959)  Last  Chance  and  Lee's  Fer- 
ry; W.  W.  Tanner  and  Banta  (1963)  Hole-in- 
the-Rock,  Lone  Rock,  Willow  Tank  Spring, 
Catstairs  Canyon,  Crossing-of-the-Fathers, 
Navajo  Mountain  Trading  Post,  and  Lower 
Wahweap  Creek;  BYU  (1971-1973)  Sites  1, 
14,  and  27,  and  Cottonwood  Wash.  Tanner 
and  Banta  (1977).  W.  W.  Tanner  (1980,  pers. 
comm.). 

Sauromalus  obesus  multiforaminatus  Tan- 
ner &  Avery.  Upper  Colorado  River  Chuck- 
walla.  V.  M.  Tanner  (1940a)  Warm  Creek; 
Hayward  et  al.  (1958)  Paria  Valley;  Wood- 
bury et  al.  (1959)  Rainbow  Bridge,  Rock 
Creek,  and  Last  Chance  drainage;  W.  W. 
Tanner  and  Avery  (1964)  Crossing-of-the-Fa- 
thers, Hole-in-the-Rock,  Warm  Creek,  and 
one  mile  upstream  from  Glen  Canyon  Dam; 
BYU  (1971-1972)  Navajo  Creek,  Warm 
Creek  Bay,  Glen  Canyon  Dam,  Grand  Bench, 
and  Navajo  Creek. 

Holbrookia  maculata  approximans  Baird. 
Lesser  Earless  Lizard.  BYU  (1972)  Site  20, 
Tietso  Spring,  and  13  miles  south  of  Page 
along  U.S.  Highway  89. 

Sceloporus  magister  cephaloflacus  Tanner. 
Orange-headed  Desert  Spiny-Lizard.  V.  M. 
Tanner  (1940a)  Willow  Tank  Spring  and 
Wahweap  Creek;  W.  W.  Tanner  (1954a)  Kai- 
parowits  Plateau,  Lone  Rock,  Catstairs  Can- 
yon, Escalante  River,  and  15  miles  northwest 
of  Hole-in-the-Rock;  Hayward  et  al.  (1958) 
Navajo  Mountain;  Woodbury  et  al.  (1959) 
Bridge  Canvon,  Rock  Creek,  and  Kane 
Creek;  BYU  (1971-1974)  Sites  1,  2,  8,  12,  14, 
16,  17,  and  23,  5  miles  south  of  Page,  Tibbet 
Canyon,  Tibbet  Spring,  Wahweap  Bay  area, 
Cottonwood  Wash/U.S.  89,  and  Three  Gar- 
den. 

Sceloporus  undulatus  elongatus  Stejnegar. 
Northern  Plateau  Lizard.  V.  M.  Tanner 
(1940a)  Cannonville,  Escalante,  and  Calf 
Creek/Escalante  River;  W.  W.  Tanner 
(1954d)  Paria  River  drainage  north  of  U.S. 
Highway  89;  Hayward  et  al.  (1958)  Navajo 
Mountain;  Russell  and  Thompson  (1964) 
Bryce  Canyon;  BYU  (1973)  Tibbet  Canyon. 

Sceloporus  graciosus  graciosus  Baird  &  Gi- 
rard.  Great  Basin  Sagebrush  Lizard.  V.  M. 
Tanner  (1930),  Weight  (1932),  and  Presnall 
(1935)  Bryce  Canyon;  V.  M.  Tanner  (1940a) 
Calf  Creek/Escalante  River,  Tropic,  and  Es- 
calante/Colorado  Rivers;   Havward  et  al. 


(1958)  Navajo  Mountain;  BYU  (1971-1974) 
south  base  of  Navajo  Mountain  and  at  Sites  1, 
10,  13,  14,  21,23,  and  27. 

Ufa  stansburiana  uniformis  Pack  &  Tan- 
ner. Upper  Colorado  Basin  Side-blotched 
Lizard.  V.  M.  Tanner  (1940)  Escalante,  Wil- 
low Tank  Spring,  and  Calf  Creek/Escalante 
River;  W.  W.  Tanner  (1954d)  Paria  River 
drainage  north  of  U.S.  Highway  89;  Hayward 
et  al.  (1958)  Navajo  Mountain;  Woodbury  et 
al.  (1959)  Rainbow  Bridge  Trail  and  lower 
Bridge  Canyon;  Russell  and  Thompson  (1964) 
Brvce  Canvon;  BYU  (1971-1974)  Sites  1,  2,  4, 
6,  7,  8,  9,  10,1  3,  14,  17,  18,  19,  20,  21,  22, 
23,  27,  28,  and  30,  Glen  Canyon  City,  Four 
Mile  Bench,  Three  Garden,  and  Tibbet  Can- 
yon. 

Urosaurus  ornata  wrighti  Schmidt.  Colo- 
rado Tree  Lizard.  W.  W.  Tanner  (1954d) 
Paria  River  drainage  north  of  U.S.  Highway 
89;  Hayward  et  al.  (1958)  Navajo  Mountain; 
Woodbury  et  al.  (1959)  Bridge  Canyon,  Rock 
Creek,  and  Last  Chance;  BYU  (1971)  three 
miles  west  of  Site  19. 

Phrynosoma  douglasii  hernandesi  Girard. 
Short-horned  Lizard.  V.  M.  Tanner  (1930), 
Weight  (1932),  and  Presnall  (1935)  Bryce 
Canyon;  BYU  (1972)  Sites  14  and  23,  and 
Warm  Creek  Bay.  W.  W.  Tanner  (1975). 

Phrynosotna  platyrhinos  calidairum  Cope. 
Sonoran  Desert  Horned  Lizard.  W.  W.  Tan- 
ner (1954d)  Paria  River  drainage  north  of 
U.S.  89;  Woodbury  et  al.  (1959)  Rock  Creek 
and  Lee's  Ferry;  BYU  (1971)  Utah-Arizona 
state  line  north  of  Page  along  U.S.  Highway 
89. 


Xantusidae  (Night  Lizards) 

Xantusia  vigilis  utahensis  Tanner.  Utah 
Night  Lizard.  W.  W.  Tanner  (1957)  records 
topotvpes  taken  from  Trachyte  Creek,  Gar- 
field Co.,  Utah;  W.  W.  Tanner  (1958b)  in- 
dicates this  species  possibly  is  found  within 
the  area  defined  as  the  Kaiparowits  Basin. 

Teidae  (Whiptail  Lizards) 

Cnemidophorus  tigris  Baird  &  Girard. 

Western  Whiptail.  Two  subspecies  occur  in 
the  Kaiparowits  Basin,  viz.  C.  t.  tigris  (Baird 
&  Girard)  reported  by  V.  M.  Tanner  (1930), 
Weight    (1932),    and' Presnall    (1935)    Bryce 


December  1980 


Atwood  et  al.:  Kaiparowits  Vertebrates 


311 


Canyon  and  V.  M.  Tanner  (1940a)  Calf 
Creek /Escalante  and  Paria  River  drainage; 
and  C.  t.  septentrionalis  Burger  reported  by 
Woodbury  et  al.  (1959)  from  Rainbow  Bridge 
Trail,  Beaver  Creek,  and  lower  Bridge  Can- 
yon, BYU  (1971-1973)  Sites  1,  2,  6,^^13,  16, 
17,  19,  20,  22,  23,  27,  and  30,  and  the  south 
base  of  Navajo  Mountain. 

Cnemidophorus  velox  Springer.  Plateau 
Whiptail.  Woodbury  et  al.  (1959)  Rock 
Creek;  Schmidt  (1953)  lists  C.  sacki  innotatus 
from  Kanab,  Kane  Countv. 

Cnemidophorus  sexlineatus  perplexus 
Baird  &  Girard,  Six  Lined  Racerunner.  V.  M. 
Tanner  (1940a)  Canonville  and  Escalante. 

Scincidae  (Skinks) 

Eumeces  skiltonianus  utahensis  Tanner. 
Great  Basin  Skink.  Russell  and  Thompson 
(1964)  Bryce  Canyon  beneath  logs  and  stones. 

Colubridae 

Thamnophis  cyrtopsis  cyrtopsis  Kennicott. 

Western  Black-necked  Garter  Snake. 
Schmidt  (1953)  lists  this  species  for  Utah,  Ari- 
zona, southern  Colorado,  New  Mexico,  and 
southward. 

Thamnophis  elegans  vagrans  Baird  &  Gi- 
rard. Wandering  Garter  Snake.  V.  M.  Tanner 
(1930),  Weight  (1932),  and  Presnall  (1935) 
Bryce  Canyon;  V.  M.  Tanner  (1940a)  Tropic 
and  confluence  Calf  Creek /Escalante  River; 
Woodbury  et  al.  (1959)  Rock  Creek. 

Masticophis  taeniatus  taeniatus  Hallow- 
ell.  Desert  Striped  Whipsnake.  W.  W'.  Tan- 
ner (1954d)  Paria  drainage  north  of  U.S. 
Highway  89;  Woodbury  et  al.  (1959)  Beaver 
Creek,  Kaiparowits  Plateau,  and  Hole-in-the- 
Rock;  Russell  and  Thompson  (1964)  Bryce 
Canyon. 

Salvadora  hexalepis  mojavensis  Bogert. 
Mojave  Patch-nosed  Snake.  W.  W.  Tanner 
(1953)  near  the  old  town  site  of  Adairville, 
Kane  County,  Utah;  W.  W.  Tanner  (1954c) 
Wahweap  Creek,  one  mile  southeast  of  Lone 
Rock;  Hayward  et  al.  (1958)  Paria  Valley; 
Woodbury  et  al.  (1959)  Rock  Creek;  BYU 
(1971)  Paria  River  three  miles  south  of  U.S. 
Highway  89  and  at  Glen  Canyon  City. 

Arizona  elegans  philipi  Klauber.  Painted 
Desert  Glossy  Snake.  W.  W.  Tanner  (1964) 


14  miles  south  of  Page,  9  miles  west  of  Page, 
and  2  and  5  miles  west  of  the  Paria  River  all 
along  U.S.  Highway  89;  BYU  (1972)  Sites  1 
and  3,  Arizona-Utah  border  between  Page 
and  Glen  Canyon  City,  and  5  miles  northwest 
of  Page. 

Pituophis  meUinoleucus  deserticola  Stejne- 
gar.  Great  Basin  Gopher  Snake.  Presnall 
(1935)  Bryce  Canyon;  V.  M.  Tanner  (1940a) 
Tropic,  Escalante,  Wahweap  Creek,  and 
W'illow  Tank  Spring;  W.  W.  Tanner  (1954d) 
Paria  River  drainage  north  of  U.S.  Highway 
89;  Hayward  et  al.  (1958)  Navajo  Moimtain; 
BYU  (1971-1973)  Sites  1,  2,  14,  17,  and  20. 

Lampropeltis  getulus  californiac  Blain- 
ville.  California  King  Snake.  \'.  M.  Tanner 
(1940a)  30  miles  south  of  Escalante;  W.  W. 
Tanner  (1958b)  undoubtedly  extending  well 
into  the  upper  Colorado  Basin;  BYU 
(1971-1973)  one  observed  dead  on  U.S.  High- 
way 89  by  Glen  Canyon  City  and  another 
collected  west  of  Cockscomb  Ridge  near  U.S. 
Highway  89. 

Rhinocheilus  lecontei  lecontei  Baird  &  Gi- 
rard. W'estern  Long-nosed  Snake.  W.  W. 
Tanner  (1964)  Wahweap  road  at  junction 
with  U.S.  89  and  Buck  Tank  Draw. 

Sonora  semiannulata  isozona  Cope.  West- 
ern Ground  Snake.  BYU  (1972)  seven  miles 
southeast  of  Glen  Canyon  City  on  U.S.  High- 
way 89.  The  specimens  collected  by  BYU  ex- 
tend the  range  of  this  subspecies  into  Kane 
County  and  east  almost  to  the  Colorado 
River. 

Hypsiglena  torquata  deserticola  Tanner. 
Desert  Night  Snake.  W.  W.  Tanner  (1954b) 
northeastern  Kane  County,  Utah,  in  the  area 
south  and  east  of  the  Vermillion  Cliffs;  Rus- 
sell and  Thompson  (1964)  Bryce  Canyon. 

Hypsiglena  torquata  loreala  Tanner. 
Plateau  Spotted  Night  Snake.  Woodbury  et 
al.  (1959)  Labyrinth  Canyon;  BYU  (1973)  Re- 
flection Canyon. 

Tantilla  planiceps  utahensis  Blanchard. 
Utah  Black-headed  Snake.  W.  W.  Tanner 
(1954c)  Paria  River  drainage  38  miles  east  of 
Kanab  in  Catstairs  Canvon. 


Crotalidae  (Rattlesnakes) 

Crotalus  viridis  lutosus  Klauber.  Great 
Basin  Rattlesnake.  V.  .\1.  Tanner  (1930), 
Weight    (1932),    and   Presnall    (1935)    Bryce 


312 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


Canyon;  W.  W.  Tanner  (1958a)  indicates  that 
this  subspecies  ranges  east  at  least  to  the 
Paria  River.  Pritchett  (1962)  extended  the 
range  to  the  plateau  east  of  the  Paria. 

Crotalus  viridis  nuntius  Klauber.  Hopi 
Rattlesnake.  W.  W.  Tanner  (1958a)  southeast 
portion  of  Navajo  Mountain  near  the  Utah- 
Arizona  border  and  at  the  confluence  of  Es- 
calante  River,  Willow  Tank  Spring,  and 
Wahweap  Creek;  W.  W.  Tanner  (1958a) 
north  Escalante/ Colorado  River  junction; 
BYU  (1972-1974)  Sites  2,  3,  10,  14,  22,  and 
28,  Tibbet  Canyon,  two  miles  southeast  of 
Glen  Canyon,  Tibbet  Spring,  and  Paria, 
Utah. 

Crotalus  viridis  concolor  Woodbury. 
Midget-faded  Rattlesnake.  V.  M.  Tanner 
(1940a)  Tropic,  Escalante  River,  Willow 
Tank  Spring,  and  Wahweap  Creek;  W.  W. 
Tanner  (1958a)  north  of  Escalante/Colorado 
River  junction;  BYU  (1972-1974)  Sites  2,  3, 
10,  14,  22,  and  28,  Tibbet  Canyon,  two  miles 
southeast  of  Glen  Canyon  City,  Tibbet 
Spring,  and  Paria,  Utah. 

Birds 

Present  avian  classification  systems  have 
divided  Fringillidae  into  two  families  and 
have  separated  them  in  their  positions  in  the 
phylogenetic  order  and  have  removed  other 
species  such  as  the  Bushtits  {Psaltripanis) 
from  tlie  families  where  they  have  been  well 
established.  Additionally,  new  positions  in 
the  phylogenetic  order  of  a  number  of 
Passerine  families  have  been  proposed.  Hay- 
ward  et  al.  (1976)  have  discussed  these  di- 
verse taxonomic  changes  in  considerable  de- 
tail and  have  deviated  from  the  Fifth  Edition 
of  the  AOU  Check-list  (1957)  to  conform 
with  some  of  the  proposed  changes.  In  view 
of  the  present  diversity  of  opinion  regarding 
avian  classification  and  because  the  AOU 
Check-list  committee  has  not  yet  made  a  de- 
cision on  the  proposed  changes,  we  have  fol- 
lowed the  Fifth  Edition  of  the  AOU  Check- 
list and  its  supplements. 

Our  ornithological  data  were  collected 
from  July  1971  through  Febniary  1974;  the 
number  of  individual  birds  seen  on  a  monthly 
or  seasonal  basis  and  the  number  seen  in  vari- 
ous vegetational  associations  on  a  seasonal 
basis  are  biased,  since  only  two  years  data 


were  collected  for  the  months  of  March 
through  June.  Also  no  data  were  collected  in 
December  1972.  The  data  are  biased  further 
by  the  fact  that  not  every  observer  deter- 
mined numbers,  nor  did  they  always  report 
the  vegetational  associations  in  which  they 
saw  the  species.  Furthermore,  the  vegeta- 
tional associations  were  not  sampled  equally 
for  bird  species  composition  and  numbers; 
some  were  sampled  more  intensively  than 
others.  Therefore,  our  data  pertaining  to 
avian  seasonal  population  trends  and  their 
preferences  for  certain  vegetational  associ- 
ation are  only  suggestive.  Additionally,  since 
birds  usually  show  decided  preferences  for 
the  architectural  structure  of  the  vegetation 
rather  than  the  plant  species  composition  of 
specific  associations,  many  of  the  plant  asso- 
ciations have  been  lumped  into  Pi7ius-Juni- 
perus  or  Juniper  us  associations,  often  referred 
to  as  woodland,  riparian  {Populus  fremontii, 
Salix,  Tamarix)  associations,  and  desert 
shrubs.  Additional  data  are  available  on  habi- 
tat relationships  of  the  vertebrate  animals 
discussed  here  by  comparing  the  sites  (Fig.  1, 
Table  1)  where  the  animal  was  seen  or  col- 
lected with  the  vegetational  associations 
found  at  that  site.  All  river  miles  listed  for 
the  Colorado  River  are  reckoned  from  Lee's 
Ferry. 

In  the  following  species  accounts,  numbers 
followed  by  localities  and  dates  refer  to  the 
number  of  specimens  taken. 

Podicipedidae  (Grebes) 

Podiceps  nigricollis  C.  L.  Brehm.  Eared 
Grebe.  BYU  (1971)  50  seen  at  Warm  Creek 
Bay,  8  Nov.;  BYU  (1972)  12  seen  at  river  mile 
56,  7  on  Lake  Powell  near  mouth  of  San  Juan 
River,  and  17  at  Three  Garden  on  Lake  Pow- 
ell, 2  Nov.  Behle  et  al.  (1958)  regarded  the 
species  as  a  regular  spring  migrant  through 
the  Kanab  area.  Late  fall  resident  on  Lake 
Powell. 

Aechmophorus  occidentalis  (Lawrence). 
Western  Grebe.  BYU  (1971)  Lake  Powell  6 
seen  on  Warm  Creek  Bay,  8  Nov.  and  one 
seen  on  Wahweap  Bay,  1  Dec.  A  total  of  80 
Western  Grebes  were  counted  at  the  follow- 
ing localities  of  Lake  Powell  2  Nov.  1972: 
river  miles  12,  14,  17,  18  and  20,  Dangling 
Rope    Canyon,    Driftwood    Canyon,    and 


December  1980 


Atwood  et  al.:  Kaiparowits  Vertebrates 


313 


Navajo  Creek.  A  29  June  1973  observation  on 
Lake  Powell  suggests  either  a  straggler  or 
possible  nesting.  Late  fall  transient  south- 
eastern Utah  (Behle  1960). 

Podilymbus  podiceps  (Linnaeus).  Pied- 
billed  Grebe.  BYU  (1971)  on  Lake  Powell, 
one  seen  at  Warm  Creek  Bay,  8  Nov.  and  an- 
other at  Wahweap  Bay,  1  Dec;  BYU  (1972) 
36  were  recorded  at  the  following  localities 
on  Lake  Powell:  Gregory  Butte  near  Rock 
Creek  Bay,  Dangling  Rope  Bay,  and  river 
miles  12  and  15,  2  Nov.  Late  fall  resident  on 
Lake  Powell. 

Pelecanidae  (Pelicans) 

Pelecanus  erythrorhynchos  Gmelin,  White 
Pelican.  Presnall  (1937)  Bryce  Canyon; 
Woodbury  and  Russell  (1945)  three  miles  be- 
low Rock  Creek  on  Colorado  River,  28  July 
1937.  Rare  transient. 

Ardeidae  (Herons  and  Bitterns) 

Ardea  herodias  Linnaeus.  Great  Blue  He- 
ron. Behle  (1948)  Aztec  Creek,  Rock  Creek, 
Last  Chance,  and  river  miles  3,  6,  and  21; 
Woodbury  and  Russell  (1945)  one  at  Lee's 
Ferry,  Aug.  1909  as  recorded  by  the  Nelsen 
and  Birdseye  trip;  Behle  and  Higgins  (1959) 
from  previously  unpublished  observations  of 
Woodbury  and  Russell  on  the  Rainbow 
Bridge-Monument  Valley  expedition,  23 
herons  seen  from  1  to  11  Aug.  1938;  they 
were  seen  from  2V2  miles  below  Lee's  Ferry 
to  river  mile  69,  3100-3200  ft;  Behle  (1960) 
common  from  Hite  to  Lee's  Ferry,  May 
through  Oct.;  BYU  one  at  Warm  Creek/Lake 
Powell,  8  Nov.  1971  and  3  May  1972;  BYU 
(1973)  one  at  Three  Garden  alcove,  21  Mar. 
The  presence  of  a  nesting  colony  at  river 
mile  117  (Woodbury  et  al.  1959)  indicates 
that  the  species  also  nests  in  the  area.  Not  un- 
common resident  between  21  Mar.  and  8 
Nov. 

Egretta  thiila  (Molina).  Snowy  Egret. 
Woodbury  and  Russell  recorded  in  their  un- 
published field  notes  observing  this  species  at 
Forbidding  Canyon  along  the  Colorado  River 
18  July  1937  and  seeing  tracks  on  shore  of 
Colorado  River  between  river  miles  63-69 
(Behle  and  Higgins  1959);  Behle  (1960)  Hig- 
gins saw   17  at  Wahweap  Creek  and  river 


mile  17  and  one  at  the  dam  site,  7  .\ug.  1958; 
BYU  (1972)  one  seen  in  Populus  at  Navajo 
Creek,  27  Apr.;  BYU  (1973)  one  seen  by  Rob- 
ert Whitmore  at  junction  of  Paria  Riv- 
er/Colorado River,  6  June  (riparian  vegeta- 
tion), and  2  more  were  seen  bv  Whitmore 
feeding  from  a  sandbar  in  the  Colorado  River 
one  mile  north  of  Lee's  Ferrv,  15  June. 
Spring  and  summer  resident,  possiblv  breeds. 
Nycticorax  nycticorax  (Linnaeus).  Black- 
crowned  Night  Heron.  \\'oodbur\-  and  Rus- 
sell (1945)  specimen:  river  mile  64,  4  Aug. 
1938  and  six  were  seen  in  65  miles,  i.e.,  be- 
tween river  mile  63  and  2'/2  miles  below 
Lee's  Ferry,  between  4  and  11  .\ug.  1938; 
Behle  (1948)  two  were  seen  16  .\pr.  1947  at 
Crossing-of-the-Fathers  near  Kane  Creek  and 
another  pair  seen  at  Wahweap  Creek.  None 
were  seen  during  the  present  investigation. 
Summer  and  spring  records  suggest  breeding. 

Threskiornithidae  (Ibises  and  Spoonbills) 

Plegadis  chihi  (Vieillot).  White-faced  Ibis. 
Behle  and  Higgins  (1959)  reported  that  ood- 
bury  saw  two  at  river  mile  13,  13  Sept.  1957. 
Uncommon  transient  along  the  Colorado  Riv- 
er. 

Anatidae  (Swans,  Geese,  and  Ducks) 

Branta  canadensis  (Linnaeus).  Canada 
Goose.  Grater  (1947)  Bryce  Canyon;  Behle 
and  Higgins  (1959)  noted  that  Woodbury  saw 
two  at  river  mile  50,  12  Sept.  1957;  Behle 
and  Higgins  (1959)  observed  a  pair  of  adults 
with  six  young  that  were  unable  to  fly  at  riv- 
er mile  106,  16  July  1958;  they  considered 
the  species  a  common  sinnmer  resident  and 
transient  in  Glen  Canyon.  Sununer  resident, 
possiblv  nesting  near  Lake  Powell. 

Anas  platyrhynchos  Linnaeus.  .Mallard. 
Behle  (1948)  six  were  seen  near  Hole-in-the- 
Rock;  BYU  (1971)  three  were  seen  on  a  small 
islet  at  Warm  Creek/Lake  Powell,  8  Nov. 
Uncommon  transient  along  the  Colorado  Riv- 
er and  Lake  Powell. 

Anas  strepera  Linnaeus.  Gadwall.  Behle 
(1948)  saw  a  pair  at  Hole-in-the-Rock,  five  at 
the  mouth  of  San  Juan  River,  and  two  at 
Wahweap  Creek;  BYU  (1972)  five  were  seen 
on  a  pond  at  Grosvenor's  Arch,  2  Mar.  and 
two  at  Lake  Powell /Cottonwood  Wash.  22 


314 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


July  (Ephedra-gr ass);  BYU  (1973)  three  on  a 
pond  near  Coyote  Creek,  5  June.  Uncommon 
spring  and  summer  resident. 

Anas  acuta  (Linnaeus).  Pintail.  Behle 
(1948)  six  near  Hole-in-the-Rock;  BYU  (1973) 
two  on  a  pond  at  Wiregrass  Spring,  5  Nov. 
Spring  and  fall  transient. 

Anas  crecca  (Linnaeus).  Green-winged 
Teal.  Woodbury  and  Russell  (1945)  one,  Col- 
orado River  between  Rock  Creek  and  Lee's 
Ferry,  17  Aug.  1937;  Behle  and  Higgins 
(1959)  Woodbury  saw  seven  at  river  mile  82, 
11  Sept.  1957;  BYU  (1972)  three  at  Wahweap 
Creek  Bay  on  a  rain  pond,  30  Oct.  Transient 
Lake  Powell  and  the  Colorado  River. 

Anas  discors  Linnaeus.  Blue-winged  Teal. 
Behle  (1948)  near  Klondike  Bar;  Behle  and 
Higgins  (1959)  Higgins  saw  six  near  mouth  of 
Rock  Creek  at  river  mile  58,  28  July  1958. 
Uncommon  spring  and  summer  transient  Col- 
orado River  and  Lake  Powell. 

Anas  cyanoptera  Vieillot.  Cinnamon  Teal. 
Woodbury  and  Russell  (1945)  one, 
Paria /Colorado  Rivers,  11  Aug.  1938;  Behle 
(1948)  mouth  of  Wahweap  Creek;  Behle  et 
al.  (1958)  found  one  dead  on  highway  south 
of  Escalante,  8  May  1954;  Behle  and  Higgins 
(1959)  reported  that  in  1958  Woodbury  saw 
one  between  river  mile  63  and  50,  5  Aug.  and 
one  at  V2  mile  below  Lee's  Ferry,  11  Aug. 
Spring  and  summer  transient. 

Anas  americana  Gmelin.  American  Wid- 
geon. BYU  (1972)  two  at  Rainbow  Landing 
on  Lake  Powell,  2  Nov.  Uncommon  transient 
on  Lake  Powell. 


Aythyinae  (Diving  Ducks) 

Aythya  valisineria  (Wilson).  Canvasback. 
Behle  (1948)  near  mouth  of  Last  Chance 
Creek.  Uncommon  transient. 

Bucephala  clangula  (Linnaeus).  Common 
Goldeneye.  Behle  (1948)  saw  12  at  Crossing- 
of-the-Fathers,  12  at  river  mile  36,  and  16  at 
river  mile  31;  BYU  (1972)  one  at  Coyote 
Creek  Pond,  8  Dec.  Spring  and  winter  tran- 
sient. 

Bucephala  albeola  (Linnaeus).  Bufflehead. 
BYU  (1972)  five  on  Lake  Powell  at  Warm 
Creek,  7  Feb.;  three  on  gravel  pit  pond  at 
Wahweap  Bay,  27  Mar.;  and  a  female  on 
Colorado  River  at  Last  Chance  Bay,  5  July. 
Uncommon  spring  and  summer  transient. 


Cathartidae  (American  Vultures) 

Cathartes  aura  teter  Friedmann.  Turkey 
Vulture.  Presnall  (1934)  Bryce  Canyon;  Behle 
and  Higgins  (1959)  reported  that  Woodbury 
saw  two  along  the  Colorado  River  between 
miles  13  and  zero,  10  Aug.  1938;  Woodbury 
and  Russell  (1945)  one,  Navajo  Mountain 
Trading  Post  on  26  July  1936  and  another  on 
8  Aug.  1936  near  Lee's  Ferry,  and  observed 
on  Kaiparowits  Plateau;  BYU  (1971)  two  seen 
12  miles  south  Page,  Ariz.,  24  July;  BYU 
(1972)  Site  2  (flying),  23  July;  Site  12,  16  Apr. 
1972  (Coleogyne);  Site  20;  Driftwood  Can- 
yon, 24  June;  three  seen  at  Dry  Rock  Creek, 
3  May;  and  one  the  confluence  of  the  San 
Juan  River/Lake  Powell,  17  July.  Woodbury 
and  Russell  (1945)  reported  the  Turkey  Vul- 
ture as  a  common  summer  resident  almost 
everywhere  in  the  Navajo  country  area  in  the 
1930s.  They  observed  it  nearly  every  day 
from  1  June  through  15  Sept.  over  a  period 
of  years.  Yet,  present  observations  covering  a 
three-year  span  yielded  fewer  than  a  dozen 
sightings,  suggesting  a  drastic  decline  in  num- 
bers. This  is  particularly  significant  when  one 
considers,  as  noted  by  Woodbury  and  Russell 
(1945),  "that  this  conspicuous  bird  is  seldom 
missed  in  observations  in  both  time  and 
space,"  a  factor  that  tends  to  overemphasize 
its  occurrence  by  comparison  with  less  con- 
spicuous birds  (Woodbury  and  Russell  1945). 
Uncommon  summer  resident  from  28  Mar.  to 
1  Oct.  (in  the  general  area). 


Accipitridae  (Eagles,  Hawks,  and  Harriers) 

Accipiter    gentilis    atricapillus    (Wilson). 

Goshawk.  Woodbury  and  Russell  (1945)  Nav- 
ajo Mountain,  i.e..  War  God  Spring,  23  July 
1936;  Russell  and  Thompson  (1964)  Bryce 
Canyon.  Uncommon  summer  resident. 

Accipiter  striatus  velox  (Wilson).  Sharp- 
shinned  Hawk.  Presnall  (1934)  Bryce  Point; 
Woodbury  and  Russell  (1945)  Navajo  Moun- 
tain, 13  July  1933,  and  Kaiparowits  Plateau, 
30  July  1937,  and  young  birds  out  of  nest 
being  fed  by  parents  on  Kaiparowits  Plateau, 
5-6  Aug.  1937;  Benson  (1935)  base  of  Navajo 
Mountain,  mid- June  1933;  BYU  (1971)  Site  8, 
1  Aug.  and  one  Navajo  Mountain,  13  Oct. 
Fairly  common  summer  resident  and  spring 
and  fall  transient. 


December  1980 


Atwood  et  al.:  Kaiparowits  Vertebrates 


315 


Accipiter  cooperii  (Bonaparte).  Cooper's 
Hawk.  Presnall  (1934)  and  Grater  (1947) 
Bryce  Canyon;  Woodbury  and  Russell  (1945) 
specimens:  Navajo  Mountain,  9  Aug.  1935 
and  25  July  1936  and  two  seen  nesting  on 
Kaiparowits  Plateau,  5  Aug.  1937;  Behle  and 
Higgins  (1959)  Rock  Creek  river  mile  58,  28 
July  1958;  BYU  (1971)  two  near  Site  1  in 
Jiinipenis,  14  and  18  July;  BYU  (1973)  one  at 
Wahweap  Creek,  30  Apr.  {Tamarix  in  Salt- 
wash).  Summer  resident,  spring  and  fall  tran- 
sient. 

Buteo  jamaicensis  calurus  (Cassin).  Red- 
tailed  Hawk.  Presnall  (1934)  Bryce  Canyon; 
Woodbury  and  Russell  (1945)  two  at  Navajo 
Mountain,  7  July  1936  at  3250  ft  and  seen  at 
Beaver  Creek,  War  God  Spring,  Soldier  Seep 
(Navajo  Mountain),  and  on  Kaiparowits 
Plateau;  Behle  and  Higgins  (1959)  reported 
that  Woodbury  found  them  common  along 
Colorado  River,  4-23  July  1936,  and  one  nest 
with  young  hawks  was  also  noted;  Behle  and 
Higgins  (1959)  abundant  in  Glen  Canyon,  1 
July  to  9  Aug.  1958.  BYU  (1971-1973")  Sites 
1,  2,  6,  12,  13,  15,  18,  20,  22,  23,  27,  and  30, 
and  Cottonwood  Wash,  Tibbet  Canyon,  Ce- 
dar Mountain,  Grosvenor  Arch,  Wahweap 
Creek,  and  specimen:  Cockscomb /Highway 
89,  14  Jan.  1972.  Recorded  every  month; 
April-July  (21  seen)  and  November-Januarv 
(17  seen).  The  months  January-June  are  un- 
fairly represented  since  no  records  were  ob- 
tained for  these  months  in  1971.  During  the 
winter  months  most  observations  were  in 
Juniperus  and  associated  species  (17),  and  the 
remainder  were  in  grass  (2),  grass-shrub  (2), 
perched  on  a  ledge  (2),  and  in  Tamarix  (1). 
During  the  spring  and  summer  nesting  season 
they  were  seen  flying  over  desert  shrubs  (9) 
consisting  of  Atriplex,  Vanclevea,  Chryso- 
thamnus,  Artemisia,  and  shrub-grass.  Four  of 
the  nine  were  in  Coleogijne;  three  others 
were  in  Juniperus  and/or  woodland,  and  one 
each  in  grass  and  washbottom  situations.  Per- 
manent resident. 

Buteo  swainsoni  Bonaparte.  Swainson's 
Hawk.  Presnall  (1934)  reported  this  species 
to  be  common  at  Bryce  Canyon  during  the 
summer. 

Buteo  lagopus  (Pontoppidan).  Rough- 
legged  Hawk.  In  1935  Long  (1937)  observed 
one  in  Bryce  Canyon.  Sparse  winter  resident. 


Buteo  regalis  (Gray).  Ferruginous  Hawk. 
Long  (1937)  and  Russell  and  Thompson 
(1964)  Bryce;  BYU  (1973)  one  perched  on 
power  lines  south  of  Glen  Canyon  near 
Warm  Creek,  27  Apr.  Uncommon  transient. 

Aquila  chrysaetos  canadimsis  (Linnaeus). 
Golden  Eagle.  Presnall  (1934)  Rainbow 
Point;  Grater  (1947)  Bryce  Canyon;  Wood- 
bury and  Russell  (1945)  Navajo  Mountain  and 
Glen  Canyon;  Behle  and  Higgins  (1959) 
sighted  adults  and  immatures  near  mouth  of 
Aztec  Creek,  26  July  1958;  BYU  (1971-1974) 
Sites  1,  6,  12,  and  23,  Glen  Canyon  City. 
Church  Wells,  Cockscomb,  Grosvenor  Arch, 
Last  Chance  Creek,  Tibbet  Canyon,  Wah- 
weap Creek,  and  Warm  Creek.  Recorded  42 
times  at  26  locations  during  every  month  of 
the  year  except  October.  Fewest  birds  were 
seen  October  to  November  and  .March  to 
April,  and  most  May- August  followed  bv  De- 
cember-Febniary.  Only  one  eagle  was  seen 
during  our  study,  in  January.  The  vegeta- 
tional  type  over  which  they  were  flving  or  in 
which  they  were  perched  during  Janu- 
ary-June consisted  of  desert  shrubs  (6),  cliffs 
or  rock  ledges  (4),  grasslands  (2),  woodland 
(1),  and  washbottom  (1).  In  August,  Septem- 
ber, and  December  they  were  recorded  in 
mixed  shrubs  and  cliffs  (4).  Common  per- 
manent resident. 

Haliaeetus  leucocephalus  (Linnaeus).  Bald 
Eagle.  Behle  and  Higgins  (1959)  report  that 
Woodbury  observed  one  flying  on  the  east 
side  of  the  Kaiparowits  Plateau  below  the 
cliffs  north  of  Glen  Canyon,  4  Aug.  1938; 
BYU  (1972)  one  seen  near  Rock  Creek  Bay 
and  another  at  confluence  of  San  Juan  Riv- 
er/Lake Powell,  2  Nov.;  BYU  (1975)  Last 
Chance  Wash,  Dec.  Sparse  fall  transient;  the 
4  Aug.  bird  seen  by  Woodbury  may  haNC 
been  a  postnesting  eagle  from  the  small  nest- 
ing populations  in  central  Arizona. 

Circus  cyaneus  hudsonius  (Linnaeus). 
Marsh  Hawk.  Woodbury  and  Russell  il945^ 
one,  Kaiparowits  Plateau,  9  Aug.  1937;  BYU 
(1971)  two  at  Site  17,  27  Aug.  (gras.slands); 
two  at  Site  20,  6  Nov.  (grassland):  BYU  (1972) 
specimen  (male):  Wahweap  Greek.  17  Jan.; 
three  seen  at  Wahweap  Creek  near  Glen 
Canyon  City,  7  Feb.,  18  Apr.,  and  3  July 
(washbottom).  Uncommon  permanent  resi- 
dent; more  common  spring  and  fall  transient. 


316 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


Falconidae  (Falcons) 

Falco  mexicanus  Schlegel.  Prairie  Falcon. 
Presnall  (1934)  and  Russell  and  Thompson 
(1964)  Bryce  Point;  Woodbury  and  Russell 
(1945)  Navajo  Mountain;  one  seen  in  Glen 
Canyon  in  1938  between  river  miles  41  and 
25,  8  Aug.;  and  three  on  cliffs  between  miles 
13  to  zero,  10  Aug.;  Behle  and  Higgins  (1959) 
report  that  in  1938  the  species  was  seen  by 
Woodbury  in  two  places  between  river  miles 
78  and  zero,  4  July;  Behle  (1948)  pairs  (prob- 
ably nesting)  near  mouth  of  Escalante  River 
and  junction  of  Bridge  and  Aztec  Canyons; 
BYU  (1971)  five  sightings;  Site  3,  16  Aug. 
(grasslands);  Site  15,  no  date;  Glen  Canyon 
City/Wahweap  Creek,  1  June  1972  (grass- 
lands); Church  Wells,  14  Nov.  1971  (grass- 
lands); and  Nipple  Creek  (chasing  dove),  23 
May  1973  (washbottom).  Common  summer 
resident,  less  common  in  spring  and  fall; 
probably  a  permanent  resident. 

Falco  peregrinus  anatum  (Bonaparte). 
Peregrine  Falcon.  Russell  and  Thompson 
(1964)  Bryce  Canyon;  Woodbury  and  Russell 
(1945)  saw  peregrines  at  Navajo  Mountain  at 
three  different  sites,  4-10  July  1936,  and  at 
Beaver  Creek,  Navajo  Mountain,  2-16  Aug. 
1936;  Behle  (1960)  Glen  Canyon  near  Wah- 
weap  Creek,  river  mile  17,  6  Aug.  1958;  BYU 
(1971)  two  observations  believed  to  be  per- 
egrines (both  uncertain)  at  Site  15,  8  Oct.  Un- 
doubtedly the  species  formerly  nested  in 
Glen  Canyon,  along  the  Colorado  River,  and 
its  side  canyons  as  well  as  on  Navajo  Moun- 
tain. With  the  construction  of  Lake  Powell, 
the  area  may  now  be  more  suited  for  pere- 
grines than  formerly  (Porter  and  White 
1973).  Since  early  investigators  did  not  exam- 
ine the  area  specifically  for  peregrines,  they 
probably  underestimated  the  size  of  the 
population.  The  entire  area  now  needs  to  be 
thoroughly  surveyed,  especially  Lake  Powell. 

Falco  columbarius  bendirei  Swann.  Mer- 
lin. Grater  (1947)  Bryce  Canyon. 

Falco  sparverius  sparverius  Linnaeus. 
American  Kestrel.  Presnall  (1934)  Bryce  Can- 
yon; Benson  (1935)  Navajo  Mountain;  Wood- 
bury and  Russell  (1945)  two  4  Aug.  1935  and 
11  July  1936  (9,500-10,000  ft,  Picea-Ahies 
and  Pinus  flexilis),  and  seen  on  Kaiparowits 
Plateau  and  at  Lee's  Ferry;  Behle  et  al. 
(1958)  one  seen  10  miles  south  of  Escalante  in 


stand  of  Juniperus,  6  May  1954;  Hayward  et 
al.  (1958)  Escalante  drainage  along  stream- 
sides;  BYU  (1971-1974)  Sites  1,  2,  3,  7,  10, 
12,  14,  16,  19,  and  26;  one  mile  south  of  Glen 
Canyon  City,  two  specimens:  26  Aug.  and  13 
Sept.  1971;  Brigham  Plains,  Cottonwood 
Wash,  Grosvenor  Arch,  Smoky  Mountain, 
Driftwood  Canyon,  Tibbet  Canyon,  Church 
Wells,  Crosby  Canyon  (nesting  on  cliff  face, 
28  Apr.  1973),  Lee's  Ferry,  and  Wahweap 
Creek.  Very  common  (over  75  birds  seen  at 
31  sites);  earliest  spring  sightings,  20  March 
1972  (1  Apr.  1973)  and  latest  fall  sightings,  7 
Nov.  1971  (3  Aug.  1972  and  9  Aug.  1973). 
The  greatest  numbers  were  seen  in  April  (16), 
June  (17),  and  July  (22).  Most  observations 
were  in  desert  shrubs  in  March-July  (29). 
They  were  also  seen  in  Tamarix  (2),  Juniperus 
(1),  saltwash  (1),  Juniperus  in  March  and 
April  (4),  grassland  (4),  and  Tamarix  in 
May- July.  The  shrubs  represented  among  the 
desert  shrubs  were  Vanclevea,  Atriplex,  Graij- 
ia,  Chrifsothamnus,  Artemisia  tridentata,  Co- 
leogyne,  and  Epiiedra.  Summer  resident,  a 
few  may  winter. 

Tetraonidae  (Grouse) 

Dendragapus  obscurus  (Say).  Blue  Grouse. 
Presnall  (1934),  Grater  (1947),  and  Russell 
and  Thompson  (1964)  Bryce  Canyon.  Per- 
manent resident. 

Centrocercus  urophasianus  urophasianus 
(Bonaparte).  Sage  Grouse.  Russell  and 
Thompson  (1964)  Bryce  Canyon.  Permanent 
resident. 

Phasianidae  (Chukars,  Pheasants,  and  Quail) 

Lophortyx  garnbelii  gambelii   Gambel. 

Gambel's  Quail.  Hayward  et  al.  (1958)  Calf 
Creek  area;  BYU  (1971)  one  at  Site  13,  15 
Oct.  {Coleogyne)  and  five  birds  in  Cotton- 
wood Wash;  also  specimen,  16  Dec.  Uncom- 
mon permanent  resident. 

Phasianus  colchicus  Linnaeus.  Ring- 
necked  Pheasant.  Behle  et  al.  (1958)  observed 
and  heard  in  the  fields  around  Escalante,  7 
and  8  May  1954.  Uncommon  permanent  resi- 
dent near  areas  of  cultivation. 

Alectoris  chukar  (Gray).  Chukar.  BYU 
(1972)  Cottonwood  Wash.  Sparse  permanent 
resident. 


December  1980 


Atwood  et  al.:  Kaiparowits  Vertebrates 


317 


Rallidae  (Rails,  Gallinules,  and  Coots) 

Fulica  americana  Gmelin.  American  Coot. 
Woodbury  and  Russell  (1945)  below  mouth 
of  Bridge  Canyon,  15  July  1936;  Behle  and 
Higgins  (1959)  river  mile  47,  flushed  from  S«- 
hx,  30  July  1958;  BYU  (1971)  on  Wahweap 
Creek,  three  miles  east  of  Glen  Canyon  Citv, 
6  Nov.  {Tamarix  around  pond);  BYU  (1973) 
Site  14,  9  May.  Uncommon  summer,  spring, 
and  fall  resident. 


Charadriidae  (Plovers) 

Charadrius  vociferus  vociferus  Linnaeus. 

Killdeer.  Woodbury  and  Russell  (1945)  one, 
Kaiparowits  Plateau,  11  Aug.  1937  (pond, 
7000  ft);  Behle  et  al.  (1958)  confluence  Calf 
Creek/Escalante  River;  BYU  (1971)  two, 
north  of  Church  Wells,  28  Sept.;  BYU  (1972) 
two  near  Glen  Canyon  City,  14  July  and  two 
at  Wahweap  Creek,  13  June  (Tamarix);  BYU 
(1973)  one  at  Coyote  Creek,  5  June.  Uncom- 
mon summer  resident  and  spring  and  fall 
transient. 


Scolopacidae  (Sandpipers  and  Willet) 

Actitis  macularia  (Linnaeus).  Spotted 
Sandpiper.  Woodbury  and  Russell  (1945)  one. 
Rock  Creek /Colorado  River,  20  July  1937 
(3300  ft),  and  as  reported  from  their  impub- 
lished  field  notes  (Behle  and  Higgins  1959) 
they  observed  17  individuals  from  river  mile 
44  to  Lee's  Ferry,  7-10  Aug.  1938;  Behle  and 
Higgins  (1959)  common  along  Colorado 
throughout  Glen  Canyon  during  summer  of 
1958  and  nesting  along  many  of  the  smaller 
streams;  Behle  et  al.  (1958)  confluence  Calf 
Creek/Escalante  River;  BYU  (1973)  one  at 
Coyote  Creek  Pond,  5  June  and  2  at  junction 
Paria/ Colorado  Rivers,  6  June.  Formerly  a 
common  summer  resident,  this  species  seem- 
ingly has  declined  in  numbers  since  the  con- 
struction of  the  Glen  Canyon  Dam  and  the 
filling  of  the  reservoir. 

Catoptrophorus  semipalmatus  inornatus 
(Brewster).  Willet.  Presnall  (1937)  Bryce 
Canyon;  BYU  (1972-1973)  Four  Mile  Bench, 
no  date.  Sparse  transient. 

Calidris  mauri  (Cabanis).  Western  Sand- 
piper. Woodbury  and  Russell  (1945)  near 
Lee's  Ferry,  1 1  Aug.  1938.  Sparse  transient. 


Phalaropodidae  (Phalaropes) 

Steganopus  tricolor  \'ieilIot.  Wilson's 
Phalarope.  Woodbury  and  lUissell  (1945) 
specimen  from  flock  on  Kaiparowits  Plateau. 
2-3  Aug.  1938.  Sparse  fall  transient. 


Laridae  (Gulls  and  Terns) 

Larus  californicus  Lawrence.  California 
Gull.  BYU  (1971)  Warm  Ocek  Bav.  8  .Nov.; 
BYU  (1973)  Lone  Rock  Ba>,  8  Aug.  Sparse 
fall  transient. 

Lams  delawaremis  Ord.  Ring-bilkd  (iiili. 
Behle  (1948)  three  at  Last  Chance  Creek  and 
again  at  the  mouth  of  Kane  Creek,  on  the 
Colorado  River.  Sparse  spring  transient. 

Columbidae  (Doves  and  Pigeons) 

Columba  fasciata  fasciata  Say.  Band- 
tailed  Pigeon.  Benson  (1935)  Navajo  Moun- 
tain; Behle  et  al.  (1958)  Bryce  Canvon.  Sum- 
mer resident. 

Zenaida  macroura  (Linnaeus),  .Mourning 
Dove.  Presnall  (1934)  Br\ce  Canvon;  Benson 
(1935)  Navajo  Mountain;  Tanner  (1940a)  Es- 
calante  River  drainage;  Woodburv  and  Rus- 
sell (1945)  Rock  Creek,  24-26  July;  Kaiparo- 
wits Plateau,  3  Aug.  (1937);  and  War  (iod 
Springs  at  Navajo  Mountain,  16  .Vug.  1935. 
common  breeder  up  to  9000  ft  elevation  on 
Navajo  Mountain;  Behle  et  al.  (1958)  con- 
fluence of  Calf  Oeek/Escalante  River;  Behle 
(1960)  abundant  Coyote  Gulch  (Whitney) 
north  side  of  Kaiparowits  Plateau.  Davis 
Gulch,  and  along  Escalante  River  55  miles 
southeast  of  Escalante,  Aug.  1957;  Behle  and 
Higgins  (1959)  abundant  in  Glen  (.'anyon. 
summer  1958;  BYU  (1971-1973)  specinK-n; 
one  mile  west  Site  2  on  23  July  1971,  Sites  1. 
6,  10,  13,  14,  19,  28,  30.  and  34.  Glen  Canyon 
City,  Cannonville,  Cottonwood  Wash. 
Grosvenor  Arch,  Tibbet  Sprinu,  Lee's  Ferry. 
Nipple  Creek,  Wahweap  Creek.  Warm 
Creek,  Driftwood  Canyon,  and  Hiree  (harden 
one  mile  above  confluence  of  .San  Juan  with 
Lake  Powell.  During  the  .\pril  and  .May  mi- 
gration period  most  Mourning  Doves  were 
encountered  in  washbottoins  102'.  5  were 
seen  in  grasslands,  14  in  desert  shrubs,  and  4 
were  distributed  more  evenly  among  varioas 
vegetational   types;    in   contrast,  during  fall 


318 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


migration  fewest  were  seen  in  washbottoms. 
For  example,  24  were  seen  in  grasslands,  22 
on  desert  shrubs,  21  in  Popidus,  12  in  Tama- 
rix,  15  in  woodland,  6  in  hanging  gardens,  3 
in  open  fields,  2  in  rocks,  and  2  in  washbot- 
toms. In  AugiLst-November  grasslands  were 
most  used  where  39  individuals  were  seen, 
followed  by  desert  shrubs  (27)  and  woodland 
(2).  This  species  was  sighted  155  times  with 
1335  individuals  recorded.  The  earliest  spring 
record  was  15  Apr.  (1972)  and  the  latest  fall 
record  was  1  Nov.  (1973).  The  peak  of  spring 
and  fall,  passage  through  the  area  was  in  May 
and  in  August,  respectively.  The  monthly  to- 
tals for  1972  and  1973  combined  are  as  fol- 
lows: Apr.  (29),  May  (121),  June  (72),  July 
(68),  Aug.  (463),  Sept.  (109),  Oct.  (197),  and 
Nov.  (2).  Abundant  summer  resident. 


Strigidae  (Typical  Owls) 

Otus  asio  (Linnaeus).  Screech  Owl.  Russell 
and  Thompson  (1964)  Bryce  Canyon.  Per- 
manent resident. 

Otus  flammeolus  (Kaup).  Flam mula ted 
Owl.  Woodbury  (1939)  indicates  that  during 
July  1936  H.  N.  Russell,  Jr.,  collected  a  speci- 
men in  Salix  near  War  God  Spring  on  Navajo 
Mountain,  6  July  1936.  Others  were  seen  or 
heard  calling  from  6,  12,  20,  and  21  July; 
they  made  additional  observations  at  Beaver 
Creek,  eight  miles  north  of  Navajo  Mountain, 
8  and  9  Aug.  Resident,  breeding  in  Piniis 
ponderosa  and  pygmy  conifers. 

Bubo  virginianus  pallescens  Stone.  Great 
Homed  Owl.  Presnall  (1934)  and  Behle  et  al. 
(1958)  Bryce  Canyon;  Woodbury  and  Russell 
(1945)  Navajo  Mountain  at  9000  ft;  BYU 
(1971-1973)  specimen:  Site  1,  near  Glen 
Canyon  City,  29  Aug.  1971;  Site  1,  16  Feb. 
1972  (grasslands);  Tibbet  Spring,  15  Apr. 
1972  {Populus  fremontii,  washbottom);  BYU 
(1973)  Four  Mile  Bench,  13  May.  Permanent 
resident. 

Glaucidium  gnoma  californicum  Sclater. 
Pygmy  Owl.  Russell  and  Thompson  (1964) 
Bryce  Canyon.  Permanent  resident. 

Athene  cunicularia  hypugaea  (Bonaparte). 
Burrowing  Owl.  Phillips  et  al.  (1964)  south- 
east portion  of  the  Kaiparowits  Basin;  BYU 
(1972)  Cottonwood  Wash,  9  Aug.  (grassland); 
five  miles  west  of  Glen  Canyon  City,  4  June 
and  21  July;  four  miles  west  of  Glen  Canyon 


City  {Artemisia  fUifoIio)  sitting  in  burrow  on 
side  of  road;  U.S.  89/Cottonwood  Wash 
Road,  22  and  23  July;  BYU  (1973)  three  to 
four  miles  west  of  Glen  Canyon  City,  5  June 
(open  grassland)  nest  under  construction 
nearby;  BYU  (1974)  Utah/Arizona  state  line 
north  of  Page  on  U.S.  89.  Uncommon  sum- 
mer resident. 

Strix  occidentalis  lucida  (Nelson).  Spotted 
Owl.  Woodbury  (1939)  reported  that  this 
species  was  collected  13  Aug.  1936  in  Navajo 
Canyon  by  H.  N.  Russell,  Jr.;  Behle  (1960) 
Davis  Gulch,  a  tributary  of  the  Escalante 
River,  55  miles  southeast  of  Escalante,  Aug. 
1957;  two  were  seen  several  times  in  a  small 
side  canyon  of  Glen  Canyon  near  river  mile 
101,  17  July,  and  another  at  the  mouth  of  the 
Escalante  River,  19  July  1958.  Uncommon 
permanent  resident. 

Asio  otus  (Linnaeus).  Long-eared  Owl. 
BYU  (1973)  Site  30,  9  June  (grassland).  Per- 
manent resident. 


Caprimulgidae  (Goatsuckers) 

Phalaenoptilus  nuttallii  nuttallii  (Audu- 
bon). Poorwill.  Woodbury  and  Russell  (1945) 
Navajo  Mountain  Trading  Post,  22  July  1936, 
and  Kaiparowits  Plateau,  29  July  and  3  Aug. 
1937  (all  in  pygmy  conifers);  common  in 
Aug.  On  Navajo  Mountain,  nests  were  found 
containing  two  downy  young  23  July  1936  in 
pygmy  conifers,  a  family  of  three  poorly 
feathered  juveniles  in  pygmy  forest,  Kaiparo- 
wits Plateau,  7000  ft,  29  July  1937,  and  sim- 
ilar family  on  3  Aug.  not  far  away  in  Arte- 
uiisio-Quercus;  Russell  and  Thompson  (1964) 
Bryce  Canyon;  BYU  (1971)  Paria  Plateau,  29 
Sept.;  BYU  (no  dates)  Site  3  and  Church 
Wells.  A  recent  decline  in  numbers  is  evi- 
denced by  the  paucity  of  sightings  in  the 
present  study  as  compared  to  the  relative 
abundance  of  the  species  in  earlier  studies. 
Uncommon  summer  resident. 

Chordeiles  minor  henryi  Cassin.  Common 
Nighthawk.  Presnall  (1934)  Bryce  Point;  Ben- 
son (1935)  Navajo  Mountain;  Tanner  (1940a) 
Escalante  drainage;  Woodbury  and  Russell 
(1945)  top  Navajo  Mountain,  14  July  1936; 
BYU  (1971-1973)  Sites  2,  3,  4,  and  28;  Glen 
Canyon  Dam;  one  mile  west,  six  miles  east, 
and  2.5  miles  southwest  of  Grosvenor  Arch; 
and  Four  Mile  Bench.  Nighthawks  were  seen 


December  1980 


Atwood  et  al.:  Kaiparowits  Vertebrates 


319 


IS  early  as  19  May  (1972)  and  as  late  as  5 
3ct.  (1972).  Monthly  distribution  of  observa- 
:ions  during  our  study  is  one  in  May,  8  in 
[une,  26  in  July,  37  in  Aug.,  none  in  Sept., 
ind  one  in  Oct.  During  June  and  Julv  night- 
lawks  were  recorded  in  the  following  vege- 
:ation  types  and  in  the  following  niuubers: 
3oleogyne  (2),  mixed  shrub  (1),  Graijia  (1), 
'iiniperus  (6),  and  grassland  (7);  from  August 
^n  they  were  distributed  as  follows:  Coleog- 
/tie  (1),  grass  (35),  Juniperus  (9),  and  Arte- 
nisia  (1).  Common  summer  resident. 


Apodidae  (Swifts) 

Aeronatites  saxatalis  saxatalis  (Wood- 
louse).  White-throated  Swift.  Presnall  (1934) 
3ryce  Canyon;  Benson  (1935)  Navajo  Moun- 
:ain;  Tanner  (1940a)  Kaiparowits  Basin; 
^oodburv  and  Russell  (1945)  one,  Kaiparo- 
wits Plateau,  27  Julv  1937  (7000  ft);  Behle  et 
il.  (1958)  Escalante  River/Calf  Creek;  Hay- 
A'ard  et  al.  (1958)  Escalante  drainage;  Behle 
1960)  Coyote  Gulch  and  Kaiparowits 
Plateau,  Aug.  1957;  Behle  and  Higgins  (1959) 
^ive  numerous  observations  for  lower  Glen 
Hanyon  from  the  field  notes  of  Woodbury 
ind  Russell.  In  these  notes  a  colony  of  swifts 
was  observed  in  a  crevice  of  a  cliff  at  the 
iiouth  of  Rock  Creek,  20-26  July  1937;  six  at 
iiile  2  on  the  San  Juan  River,  1  Aug.  1938, 
md  others  were  seen  between  miles  41  and 
15,  8  Aug.,  and  at  mile  25,  9  Aug.  BYU 
1971-1973)  Sites  1,  2,  3,  4,  10,  and  23,  Cot- 
:onwood  Wash,  Hackberry  Canyon,  Lee's 
Ferry,  Kelly  Grade,  Tibbet  Spring,  Wiregrass 
spring,  Reflection  Canyon,  Ribbon  Canyon, 
md  Three  Garden  one  mile  above  San  Ju^n 
•onfluence  with  Lake  Powell.  Although  most 
;wifts  were  seen  near  cliffs,  those  migrating 
were  seen  away  from  cliffs  and  over  various 
<inds  of  vegetation,  including  Coleogyne, 
^rass,  Grayia,  woodland,  and  Tamarix. 
Where  cliffs  were  near  rivers  or  streams  they 
kvere  seen  above  riparian  streamside.  In  1972 
:he  earliest  spring  sighting  was  28  Apr.;  in 
1973  it  was  1  May.  The  latest  fall  sighting 
was  8  Aug.  in  1972,  when  150  birds  were 
»een.  Sightings  in  April  and  August  contain 
large  numbers  of  swifts  (50-150).  In  May  and 
[une  they  were  usually  seen  singly  or  in  twos 
3r  threes;  9  were  seen  in  one  flock.  In  July 
they  were  seen  mostly  in  groups  containing 


manv  birds.  In  earlv  julv  some  sightings  con- 
tained 1,  2,  5,  and  10  birds.  From  1971-1973 
the  species  was  sighted  37  times  and  324  in- 
dividuals were  recorded.  Common  summer 
resident. 

Trochilidae  (Hummingbirds) 

Archilochns  alcxandri  (Bourcier  &  Mul- 
sant).  Black-chinned  Hununingbird.  Presnall 
(1934)  Bryce  Canyon;  Benson  (1935)  one, 
Navajo  Mountain;  Woodbury  and  Russell 
(1945)  one,  Navajo  Mountain,  9  July  1936 
{Piniis  ponderosa)  and  seen  on  the  Kaiparo- 
wits Plateau;  Behle  et  al.  (1958)  nest  with 
eggs  confluence  of  Calf  Creek /Escalante  Riv- 
er; Behle  (1960)  Kaiparowits  Plateau,  Aug. 
1957;  BYU  (1972)  Tliree  Garden,  25  June. 
Summer  resident. 

Selasphorus  platycercus  platycercus 
(Swainson).  Broad-tailed  Hununingbird.  Pres- 
nall (1934)  and  Russell  and  Thompson  (1964) 
Bryce  Canyon;  Benson  (1935)  Navajo  Moun- 
tain; Woodbury  and  Russell  (1945)  three, 
Navajo  Mountain,  17  July  1936  {Populiis 
treinulokles),  9  Aug.  1935  {Finns  ponderosa, 
8,500-10,500  ft),  and  Kaiparowits  Plateau,  1 
Aug.  1937  (deciduous  shrub,  7000  ft);  Behle 
(1948)  Aztec  Creek;  Behle  et  al.  (1958)  con- 
fluence of  Calf  Creek /Escalante  River;  Hay- 
ward  et  al.  (1958)  Calf  Creek  area;  Behle  and 
Higgins  (1959)  Aztec  Creek,  26  July  1958; 
BYU  (1972)  Reflection  Canyon  and  Three 
Garden.  Summer  resident;  the  species  is 
likely  more  common  than  these  few  sightings 
suggest  because  a  number  of  hummingbirds 
were  not  identified  and  others  were  mis- 
identified. 

Selasphorus  rtifus  (Gmelin).  Rufous  Hum- 
mingbird. Woodbury  and  Russell  (1945)  the 
Nelson  and  Birdseye  trip  noted  the  species 
from  Fort  Defiance  to  Lee's  Ferry;  Wood- 
bury and  Russell  (1945)  four,  Navajo  Moun- 
tain, 9  Aug.  1935,  and  11-17  July  1936,  pon- 
derosa zone,  7000-10,400  ft);  Russell  and 
Thompson  (1964)  Bryce  Canyon;  Woodbury 
and  Russell  (1945)  found  it  mostly  at  altitudes 
above  5300  feet.  Summer  resident. 

Stellida  calliope  (Gould).  C:alliope  Hum- 
ingbird.  Behle  et  al.  (1958)  five  miles  north- 
west of  E.scalante;  Behle  (1960)  one  mile  up 
Aztec  Creek  from   river  mile  68.5.  26  July 


320 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


1958   (believed   to   be   this   species).    Sparse 
migrant. 

Alcedinidae  (Kingfishers) 

Megaceryle  alcyon  (Linnaeus).  Belted 
Kingfisher.  Behle  (1960)  one  seen  at  the 
Crossing-of-the-Fathers  (mouth  of  Kane 
Creek);  river  mile  41,  by  Clayton  White,  2 
Apr.  1954.  Permanent  resident,  more  com- 
mon in  summer. 


Russell  (1945)  three,  Navajo  Mountain,  18 
Aug.  1935;  six,  16  July  1936  {Picea-Abies  and 
Pinus  ponderosa  between  8500-10,000  ft); 
this  expedition  had  nearly  50  records,  chiefly 
from  Navajo  Mountain,  distributed  through 
June,  July,  and  August.  Summer  resident. 

Picoides  pubescens  leucurus  (Hartlaub). 
Downy  Woodpecker.  Long  (1937),  Grater 
(1947),  and  Russell  and  Thompson  (1964) 
Bryce  Canyon.  Summer  resident,  descending 
to  river  bottoms  in  winter  (Behle  1960). 


Picidae  (Woodpeckers) 

Colaptes  auratus  cafer  (Linnaeus).  Red- 
shafted  Flicker.  Presnall  (1934)  Bryce  Can- 
yon; Benson  (1935)  Navajo  Mountain;  Wood- 
bury and  Russell  (1945)  Navajo  Mountain,  5 
Aug.  1934,  10  Aug.  1935,  and  22  July  1936 
(6700-9500  ft  including  pygmy  forest,  Pinus 
ponderosa,  and  Picea-Abies  zones;  most  com- 
monly observed  from  1  Mar.  to  31  Oct.); 
Behle  et  al.  (1958)  eight  miles  south  of  Esca- 
lante,  20  Sept.  1935;  BYU  (1971)  two  speci- 
mens: Site  1,  9  and  15  Oct.:  lower  Cedar 
Moimtain,  7  Oct.  (woodland);  BYU  (1972) 
Site  2,  one  seen  11  Apr.  {Jtinipems);  one  mile 
east  of  Glen  Canyon  City,  28  Jan.  (wood- 
land); Cottonwood  Wash,  29  Sept.  (wash- 
bottom);  BYU  (1973)  Site  28,  13  June  {Jiini- 
penis);  Four  Mile  Bench;  Lee's  Ferry; 
Wiregrass  Spring;  and  Tibbet  Spring,  8  Feb. 
(washbottom).  Permanent  resident,  uncom- 
mon. 

Melanerpes  lewis  (Gray).  Lewis'  Wood- 
pecker. Presnall  (1934)  Bryce  Canyon.  Sum- 
mer resident. 

Sphyrapicus  variiis  (Linnaeus).  Yellow- 
bellied  Sapsucker.  Presnall  (1934)  and  Russell 
and  Thompson  (1964)  Bryce  Canyon;  Wood- 
bury and  Russell  (1945)  one,  Navajo  Moun- 
tain, 23  July  1936,  with  other  observations, 
11-23  July;  BYU  (1971)  Nipple  Spring,  15 
Oct.  (riparian  woodland);  Site  12;  specimen: 
Site  15,  9  Oct.  Summer  reident  and  migrant. 

Sphyrapicus  thyroideus  (Cassin).  William- 
son's Sapsucker.  Presnall  (1934)  and  Behle  et 
al.  (1958)  Bryce  Canyon.  Summer  resident. 

Picoides  villosus  leucothorectis  (Oberhol- 
ser).  Hairy  Woodpecker.  Presnall  (1934) 
Bryce  Canyon;  Benson  (1935)  three,  Navajo 
Mountain;  Behle  et  al.  (1958)  confluence  of 
Calf  Creek /Escalante  River;  Woodburv  and 


Tyrannidae  (Tyrant  Flycatchers) 

Tyrannus  verticalis  Say.  Western  King- 
bird. Woodbury  and  Russell  (1945)  the  Nel- 
son and  Birdseye  trip  recorded  a  few  at  Lee's 
Ferry,  23-24  Aug.  1909;  Woodbury  and  Rus- 
sell (1945)  two.  Rock  Creek /Colorado  River, 
22  July  1937  (streamside  trees,  3300  ft);  Kai- 
parowits  Plateau,  13  Aug.  1937  {Salix,  8000 
ft);  Hay  ward  et  al.  (1958)  Escalante  drainage; 
Behle  and  Higgins  (1959)  several  sightings  in 
lower  Glen  Canyon,  early  Aug.  1938;  Behle 
and  Higgins  (1959)  field  notes  of  Woodbury 
and  Russell  for  1938,  give  the  observation  of 
eight  kingbirds  on  Colorado  River  between 
miles  63  and  2V2  miles  below  Lee's  Ferry, 
5-11  Aug.;  BYU  (1971)  Site  3,  13  July;  BYU 

(1972)  Site  1,  3  Aug.  {Jiiniperus);  between 
Sites  1  and  2,  3  June  (Jtinipems);  Site  3,  3 
June  (grass).  Site  2,  two  seen  4  June  {Jiini- 
perus);   Site   21,   24   June   (woodland);    BYU 

(1973)  Site  12,  9  May;  Nipple  Bench,  1  Aug.; 
and  Tibbet  Spring,  16  June  (Popuhis-Tama- 
rix);  Glen  Canyon  City,  28  Apr.;  Cottonwood 
Wash,  5  June  (Taniarix);  and  Paria  River/U.S. 
89,  three  seen  6  June  (stand  of  Populus). 
Earliest  spring  date,  28  Apr.  (1973);  latest  fall 
date,  3  Aug.  (1972).  Common  summer  resi- 
dent. 

Tyrannus  vociferous  vociferous  Swainson. 
Cassin's  Kingbird.  Presnall  (1934)  and  Russell 
and  Thompson  (1964)  Bryce  Canyon;  Wood- 
bury and  Russell  (1945)  two,  Kaiparowits 
Plateau,  13  Aug.  1937  (chaparral  at  7000  ft) 
and  Rock  Creek/Colorado  River,  26  July 
1937  (Querciis  at  3300  ft);  Behle  et  al.  (1958) 
one,  10  miles  south  of  Escalante,  9  May  1937; 
Behle  (1960)  frequently  seen  in  side  canyons 
of  Glen  Canyon,  1  July  to  9  Aug.  1958;  BYU 
(1973)  Site  3,  three  believed  to  be  this  species 
seen  6  Aug.;  Cottonwood  Wash,  two  mated 


December  1980 


Atwood  et  al.:  Kaiparowits  Vertebrates 


321 


pairs  seen  each  day,  5  and  6  June  (Popuhis)-, 
Paria  River/ U.S.  Highway  89  and  Lee's  Fer- 
ry, two  seen  each  day,  6  and  7  June  {PopuUis- 
Salix-Tamarix).  Summer  resident. 

Myiarchus  tyrannulus  (Muller).  W'ied's 
Crested  Flycatcher.  BYU  (1973)  this  rare  spe- 
cies was  seen  in  semiarid  grassland  several 
times  at  Site  3  by  Clyde  Pritchett  betwen  1 
and  9  Aug.:  two  were  seen  1  Aug.,  three  on  3 
Aug.,  one  on  4  Aug.,  three  on  6  Aug.,  and 
two  on  9  Aug.  Behle  and  Perry  (1975)  report 
this  species  as  a  rare  summer  resident  only  in 
extreme  southwestern  Utah.  Hayward  et  al. 
(1976)  cite  only  those  records  given  bv  Behle 
and  Perry.  Thus,  Pritchett's  observations 
seemingly  extend  the  range  of  this  species  to 
southeastern  Utah.  Sparse  migrant. 

Myiarchus  cinerascens  cinerascens  (Law- 
rence). Ash-throated  Flycatcher.  Presnall 
(1934)  Bryce  Canyon;  Benson  (1935)  Navajo 
Mountain;  Woodbury  and  Russell  (1945)  five. 
Rock  Creek/Colorado  River,  21-26  July 
1937  (canyon  shrubs,  3300  ft)  and  three  miles 
north  Navajo  Mountain  Trading  Post,  12  July 
1934  and  27  July  1936  (pygmy  forest,  6000 
ft);  Behle  et  al.  (1958)  10  miles  south  of  Esca- 
lante,  9  May  1937,  and  confluence  of  Calf 
Creek/Escalante  River;  Behle  and  Higgins 
(1959)  give  at  least  13  observations  from 
Woodbury  and  Russell's  field  notes  for  the 
Colorado  River  between  river  mile  75  and  Vz 
mile  below  Lee's  Ferry,  3-10  Aug.  1938; 
Higgins  (Behle  and  Higgins  1959)  found  this 
flycatcher  to  be  very  abundant  along  the 
length  of  Glen  Canyon  and  side  canyons,  1 
July  to  9  Aug.  1958;  Behle  (1960)  junction 
Colorado  and  Escalante  Rivers,  river  mile  88, 
19  July  1958;  BYU  (1972)  Site  1,  27  June  and 
31  July  (Atriplex);  Reflection  Canyon,  4  July 
(washbottom);  Three  Garden,  3  July  (Hang- 
ing Garden);  confluence  of  San 
Juan/Colorado  Rivers,  24  June;  BYU  (1973)  8 
miles  north  of  U.S.  89  on  Cottonwood  Wash 
road,  5  June  (possibly  nesting  in  Tdiiuirix)-  13 
miles  north  U.S.  Highway  89  on  Cottonwood 
Wash  road,  8  June  (Populu.s);  Brigham  Plains 
Road,  8  June  {Populus);  Cow  Camp  on  Four 
Mile  Bench,  13  June;  Hackberry  Canyon,  8 
June  (scattered  bnish);  2.5  miles  north  Lee's 
Ferry,  14  June  (scattered  brush);  Paria  River 
(1  mile  south  of  Paria),  7  June  (dense  Salix); 
Tibbet  Spring,  three  seen  5  June  (open  shrubs 


and  Populus)..  and  Wiregrass  Spring,  two  seen 
5  June  (Taniarix).  Common  summer  resident. 

Sayornis  nigricans  (Swainson).  Black 
Phoebe.  Woodbury  and  Russell  (1945)  Lee's 
Ferry,  observed  bv  E.  W.  Nelson,  23-26  Aug. 
1909;  Phillips  et  al.  (1964)  south  central  por- 
tion of  the  Kaiparowits  Basin.  Occasional  mi- 
grant or  summer  resident. 

Sayornis  saya  saya  (Bonaparte).  Say's 
Phoebe.  Benson  (1935)  Navajo  Mountain, 
21-24  June  1933;  Tanner  (1940a)  Escalante 
drainage;  Woodbury  and  Russell  (1945)  sev- 
en, Navajo  Mountain,  13  Aug.  1935;  Warm 
Creek,  16  July  1936;  Rock  Creek,  21  July 
1937;  Kaiparowits  Plateau,  4  Aug.  1937; 
Navajo  Mountain  Trading  Post,  27  July  1936 
(pygmy  forest,  Salix,  desert  bnish,  Populus 
tremuioides,  and  Salix,  3190-9000  ft);  Behle 
et  al.  (1958)  Escalante  drainage;  Behle  and 
Higgins  (1959)  three,  Colorado/Escalante 
River,  19  July  1958;  common  all  along  Glen 
Canyon,  summer  1958;  they  believed  this  tax- 
on  nested  in  cliffs  and  larger  trees;  one  in 
Warm  Creek  Canyon,  18  Oct.  1958  {Salix, 
Tamarix,  Baccharis,  and  Pltichea);  in  addition, 
Behle  and  Higgins  (1959)  reported  from  the 
Rainbow  Bridge-Monument  Valley  expedi- 
tion field  notes,  common  along  Colorado  Riv- 
er and  tributaries,  4-22  July  1938,  and  over 

29  seen  between  mile  75  and  V2  mile  below 
Lee's  Ferry,  4-11  Aug.  1938;  Russell  and 
Thompson  (1964)  Bryce  Canyon;  BYU  (1961) 
specimen  (male):  Paria,  20  May;  BYU 
(1971-1973)  one  mile  .south  Site  2,  28  Aug. 
1971;  Sites  3,  7,  12,  13,  and  23,  Church 
Wells,  Cottonwood  Wash  at  miles  5,  8,  and 
11  north  of  U.S.  Highway  89,  Paria  River, 
and  Nipple  Spring;  one,  four  miles  north  of 
Church  Wells.  Phoebes  were  sighted  13  times 
(17  individuals)  at  10  localities.  Habitat  in 
which  phoebes  were  seen,  based  on  only  six 
of  13  sightings,  consisted  of  a  rocky  cliff, 
rockv  areas,  scattered  bru.sh,  Chrysothamnus, 
and  Populus.  The  earliest  spring  date  was  5 
June  (1973).  The  latest  fall  observation  was 

30  Sept.  (1971).  Common  summer  resident. 
Empidonax  traillii   (Audubon).  Willow 

Flycatcher.  Woodbury  and  Russell  (1945) 
five,  Navajo  Mountain,  26  July  1936;  Kai- 
parowits Plateau,  3  Aug.  1937;  Lee's  Ferry 
(nest),  11  Aug.  1938,  and  25  Aug.  1909;  and 
two  miles  below  San  Juan/Colorado  Rivers,  3 
Aug.    1938  (Salix,    Tamarix,   and  Quercus, 


322 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


3200-7000  ft);  Phillips  et  al.  (1964)  south 
central  portion  of  the  Kaiparowits  Basin; 
Higgins  (Behle  1960)  considered  this  species 
common  to  stands  of  Salix,  Tamarix,  Bac- 
charis,  and  Pluchea  along  Colorado  River  in 
Glen  Canyon,  summer  1958;  Woodbury  and 
Russell  (1945)  considered  it  primarily  as  a 
bird  of  .streamside  thickets;  BYU  (1973) 
Hackberry  Canyon,  8  June,  male  displaying 
(Tamarix);  Lee's  Ferry,  7,  14,  15  June,  singing 
(Tamarix  with  Salix);  and  Paria /Colorado 
Rivers,  7  and  14  June  (dense  Salix  and  Tama- 
rix). Summer  resident. 

Empidonax  oberholseri  Phillips.  Dusky 
Flycatcher.  Woodbury  and  Russell  (1945) 
several,  Kaiparowits  Plateau,  31  July;  6  Aug. 
1937  (also  female  seen  with  three  half-grown 
nestlings);  and  13  Aug.  1936  (nest  in 
Quercus);  BYU  (1973)  one  (female),  Tibbet 
Spring,  2  May.  Summer  resident. 

Empidonax  wrightii  Baird.  Gray  Fly- 
catcher. Woodbury  and  Russell  (1945)  one, 
Navajo  Mountain  Trading  Post,  26  July  1936 
(pygmy  conifers,  6500  ft);  BYU  (1973)  two, 
female,  two  miles  east  Glen  Canyon  City,  29 
Apr.  and  male,  Tibbet  Spring,  May.  Summer 
resident. 

Empidonax  difficilis  hellmayri  Brodkorb. 
Western  Flycatcher.  Woodbury  and  Russell 
(1945)  Beaver  Creek/Navajo  Mountain  area, 
7  Aug.  1936  (Quercus,  6000  ft);  Behle  and 
Higgins  (1959)  one,  Kane  Creek /Colorado 
River,  2  Aug.  1958.  Summer  resident. 

Contopus  sordidulus  veliei  Coues.  West- 
em  Wood  Pewee.  Presnall  (1934)  Bryce  Can- 
yon; Woodbury  and  Russell  (1945)  two,  Nav- 
ajo Mountain,  12  Aug.  1935  and  13  Aug.  1936 
(Pinus  ponderosa,  8500-9000  ft);  Hayward  et 
al.  (1958)  Escalante  drainage;  BYU  (1971) 
Site  4  (Paria  Narrows),  30  Sept.  Uncommon 
summer  resident  and  migrant. 

Nuttallornis  borealis  (Swainson).  Olive- 
sided  Flycatcher.  Presnall  (1934)  Bryce  Can- 
yon; Woodbury  and  Russell  (1945)  several, 
Beaver  Creek /Navajo  Mountain  area,  6  and  9 
Aug.  1936,  and  War  God  Spring  on  Navajo 
Mountain,  13  Aug.  1935;  seen  near  Lee's  Fer- 
ry, 10  Aug.  1938;  six  birds  observed  at  War 
God  Spring,  13  Aug.  1937  (Pinus  ponderosa, 
6500-9000  ft);  Behle  and  Higgins  (1959)  seen 
at  river  mile  3.5  and  V2  mile  below  Lee's  Fer- 
ry; and  BYU  (1961)  one,  Paria,  Utah,  20  May. 
Summer  resident. 


Alaudidae  (Larks) 

Eremophila    alpestris    leucolaema    Coues. 

Horned  Lark.  Behle  et  al.  (1958)  10  miles 
southeast  of  Escalante,  8  May  1954;  Russell 
and  Thompson  (1964)  Bryce  Canyon;  Behle 
and  Higgins  (1959)  Higgins  saw  flock  near 
mouth  Kane  Creek,  river  mile  40,  2  Aug. 
1958;  one,  between  Warm  and  Wahweap 
Creeks,  25  March  1958  (3800  ft);  BYU 
(1971-1974)  4,  (males)  T  43S  R2W  S24,  near 
tank,  7  Dec.  1971;  Sites  1,  3,  10,  12,  13,  15, 
and  23;  Warm  Creek;  Church  Wells;  Cotton- 
wood Wash;  Smoky  Mountain;  and  Tibbet 
Canyon;  Summit  Navajo  Mountain;  Esca- 
lante Valley/Little  Valley  Road;  three  miles 
north  U.S.  Highway  89  on  Cottonwood  Wash 
Road.  A  total  of  1639  larks  or  sightings  were 
seen  during  the  study,  at  over  20  locations 
and  from  97  sightings.  A  number  of  sightings 
were  recorded  as  many  or  common.  The  spe- 
cies is  decidedly  more  frequent  in  the  late 
fall/winter  months,  i.e.,  November  through 
February,  when  at  least  1220  were  recorded. 
In  March  through  June  only  87  individuals 
were  recorded,  with  only  3  in  April  and  one 
in  May.  The  numbers  increased  to  32  in  June. 
In  July  and  August  they  greatly  increased 
(165  individuals  plus  five  flocks  containing 
many  individuals).  Only  11  were  seen  in  Sep- 
tember and  156  in  October.  A  January  flock 
contained  600  birds.  The  species  shows  a 
marked  affinity  for  grasslands.  During  the 
nonbreeding  season  of  September-April,  482 
individuals  were  recorded  in  grassland  vege- 
tation, compared  to  75  in  desert  shrubs  and 
15  in  saltwash.  In  May  and  June,  4  were  seen 
in  grass  and  18  in  desert  shrubs.  In  July  and 
August,  45  (one  flock)  were  seen  in  Juni- 
penis/ grass,  and  12  in  grass. 

In  April  through  June,  Honied  Larks  were 
seen  either  singly  (11  of  16  sightings),  in  pairs 
(2),  or  in  small  flocks  containing  6  (2)  or  8  (1) 
birds.  They  congregated  into  larger  groups  or 
flocks  in  July,  half  of  which  contained  10-15 
birds;  one,  about  45  birds;  one,  5  birds,  and 
three,  one  bird  each.  Additionally,  four  flocks 
containing  many  birds  were  also  encountered 
for  which  numbers  were  not  obtained.  As  the 
season  progressed  the  Honied  Larks  were 
seen  in  increasingly  larger  flocks.  For  ex- 
ample, from  July  through  September  the  av- 
erage number  of  birds  per  sighting  was  9.6 


December  1980 


Atwood  et  al.:  Kaiparowits  Vertebrates 


323 


range  1-45);  for  October  and  November  it 
A'as  18.1  (range  10-30);  and  for  December 
;hrough  March  it  was  53.2  (range  1-600). 
During  the  non-nesting  season,  July  through 
Vlarch,  only  7  larks  were  seen  singly,  and 
)nly  one  sighting  each  contained  2  and  3 
)irds;  five  contained  4  birds;  three,  5  birds; 
wo,  6  birds;  one,  7  birds;  two,  9  birds,  seven, 
10  birds;  eight,  20  birds;  one,  23  birds;  one, 
15  birds;  four,  30  bird;  three,  50  birds;  two, 
LOO  birds;  and  one,  600  birds.  Additionallv, 
here  were  five  flocks  for  which  no  coimt  was 
iiade.   Common  vear-roimd  resident  of  the 


Hinrndinidae  (Swallows) 

Tachycineta    thalassina    lepida    Mearns. 

/iolet-green  Swallow.  Presnall  (1934)  Bryce 
Ilanyon;  Benson  (1935)  Navajo  Mountain; 
^Voodbury  and  Russell  (1945)  one,  four  miles 
lorth  of  Navajo  Mountain  Trading  Post,  27 
uly  1936  (pygmy  conifers,  6000  ft)  and  nest 
bund  in  old  woodpecker  hole,  Navajo  Moun- 
ain;  Behle  (1948)  Aztec  Creek;  Behle  et  al. 
1958)  confluence  Calf  Creek /Escalante  Riv- 
M-,  1954;  Behle  and  Higgins  (1959)  Rainbow 
3ridge-Monument  Valley  expedition  field 
lotes,  seen  frequently  along  Colorado  River, 
1-22  July;  over  28  seen  from  river  mile  69  to 
iver  mile  41,  4-8  Aug.  1938;  seen  frequently 
hroughout  Glen  Canyon;  Phillips  et  al. 
1964)  eastern  part  of  the  Kaiparowits  Basin; 
3YU  (1972)  Site  30,  23  Julv;  Cedar  Mountain, 
U  July  {Ephedra);  Site  20',  21  July;  Site  16,  2 
lime  {Tcunarix);  Cottonwood  Wash,  26  June 
Populus);  Nipple  Bench,  1  Aug.;  San 
uan/Colorado  Rivers,  24  June;  Tibbet 
spring,  17  June  (Tamarix);  Four  Mile  Bench, 
18  June  {Artemisia);  BYU  (1973)  Lee's  Ferry, 
15  June;  7  at  Paria /Colorado  Rivers,  14  June 
Tamarix);  Site  7,  4  July;  Site  14,  10  May; 
lear  Site  28,  13  June  {Juniperus/grsLSs);  and 
site  34,  5  June  (open  bnish).  Summer  resi- 
dent. 

Iridoprocne  bicolor  (Vieillot).  Tree  Swal- 
ow.  BYU  (1973)  Lee's  Ferry,  one  seen  6 
fune.  Sparse  migrant. 

Riparia  riparia  riparia  (Linnaeus).  Bank 
5wallow.  Russell  and  Tliompson  (1964)  Bryce 
Canyon.  Summer  resident. 

Stelgidopteryx  ruficollis  (Vieillot).  Rough- 
winged   Swallow.   Woodbury   and   Russell 


(1945)  specimens  at  river  mile  63,  4  Aug. 
1938  and  Lee's  Ferry,  23-26  Aug.  1909; 
Behle  and  Higgins  (1959)  Woodbury  and 
Russell's  field  notes  from  1938,  six  observed 
between  San  Juan /Colorado  Rivers  and  five 
miles  up  the  San  Juan,  2  Aug.;  nine  seen 
along  the  Colorado  River,  3-4  Aug.;  BYU 
(1971)  Wahweap  Creek,  10  Aug.;  BYU  (1973) 
Paria/Colorado  Rivers,  two  seen  in  Tamarix, 
14  June.  Summer  resident. 

Petrochelidon  pyrrhonota  pyrrhonota 
(Vieillot).  Cliff  Swallow.  Presnall  (1934) 
Bryce  Canyon;  Tanner  (1940a)  Kaiparowits 
Basin;  Behle  et  al.  (1958)  confluence  Calf 
Creek/Escalante    River;    Hayward    et    al. 

(1958)  Escalante  drainage;  Behle  and  Higgins 

(1959)  Higgins  found  it  common  in  Glen 
Canyon,  nests  seen  attached  high  canyon 
walls,  July  1958;  BYU  (1973)  Site  13,  11  June; 
Site  .30  nine  seen,  8  June  (nesting  in  rocks  and 
Juniperus);  Nipple  Spring,  12  June;  and  Tib- 
bet  Spring,  10  June.  Fairly  common  summer 
resident. 


Corvidae  (Jays,  Magpies,  Ravens, 
and  Nutcrackers) 

Perisoreus  canadensis  capitalis  (Ridgway. 
Gray  Jay.  Presnall  (1934)  and  Russell  and 
Thompson  (1964)  Bryce  Canyon.  Permanent 
resident. 

Cyanocitta  stelleri  macrolopha  Baird.  Stel- 
ler's  Jay.  Presnall  (1934)  Bryce  Canyon;  Ben- 
son (1935)  four,  Navajo  Mountain,  13-20  July 
1933;  Woodbury  and  Russell  (1945)  several, 
Navajo  Mountain,  13  July  1933,  12  Aug. 
1935,  10  July  and  10  Aug.  1936,  (numerous  in 
Pintis  ponderosa,  9000  ft);  Grater  (1947)  and 
Russell  and  Thompson  (1964)  Bryce  Canyon. 
Permanent  resident. 

Aphelocoma  coerulescens  woodhouseii 
(Baird).  Scrub  Jay.  Benson  (1935)  Navajo 
Mountain  area;  Long  (1937)  and  Russell  and 
Thompson  (1964)  Bryce  Canyon;  Woodbury 
and  Russell  (1945)  four,  Navajo  Mountain,  2 
and  25  Julv  1936  and  Kaiparowits  Plateau,  21 
and  31  July  1937  {Pinus  ponderosa,  Populus 
tremuloides,  pygmy  conifers,  and  Amelan- 
chier,  6500-8500  ft);  Tanner  (1940a)  Kaiparo- 
wits Basin;  Behle  et  al.  (1958)  Calf 
Creek/Escalante  River;  Hayward  et  al. 
(1958)  Escalante  drainage;  BYU  (1971)  ^k 


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Great  Basin  Naturalist 


Vol.  40,  No.  4 


mile  south  Site  1;  BYU  (1972)  one  mile  south 
Glen  Canyon  City.  Permanent  resident. 

Pica  pica  hudsonia  (Sabine),  Black-billed 
Magpie.  Russell  and  Thompson  (1964)  Bryce 
Canyon;  BYU  (1971)  near  Nipple  Spring,  16 
Oct.  (riparian)  and  Wahweap  Creek,  four 
seen  18  Dec.  (Tatnanx);  BYU  (1972)  Wah- 
weap Creek,  four  seen  17  Jan.  (salt  wash)  and 
Glen  Canyon  City,  13  Feb.  (city  dump).  The 
magpie  seemingly  is  absent  during  the  spring 
and  summer,  with  a  few  moving  into  the  area 
during  fall  and  winter. 

Corvus  corax  sinuatus  Wagler.  Common 
Raven.  Presnall  (1934)  Bryce  Canyon;  Benson 
(1935)  one,  Navajo  Mountain;  Woodbury  and 
Russell  (1945)  up  to  10,000  feet  elevation  on 
Navajo  Mountain;  Behle  (1948)  river  mile  31; 
Behle  et  al.  (1958)  10  miles  south  of  Esca- 
lante,  8  May  1954;  BYU  (1971-1973)  Sites  1, 
2,  3,  4,  5,  10,  11,  19,  20,  21,  and  22,  Page, 
Cottonwood  Wash,  Buckskin  Gulch,  Warm 
Creek,  Church  Wells,  Grand  Bench,  Grosve- 
nor  Arch,  Smoky  Mountain,  Tibbet  Canyon, 
Wahweap  Creek,  Driftwood  Canyon,  San 
Juan/Colorado  Rivers,  Reflection  Canyon, 
Lee's  Ferry,  Three  Garden,  one  mile  above 
confluence  of  San  Juan  River  and  Lake  Pow- 
ell, and  east  of  Lone  Rock/Lake  Powell. 
Sight  records  are  distributed  throughout 
every  month  of  the  year.  The  greatest  num- 
ber of  individuals  was  seen  during  December 
(85)  and  February  (97)  and  the  least  during 
January  (6),  April  (7),  and  May  (10).  A  total 
of  407  individuals  were  counted  during  the 
study.  Of  164  sightings  of  ravens,  82  (50  per- 
cent) of  them  were  of  single  birds,  43  (26  per- 
cent) were  of  pairs,  11  (7  percent)  of  triples, 
and  15  (9  percent)  were  quadruples.  Four 
sightings  contained  5  birds  and  four,  6  birds, 
and  one  each  consisted  of  flocks  containing  7, 
10,  25,  and  60.  The  flocks  containing  25  and 
60  birds  were  at  city  dumps.  Ravens  were 
seen  in  all  vegetational  types,  but  the  obser- 
vations were  not  distributed  evenly  among 
them;  of  178  individuals  57  percent  were 
seen  in  grasslands  and  25  percent  were  seen 
in  desert  shnibs.  The  remaining  32  birds  were 
seen  in  Juniperus  (5  percent),  washbottoms  (4 
percent),  Populus  (1  percent)  and  mis- 
cellaneous situations  (8  percent).  Ninety- 
eight  additional  ravens  were  seen  at  city 
dumps.  Permanent  resident. 


Gymnorhinus  cyanocephaliis  Wied.  Pin- 
yon  Jay.  Presnall  (1934)  Bryce  Canyon; 
Woodbury  and  Russell  (1945)  three,  Kaiparo- 
wits  Plateau,  2  and  3  Aug.  1937,  and  Navajo 
Mountain  Trading  Post,  31  July  1936  (pygmy 
forest,  6500-7000  ft);  Woodbury  encountered 
a  flock  of  100  birds  on  Kaiparowits  Plateau,  4 
Aug.  1937;  Behle  et  al.  (1958)  eight  miles 
south  of  Escalante,  19  Sept.  1935;  Hayward 
et  al.  (1958)  Escalante  drainage;  BYU 
(1971-1973)  Sites  1,  2,  3,  4,  14,  21,  27,  28,  29, 
and  30;  Cannonville;  Tibbet  Spring;  and  Cot- 
tonwood Wash;  Paria  Plateau,  29  Sept.; 
lower  Cedar  Mountain;  five  miles  west  Site 
27.  Pinyon  Jays  were  recorded  every  month 
of  the  year  with  the  largest  numbers  in  Aug. 
(310).  Numbers  observed  monthly  were  as 
follows:  5  were  seen  in  Sept.,  14  in  Oct.,  6  in 
Nov.,  16  in  Dec,  3  in  Jan.,  35  in  Feb.,  31  in 
Mar.,  22  in  Apr.,  8  plus  one  flock  in  May,  14 
plus  one  flock  in  June,  and  39  in  July.  A  total 
of  503  individuals  (plus  two  flocks  un- 
counted) were  seen  from  1971  to  February 
1974.  Most  sightings  (25)  and  individuals 
(254)  where  vegetational  type  was  recorded 
were  in  pygmy  conifers.  Six  sightings  and  16 
individuals  were  in  desert  shrubs  consisting  of 
Coleogyne,  mixed  grass-shrubs,  Grayia-grass, 
and  Artemisia.  A  sighting  of  30  individuals 
was  in  reseeded  grass.  Fourteen  of  the  52 
sightings  were  of  single  birds;  4  were  of 
doubles;  6  contained  3  to  5  birds;  8,  6  to  10 
birds;  3,  15  birds;  2  each,  20  and  30  birds; 
and  one  each  consisted  of  16,  17,  35,  37,  and 
100  birds.  Permanent  resident. 

Nucifraga  columbiana  (Wilson).  Clark's 
Nutcracker.  Presnall  (1934)  Bryce  Canyon; 
Benson  (1935)  specimen:  Navajo  Moun- 
tain/Bridge Canyon,  13  July  1933  (seen 
everyday  in  mid-June  1933);  Woodbury  and 
Russell  (1945)  six  specimens:  Navajo  Moun- 
tain, July  1936-37;  Colorado  River  side  can- 
yon next  below  Bridge  Canyon,  13  July  1936 
{Popiihis,  Pinus  ponderosa,  and  Picea-Ahies 
forest  from  3200  to  10,000  ft);  BYU  (1971)  in 
Picea-Ahies  at  summit  Navajo  Mountain,  13 
Oct.  Permanent  resident  higher  mountains, 
moves  lower  after  nesting. 

Paridae  (Bushtits,  Chickadees,  and  Titmice) 

Parus  atricapiUus  garrintis  Behle.  Black- 
capped    Chickadee.    Russell    and   Thompson 


December  1980 


Atwood  et  al.:  Kaiparovvits  Vertebrates 


325 


(1964)  Bryce  Canyon;  Presnall  (1934)  also  re- 
ported P.  a.  septentrionalis  in  Bryce  Canyon. 
However,  Behle  et  al.  (1958)  and  Behle  and 
Perry  (1975)  place  the  area  of  Utah  jnst  west 
of  the  Kaiparowits  Plateau  in  a  zone  of  in- 
tegradation  between  P.  a.  nevadensis  of  west- 
ern Utah  and  P.  a.  gairinus  of  extreme  east- 
em  Utah.  Permanent  resident. 

Pariis  gamheli  Ridgway.  Mountain  Chick- 
adee. Presnall  (1934)  Bryce  Canyon;  Wood- 
bury and  Russell  (1945)  three  at  Kaiparowits 
Plateau,  5-6  and  12  Aug.  1937,  and  one  at 
Navajo  Mountain,  10  July  1936  {Populus 
tremuhides,  Pinus  ponderosa,  and  pvgmv 
conifers,  7000-9500  ft);  BYU  (1971)  Lake 
Powell /Wann  Creek  inlet,  8  Nov.  and  sum- 
mit of  Navajo  Mountain,  12  Oct.  Behle 
(1960)  described  the  race  wasatchensis, 
which  included  the  population  of  the  Kai- 
parowits Basin.  Hayward  et  al.  (1976)  follow 
Snow  (1967a),  who  considers  wasatchensis  to 
be  a  synonym  of  inyoensis.  Permanent  resi- 
dent of  mountains,  moves  lower  in  winter. 

Parus  iyiornatus  ridgwayi  Richmond. 
Plain  Titmouse.  Behle  et  al.  (1958)  con- 
fluence of  Calf  Creek /Escalante  River;  Rus- 
sell and  Thompson  (1964)  Bryce  Canyon; 
BYU  (1971)  specimen:  Paria  Plateau!  29 
Sept.;  BYU  (1972)  Sites  3  and  21,  25  June 
(pygmy  conifers)  and  Navajo  Mountain.  P.  i. 
griseus  which  is  now  a  synonym  of  P.  i.  ridg- 
wayi was  listed  by  Presnall  (1934)  for  Bryce 
Canvon  and  collected  bv  Woodburv  and  Rus- 
sell (1945)  four,  Kaiparowits  Plateau,  31  July 
1937  and  the  Navajo  Mountain  area,  26  Julv 
1936  (pygmy  conifers,  6500-7000  ft).  This 
species  should  be  more  common  in  the  pyg- 
my conifers  than  our  few  observations  sug- 
gest. Woodbury  and  Russell  (1945)  reported 
it  in  pygmy  conifers  every  month  of  the  year; 
they  considered  it  a  permanent  resident. 

Psaltriparus  minimum  (Townsend).  Com- 
mon Bushtit.  Long  (1937)  and  Russell  and 
Thompson  (1964)  Bryce  Canyon;  Woodbury 
and  Russell  (1945)  six,  Kaiparowits  Plateau,  1 
Aug.  1937,  and  Navajo  Mountain,  8,  24,  and 
31  July  1936  and  14  Aug.  1937;  Hayward  et 
al.  (1958)  benches  of  the  Escalante  drainage; 
BYU  (1971)  one,  T43S,  R2W  Sec  24,  7  Dec; 
Site  2,  30  .seen  7  Dec.  (woodland);  also  seen 
in  Tibbet  Canyon,  11  Dec.  (ChrysotJuimniis). 
Permanent  resident.  This  taxon  is  placed  in 
Aegithalidae  hy  Snow  (1967b). 


Sittidae  (Nuthatches) 

Sitta  carolinensis  nelsoni  Mearns.  White- 
breasted  Nuthatch.  Presnall  (1933,  1934,  and 
1936)  and  Russell  and  Thompson  (1964) 
Bryce  Canyon;  Benson  (1935)  three,  Navajo 
Mountain,  13  Aug.  1935  (mixed  coniferous 
forest,  9()()()  ft).  Woodbury  and  Russell  (1945) 
found  this  species  a  regular,  but  not  abun- 
dant, inhabitant  of  the  pygmy  forest,  Pinus 
ponderosa  zone,  and  more  sparingly  in  the 
Picea-Ahics  forests.  Permanent  resident. 

Sitta  canadensis  Linnaeus.  Red-breasted 
Nuthatch.  Presnall  (1934  and  1936)  Bryce 
Canyon  and  Woodbury  and  Rus.sell  (1945) 
three,  Navajo  Mountain,  14  July  1936,  and 
Kaiparowits  Plateau,  11  Aug.  1937  {Picea- 
Ahies-Populus  zone  at  7000-10,400  ft);  saw 
one  to  six  birds  each  day  in  Picea-Abies  forest 
on  Navajo  Mountain,  6' and  13-18  July  1936. 
Woodbury  and  Russell  (1945)  believed  that 
the  species  probably  nests  in  Picea-Abies  for- 
est of  mountain  tops  and  canyon  heads  and 
spreads  to  other  habitats  following  nesting 
season.  Permanent  resident. 

Sitta  pygmaea  melanotis  van  Rossem.  Pyg- 
my Nuthatch.  Presnall  (1934)  Bryce  Canyon; 
Benson  (1935)  two,  Navajo  Mountain  and  ob- 
served daily;  Woodbury  and  Russell  (1945) 
three.  Navajo  Mountain,  11  Aug.  1935  and  10 
and  21  July  {Pinus  ponderosa,  8000-8500  ft); 
very  common  and  abundant  in  Pinus  ponde- 
rosa forest,  but  not  in  Picea-Abies  forest  on 
Navajo  Mountain,  1  July- 13  Aug.  1936; 
adults  feeding  well-grown  voung,  4  Jidv; 
Woodburv  and  Russell  foimd  it  very  abun- 
dant and  primarily  in  the  Pinus  ponderosa 
zone,  restricted  to  elevations  between 
7500-9820  ft.  Permanent  resident. 


Certhiidae  (Creepers) 

Certhia  familiaris  Linnaeus.  Brown 
Creeper.  Presnall  (1934)  Bryce  Rim;  Wood- 
burv and  Russell  (1945)  one,  Navajo  Moun- 
tain, 12  Aug.  1935  (Pinus  ponderosa,  9000  ft); 
seen  at  Navajo  Mountain,  18  Julv  1933  (dense 
forest)  and  6-13  July  to  13  Aug'.  1936  (Picea- 
Abies  forest).  Woodbury  and  Ru,sse!l  found  it 
there  during  June,  July,  and  Augu.st  in  the 
Picea-Abies  forest  with  winter  records  at 
lower  elevations.  Permanent  resident. 


326 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


Cinclidae  (Dippers) 

Cinclus  mexicanus  unicolor  Bonaparte. 
Dipper.  Benson  (1935)  Bridge  Canyon  at 
northwest  base  of  Navajo  Mountain;  Tanner 
(194()a)  Kaiparowits  Basin.  Woodbury  and 
Russell  (1945)  believed  that  the  dipper  did 
not  occur  on  Navajo  Mountain  and  that  it 
had  a  definitely  limited  distribution  in  Nav- 
ajo country;  they  gave  sight  records  for 
Bridge  Canyon,  6  July  1933;  several  records 
in  1934  and  one  record  7  Aug.  1935.  Per- 
manent resident. 


Troglodytidae  (Wrens) 

Troglodytes  aedon  parkmanii  Audubon. 
House  Wren.  Presnall  (1934)  Navajo  Trail; 
Benson  (1935)  one,  Navajo  Mountain,  14  July, 
and  seen  daily  in  fallen  Pinus  ponderosa, 
July;  Woodbury  and  Russell  (1945)  three, 
Navajo  Mountain,  14  July  1933,  16  Aug. 
1935,  and  9  July  1936  (common  in  thickets  of 
Arctostaphylos,  Ceanothiis,  and  Roso  under 
Finns  ponderosa,  8500-9000  ft);  Tanner 
(1940a)  Kaiparowits  Basin.  Summer  resident. 

Thryomanes  bewickii  eremophilus  Ober- 
holser.  Bewick's  Wren.  Woodbury  and  Rus- 
sell (1945)  one,  two  miles  north  of  Navajo 
Mountain  Trading  post,  27  July  1936  (pygmy 
conifers,  6500  ft);  they  reported  its  regular, 
though  not  abundant,  occurrence  in  pygmy 
conifers  on  the  lower  slopes  of  Navajo  Moun- 
tain from  22  June  to  9  Aug.,  1934-1938. 

Catherpes  mexicanus  conspersus  Ridgway. 
Canyon  Wren.  Presnall  (1934)  Bryce  Canyon; 
Woodbury  and  Russell  (1945)  six,  Lee's  Fer- 
ry, 26  Aug.  1909  (Nelson  and  Birdseye  trip) 
and  18  July  1936;  Rock  Creek /Colorado  Riv- 
er, 21  July  1936;  two  miles  north  of  Navajo 
Moimtain  Trading  Post,  2  Aug.  1936;  Navajo 
Mountain,  6  Aug.  1935  and  2  Aug.  1936  (in 
cliffs  of  some  size,  bearing  cracks  for  nesting, 
3100  ft  to  10,000  ft);  Behle  (1948)  Hidden 
Passage  Canyon,  Lee's  Ferry,  and  mouth  of 
Aztec  Canyon;  Behle  et  al.  (1958)  one,  con- 
fluence Calf  Creek/  Escalante  River,  1954; 
Hayward  et  al.  (1958)  Escalante  drainage; 
Behle  and  Higgins  (1959)  recorded  a  consid- 
erable number  of  individuals  in  the  Glen 
Canyon  and  Navajo  Mountain  areas,  21-23 
June  and  July  1933,  4-22  July  1936,  and  3-10 
Aug.  1938.  In  summer  of  1958  they  found  the 


species  abundant  along  the  full  length  of 
Glen  Canyon;  BYU  (1971)  Cockscomb  Ridge; 
29  Sept.:  BYU  (1972)  mouth  of  Escalante  Riv- 
er, 25  June  (hanging  garden);  BYU  (1973) 
Hackberry  Canyon,  8  June  (open  cliffs);  Paria 
River,  14  June  (cliffs);  Lees  Ferry  eight  seen 
7,  14,  and  15  June  (rocks);  Driftwood  Can- 
yon; Three  Garden;  and  Paria/Colorado  Riv- 
ers, 15  June  (rocks).  Permanent  resident. 

Salpinctes  ohsoletus  obsoletus  (Say).  Rock 
Wren.  Benson  (1935)  several  seen,  Navajo 
Mountain;  Woodbury  and  Russell  (1945) 
four,  Navajo  Mountain,  11  Aug.  1935  and  13 
July  1936;  above  Lee's  Ferry,  19  July  1936 
{Artemisia,  pygmy  forest,  Artemisia-Arctosta- 
phijlos  under  Pinus  ponderosa,  3100-10,000 
ft);  seen  at  Kaiparowits  Plateau  and  Beaver 
Creek /Navajo  Mountain  area,  8  Aug.  1936; 
Behle  (1948)  Lee's  Ferry;  Behle  et  al.  (1958) 
Escalante  River,  7  May  1954  and  10  miles 
south  of  Escalante,  8  May  1954;  Hayward  et 
al.  (1958)  Escalante  drainage;  Woodbury  (in 
Behle  and  Higgins  1959)  one  on  23  July  1937, 
and  20  in  August  1938  at  Rock  Creek,  Last 
Chance  Creek,  Lee's  Ferrv,  and  various  dis- 
tances along  the  Colorado  River  from  river 
mile  50  to  river  mile  18.  Russell  and  Thomp- 
son (1964)  Bryce  Canyon;  BYU  (1958)  one 
(male),  Paria  Basin,  9  June;  BYU  (1961)  one 
(male),  Paria,  20  May;  BYU  (1971)  Sites  2  and 
15;  Church  Wells  area,  28  Sept.;  Grand 
Bench,  9-10  Oct.;  Cottonwood  Wash;  Hack- 
berry  Canyon;  Lee's  Ferry;  Nipple  Spring; 
Smoky  Mountain;  Tibbet  Spring;  Ribbon 
Canyon;  and  Cockscomb  Ridge,  29  Sept.; 
BYU  (1973)  82  wrens  of  this  species  were 
seen  at  15  localities  in  June  and  July.  The 
earliest  spring  record  was  5  June  1973  with 
nesting  on  8  June.  The  latest  fall  record  was 
10  Oct.  1971.  The  Rock  Wren  is  nearly  obli- 
gate to  cliffs  and  rocks.  Seventeen  of  the  indi- 
viduals observed  were  in  rocks  or  cliffs  and 
one  in  Popuhis.  Summer  resident,  mav  winter 
sparingly  (Woodbury  and  Russell  1945)  in 
Kaiparowits  Basin. 

Mimidae  (Mockingbirds  and  Thrashers) 

Mimus  polyglottos  leucopterus  (Vigors). 
Mockingbird.  Benson  (1935)  Navajo  Moun- 
tain area;  Tanner  (1940a)  base  Kaiparowits 
Plateau,  June  1936;  Woodbury  and  Russell 
(1945)  one,  five  miles  south  Navajo  Mountain 


December  1980 


Atwood  et  al.:  Kaiparowits  Vertebrates 


327 


Trading  Post,  30  July  1965  {Artemisio,  6500 
ft);  few  seen  at  Lee's  Ferry  and  flats  south  of 
Navajo  Mountain;  Behle  et  al.  (1958)  one,  10 
miles  south  of  Escalante,  8  May  1954  (fairly 
common);  Higgins  saw  one  in  Atriplex  confer- 
tifolia  at  mouth  of  Escalante  Canvon,  19  julv 
1958;  Russell  and  Thompson  (1964)  Bryce 
Canyon;  B\TJ  (1971)  Last  Chance  Wash,  25 
Sept.  {Clmjsothammis);  BYU  (1972)  Site  27,  1 
Jime  (woodland);  Cedar  Mountain,  28  July 
{Epliedm-grdss);  Site  24,  28  June  (shrubs); 
BYU  (1973)  near  Site  1,  9  and  21  May  {Juni- 
perus);  Site  2,  2  seen  30  April  (shrubs)  and  3 
July  {Juniper us);  near  Site  3,  28  May  (wood- 
land) and  2  July  (grass);  Cottonwood  Wash,  5 
June  (riparian);  Nipple  Spring,  12  June  {Pop- 
ulus);  Tibbet  Spring,  5  and  10  June  (open 
brush);  Site  34,  4  seen,  10  July;  and  between 
Sites  6  and  8  on  Smoky  Mountain,  20  were 
seen  24  July  {Grayia-Coleogync).  Summer 
resident  between  30  April  and  25  September. 

Dumetella  carolinensis  (Linnaeus).  Gray 
Catbird.  Woodbury  and  Russell  (1945)  Lee's 
Ferry,  as  recorded  by  the  Nelson  and  Birds- 
eye  trip  of  1909;  Phillips  et  al.  (1964)  Lee's 
Ferry.  Summer  resident. 

Toxostoma  bendirei  (Coues).  Bendire's 
Thrasher.  Woodbury  (1939)  near  Escalante,  9 
May  1937;  Behle  et  al.  (1958)  give  the  local- 
ity of  the  aforementioned  specimen  as  five 
miles  south  of  Escalante;  however,  they  did 
not  observe  this  species  at  this  site,  8  May 
1954;  Hayward  (1967)  Wahweap  Creek,  20 
May  1956;  BYU  (1971)  Site  15,  25  Sept. 
{Chnjsothamnus).  Summer  resident. 

Eoroscoptes  montanus  (Townsend).  Sage 
Tlirasher.  Woodbury  and  Russell  (1945)  one, 
Lee's  Ferry,  26  Aug.  1909,  as  recorded  by 
Nelson  and  Birdseye's  trip  of  1909;  Behle  et 
al.  (1958)  two  miles  south  of  Escalante,  9 
May  1954,  and  common  in  Escalante  Valley, 
May  1954;  Russell  and  Thompson  (1964) 
Bryce  Canyon;  BYU  (1971)  Site  12,  15  Oct.; 
Site  14,  15' Oct.;  Tibbet  Spring,  13  Aug.;  Site 
15,  11  Oct.;  and  Grand  Bench,  9  Oct.  Un- 
common migrant. 

Turdidae  (Thnishes,  Solitaires,  and  Bluebirds) 

Turdus  migratorius  propinquus  Ridgway. 

Robin.  Presnall  (1933  and  1934)  Bryce  Can- 
yon; Benson  (1935)  Navajo  Mountain;  Wood- 
bury and  Russell  (1945)  three,  Navajo  Moun- 


tain, 21  July  and  16  Aug.  1936,  and 
Kaiparowits  Plateau,  11  Aug.  1937  {Pinus 
ponderosa,    7000-9000   ft);    Hayward   et   al. 

(1958)  Escalante  drainage;  Behle  and  Higgins 

(1959)  seen  at  many  places  in  Glen  Canvon; 
most  abundant  in  large  vegetation  at  mouths 
of  side  canyons;  BYU  (1972)  Tibbet  Canyon, 
1  Aug.  (Coleogyne);  BYU  (1973)  two,  Wah- 
weap Lodge,  31  Jan.  (grass);  Woodbury  and 
Russell  (1945)  consider  Navajo  Mountain  and 
the  Kaiparowits  Plateau  to  be  chief  breeding 
area  of  this  species  in  the  Navajo  country  of 
Utah.  They  list  a  juvenile  from  Navajo  Moun- 
tain taken  16  Aug.  1936.  Summer  resident,  a 
few  apparently  winter. 

Catharus  guttatus  auduboni  (Baird).  Her- 
mit Thrush.  Presnall  (1934)  and  Grater  (1947) 
Bryce  Canyon;  Benson  (1935)  two  specimens: 
Navajo  Mountain  (10,000  ft,  species  com- 
mon); Woodbury  and  Russell  (1945)  speci- 
men: Navajo  Mountain,  14  July  1936  (Picea- 
Abies  forest  10,000  ft);  common  (estimates 
100  pairs)  in  Picea-Abies  forest  6  and  11-18 
July  1936  (nesting  16  July;  absent  13  Aug.); 
BYU  (1971)  summit  of  Navajo  Mountain,  13 
Oct.  Summer  resident  and  transient  in  higher 
mountains  and  transient  in  lowland. 

Sialia  mexicana  bairdi  Ridgway.  Western 
Bluebird.  Presnall  (1933  and  1934)  and  Rus- 
sell and  Thompson  (1964)  Bryce  Canvon; 
Benson  (1935)  and  Woodbury  and  Russell 
(1945)  specimens  at  Navajo  Mountain,  16 
Aug.  1935  and  4  July  1936  (Pinus  ponderosa); 
Behle  et  al.  (1958)  one,  confluence  of  Calf 
Creek/Escalante  River,  1954;  BYU  (1971) 
one  (male),  Cottonwood  Wash,  7  Nov.;  BYU 
(1972)  Site  23,  25  .seen  3  Mar.  (mixed  shrubs); 
and  11  miles  east  Grosvenor  Arch,  one  seen 
(not  identified  to  species),  17  June  (wood- 
land). Permanent  resident. 

Sialia  currucoides  (Bechstein).  Mountain 
Bluebird.  Presnall  (1933  and  1934)  and  Grat- 
er (1947)  Bryce  Canyon  (also  late  summer  to 
late  winter  at  Cedar  Breaks);  Tanner  (1940a) 
Escalante  drainage;  Behle  et  al.  (1958)  Paria, 
6  March  1946;  Hayward  et  al.  (1958)  Esca- 
lante drainage;  BYU  (1971)  one,  Paria 
Plateau,  29  Sept.;  BYU  (1972)  one,  one  mile 
south  Glen  Canyon  City,  7  Feb.;  Sites  3  and 
6;  Site  23,  three  seen  on  24  Feb.  (Coleogyne); 
Four  Mile  Bench,  25  seen  26  Oct.  (wood- 
land); BYU  (1973)  one  (ad.  male),  eight  miles 
east  Glen  Canyon  Citv,  2  May;  Warm  Creek 


328 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


Bay,  two  seen  10  June;  and  Four  Mile  Bench, 
13  June  (near  a  spring).  Simnmer  resident,  a 
few  may  winter. 

Myadestes  townsendi  townsendi  (Audu- 
bon). Townsend's  Solitaire.  Presnall  (1933 
and  1934)  and  Grater  (1947)  Bryce  Canyon; 
Behle  et  al.  (1958)  five  miles  west  of  Esca- 
lante,  20  Sept.  1935.  Transient. 


Sylviidae  (Old  World  Warblers, 
Gnatcatchers,  and  Kinglets) 

Polioptila  caerulea  amoenissinia  Grinnell. 

Blue-gray  Gnatcatcher.  Presnall  (1934)  and 
Russell  and  Thompson  (1964)  Bryce  Canyon; 
Benson  (1935)  Navajo  Mountain;  Tanner 
(1940a)  Kaiparowits  Basin;  Woodbury  and 
Russell  (1945)  five,  Navajo  Mountain  Trading 
Post,  25  July  1936;  Rock  Creek/Colorado 
River,  26  July  1937;  Kaiparowits  Plateau, 
30-31  July  and  10  Aug.  {Artemisia,  wood- 
land, Quercus,  Amelanchier,  3300-7000  ft); 
Hayward  et  al.  (1958)  Escalante  drainage; 
BYU  (1971)  Last  Chance,  25  Sept.  (Tamarix); 
BYU  (1973)  Sites  15  and  28  {Junipems-grass)- 
Lee's  Ferry,  14-15  June  {Tamarix  and  Salix); 
Cottonwood  Wash/Paria  River,  6  July;  Rib- 
bon Canyon;  Paria /Colorado  Rivers,  7  and 
14  June  (Tamarix);  Tibbet  Spring,  two  seen  5 
June  (Popuhis);  and  Wahweap  Creek,  30  Apr. 
and  7  Jime.  Summer  resident  (30  April  to  25 
Sept.). 

Regulus  satrapa  Lichtenstein.  Golden- 
crowned  Kinglet.  Russell  and  Thompson 
(1964)  Bryce  Canyon;  BYU  (1971)  north  of 
Church  Wells;  one,  summit  of  Navajo  Moun- 
tain, 13  Oct.  Permanent  resident. 

Regulus  calendula  (Linnaeus).  Ruby- 
crowned  Kinglet.  Presnall  (1934)  and  Russell 
and  Thompson  (1964)  Bryce  Canyon;  Behle 
(1948)  Lee's  Ferry;  Woodbury  and  Russell 
(1945)  two,  Navajo  Mountain,  14  Aug.  1935 
and  15  July  1936  {Picea-Ahies,  Pinus  ponde- 
rosa  forest,  9000-10,000  ft);  breeds  on  top  of 
Navajo  Mountain.  Permanent  resident. 


Bombycillidae  (Waxwing) 

Bombycilki  cedrorum  Vieillot.  Cedar  Wax- 
wing.  Behle  et  al.  (1958)  Bryce  Canyon.  Un- 
common transient,  some  may  breed  (Behle  et 

al.  (1958). 


Ptilogonatidae  (Silky  Flycatchers) 

Phainopepla    nitens   lepida    Van   Tyne. 

Phainopepla.  BYU  (1973)  near  Tibbet  Spring, 
5  June;  seen  by  Robert  Whitmore  in  open 
brush.  This  species  was  previously  known  in 
southern  Utah  only  from  lower  Santa  Clara 
Valley,  Washington  County,  to  Kanab,  Kane 
County  (Hayward  et  al.  (1976).  Summer  resi- 
dent. 


Laniidae  (Shrikes) 

Lanius  excubitor  Linnaeus.  Northern 
Shrike.  BYU  (1973)  Site  3,  3  Aug.  Rare  tran- 
sient. 

Lanius    ludovicianus    Linnaeus.     Log- 
gerhead Shrike.  Benson  (1935)  Navajo  Moun- 
tain area;  Tanner  (1940a)  Escalante  drainage; 
Woodbury  and  Russell  (1945)  one,  Kaiparo- 
wits     Plateau,      12      Aug.      1937;      Rock 
Creek/Colorado  River,  26  July  1937;  Navajo 
Mountain  Trading  Post,  31  July  1936  {Popii- 
his,  Artemisia,  and  Amelanchier,  4000-7000 
ft);  also  in  Artemisia  flats,  Sarcobatus,  Xan- 
thocephalum,    Atriplex   confertifolia.    Yucca, 
and  Ephedra  types  of  cover  (Woodburv  and 
Russell   1945);  Behle  et  al.   (1958)   10 'miles 
southeast  of  Escalante,  8  May  1954;  Russell 
and  Thompson  (1964)  Bryce  Canyon;   BYU 
(1971)  Sites  1  and  2;  specimens  at  two  miles 
southwest  of  Site  3,  28  Aug.  and  two  miles 
west  of  Site  3,  20  Aug.;  Sites  6  and  10;  Cot- 
tonwood Wash,   7  Nov.;   Four  Mile  Bench; 
Glen   Canyon   City;    Pump   Canyon   Spring; 
Tibbet  Spring;  one.  Warm  Creek,  two  miles 
below  road  to  Escalante,  6  Dec;  BYU  (1973) 
one   (ad.   female),   two   mile  southwest  Glen 
Canyon  City,  29  Apr.  It  is  a  permanent  resi- 
dent of  the  area  according  to  Woodburv  and 
Russell  (1945).  We  saw  155  individuals  be- 
tween  1971   and   1973.   Sightings  (73)  were 
distributed  throughout  the  year,  except  for 
Feb.  and  Mar.  when  none  were  seen.  Wood- 
bury and  Russell  (1945)  believed  the  species 
to  be  more  plentiful   in   late  summer  after 
emergence  of  young.   This  agrees  with  our 
findings.  We  found  the  species  very  sparse  in 
Nov.  (2),  Dec.  (1),  and  Jan.  (1);  absent  in  Feb. 
and  Mar.;   and  somewhat  more  frequent  in 
Apr.  (10),  May  (3),  and  June  (3).  It  was  com- 
mon in  July  (28),  reaching  a  peak  in  Aug. 
(56),  and  declining  in  numbers  in  Sept.  (27) 


i 


December  1980 


Atwood  et  al.:  Kaiparowits  Vertebrates 


329 


and  Oct.  (16).  It  was  most  frequently  encoun- 
tered in  woodlands,  Vanclevea-grdss,  Coleog- 
ijne,  Ephedra-grsiss,  and  Atriplex-grass  in  this 
order  of  abundance.  Common  permanent 
resident. 


Stumidae  (Starlings) 

Sturnus  vulgaris  vulgaris  Linnaeus.  Star- 
ling. Russell  and  Thompson  (1964)  Bryce 
Canyon;  BYU  (1971)  two  (males),  Glen  Can- 
yon City,  2  Dec;  two  seen  8  Dec;  BYU 
(1972)  Glen  Canyon  City,  two  seen  5  Mar.; 
BYU  (1973)  Paria  River /U.S.  89,  6  June.  The 
starling  is  a  newcomer  to  the  area.  Behle  et 
al.  (1958)  reported  sightings  of  this  species  in 
Jan.  1941  and  Sept.  1948  for  Kane  County, 
but  the  observations  by  Russell  and  Thomp- 
son (1964)  appear  to  be  the  first  published 
observations  of  this  species  in  the  area  of  the 
current  report.  Earlier  observers  (Behle  1960, 
Woodbury  and  Russell  1945,  Hayward  1976) 
did  not  report  the  species.  The  species  is 
known  in  Utah  to  first  occur  as  a  migrant  or 
winter  visitant,  followed  later  by  permanent 
residency  (Behle  and  Perry  1975).  A  similar 
pattern  of  occurrence  in  the  Kaiparowits 
area  suggests  that  the  species  is  establishing 
itself  here  as  a  breeder  also.  Permanent  resi- 
dent. 


Vireonidae  (Vireos) 

Vireo  bellii  Audubon.  Bell's  Vireo.  BYU 
(1973)  Lee's  Ferry,  7  June.  Until  this  vireo 
was  seen  at  Lee's  Ferry  by  Robert  Whitmore 
it  was  known  in  Utah  only  from  southwestern 
Utah  (Hayward  et  al.  (1976),  where  it  is  an 
imcommon  summer  resident  (Behle  and  Perry 
1975).  Sparse  summer  resident. 

Vireo  vicinior  Coues.  Gray  Vireo.  Behle  et 
al.  (1958)  specimen:  confluence  (Calf 
Creek /Escalante  River,  1954;  BYU  (1973) 
Site  28,  13  June  by  Lloyd  Pack  in  Juniperiis- 
grass  association.  Behle  et  al.  (1958)  report 
this  species  as  an  uncommon  resident  of 
woodlands  of  the  Kanab  area  and  adjacent 
high  plateaus.  Although  Behle  and  Perry 
(1975)  consider  it  a  common  summer  resident 
of  southern  Utah,  in  the  area  of  the  Kaiparo- 
wits Plateau  it  seems  to  be  rather  sparse,  as 
suggested  by  only  one  sighting  in  2V2  years. 
Sparse  summer  resident. 


Vireo  solitarius  (Wilson).  Solitary  Vireo. 
Presnall  (1934)  and  Russell  and  Thompson 
(1964)  Bryce  Canyon;  Behle  et  al.  (1958)  con- 
sider V.  s.  plumbeus  to  be  a  fairly  common 
summer  resident  in  the  lower  reaches  of 
Bryce  Canyon.  They  observed  this  taxon  in 
canyons  vegetated  with  Quercus  gainheUii, 
Acer  negundo,  and  Populus  frernontii;  Wood- 
bury and  Russell  (1945)  three  (all  V.  s.  cas- 
sinii),  Navajo  Mountain,  7  Aug.  1935,  11 
Aug.  1936,  and  Lee's  Ferry,  25  Aug.  1909,  as 
recorded  by  the  Nelson  and  Birdseye  trip 
{Populus  tre7nuloides  and  riparian  forest, 
Quercus  and  Salix,  3100-9500  ft);  it  is  prob- 
able that  V.  s.  plwnheus  breeds  in  the  Kai- 
parowits Plateau  and  Navajo  Moimtain  areas, 
although  none  have  been  seen.  Vireo  s.  cas- 
sinii,  on  the  other  hand,  is  a  migrant.  Wood- 
bury and  Russell  (1945)  considered  V.  s. 
plumbeus  as  a  siunmer  resident  of  Pinus  pon- 
derosa,  Quercus,  and  woodlands  of  the  mesa 
tops  and  slopes  of  the  Navajo  country.  Sum- 
mer resident  and  transient. 

Vireo  gilvus  (Vieillot).  Warbling  Vireo. 
Benson  (1935)  and  Woodbury  and  Russell 
(1945)  Navajo  Mountain  area,  17  Aug.  1935 
and  4  Aug.  1936,  and  Kaiparowits  Plateau,  2 
Aug.  1937;  Tanner  (1940a)  Kaiparowits  Ba- 
sin; Behle  et  al.  (1958)  Bryce  Canyon.  Al- 
though specimens  from  Garfield  County, 
Utah,  have  been  assigned  to  V.  g.  swainsonii, 
Worthen  (1968)  has  questioned  the  presence 
of  the  race  swainsonii  in  Utah  (also  see  Hay- 
ward et  al.  1976);  it  is  probablv  a  migrant 
(Behle  and  Perry  1975).  The  breeding  popu- 
lation represents  the  race  leucopolius  (Behle 
and  Perry  1975).  Woodbury  and  Russell 
(1945)  found  this  latter  race  in  stands  of 
Qiwrcus,  streamside  Populus  fremontii  and 
Salix,  and  Populus  tremuloides.  On  Navajo 
Mountain  they  encountered  it  in  Populus 
tremuloides  at  9000-10,500  ft.  They  also 
found  breeding  adults  and  two  young,  16  to 
18  July  1936.  We  did  not  encounter  it.  Sum- 
mer resident. 


Parulidae  (Wood  Warblers) 

Vermivora  celata  (Say).  Orange-crowned 
Warbler.  Behle  and  Higgins  (1959)  Hole-in- 
the-Rock/Colorado  River,  20  Oct.  1958  (ri- 
parian, 3266  ft);  BYU  (1971)  Site  15,  9  Oct. 
Uncommon  migrant. 


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Great  Basin  Naturalist 


Vol.  40,  No.  4 


Vermivora  ruficapilla  ridgwayi  van  Ros- 
sem.  Nashville  Warbler.  Woodbury  and  Rus- 
sel  (1945)  one,  Navajo  Mountain,  11  Aug. 
1935  {Pinus  pundewsa,  8500  ft).  Transient. 

Vermivora  virginiae  (Baird).  Virginia's 
Warbler.  Presnall  (1934)  Bryce  Canyon; 
Woodbury  and  Russell  (1945)  Navajo  Moun- 
tain, 17  July  1936  {Populus  tremuloides)  and 
Kaiparowits  Plateau,  9  Aug.  1937  (pygmy 
forest,  7000-10,000  ft);  Behle  et  al.  (1958) 
confluence  Calf  Creek/Escalante  River, 
1954;  Russell  and  Thompson  (1964)  Bryce 
Canyon;  BYU  (1971)  Site  4;  Vi  mile  south 
Paria  River  Bridge,  3  Sept.  According  to 
Woodbury  and  Russell  (1945),  this  warbler  is 
a  breeder  of  the  deciduous  bnish  and  tree 
zones  between  6000-10,000  ft.  Summer  resi- 
dent. 

Vermivora  luciae  (Cooper).  Lucy's  War- 
bler. Woodbury  (1939)  and  Woodbury  and 
Russell  (1945)  two,  river  mile  41,8  Aug.  1938 
{Salix  and  Rhus,  3180  ft);  Colorado  River  side 
canyon,  13  July  1936  {Salix  thickets,  3200  ft); 
seen  at  mouth  of  Paria  River,  1938;  V2  mile 
below  Lee's  Ferry,  30  July  to  11  Aug.  1938; 
yoimg  out  of  nest,  banks  Colorado  River,  two 
miles  below  San  Juan  River,  17  July  1931; 
and  single  birds  on  the  Colorado  at  river  mile 
41,  8  Aug.  and  one  mile  above  Lee's  Ferry, 
10  Aug.;  Woodbury  and  Russell  (1945)  be- 
lieve this  species  to  be  a  sparse  inhabitant  of 
Salix  and  brush  thickets  along  the  Colorado 
and  San  Juan  Rivers;  Behle  (1948)  Lee's  Fer- 
ry; Behle  et  al.  (1958)  one,  junction  Calf 
Creek/Escalante  River,  4  July  1938;  BYU 
(1973)  Lee's  Ferry  and  one  mile  south  of 
Paria,  Utah,  7  June  (Salix  and  Tamarix).  Sum- 
mer resident. 

Dendroica  petechia  morcomi  Coale.  Yel- 
low Warbler.  Woodbury  and  Russell  (1945) 
four,  three  miles  below  confluence  San 
Juan/Colorado  Rivers,  3  Aug.  1938;  six  at 
Rock  Creek/Colorado  River,  20-23  July 
1937;  one  from  Kaiparowits  Plateau,  4  Aug. 
1937;  one  from  Navajo  Mountain,  13  Aug. 
1935;  one  from  Lee's  Ferry,25  Aug.  1909; 
two  along  Colorado  River,  11  July  1936;  one 
and  nest  at  river  mile  63,  5  Aug.  1938;  and 
two  at  river  mile  50,  6  Aug.;  (Quercus,  Salix, 
riparian  thickets,  Artemisia,  Populus  tremu- 
loides, 3120-7000  ft;  nested  below 
6500-7000  ft  in  riparian  Salix,  brush  or  Popu- 
lus); Behle  and  Higgins  (1959)  give  the  addi- 


tional observational  records  from  Wood- 
bury's notes  as  follows:  "many  were  seen  at 
several  points,  six  at  two,  and  two  at  another, 
plus  three  family  groups  at  one  and  six  at  an- 
other, from  river  mile  75  to  the  mouth  of 
Paria  Creek;  common  breeder  1  July  to  9 
Aug.  1958  in  riparian  vegetation  along  Glen 
Canyon";  Hayward  et  al.  (1958)  Escalante 
drainage;  Russell  and  Thompson  (1964)  Bryce 
Canyon;  BYU  (1972)  Reflection  Canyon,  4 
July  and  confluence  San  Juan/Colorado  Riv- 
ers, 24  June  (Fop«/MS-riparian);  BYU  (1973) 
Lee's  Ferry,  three  seen  7  June  (Tamarix)  and 
one  seen  15  June  (Tamarix);  one  mile  south  of 
Paria  near  Paria  River,  7  June  (dense  Salix); 
and  Paria  River/Colorado  River,  7  June 
(Tamarix-Salix).  Summer  resident. 

Dendroica  coronata  auduboni  (Town- 
send).  Yellow-rumped  (Audubon's)  Warbler. 
Presnall  (1933)  Bryce  Canyon;  Benson  (1935) 
Navajo  Mountain;  Woodbury  and  Russell 
(1945)  three,  Navajo  Mountain,  11  Aug. 
1935;  nests  in  Pinus  ponderosa,  2  July  1936 
(8500  ft;  breeds  8000-10,000  ft,  spring  migra- 
tion dates  in  general  area  of  Navajo  Moun- 
tain are  13  Apr. -25  Mav;  and  fall,  20 
Aug.-14  Oct.);  Behle  (1948)  Lee's  Ferry; 
Behle  et  al.  (1958)  10  miles  west  of  Escalante, 
7  May  1941;  BYU  (1971)  one,  summit  Navajo 
Mountain,  13  Oct.;  and  25  at  Site  15,  9-10 
Oct.;  BYU  (1973)  Crosby  Canyon  Bay,  two 
seen  courting,  28  Apr.  (Tamarix);  and  two 
seen  5V2  miles  south  of  Tibbet  Springs,  1 
May.  The  intermountain  race  D.  auduboni 
memorabilis  is  now  in  synonymy  with  D.  a. 
auduboni  (Townsend)  and  the  species  has 
been  placed  in  Dendroica  coronata.  Summer 
resident  and  fall  and  spring  transient. 

Dendroica  nigrescens  (Townsend).  Black- 
throated  Gray  Warbler.  Presnall  (1934)  Bryce 
Canyon;  Benson  (1935)  Navajo  Mountain; 
Woodbury  and  Russell  (1945)  six  specimens: 
Kaiparowits  Plateau,  31  Julv  and  4,  5,  and  9 
Aug.  1937;  Navajo  Mountain,  14  Aug.  1935 
and  2  Aug.  1936  (pygmy  forest,  6000-9000 
ft;  a  nearly  obligate  breeder  in  pygmy  forest, 
migrating  through  most  other  vegetational 
types);  BYU  (1971)  Site  12  and  specimen 
(male):  V2  mile  north  Nipple  Spring,  16  Oct. 
Summer  resident  and  transient  spring  and 
fall. 

Dendroica  townsendi  (Townsend).  Towns- 
end's  Warbler.   Presnall  (1933  and   1934) 


December  1980 


Atwood  et  al.:  Kaiparowits  Vertebrates 


331 


Bryce  Canyon;  Woodbury  (1939)  two,  Nav- 
ajo Mountain,  10  Aug.  1936  (also  seen  13 
Aug.  1935);  Kaiparowits  Plateau,  12  Aug. 
1927  (pvgmv  forest,  Piniis  ponderosa, 
6000-700'(Ht).'  Transient. 

Dendroica  occidentalis  (Townsend).  Her- 
mit Warbler.  Woodbury  and  Russell  (1945) 
two,  Navajo  Mountain,  11  Aug.  1935  and  13 
Aug.  1936  [Pinits  ponderom,  8500-10,000  ft). 
In  1935  this  normally  rare  species  was  a  com- 
mon member  of  the  band  of  warblers  on 
Navajo  Mountain  during  Aug.  (Woodbury 
and  Russell  1945).  Transient. 

Dendroica  graciae  graciae  Baird.  Grace's 
Warbler.  Presnall  (1934)  Bryce  Canyon; 
Woodbury  and  Russell  (1945)  seen  at  Navajo 
Moimtain,  15-16  June  1938,  and  one  collect- 
ed but  discarded,  15  June;  Benson  (1935)  saw 
and  heard  them  in  Pinus  ponderosa,  but  not 
numerous  (Woodbury  and  Russell  1945). 
Summer  resident. 

Oporornis  tolmiei  (Townsend).  MacGil- 
livray's  Warbler.  Woodbury  and  Russell 
(1945)  one,  Navajo  Mountain,  O.  t.  tolmiei; 
Behle  (1960)  5  Aug.  1936  {Salix  in  canyon 
bottoms);  BYU  (1971)  Glen  Canyon  City; 
Kaibab  Wash,  29  Sept.  Summer  resident. 

Geothlypis  trichas  occidentalis  Brewster. 
Common  Yellowthroat.  Woodbury  and  Rus- 
sell (1945)  two  at  Rock  Creek/Colorado  Riv- 
er, 20-26  July  1927  (large  family  flocks  also 
seen  in  streamside  thickets);  one  near  Bridge 
Canyon,  12  July  1936;  one  from  river  mile 
60,  6  Aug.  1938;  and  one  from  river  mile  25, 
9  Aug.  1938  (including  nest  );  nest  observed 
at  river  mile  63,  5  Aug.  1938  (Salix);  family 
flocks  mouth  of  Rock  Creek,  20-26  July 
1937;  usually  found  in  Salix  reeds,  canes, 
brush,  or  Ti/phits  in  canyons  or  valleys  below 
5500-6000  ft;  Behle  et  al.  (1958)  confluence 
Calf  Creek /Escalante  River,  2  May  1954; 
Behle  and  Higgins  (1959)  165  individuals 
seen,  plus  many  others  including  12  families 
(out  of  nest)  at  various  points  along  the  Colo- 
rado River  by  Woodbury  and  others  during 
the  Rainbow  Bridge-Monument  Valley  expe- 
dition. Higgins  (Behle  and  Higgins  1959)  saw 
both  adults  and  immatures  in  the  Salix  fringe 
at  Lee's  Ferry,  8  Aug.  1958,  and  found  the 
species  nesting  abundantly  during  the  sum- 
mer of  1958  (Behle  1960).  Woodbury  and 
Russell  (1945)  noted  that  the  habitats  of  the 
Yellow   Warbler   and   Yellowthroat   overlap. 


but  the  warbler  extended  landward  into  Pop- 
ulus  fremontii,  whereas  the  yellowthroat  ex- 
tended toward  the  moist  areas  containing 
Typhus  and  Jiincus.  It  is  interesting  that  none 
were  seen  during  this  study.  It  is  not  known  if 
this  is  because  of  the  creation  of  Lake  Powell 
or  due  to  insufficient  observations. 

Icteria  virens  (Linnaeus).  Yellow-breasted 
Chat.  Woodbury  and  Russell  (1945)  three, 
Colorado  River,  11  July  1936  and  river  miles 
41  and  50,  7  and  8  Aug.  1938  (stream  bank 
thickets,  Salix,  and  Rhii.s);  Behle  et  al.  (1958) 
confluence  of  Calf  Creek /Escalante.  1954, 
i.e.,  /.  f.  auricollis  in  dense  vegetation.  Behle 
and  Higgins  (1959)  give  the  following  addi- 
tional records  from  the  field  notes  of  Wood- 
bury and  Russell  for  1938  for  the  Colorado 
River  from  the  Rainbow  Bridge-Monument 
Valley  expedition:  80  were  counted  from  riv- 
er mile  50  to  V2  mile  below  Lee's  Ferry  near 
the  mouth  of  Paria  Creek,  between  7  and  1 1 
Aug.  Higgins  found  them  to  be  abundant 
breeders  along  the  length  of  Glen  Canyon. 
He  saw  them  every  day  of  the  trip  in  the 
densest  streamside  vegetation.  Russell  and 
Thompson  (1964)  Bryce  Canyon;  BYU  (1973) 
Lee's  Ferry,  a  total  of  12  were  seen  6,  7,  and 
14-15  June;  and  along  the  Paria  River  near 
Paria,  Utah,  three  were  seen  7  and  14  June. 
We  saw  them  in  the  Tamarix  and  Salix  along 
the  rivers.  Summer  resident. 

Wilsonia  pusilla  pilcolata  (Pallas).  Wil- 
son's Warbler.  \Voodbur\  and  Russell  (1945) 
Navajo  Mountain,  one  or  two  seen  daily  in 
Populus  tremuloides,  9-16  Aug.  1935,  and 
one,  1  Sept.  1934;  Russell  and  Thompson 
(1964)  Bryce  Canyon;  BYU  (1973)  lower 
Wahweap  Creek,  30  Apr.  Woodbury  and 
Russell  give  the  migration  date  through  the 
area  as  4-26  May  in  the  spring  and  9 
Aug. -26  Oct.  in  the  fall.  Transient. 


Ploceidae  (Weaver  Finches) 


Passer  domesticus  (Linnaeus).  House  Spar- 
row. Woodbury  and  Russell  (1945)  Lee's  Fer- 
ry/Paria  River,  several  birds  seen  10  Aug. 
1938;  Behle  et  al.  (1958)  Escalante;  BYU 
(1971)  Glen  Canyon  City,  .30  seen  2  Dec.  and 
5  seen  8  Dec.  Permanent  resident. 


332 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


Icteridae  (Meadowlarks,  Blackbirds, 
and  Orioles) 

Sturnella    neglecta    neglect  a    Audubon. 

Western  Meadowlark.  Presnall  (1934)  Bryce 
Canyon;  Behle  et  al.  (1958)  near  Escalante  in 
cultivated  fields;  Hay  ward  et  al.  (1958)  Esca- 
lante drainage;  BYU  (1971)  Site  3  (no  date, 
semiarid  grasslands);  and  Paria  Plateau,  29 
Sept.  Uncommon  permanent  resident. 

Xanthocephalus  xanthocephalus  (Bona- 
parte). Yellow-headed  Blackbird.  Woodbury 
and  Russell  (1945)  Lee's  Ferry  as  recorded  by 
Nelson  and  Birdseye's  trip,  23-26  Aug.  1909; 
Russell  and  Thompson  (1964)  Bryce  Canyon; 
Behle  and  Higgins  (1959)  river  mile  25,  three 
females  seen,  6  Aug.  1958;  BYU  (1972)  Warm 
Creek  Bay,  one  pair  seen  on  shore  of  Lake 
Powell,  3  May.  Migration  through  the  area  is 
given  by  Woodbury  and  Russell  (1945)  as 
April-May  passing  north  and  12  July-11  Oct. 
passing  south.  Transient. 

Agelaius  phoenicetis  (Linnaeus).  Red- 
winged  Blackbird.  Behle  (1948)  Aztec  Creek, 
two  seen;  Russell  and  Thompson  (1964) 
Bryce  Canyon;  Hay  ward  et  al.  (1958)  Esca- 
lante drainage;  BYU  (1973)  one  mile  above 
Lee's  Ferry  (flying),  15  June.  Transient. 

Icterus  parisorum  Bonaparte.  Scott  s  Ori- 
ole. Benson  (1935)  south  of  Navajo  Mountain; 
BYU  (1972)  Site  4,  25  June  {Coleogijne)-  BYU 
(1973)  three  miles  northwest  of  Site  3,  29 
Apr.  (Pintis  edulis);  two  miles  north  Site  3,  28 
May  (woodland);  Site  6,  3  May  {Junipenis); 
one  mile  east  of  Site  7,  4  July;  and  Site  28,  13 
June  {Juniperus-grsLSs).  All  1973  observations 
except  the  one  July,  were  in  Juniperus-grass 
or  pygmy  forest.  No  vegetation  was  given  for 
the  July  sighting.  Summer  resident. 

Icterus  galhula  bullockii  (Swainson). 
Northern  Oriole.  Benson  (1935)  Beaver 
Creek/Navajo  Mountain;  Woodbury  and 
Russell  (1945)  two,  north  foot  and  southwest 
foot  of  Navajo  Mountain,  6  Aug.  1935  and  1 1 
Aug.  1936  {Populus  fremontii  near  woodland 
and  Quercus  community,  5500-6000  ft); 
Behle  et  al.  (1958)  confluence  Calf  Creek/ 
Escalante  River,  1954;  Hay  ward  et  al.  (1958) 
Escalante  drainage;  Behle  and  Higgins  (1959) 
field  notes  of  Rainbow  Bridge-Monument 
Valley  expedition  of  1938:  seven  were  seen 
along  the  Colorado  River  from  river  mile  75 
to  '/2  mile  below  Lee's  Ferry,  3-11  Aug.;  Rus- 


sell and  Thompson  (1964)  Bryce  Canyon; 
BYU  (1961)  one  male  and  three  females, 
Paria,  20  May;  BYU  (1973)  Paria  River/U.S. 
89,  two  seen  6  June;  and  Cottonwood  Can- 
yon, 8  June;  Hackberry  Canyon,  8  June  (male 
displaying);  and  Lee's  Ferry,  adult  and  fledg- 
ling seen  in  Tamarix  and  Salix  15  June.  Ex- 
cepting the  15  June  sighting,  all  others  were 
in  stands  of  Populus  fremontii.  Summer  resi- 
dent. 

Euphagus  cyanocephalus  (Wagler). 
Brewer's  Blackbird.  Presnall  (1934)  Bryce 
Canyon;  Tanner  (1940a)  Escalante  drainage; 
Behle  et  al.  (1958)  confluence  of  Calf 
Creek/Escalante    River;    Hayward    et    al. 

(1958)  Escalante  drainage;  Behle  and  Higgins 

(1959)  5  were  seen  at  mouth  of  Kane  Creek, 
river  mile  41,  1  Aug.  1958;  BYU  (1973)  Lee's 
Ferry,  three  sightings  totaled  53  birds,  7  June 
(Tamarix);  Glen  Canyon  City,  50  seen  28 
Apr.  (grasslands);  and  Cottonwood 
Wash/Brigham  Plains  road,  2  seen  8  June  (ri- 
parian vegetation).  Transient,  a  few  may  nest 
and  a  few  may  winter. 

Molothrus  ater  obscurus  (Gmelin).  Brown- 
headed  Cowbird.  Woodbury  and  Russell 
(1945)  specimen:  Colorado  River  at  Rock 
Creek,  22  July  1937  (riparian  thicket  of 
Tamarix  and  Salix,  3100-3300  ft)  and  one  egg 
found  in  the  nest  of  Willow  Flycatcher  near 
Lee's  Ferry,  11  Aug.  1938;  Behle  et  al.  (1958) 
Bryce  Canyon;  Behle  and  Higgins  (1959)  Col- 
orado River/Rock  Creek,  28  July  1958;  BYU 
(1972)  Site  3  on  Cedar  Mountain,  25  July 
(Ephedra-grsiss);  BYU  (1973)  Cottonwood 
Wash,  5  June  (Popuhis);  and  Lee's  Ferry,  one 
seen  7  June  (Tamarix)  and  two  seen  15  June 
(brush).  Summer  resident. 

Thraupidae  (Tanagers) 

Piranga  ludoviciana  (Wilson).  Western 
Tanager.  Benson  (1935)  one,  War  God  Spring 
on  Navajo  Mountain,  14  June  1933;  Wood- 
bury and  Russell  (1945)  six,  two  at  Navajo 
Mountain  (War  God  Spring  and  Beaver 
Creek  Canyon),  13  July,  7  Aug.  1936;  one  at 
Navajo  Mountain  Trading  Post,  26  July  [no 
vear];  one  at  Colorado  River/Rock  Creek,  27 
July  1937;  one  at  Rock  Creek,  28  July  1937; 
and  one  at  Kaiparowits  Plateau,  6  Aug.  1937; 
usually  found  in  Piniis  pondcrosa,  Populus 
fremontii  and  other  riparian  trees,  chaparral. 


December  1980 


At  WOOD  ET  AL.:  Kaiparowits  Vertebrates 


333 


and  Quercus.  3300-8500  ft;  Behle  and  Hig- 
gins  (1959)  Aztec  Canyon  at  river  mile  68.5, 
7  July  1958;  mouth  of  Kane  Creek  at  river 
mile  41,  1  Aug.;  and  one  mile  upstream  from 
mouth  of  Escalante  River,  several  seen  of 
both  sexes,  19  July  1958.  Woodburv  and  Rus- 
sell (1945)  saw  this  species  during  the  breed- 
ing season  in  stands  of  Pinus  ponderosa  and 
Picea-Abies  on  Navajo  Mountain;  Russell  and 
Thompson  (1964)  Brvce  Can  von;  BYU  (1971) 
Site  16  in  Tamarix,  \H  Aug.;' BYU  (1972)  one 
mile  south  of  Site  1,  8  June.  Summer  resident. 


Fringillidae  (Grosbeaks,  Finches, 
Sparrows,  and  Buntings) 

Pheucticus  melanocephalns  melanoce- 
phalus  (Swainson),  Black-headed  Grosbeak. 
Benson  (1935)  Navajo  Mountain;  Woodbury 
and  Russell  (1945)  two,  Navajo  Mountain,  15 
Aug.  1935  and  4  Aug.  1936  {Salix  and  Picea- 
Abies  forest,  6500-10,300  ft);  they  give  post- 
breeding  dates  23  July  to  15  Aug.  1934-38  at 
Navajo  Mountain  Trading  Post,  Beaver 
Creek  Canyon,  and  Lee's  Ferry.  They  in- 
dicated that  it  breeds  at  canyon  heads  and 
moimtains  mostly  below  the  coniferous  for- 
est, among  Querciis,  but  mostly  above  the 
pygmy  forest;  it  may  descend  into  the 
streamside  fringes  of  lower  elevations.  Wood- 
bury and  Russell  (1945)  record  extreme  oc- 
currence dates  as  8  May  and  22  September; 
Hayward  et  al.  (1958)  Calf  Creek  area;  Behle 
and  Higgins  (1959)  mouth  of  Aztec  Creek  at 
river  mile  68.5  in  stands  of  Quercus,  26  July 
1958;  river  mile  46  in  Salix,  30  July  1958; 
and  from  Woodbury  and  Russell's  field  notes 
(Behle  and  Higgins  1959)  between  river  miles 
63  and  50,  5  Aug.  1938;  Russell  and  Thomp- 
son (1964)  Bryce  Canyon;  BYU  (1971)  Site  4 
and  V2  mile  south  of  Paria  Bridge,  30  Sept. 
Summer  resident. 

Guiraca  caerulea  interfusa  (Dwight  & 
Griscom).  Blue  Grosbeak.  Presnall  (1934) 
Bryce  Canyon;  Tanner  (1940a)  Kaiparowits 
Basin;  Woodbury  and  Russell  (1945)  2  at  riv- 
er mile  13,  9-10  Aug.  1938;  3  near  Lee's  Fer- 
r\',  17  July  1936  and  9-11  Aug.  19.38  (Tama- 
rix  and  Salix  3100  ft);  in  addition  14  birds 
seen  including  a  nest  and  two  nestlings, 
10-11  Aug.  1938;  Higgins  (Behle  and  Higgins 
1959)  foimd  it  an  abundant  breeder  in  dense 
streamside    vegetation    along    the    length    of 


Glen  Canyon;  BYU  (1971)  Site  16,  8  Aug. 
(Tamarix);  BYU  (1972)  3  .seen  in  Cottonwood 
Wash,  22  July  (Tamarix);  BYU  (1973)  Lee's 
Ferry,  a  total  of  5  birds  seen  in  Tamarix, 
6-15  June.  Common  summer  resident  in 
riparian  vegetation. 

Passenna  cyanea  (Linnaeus).  Indigo  Bun- 
ting. BYU  (1973)  Cottonwood  Wash,  6  June. 
One  was  seen  by  Robert  Whitmore  in  Tama- 
rix-Populiis  fremontii  association  about  six 
miles  up  Cottonwood  Wash  from  U.S.  High- 
way 89.  Lazuli  Buntings  were  also  present. 
Whitmore  (1975)  noted  apparent  com- 
petition between  males  of  the  two  species  in 
the  Paria  River  area.  Other  sightings  include 
Lee's  Ferry  where  Whitmore  (1975)  observed 
the  species  in  Tamarix.  As  noted  bv  Whit- 
more, the  species  apparently  has  moved  into 
the  Paria  River  drainage  since  the  in- 
vestigations of  Woodbury  and  Russell  (1945) 
and  Behle  et  al.  (1958).  VVhitmore  (1975)  also 
summarized  the  expansion  of  the  species  into 
Utah.  Summer  resident. 

Passerina  amoena  (Say).  Lazuli  Bunting. 
Tanner  (1940a)  Kaiparowits  Basin;  Wood- 
bury and  Russell  (1945)  two,  Navajo  Moun- 
tain, 13  Aug.  1935,  and  Colorado  River/ Rock 
Creek,  20  July  1937  (riparian  thicket,  under 
shrubs,  Populus  tremuloides,  and  Pinus  pon- 
derosa, 3300-9000  ft);  Russell  fovuid  it  fairly 
conniion  on  lower  Navajo  Mountain,  10-16 
Aug.  1936;  Behle  and  Higgins  (1959)  Kane 
Creek  and  river  mile  41,  1  Aug.  1958;  BYU 
(197.3)  in  Cottonwood  Wash  at  six  (five  seen 
5-6  June),  eight  (two  seen  6  June),  and  nine 
(six  seen  6  June)  miles  north  of  U.S.  Highway 
89,  Kane  County,  Utah.  Two  sightings  were 
in  Atriplcx  canescens,  one  was  in  Juniperus- 
Tainarix.  and  the  remainder  were  in  Tamarix. 
Whitmore  (1975)  noted  the  Lazuli  Bunting 
nesting  in  the  Paria  River  area.  Extreme 
dates  of  occurrence  are  15  May  and  28  Aug. 
(Woodbury  and  Russell  1945).  Summer  resi- 
dent. 

Hesperiphona  vespertina  (Cooper).  Eve- 
ning Grosbeak.  Presnall  (1934)  Bryce  Point; 
BYU  (1972)  Cottonwood  Wash,  four  seen  in 
stands  of  Piniis-Jiiniperus,  26  Oct.  Fall  and 
winter  visitant. 

Carpodacus  cassinii  Baird.  Cassin's  Finch. 
Woodbury  and  Russell  (1945)  two  specimens: 
Navajo  Mountain  9  and  14  Aug.  1935  (9000 
ft);  it  is  a  sparse  breeder  in  stands  of  Pinus 


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Great  Basin  Naturalist 


Vol.  40,  No.  4 


ponderosa;  Russell  and  Thompson  (1964) 
Brvce  Canyon.  Summer  resident. 

Carpodacus  mexicanus  frontalis  (Say). 
House  Finch.  Woodbury  and  Russell  (1945) 
two,  Navajo  Mountain  Trading  Post,  26  July 
1936  and  mouth  of  Rock  Creek,  23  and  26 
July  1937  {Populus  frernontii  and  riparian 
vegetation,  33()()-65()0  ft);  Behle  (1948)  Az- 
tec Creek  and  Lee's  Ferry;  Behle  et  al.  (1958) 
confluence  Calf  Creek /Escalante  River, 
1954,  and  Paria  River,  6  Mar.  1946;  Hayward 
et  al.  (1958)  Escalante  drainage;  Russell  and 
Thompson  (1964)  Bryce  Canyon;  Behle  and 
Higgins  (1959)  Higgins  found  them  abundant 
in  Glen  Canyon  during  summer  1958;  they 
were  seen  nearly  every  day  in  streamside 
vegetation,  on  the  terraces,  and  on  the  hill- 
sides; Behle  and  Higgins  (1959)  give  the  fol- 
lowing additional  unpublished  data  from 
Woodbury  and  Russell's  field  notes:  "species 
common  and  conspicuous  between  river 
miles  78  and  zero  of  the  Colorado  River, 
4-22  July  1936;  along  the  Colorado  River,  22 
were  comited  and  many  others  seen  between 
river  mile  75  and  Paria  Creek,  3-11  Aug. 
1938";  BYU  (1971)  one,  Warm  Creek,  8 
Nov.;  Wiregrass  Spring;  Lee's  Ferry;  three 
(ad.  males),  Tibbet  Spring,  16  Oct.;  Wah- 
weap  Creek;  Escalante  River  Bay;  Driftwood 
Canyon;  T43S  R2W  Sec  19,  7  Dec;  Three 
Garden;  2  miles  south  Church  Wells,  10 
Nov.;  T43S  R2W  Sec  24,  7  Dec;  along  the 
Escalante  River;  Paria  Plateau,  29  Sept.,  and 
mouth  of  Long  Canyon  18  Dec;  BYU  (1972) 
Driftwood  Canyon;  Three  Garden;  mouth  of 
Escalante  Canyon,  24-25  June;  and  Wire- 
grass  Spring,  22  July;  BYU  (1973)  Highway 
273/Wahweap  Creek,  20  Apr.;  five  miles 
north  of  Site  34,  1  May;  left  fork  Tibbet 
Spring,  5  June;  Sites  6,  23,  28,  and  30,  6-13 
June;  Sites  2  and  3,  12  July-9  Aug.;  Cotton- 
wood Wash,  9  miles  north  Highway  89;  Lee's 
Ferry,  Paria/Colorado  Rivers,  Paria  River, 
Tibbet  Spring,  and  Wahweap  Creek,  30 
Apr.-15  June;  Glen  Canyon  City,  17  Aug. 
The  greatest  numbers  were  seen  in  the  win- 
ter. Three  large  flocks  were  seen  8  Nov. 
1971,  and  two  flocks  each  containing  about 
300  birds  and  another  containing  50  were 
seen  7-18  Dec.  1971.  From  1971-1973,  2 
finches  were  seen  in  Apr.,  46  in  June,  8  in 
July,  and  12  in  Aug.  In  April-June  1973, 
when  House  Finches  are  expected  to  nest,  8 


were  seen  in  Tamarix,  2  in  Pinus  edulis,  and 
one  each  in  Coleogijne,  Juniperus-gra.ss,  and 
Salix;  2  were  seen  during  July  in  semiarid 
grasslands  and  Tamarix.  The  4  seen  in  June 

1972  were  in  hanging  gardens.  The  Novem- 
ber and  December  1971  flocks  were  in  semi- 
arid   grasslands   and  Juniper  us.    The    August 

1973  sightings  were  in  semiarid  grassland 
(10),  Artemisia  (5),  woodland  (2),  Vanclevea- 
grass  (1),  and  Glen  Canyon  City  (1).  Per- 
manent resident;  abundant  in  winter. 

Pinicola  enucleator  (Linnaeus).  Pine  Gros- 
beak. Presnall  (1934)  Bryce  Canyon  Rim. 
Summer  resident. 

Leticosticte  tephrocotis  tephrocotis  (Swain- 
son).  Gray-crowned  Rosy  Finch.  Behle  et  al. 
(1958)  Paria  River  (see  Black  Rosy  Finch). 
Winter  resident  or  transient. 

Leticosticte  atrata  Ridgway.  Black  Rosy 
Finch.  Behle  et  al.  (1958)  Paria  River,  about 
300  mixed  with  Gray-crowned  Rosy  Finch,  6 
Mar.  1946;  BYU  (1971)  Warm  Creek;  seven 
miles  above  Escalante;  Warm  Creek  drainage 
near  Tibbet  Spring,  200-400  seen  6  Dec;  3 
females,  2  males,  Tibbet  Canyon,  7  Dec; 
Nipple  Bench  above  Tibbet  Canyon,  1000 
seen  9  Dec;  and  Vi  mile  up  long  Canyon,  200 
seen  16  Dec;  BYU  (1972)  three  miles  south 
of  Tibbet  Spring  in  Tibbet  Canyon,  200  seen 
14  Feb.  and  Grosvenor  Arch  pond,  100  seen 
26  Oct.;  BYU  (1973)  Tibbet  Canyon,  25  seen 
25  Jan.  and  Tibbet  Spring,  100  seen  1  Mar. 
The  earliest  fall  sighting  was  26  October;  the 
latest  spring  sighting  was  6  March.  They 
were  encountered  in  Chnjsothamnus  (1000), 
washes  (300),  woodlands  (200),  Artemisia 
(100),  and  salt  wash  (25).  Locally  abundant, 
winter  resident. 

Cardiielis  pinus  pintis  (Wilson).  Pine  Sis- 
kin. Presnall  (1934)  Bryce  Canyon;  Benson 
(1935)  Navajo  Mountain;  Woodbury  and  Rus- 
sell (1945)  Russell  saw  from  one  to  four  birds 
almost  daily  on  Navajo  Mountain,  13-21  July 
{Picea-Abies  forest);  BYU  (1971)  Grand 
Bench,  5  Oct.  Summer  resident  and  transient. 

Carduelis  tristis  pallida  (Mearns).  Ameri- 
can Goldfinch.  Behle  et  al.  (1958)  confluence 
of  Calf  Creek /Escalante  River  1954;  Long 
(1937)  Bryce  Canyon,  23  Nov.  1935;  Russell 
and  Thompson  (1964)  Bryce  Canyon;  BYU 
(1971)  Grand  Bench,  5  Oct.  and  three  (males) 
Nipple  Spring,  6  Dec;  BYU  (1972)  Cedar 
Mountain.  Permanent  resident. 


December  1980 


Atwood  et  al.:  Kaiparowits  Vertebrates 


335 


Carduelis  psaltria  hesperophila  (Oberhol- 

ser).  Lesser  Goldfinch.  Presnall  (1934)  and 
Behle  et  al.  (1958)  Bryce  Canyon;  Woodburv 
and  Russell  (1945)  Navajo  Mountain,  two 
adults  and  three  young  seen  on  10,  11,  and  13 
July  1936  and  Navajo  Mountain  Trading 
post,  two  or  three  in  Populus  fremontii, 
27-29  July  1936;  believed  to  nest  in  larger 
stands  of  Populus  and  Salix  (Monson  Expedi- 
tion in  Navajo  Country  as  recorded  in  Wood- 
burv and  Russell  1945).  Permanent  resident. 

Loxia  curvirostra  Linnaeus.  Red  Crossbill. 
Presnall  (1934)  and  Russell  and  Thompson 
(1964)  Bryce  Canyon;  Woodbury  (1939)  2, 
Navajo  Mountain  obtained  from  a  flock  of 
12,  i.e.,  one  L.  c.  bendierei,  13  Aug.  1935,  the 
other  L.  c.  stricklundi,  12  Aug.  1935;  Wood- 
bury and  Russell  (1945)  flock  of  30-40  cross- 
bills were  seen  many  times  by  Russell  in 
stands  of  Pinus  ponderosa  and  Picea-Ahies  on 
Navajo  Moimtain,  2-23  July  1936.  Transient. 

Pipilo  chlorurus  (Audubon).  Green-tailed 
Towhee.  Presnall  (1934)  Bryce  Canyon; 
Woodbury  and  Russell  (1945)  two,  Kaiparo- 
wits Plateau  at  7000  ft  in  bushes  of  Amekm- 
chier,  3  and  12  Aug.  1937;  seen  several  times 
in  Amelanchier  and  Querctts  in  the  heads  of 
canyons  on  Kaiparowits  Plateau,  late  July 
and  early  Aug.  1913  [1937],  believed  to  be 
nesting.  Extreme  dates  for  the  general  area 
by  Woodbury  and  Russell  (1945)  were  22 
Aug.- 13  Oct.  in  the  fall  and  29  April- 11  May 
in  the  spring.  Summer  resident. 

Pipilo  erythrophthalmus  montanus 
Swarth.  Rufous-sided  Towhee.  Benson  (1935) 
three,  Navajo  Mountain  in  thickets  beneath 
Pinus  ponderosa;  Woodbury  and  Russell 
(1945)  three,  Navajo  Mountain,  3  July  and  7 
Aug.  1936,  and  11  Aug.  1935,  and  five  from 
Kaiparowits  Plateau,  30  July- 10  Aug.  1937 
{Archtostaphylos  under  Pinus  ponderosa,  Sa- 
lix,  Populus  treniuloides,  chaparral,  i.e.,  very 
common  in  bushy  thickets,  6000-8500  ft); 
Behle  et  al.  (1958)  Escalante  drainage;  Behle 
and  Higgins  (1959)  one  of  two  seen  was  col- 
lected at  mouth  of  Aztec  Creek,  26  July 
1958;  eight  seen  Navajo  Creek  Canyon,  9 
July  1936  (Rainbow  Bridge-Monument  Val- 
ley expedition);  Russell  and  Thompson  (1964) 
BWce  Canyon;  BYU  (1972)  Three  Garden  lo- 
cated one  mile  above  the  confluence  of  San 
Juan  River  and  Lake  Powell,  4  July  (hanging 


gardens).  Summer  resident,  a  few  mav  win- 
ter. 

Passerciilus  sandwichensis  nevadensis 
Grinnell.  Savannah  Sparrow.  Presnall  (1934) 
Bryce  Canyon;  Tanner  (1940a)  Kaiparowits 
Basin;  Behie  et  al.  (1958)  10  miles  south  of 
Escalante,  1954;  Hay  ward  et  al.  (1958)  Esca- 
lante drainage.  Transient. 

Ammospiza  leconteii  (Audubon).  Le- 
Conte's  Sparrow.  [Hypothetical]  BYU  (1973) 
Site  2,  one  seen  by  Clyde  L.  Pritchett,  1  Aug. 
This  species  is  considered  bv  Behle  and  Perrv 
(1975)  and  Hay  ward  et  al.  (1976)  to  be  of  ac- 
cidental, rare  occurrence  in  Utah.  They  note 
a  specimen  from  Provo,  Utah,  24  Dec.  1927, 
and  three  observations  there  on  10  Mar. 
1928;  one  was  also  seen  at  Moab,  19  Apr. 
1966.  This  adds  one  additional  sight  record 
for  the  state.  Transient. 

Pooecetes  gramineus  confinis  Baird.  Ves- 
per Sparrow.  Presnall  (1934)  Bryce  Canyon; 
Tanner  (1940a)  Kaiparowits  Basin;  Behle  et 
al.  (1958)  10  miles  south  of  Escalante,  1954; 
BYU  (1972)  one  mile  above  confluence  of  the 
San  Juan /Colorado  Rivers,  25  June  (hanging 
garden  pool);  BYU  (1973)  one  (ad.  female), 
two  miles  southwest  Glen  Canyon  City,  30 
Apr.;  Site  1,  two  seen  30  .\pr.;  Site  8,  3  May; 
and  one  mile  north  of  Site  10,  several  flocks 
seen,  1  May;  1973  sightings  were  in  Van- 
clevea-grsLss,  Grayia-Coleogyne,  and  Ephedra- 
grass.  Siunmer  resident,  more  common  in  mi- 
gration. 

Chondestes  grammacus  strigatus  Swain- 
son.  Lark  Sparrow.  Presnall  (1934)  Bryce 
Canyon;  Tanner  (1940a)  Kaiparowits  Basin; 
Woodbury  and  Russell  (1945)  two,  Kaiparo- 
wits Plateau,  4  Aug.  1937;  Navajo  Mountain 
Trading  Post  {Artemisia  iridcniata, 
6000-7000  ft);  Behle  and  Higgins  (1959)  Col- 
orado River/San  Juan  River  and  river  mile 
75,  2-3  .\ug.  1938  (Woodbury's  field  notes  of 
Rainbow  Bridge-Monument  Valley  Expedi- 
tion, 1938);  Hayward  et  al.  (1958)' Escalante 
drainage;  Behle  et  al.  (1958)  Bryce  Canyon, 
confluence  of  Calf  Creek /Escalante  River, 
and  Hayward  et  al.  (1958)  Escalante  drain- 
age; Behle  et  al.  (1958)  Bryce  Canyon,  con- 
fluence Calf  Creek /Escalante  River,  and  10 
miles  south  of  Escalante,  7  May  1954.  Ex- 
treme dates  of  arrival  and  departure  for  the 
general  area  are  28  Apr.  and  14  Sept.,  re- 
spectivelv  (Woodbury  and  Russell    1945). 


336 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


Nesting  takes  place  on  open  flats  in  medium 
dense  brush  and  often  in  savannah  at  the 
woodland  edges  or  openings  (Woodbury  and 
Russell  1945).  Summer  resident. 

Aimophila  cassinii  (Woodhouse).  Cassin's 
Sparrow.  [Hypothetical]  BYU  (1973)  Sites  1, 
2,  and  3,  small  flocks  common,  30  Apr.; 
Wahweap  Creek,  30  Apr.;  and  Tibbet  Can- 
yon, small  flocks  common,  1  May.  The  Cas- 
sin's Sparrow  was  placed  on  the  hypothetical 
list  of  the  birds  of  Utah  by  Behle  and  Perry 
(1975).  The  present  observations  made  by 
Lloyd  Pack  are  questionable.  He  possibly  ob- 
served Brewer's  and  Chipping  Sparrows  in- 
stead of  Cassin's  Sparrows.  Transient. 

Amphispiza  bilineata  deserticola  Ridg- 
way.  Black-throated  Sparrow.  Woodbury  and 
Russell  (1945)  3,  Rock  Creek /Colorado  Riv- 
er, 21  July  1937;  Navajo  Mountain  Trading 
Post,  27  July  1936  and  6  Aug.  1935  (desert 
brush  at  3200-6500  ft  and  Artemisia  triden- 
tata  at  5800-6500  ft);  Woodbury  and  Russell 
(1945)  give  extreme  dates  of  occurrence  in 
the  Navajo  County  as  12  Apr.  and  1  Sept. 
with  exception  of  20  birds  seen  29-30  No- 
vember; Behle  and  Higgins  (1959)  report  that 
during  the  Rainbow  Bridge-Monument  Val- 
ley expedition  Woodbury  observed  this  spe- 
cies at  river  mile  63,  5  Aug.  1938,  river  mile 
50,  a  few  seen  6  Aug.  1938,  four  miles  up 
Last  Chance  Creek,  a  few  seen  7  Aug.  1938, 
river  mile  41,  2  seen  8  Aug.  1938,  and  mouth 
of  Paria  Creek,  3  seen  11  Aug.  1938;  Behle  et 
al.  (1958)  10  miles  south  of  Esca- 
lante/Kaiparowits  Plateau,  Aug.  1957;  Behle 
and  Higgins  (1959)  mouth  of  Aztec  Creek 
and  Woodbury  saw  more  than  6  along  the 
Colorado  River  from  mile  63  to  the  Paria 
Creek,  5-11  Aug.;  BYU  (1971-1973)  Sites  1, 
2,  and  30;  Glen  Canyon  City,  Cottonwood 
Wash/ Paria  River;  Tibbet  Spring;  and  one 
mile  above  confluence  of  San  Juan/Colorado 
Rivers.  Our  earliest  spring  observation  of  this 
species  is  1  May  (1973).  The  latest  in  the  fall 
is  8  Aug.  (1973).  Numbers  seen  were  greatest 
in  Aug.  (35)  and  least  in  May  (6);  in  addition, 
20  were  seen  in  June  and  12  in  July.  Fifteen 
of  the  birds  were  seen  in  scattered  bnish  con- 
.sisting  of  Artemisia  tridentata-Ephedra  and 
other  desert  species,  4  in  Juniperus  or  Juni- 
penis- Artemisia,  and  2  in  Tamanx.  Wood- 
bury and  Russell  (1945)  believe  this  species  is 


closely  associated  in  its  distribution  with  Co- 
leogyne,  from  the  lower  part  of  the  pygmy 
forest  downward.  Common  summer  resident. 

Amphispiza  belli  nevadetisis  (Ridgway). 
Sage  Sparrow.  Benson  (1935)  south  of  Navajo 
Mountain;  Woodbury  and  Russell  (1945)  2, 
Warm  Creek,  15  July  1936,  and  five  miles 
south  Navajo  Mountain  Trading  Post,  .30  July 
1936  {Salix,  Artemisia,  3100-6500  ft);  Wood- 
bury and  Russell  (1945),  commonest  bird  of 
open  Artemisia  flats  south  of  Navajo  Moun- 
tain, with  at  least  100  seen  in  a  single  day; 
Russell  and  Thompson  (1964)  Bryce  Canyon; 
BYU  (1971)  one  mile  south  Glen  Canyon 
City,  10  Nov.;  Grand  Bench,  10  Oct.;  Warm 
Creek  Inlet,  8  Nov.  and  6  Dec;  Tibbet  Can- 
yon, near  spring,  4  Dec;  and  two  miles  east 
of  Nipple  Butte,  6  Dec;  BYU  (1972)  Site  1, 
25  Jan.  and  7  Feb.;  Tibbet  Spring,  25  Jan.; 
BYU  (1971-1973)  Sites  2,  3,  6,  10,  12,  13,  14, 
15,  23,  and  28  and  Cottonwood  Wash.  Sage 
Sparrows  winter  in  the  area;  the  largest  num- 
bers were  seen  in  Dec.  (67)  and  Jan.  (34). 
None  were  seen  in  Mar.,  17  in  Apr.,  12  in 
June,  one  in  July,  18  in  Aug.,  none  in  Sept., 
and  3  in  Nov.  Although  this  species  is  ex- 
pected to  occur  in  Artemisia  during  the  nest- 
ing season  (May-June),  it  was  encountered  in 
Graijia  (2),  Coleogijne-Vanclevea  (19),  Arte- 
7nisia  tndentata  (1),  Artemisia-Juniperus  (1), 
and  Atriplex  (3).  During  the  remaining  period 
they  were  recorded  in  Mahonia  (40),  Arte- 
misia tridentata  (25),  Cercoearpiis  (10), 
Chnjsothammis  (6),  Juniperus  (2),  Juniperus- 
Artemisia  (1),  Ephedra  (1),  grass  (1),  Atriplex 
(1),  and  unidentified  shrubs  (13).  The  Dec. 
and  Jan.  birds  were  recorded  in  Cercocarpus 
(10),  Ephedra-Atriplex-Chrysothamnus  (4), 
Mahonia  (40),  Artemisia  tridentata  (25),  and 
mixed  shrub-grass  (5).  Common  permanent 
resident. 

Jtinco  hyemalis  (Linnaeus).  Dark-eyed 
Junco.  In  the  thirty-second  supplement  to  the 
American  Ornithologists  Union  Check-list  of 
North  American  Birds  (Auk  90:411-419, 
April  1973),  the  committee  on  classification 
and  nomenclature  considered  most  of  the 
Juneo  species  conspecific  with  /.  hyemalis. 
Following  this  pattern  we  are  combining  all 
the  juncos  in  the  Kaiparowits  Basin,  except 
one,  into  the  hyemalis  complex.  The  former 
specific  names  now  become  the  subspecific 
names.  Junco  caniceps  is  currently  under 


December  1980 


Atwood  et  al.:  Kaiparowits  Vertebrates 


337 


study  and  therefore  is  presently  maintained 
as  a  separate  species. 

Junco  hyemalis  oreganus  (Townsend)  (=/. 
oreganus  moiitanus  and  /.  o.  shiifeldti).  Pres- 
nall  (1934)  /.  o.  oreganus  {=}.  shufcldti)  at 
Bryce  Canyon;  Grater  (1947)  Bryce  Canyon; 
Behle  et'  al.  (1958)  confluence  Calf 
Creek/Escalante  River,  1954;  and  BYU 
(1971)  Site  12;  Site  15,  10  Oct.;  Cottonwood 
Wash,  13  Nov.;  Tibbet  Spring,  4  Dec;  and 
Wahweap  Creek,  16  Dec;  BYU  (1972)  Wah- 
weap  Creek,  28  Jan.;  Tibbet  Spring,  25  Jan.; 
and  Cockscomb,  28  Jan.  The  earliest  fall  ob- 
servations of  juncos  are  10  Oct.  1971  and  26 
Oct.  1972;  the  latest  spring  sighting  is  28  Jan. 
1972.  The  greatest  numbers  were  seen  Dec. 
(48  +  a  large  flock).  Juncos  were  observed  in 
Tamarix  (35),  Cercocai-pus  (10),  mixed  shrubs 
(10),  Mahonia  (3),  Chnjsothamnus  (3),  Popu- 
his  fremontii-Tamarix  (1).  Common  transient 
and  winter  visitant. 

Junco  caniceps  caniceps  (Woodhouse). 
Gray-headed  Junco.  Presnall  (1934)  Bryce 
Canyon;  Benson  (1935)  Navajo  Mountain, 
breeding  bird,  common  1-23  July  1933; 
Woodbury  and  Russell  (1945)  seven,  Navajo 
Mountain,  14  Aug.,  1935  and  2-18  July  1936 
{Populus  tremuloides,  Finns,  Picea-Ahies, 
8500-10,000  ft);  Behle  et  al.  (1958)  con- 
fluence Calf  Creek/Escalante  River,  1954; 
BYU  (1971)  xNavajo  Mountain  summit,  12 
Oct.;  Cottonwood  Wash,  7  Nov.,  and  a  large 
flock  was  seen,  7  Nov.  (Populus  fremontii) 
and  one  was  seen  in  Dec  [Juniperus).  Found 
on  Navajo  Mountain  as  a  breeding  bird  be- 
tween 1  to  23  July  1936  (Woodbury  and  Rus- 
sell 1945).  Common  summer  resident  of  Nav- 
ajo Mountain,  winters  in  lowlands. 

Spizella  arborea  (Wilson).  Tree  Sparrow. 
BYU  (1971)  one  (male).  Warm  Creek  Inlet,  8 
Nov.  Rare  transient  or  winter  visitant. 

Spizella  passerina  arizonae  Coues.  Chip- 
ping Sparrow.  Presnall  (1934)  Bryce  Canyon; 
Benson  (1935)  Navajo  Mountain  area;  Tanner 
(1940a)  Kaiparowits  Basin;  Woodbury  and 
Russell  (1945)  river  mile  69,  3  Aug.  1938;  3 
specimens:  Navajo  Mountain  Trading  Post, 
23  July  1936;  Navajo  Mountain,  16  Aug. 
1935,  and  Navajo  Mountain  Trading  Post,  23 
July  1936  {Pinus  ponderosa,  pygmy  forest, 
65()0-90(X)  ft);  also  observed  69  miles  above 
Lee's  Ferry,  3  Aug.  1938;  Behle  et  al.  (1958) 
confluence  Calf  Creek/Escalante   River,    13 


Jime  (stand  of  Junipcrus-grass);  Russell  and 
Thompson  (1964)  Bryce  Canyon;  BYU  (1973) 
Site  28  {Artemisia  tridentata);  Woodbury  and 
Russell  (1945)  reported  seeing  1074  individ- 
uals of  this  species  (15  sightings)  between  12 
Apr.  and  13  Oct.  1936-38.  Our  paucity  of  re- 
cords may  be  due  to  the  difficulty  of  in- 
experienced observers  in  distinguishing 
among  the  various  species  of  small  sparrows. 
It  is  probable  that  many  were  misidentified, 
e.g.,  Cassin's  Sparrow  as  previously  men- 
tioned as  well  as  Brewer's  Sparrow.  Summer 
resident,  spring,  and  fall  transient. 

Spizella  hreweri  breweri  Cassin.  Brewers 
Sparrow.  Tanner  (1940a)  Kaiparowits  Basin; 
Woodbury  and  Russell  (1945)  three,  Kaiparo- 
wits Plateau,  31  July  1937;  Navajo  Mountain, 
8  Aug.  1936;  and  Navajo  Mountain  Trading 
Post,  30  July  1936  (Artemisia  tridentata,  pyg- 
my forest,  6500-7000  ft);  observed  at  Lee's 
Ferry,  14-26  Aug.  1909,  bv  E.  W^  Nelson; 
Behle  et  al.  (1958)  confluence  Calf 
Creek/Escalante  River,  1954,  and  10  miles 
south  of  Escalante,  8  May  1954;  BYU  (1971) 
one.  Site  2,  28  Aug.;  one.  Church  wells,  28 
Sept.;  and  one  Paria  Plateau,  29  Sept.;  BYU 
(1973)  Site  4,  6  June;  Site  6,  10  seen  4  July; 
Site  23,  5  June;  Site  28,  13  June;  Cottonwood 
Wash  road,  U/2  miles  north  of  U.S.  Highway 
89,  6  June  and  13  miles  north  of  U.S.  High- 
way 89,  8  June;  Hackberry  Can- 
yon/Cottonwood Wash,  two  seen  8  June;  and 
Kelly  Grade/Smoky  Mountain,  3  July.  Al- 
though the  species  is  primarily  a  bird  of  Arte- 
misia tridentata,  most  sightings  were  in  open 
or  scattered  desert  shrub  and  one  was  in  a 
Juniperus-grass  association.  Common  summer 
resident  and  migrant. 

Zonotrichia  quenila  (Nuttall),  Harris 
Sparrow.  BYU  (1973)  one  specimen  (ad. 
male),  three  miles  east  of  Glen  Canyon  City, 
30  Apr.  Winter  visitor  of  lower  and  warmer 
vallevs. 

Zonotrichia  leucophrys  (Forster).  White- 
crowned  Sparrow.  Tanner  (1940a)  Kaiparo- 
wits Basin;  Behle  (1948)  Lee's  Ferry;  Behle  et 
al.  (1958)  specimens  at  confluence  Calf 
Creek/Escalante  River  (Z.  /.  gambelii  and 
oriantha)  and  10  miles  south  of  Escalante, 
7-8  May  1954;  Russell  and  Thompson  (1964) 
Bryce  Canyon;  Presnall  (1934)  recorded  Z.  /. 
gambelii  from  Bryce  Canyon;  it  is  a  common 
migrant    in   Bryce   Canyon    fall   and   spring. 


338 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


occurring  in  mixed  flocks  of  Z.  /.  oriantha; 
BYU  (1961)  two  (females),  old  Paria  townsite, 
20  May;  BYU  (1971)  Site  15,  9  Oct.  (salt  wash 
vegetation);  Cottonwood  Wash,  7  Nov.;  BYU 

(1972)  Tibbet  Spring,  30  seen  15  Apr.;  BYU 

(1973)  Wahweap  Creek,  30  Apr.  Transient 
and  winter  visitant. 

Passerella  iliaca  (Merrem).  Fox  Sparrow. 
BYU  (1971)  one  (not  kept),  Wahweap  Creek, 
4  Nov.  Transient. 

Melospiza  lincolnii  alticola  (Miller  & 
McCabe).  Lincoln's  Sparrow.  Behle  et  al. 
(1958)  Bryce  Canyon.  Transient. 

Melospiza  melodia  montana  Henshaw. 
Song  Sparrow.  Russell  and  Thompson  (1964) 
Bryce  Canyon;  Behle  and  Higgins  (1959) 
Kane  Creek  at  river  mile  41,  2  seen  18  Oct. 
1958;  BYU  (1971)  Site  12  (Coleogyne);  Glen 
Canyon  City;  and  one  (not  kept).  Nipple 
Spring,  15  Oct.  Transient. 


Summary  of  Birds  of  Kaiparowits  Region 

(183) 

Permanent  Residents  (.36) 

Biiteo  janiaicensis  calurtis 

Aqtiila  chnjsaetos  canadensis 

Dendragapus  ohscii nis 

Centrocerciis  uropliasianus  mopluisianiis 

Ottis  ash 

Otus  flammeolus 

Bubo  virainiantis  pallescens 

Glaucidium  gnoma  caUfornicinn 

Asio  ottis 

Perisoreus  canadensis  capitalis 

Ctjanocitta  stelleri  macrolopha 

Aphelocoma  coerulescens  woodhuuscii 

Corvus  corax  sinuattis 

Ci/ninorliinus  cijanocephalus 

Nncifraga  colwnhiana 

Pants  atricapilltis  garrinus 

Panis  gainbcli 

Partis  itiornattis  ridgivai/i 

Psaltripants  ntinim us 

Sitta  carolinensis  nelsoni 

Sitta  canadensis 

Sitta  pijgniuea  nielanotis 

Certhia  familiaris 

Cinchts  inexicanus  unicolor 

Troglodijtes  aedon  parknianii 

Catlierpes  niexicanus  consperstis 

Si(tli(t  niexicatia  hairdi 

Reguhts  satrapa 

Regultis  calenditia 

Lanitts  ludovicianus 

Sturntts  vulgaris  ntlgaris 

Passer  domestic  us 

Car}wdacits  niexicanus  frontalis 

Carduelis  tristis  pallida 


Carduelis  psaltria  hesperophila 
Aniphispiza  belli  nevadensis 

Uncommon  or  Sparse  Permanent  Residents  (7) 

Ualiaeettts  leucocephalus 
Lophortyx  gambelii  gandjelii 
Phasianus  colchicus 
Alectoris  ehukar 
Strix  occidentalis  lucida 
Colaptes  auratits  cafer 
Sturnella  neglecta  neglecta 

Spring-Siinnner  Residents-May  Be  I'ncommon  (3) 

Egretta  tint  la 
Nycticorax  n  ycficorax 
Anas  strepera 

Summer  Residents— Common  or  Abundant  (57) 

Ardca  hcrodias 

Accipiter  striatus  velox 

Buteo  sivainsoni 

Faico  niexicanus 

Actitus  macularia 

Cohimba  fasciata  fasciata 

7,enaida  macroura 

Chordeiles  minor  lienryi 

Aeronautcs  saxatalis  saxatalis 

ArcliUochits  alexandri 

Selasphorus  platycents  platycerus 

Selasphorus  rufus 

Spli yrapicus  tli  yroideus 

Picoides  villosus  leucothorectus 

Picoides  pubescens  leucttrus 

Tyrannus  verticalis 

Tyrannus  vociferans  vociferans 

Myiarchus  cinerascens  cinerascens 

Sa  yorn  is  n  igrica n s 

Sayornis  say  a  say  a 

Enipidonax  traillii 

Enipidonax  oberliolseri 

Enipidonax  icrigli tii 

Enipidonax  difficilis  lielhnaiiri 

Sutallornis  borcalis 

Ereniophila  alpcstris  leucohienui 

Tacliycineta  thalassina  lepida 

Riparia  riparia  riparia 

Stelgidopteryx  ruficollis 

Petrochclidon  pyrrhonota  pyrrhonota 

Tliryomancs  beivickii  ereniophilus 

Minius  polyglottos  leucopterus 

Duniettella  carolinensis 

Toxostonia  bendirei 

Catharus  guttatus  auduhoni 

Polioptila  caerulea  anioenissima 

Phainopepla  nitens  lepida 

Vireo  gihiis 

Vennivora  virginiae 

Virniivora  luciae 

Dcndroica  petechia  morconii 

Dendroica  graciae  gruciae 

Geothlypis  triclias  occidentalis 

Icteria  virens 

Icterus  parisorunt 

Icterus  iialbula  bullockii 


December  1980 


Atwood  et  al.:  Kaiparowits  Vertebrates 


339 


Molotlinis  ater  ohscurus 

Fimngii  ludoiiciana 

Fhciicticus  inelanoccphalits  ruchinoccpluilus 

Gtiinicd  caerulcd  interfusa 

Passerina  cijiineu 

Passerina  amociui 

Carpoddciis  cassin n 

Pinicohi  cniiclcdtor 

Pipilo  rhlorurtis 

Chondcstcs  oranuiidcus  strigattis 

Amphifipizu  hilincdld  dcsciiiroUi 

Summer  Residents,  Some  Mav  Winter  (6) 

Brdtita  cdiuidcnsis 

Falco  spdrverius  spdnciiiis 

Salpinctes  obsoletus  oh.wlctiis 

Tardus  migmtorius  propiiupitis 

Skdui  curmroides 

Pipilo  en/throphthdhmi.s  uiontdniis 

Summer  Residents,  Transient  or  Migrant  during  Spring 
and/or  Fall  (10) 

Accipitcr  cooperii 
Sphyrdpicu.s  vdiiiis 
Vireo  solitariiis 

Dendroica  cororidta  duduhoni 
Dendroicd  niorescens 
Opowrnis  tolmiei 
Cardiu'lis  piniis  pinus 
Pooecetcs  grdmineiis  con f inns 
Spizella  passerina  arizonae 
Spizella  breweri  hreweri 

Uncommon   Summer   Residents,   Mav  Be   Spring-Fall 

Transients  (12) 

Carthartes  aura  tetcr 

Accipiter  gentilis  atricapillus 

Falco  peregrinus  anatum 

Falco  colnmbarius  bcndirei 

Fulica  anwricana 

Cbaradrius  vocifcrus  lociferus 

Athene  cuniculdria  hijpugaea 

Phalaenoptilus  nnttallii  nuttaUii 

Melanerpes  lewis  , 

Contopus  sordidulus  veliei 

Vireo  bellii 

Vireo  vicinior 

Late  Fall/Fall-VVinter  Residents  (3) 

Podiceps  nigricollis 
Podih/nduts  podiceps 
Pica  pica  hudsonid 

Winter  Residents  (4) 
Buteo  logopus 
Leucosticte  dtratd 
Junco  hyenialis  oreganus 
Zonotrichia  cpicrula 

Transients,  Some  May  Nest,  Some  Mav  Winter  (20) 

Aruis  creccd 
Bucephdla  chingula 
Myadestes  towiisendi  toicnscndi 
Vennivora  ruficapilla  ridgtcayi 
Dendroica  toicnscndi 
Dendroica  occidenlalis 


Wdsoniu  pusdla  pileolata 

Xdnthocephdlus  xdnthoceplialus 

Agelaius  phocn iceus 

Euphdgus  cydnocephalus 

Leucosticte  tephrocotis  tephrocotis 

Loxia  cunirostra 

Passerculus  sandwicliensis  nevadensis 

Animonspiza  leconteii 

Airnophila  cassinii 

Jttnco  caniceps  canicc})s 

7.onotri(liia  Icucuphrys 

Passerelld  ilidca 

Melospiz^i  lincolnii  alticola 

Melospiza  melodia  niontana 

Uncommon  Spring-Summer  Transients  (3) 
Anas  discors 
Bucephdld  albeola 
Larus  delawarensis 

Spring-Fall  Transients  (2) 

Anas  dcutd 
Anas  cyanoptcra 

Fall  Transients  (3) 
Steganopus  tricolor 
Larus  californicus 
Hesperiphona  vespcrtina  vcspcrtina 

Uncommon  Transients  (8) 
Pie  gad  is  chili  i 
Anas  platyrhynchos 
Aruis  amcricana 
Aydiya  vdlisinerid 
Buteo  regdlis 
Oreoscoptes  i>i(intdnus 
Bombycilld  ccdrorum 
Vennivord  crlata 

Sparse  Transients  (5) 

Catoptrophorus  scinipalinatus  inorudtus 
Calidris  inauri 
Stellula  cdlliope 
Myiarchus  tyrannulus 
Iridoprocne  bicolor 

Rare  Transients  or  Winter  Visitants  (4) 

Accluuophorus  occidentalis 
Pelecanus  eryth rorh ynchos 
Lanius  excubilor 
S})izclla  arborcd 

Formerly    More    Abundant    Than    Indicated    h\    This 

Study  (4) 

Cathartes  dura  teter 

Actitis  macularia 

PJidldcnoptilus  nuttallii  nuttallii 

Geothylypis  trichas  occidentalis 

Range  Extensions  into  Kaiparowits  Basin  (5) 

Myiarchus  tyrannulus 
Phainopepla  nitens  lepida 
Sturnus  vulgaris  vulgaris 
Passerina  cyanea 
Anunospiza  leconteii 


340 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


Mammals 

In  general  we  have  followed  Durrant's 
(1952)  classification  with  regard  to  the  no- 
menclature and  taxonomy  of  mammals. 

Soricidae  (Shrews) 

Sorex  obscurus  obscurus  Merriam.  Dusky 
Shew.  This  species  has  been  collected  on 
Boulder  Mountain  just  north  of  Escalante 
(Durrant  1952);  it  probably  occurs  on  the 
portion  of  Boulder  Mountain  in  the  Kaiparo- 
wits  Basin.  This  is  within  the  range  of  the 
species  as  indicated  by  Durrant. 

Sorex  palustris  naingator  (Baird).  Water 
Shrew.  Reported  by  Tanner  (1940a)  from  the 
Kaiparowits  Basin. 

Sorex  merriami  leucogenys  Osgood.  Mer- 
rian  Shrew.  Benson  (1935)  reported  this  spe- 
cies as  S.  leucogenys,  but  in  1939  combined  it 
with  S.  merriami  as  a  subspecies.  This  was 
based  on  five  specimens  from  War  God 
Spring  on  Navajo  Mountain. 

Vespertilionidae  (Verpertilionid  Bats) 

Myotis  yumanensis  yumanensis  H.  Allen. 

Yuma  Bat.  Durrant  and  Dean  (1959)  collect- 
ed 10  specimens  from  along  Glen  Canyon 
and  8  from  a  large  colony  at  the  mouth  of  the 
Escalante  River. 

Myotis  volans  interior  Miller.  Hairy- 
winged  Myotis.  Reported  by  Benson  (1935) 
from  Rainbow  Bridge. 

Myotis  leibii  melanorhinus  (Merriam). 
Small-footed  Bat.  Durrant  and  Dean  (1959) 
made  a  collection  at  the  mouth  of  Kane 
Creek  on  the  Colorado  River. 

Lasionycteris  noctivagans  (LeConte).  Sil- 
ver-haired Bat.  Tanner  (1940a)  reported  this 
species  from  the  Kaiparowits  Plateau. 

Pipistrellus  Hesperus  Hesperus  H.  Allen. 
Western  Pipistrelle.  Benson  (1935)  reported  a 
collection  from  Rainbow  Bridge  and  Tanner 
(1940a)  reported  it  from  the  Kaiparowits 
Plateau.  Durrant  and  Dean  (1959)  indicate 
that  this  is  the  most  common  bat  in  Glen 
Canyon.  They  collected  two  individuals  at 
Lee's  Ferry  and  one  from  river  mile  91. 
Cockrum  (1960)  also  collected  this  species  at 
Lee's  Ferry.  Pritchett  (1962)  reports  this  spe- 
cies occurring  on  both  sides  of  the  Cocks- 


comb Ridge;  in  1961-1962  it  was  the  most 
common  bat  flying  in  the  early  evening. 

Eptesicus  fuscus  pallidus  Young.  Big 
Brown  Bat.  Benson  (1935)  reported  this  spe- 
cies from  Navajo  Mountain  Trading  Post. 

Antrozous  pallidus  pallidus  (LeConte). 
Pallid  Bat.  The  occurrence  of  the  pallid  bat 
in  the  Kaiparowits  region  has  not  been  re- 
ported in  the  literature.  Pritchett  (1962)  re- 
ports it  as  the  most  common  late-flying  bat 
(after  2300  hours)  near  the  Old  Paria  town- 
site.  He  collected  specimens  there  in  1961 
and  1972.  BYU  (1973)  observed  them  at  Sites 
2  and  3. 


Molossidae  (Free-tailed  Bats) 

Tadarida  brasiliensis  mexicana  (Saussure). 

Brazilian  Free-tailed  Bat.  Durrant  (1952)  in- 
dicates the  Brazilian  free-tailed  bat  occurs 
throughout  southern  Utah,  but  did  not  list  a 
reference  for  any  being  collected  in  the  envi- 
rons of  the  Kaiparowitz  Plateau.  Hardy 
(1941)  records  them  as  occurring  in  Zion  Na- 
tional Park.  BYU  did  not  collect  this  species 
during  this  study,  or  did  Pritchett  (1962). 
However,  they  are  one  of  the  most  common 
bats  in  Carlsbad  Caverns  in  New  Mexico  and 
we  are  including  them  as  part  of  the  fauna  of 
the  Kaiparowits  Plateau. 

Leporidae  (Hares  and  Rabbits) 

Sylvilagus  nuttallii  (Bachman).  Nuttall 
Cottontail.  This  species  is  restricted  to  the 
higher  elevation  with  S.  n.  grongeri  (Allen) 
occurring  in  Bryce  Canyon  and  vicinity  as  re- 
ported by  Presnall  (1934).  The  subspecies  S. 
n.  pinetis  (Allen)  was  reported  for  Navajo 
Mountain  by  Benson  (1935)  and  is  apparently 
restricted  to  the  mountain  ranges  east  of  the 
Colorado  River.  One  was  observed  by  BYU 
near  Site  33,  which  is  one  mile  north  of  the 
confluence  of  the  Colorado  and  San  Juan 
Rivers. 

Sylvilagus  audubonii  (Baird).  Desert  Cot- 
tontail. Durrant  (1952)  reported  S.  a.  ari- 
zonae  (Allen)  to  be  found  seven  miles  south- 
west of  Tropic,  Utah.  The  range  of  the 
subspecies  S.  a.  warreni  Nelson  in  the  Kai- 
parowits is  uncertain.  Benson  (1935)  reported 
tlie  latter  subspecies  from  the  mesa  top  south 
of  Navajo  Mountain.  The  Desert  Cottontail 


December  1980 


Atwood  et  al.:  Kaiparovvits  Vertebrates 


341 


or  their  signs  have  been  observed  throughout 
most  of  the  Kaiparowits  Basin,  but  no  speci- 
mens have  been  retained  (BYU  1971-1974). 

Lepus  americanus  bairdi  Hayden.  Snow- 
shoe  Kabliit.  Russell  and  Thompson  (1964) 
list  this  species  as  occurring  in  Brvce  Canyon. 
Durrant  (1952)  includes  the  northwest  por- 
tion of  the  Kaiparowits  Basin  within  its 
range. 

Lepus  townsendii  Bachman.  White-tailed 
Jack  Rabbit.  Durrant  (1952)  includes  the 
northwest  portion  of  the  Kaiparowits  Basin  in 
the  range  of  this  species.  Russell  and  Thomp- 
son (1964)  list  it  for  Bryce  Canyon.  We  did 
not  find  this  species  on  anv  of  our  collection 
sites. 

Lepus  californicus  Gray.  Black-tailed  Jack 
Rabbit.  It  is  common  throughout  Utah  west 
of  the  Colorado  River.  This  animal  has  been 
observed  by  BYU  (1971-1974)  crews  at 
nearly  every  site  studied.  They  have  been 
common  in  Butler  Valley,  Four  Mile  Bench, 
Smoky  Moimtain,  and  Cedar  Mountain.  The 
subspecies  L.  c.  texianus  Waterhouse  was  ob- 
served by  our  crews  east  of  the  Colorado  Riv- 
er at  Sites  19,  20,  21,  and  22  near  Navajo 
Mountain  (BYU  1971-1974).  Benson  (1935) 
reported  it  for  the  mesa  top  south  of  Navajo 
Mountain. 


Sciuridae  (Squirrels,  Prairie  Dogs) 

Eutamias    minimus    consobrinus    (Allen). 

Least  Chipmimk.  Presnall  (1934  and  1938) 
reported  this  species  for  Bryce  Canyon.  We 
have  a  single  collection  from  Brigham  Plains 
and  have  observed  it  at  Site  2  (BYU  1972  and 
1973). 

Eutamias  dorsalis  utahensis  Merriam. 
Cliff  Chipmunk.  Durrant  (1952)  cites  a  speci- 
men which  was  collected  eight  miles  south  of 
Escalante.  Pritchett  (1962)  collected  it  on  the 
Cockscomb  Ridge.  BYU  records  (1972-1973) 
are  for  Sites  2,  27,  and  28. 

Eutamias  quadrivittatus  (Say).  Say  Chip- 
mimk. Presnall  (1934)  reported  that  chip- 
niimks  were  common  at  Bryce  Canyon.  Dur- 
rant (1952)  examined  a  specimen  of  E.  q. 
adsitus  (Allen)  from  Bryce  Canyon.  Tanner 
(1940a)  collected  a  specimen  from  the  north- 
em  part  of  the  Kaiparowits  Basin.  We  have  a 
single  record  from  Site  27  (BYU  1974).  The 
subspecies   E.    q.    hopiensis    Merriam    was 


reported  by  Benson  (1935)  from  Rainbow 
Bridge  and  Navajo  Mountain.  Durrant  and 
Dean  (1959)  reported  two  collections  from 
the  mouth  of  Kane  Creek.  BYU  records 
(1971-1974)  are  from  sites  2  and  15,  near 
Page,  along  the  Colorado  River  in  Driftwood 
and  Reflection  Canyons,  and  in  Three  Gar- 
den one  mile  above  the  San  Juan  confluence 
with  Lake  Powell. 

Marmota  flaviventris  engelharti  Allen. 
Yellow-bellied  Marmot.  The  only  records  of 
this  species  for  the  Kaiparowits  Basin  are 
those  of  Patraw  and  Gray  (1932)  and  Presnall 

(1934)  for  Bryce  Canyon. 
Ammospermophilus    leucurus    (Merriam). 

Antelope  Ground  Squirrel.  According  to 
Hansen  (1955),  two  subspecies  occur  in  the 
Kaiparowits  Basin,  viz.  A.  I.  escalante  (Han- 
sen) on  the  west  .side  of  the  Colorado  and  A. 
/.  cinnamomeus  (Merriam)  on  the  east  side. 
BYU  crews  (1971-1974)  have  recorded  the 
former  subspecies  for  Sites  1,  2,  3,  4,  8,  10, 
14,  16,  18,  23,  27,  28,  and  30,  and  in  all  the 
drainages  west  of  the  Colorado.  The  latter 
subspecies  was  reported  bv  Benson  (1935) 
from  Rainbow  Lodge.  (>ockrum  (1960)  col- 
lected it  at  Lee's  Ferry,  and  BYU 
(1971-1972)  collected  it  at  Sites  19,  20,  21, 
and  22,  and  at  Three  Garden. 

SpermophiUis  spilosoma  crtjptospilotus 
Merriam.  Spotted  Ground  Squirrel.  This  spe- 
cies was  observed  by  Benson's  party  on  13 
June  1934,  five  miles  .south  of  the  summit  of 
Navajo  Mountain  (Benson  1935).  BYU  (1972) 
records  are  from  Sites  20  and  21. 

Spermophilus  variegatus  (Erxleben).  Rock 
Squirrel.  Presnall  (1934)  reported  that  S.  v. 
utali  was  frequently  seen  in  Bryce  Canyon 
and  Tanner  (1940a)  reported  it  for  the  Kai- 
parowits Basin.  BYU  (1972-1973)  records  are 
from  Cottonwood  Wa.sh,  Last  Chance,  Pet 
Hollow,  and  Grosvenor  Arch.  Durrant  and 
Dean  (1959)  observed  S.  r.  utali  sporadically 
throughout  Glen  Canyon  and  obtained  one 
.specimen  of  S.  v.  <[,raimnurus  (Say)  at  .\ztec 
Creek.  BYU  personnel  also  observed  speci- 
mens on  both  sides  of  Lake  Powell.  Benson 

(1935)  reported  that  this  species  has  been  ob- 
.served  at  Rainbow  Lodge  and  Navajo  Trad- 
ing Post. 

Spermophilus  lateralis  lateralis  (Say). 
Golden-mantled  Ground  Squirrel.  Presnall 
(1934)  reported  it  as  common  on  the  rim  of 


342 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


Bryce  Canyon.  Tanner  (1940a)  reported  it  for 
the  Kaiparowits  Basin. 

Cynomys  parvidens  Allen,  Prairie  Dog. 
Presnall  (1934)  reported  two  small  prairie 
dog  towns  in  Bryce  Canyon,  "one  near  Fairy- 
land (on  the  rim)  and  another  about  two 
miles  farther  north.  A  total  of  about  two  doz- 
en occupied  burrows."  Tanner  (1940a)  re- 
ported this  species  for  the  Kaiparowits  Basin. 

Tamiasciurus  hudsonicus  dixiensis  Hardy. 
Red  squirrel.  Presnall  (1934,  1937)  indicates 
that  this  species  was  quite  common  in  Bryce 
Canyon  during  the  1920s.  During  1933  he 
observed  only  two  families  and  six  in  1934;  in 
1935,  however,  they  were  quite  numerous. 

Glaucomys  sobrinus  lucifugus  Hall. 
Northern  Flying  Squirrel.  This  species  was 
collected  10  miles  southwest  of  Bryce  Can- 
yon by  Lowell  Hansen  and  reported  by  Tan- 
ner (1940b).  Russell  and  Thompson  (1964) 
listed  it  as  a  rare  species  in  Bryce  Canyon. 

Geomyidae  (Pocket  Gophers) 

Thomomys   bottae    (Eydoux    &    Gervais). 

Botta  Pocket  Gopher.  Benson  (1935)  referred 
the  material  described  by  Goldman  (1937)  to 
T.  h.  alexandrc  Goldman  based  on  material 
collected  on  Navajo  Mountain.  According  to 
Durrant  (1952)  and  Cockrum  (1960)  this  sub- 
species is  known  only  from  Navajo  Mountain 
and  tlie  vicinity  south  of  the  San  Juan  and 
east  of  the  Colorado  River.  Thomomys  h.  ah- 
sonus  Goldman,  according  to  Durrant  (1952), 
occurs  only  on  the  west  side  of  the  Colorado 
and  lists  a  specimen  from  Escalante.  Our 
(BYU  1971)  only  collection  is  from  Site  3  on 
Cedar  Mountain.  However,  their  tunnels  are 
scattered  throughout  the  region.  Presnall 
(1934)  reported  that  pocket  gophers  were 
abundant  in  Bryce  Canyon  and  are  probably 
referable  to  the  latter  subspecies. 

Heteromyidae  (Kangaroo  Rats,  Pocket  Mice) 

Perognathiis  flavus   hopiensis    Goldman. 

Silky  Pocket  Mouse.  Benson  (1935)  indicates 
that  four  specimens  were  collected  five  miles 
southeast  of  Navajo  Mountain  Trading  Post 
and  tliree  from  the  environs  of  the  trading 
post.  Durrant  (1952)  says  they  are  confined  to 
the  region  east  of  the  Colorado  River  in 
loose,  sandy,  sparsely  vegetated  areas. 


Perognathus  apache  Merriam.  Apache 
Pocket  Mouse.  Benson  (1935)  reports  three 
collections  of  P.  a.  apache  Merriam  from  the 
vicinity  of  Navajo  Mountain.  According  to 
Durrant  and  Dean  (1959)  and  Cockrum 
(1960),  this  subspecies  is  restricted  to  the  area 
south  of  the  San  Juan  River  on  the  east  side 
of  the  Colorado.  P.  a.  caryi  Goldman  is  re- 
stricted to  the  east  side  of  the  Colorado  River 
above  the  San  Juan  River  (Durrant  and  Dean 
1959).  The  only  specimens  of  this  subspecies 
from  the  Kaiparowits  Basin  are  those  collect- 
ed by  BYU  (1972)  at  Three  Garden  above  the 
confluence  of  the  San  Juan/Colorado  Rivers. 

Perognathus  longimembris  (Coues).  Little 
Pocket  Mouse.  This  species  is  represented  in 
the  Kaiparowits  Basin  by  two  subspecies,  one 
occurring  on  each  side  of  the  Colorado  River. 
PerognatJius  I.  arizonensis  Goldman  is  re- 
stricted to  the  west  side  of  the  river  and  was 
reported  by  Durrant  and  Dean  (1959)  from 
Kane  Creek  and  at  river  mile  34.  Our  records 
(BYU  1971-1974)  are  from  Sites  1,  2,  3,  4,  5, 
6,  7,  8,  9,  10,  11,  12,  13,  14,  15,  17,  18,  23, 
27,  28,  and  30.  Benson  (1935)  described  P.  I. 
acrus  (Benson)  from  material  collected  at 
Rainbow  Bridge  (type  #58624).  Durrant  and 
Dean  (1959)  took  one  specimen  from  river 
mile  43.  Our  records  (BYU  1971-1972)  are 
from  Sites  19,  20,  21,  and  22;  all  are  east  of 
the  Colorado  River. 

Perognathus  ampins  ammodytes  Benson. 
Arizona  Pocket  Mouse.  Cockrum  (1960)  in- 
dicates that  this  species  has  been  taken  along 
the  Echo  Cliffs  which  is  the  southern  bound- 
ary line  for  the  Kaiparowits  Basin. 

Perognathus  formosus  domisaxensis  Cock- 
rum. Long-tailed  Pocket  Mouse.  Durrant  and 
Dean  (1959)  collected  four  specimens  from 
Kane  Creek,  i.e.,  one  from  river  mile  34  and 
three  at  Lee's  Ferry,  and  indicated,  "Ecologi- 
cally, these  pocket  mice  differ  somewhat 
from  those  kinds  farther  west.  They  appear 
to  favor  more  rocky  areas  and  occur  on  hill- 
sides and  at  the  junction  of  hillsides  and  stabi- 
lized terraces."  Our  records  (BYU 
1971-1974)  are  from  Sites  4,  9,  10,  15,  27,  28, 
and  30.  In  addition,  material  from  Three 
Garden  located  one  mile  north  of  the  con- 
fluence of  the  San  Juan  on  the  east  side  of  the 
Colorado  River  has  been  tentatively  referred 
to  this  species.  When  additional  material  is 


December  1980 


Atwood  et  al.:  Kaiparowits  Vertebrates 


343 


obtained  and  analyzed,  it  may  prove,  how- 
ever, to  be  P.  intennedius  crinitus. 

Perognathus  intermedius  criyiittis  Benson. 
Rock  Pocket  Mouse.  .According  to  Durrant 
and  Dean  (1959),  this  subspecies  is  confined 
to  the  east  side  of  the  Colorado  River  south 
of  the  San  Juan  River.  No  specimens  were 
obtained  by  them  until  they  reached  Aztec 
Creek  at  river  mile  68.5.  Between  this  point 
and  river  mile  28  they  collected  15  speci- 
mens. Benson  (1934b)  reported  a  collection 
from  Navajo  Mountain  Trading  Post  and  two 
specimens  from  Rainbow  Bridge.  Cockrum 
(1960)  obtained  specimens  from  Lee's  Ferry. 
BYU  (1973)  obtained  specimens  from  Three 
Garden  near  the  confluence  of  the  San  Juan 
and  Colorado  River  Canyons.  These  speci- 
mens document  the  range  of  this  species 
north  of  the  San  Juan  River. 

Dipodotnys  ordii  Woodhouse.  Kangaroo 
Rat.  Two  subspecies  occur  in  the  Kaiparowits 
Basin  and,  of  the  two,  D.  o.  cupidineus  Gold- 
man is  restricted  to  the  west  side  of  the  Colo- 
rado south  of  the  Escalante  River.  Specimens 
have  been  collected  by  Durrant  and  Dean 
(1959)  at  Lee's  Ferry  and  Kane  Creek;  Pres- 
nall  (1934)  for  Bryce;  Tanner  (1940a)  for  the 
Kaiparowits  Basin;  and  BYU  (1971-1974)  for 
Sites  1,  2,  3,  4,  6,  7,  8,  10,  13,  14,  15,  17,  27, 
28,  30,  and  33. 

Dipodcnnys  o.  longipes  (Merriam)  is  con- 
fined to  the  area  south  of  the  San  Juan  River 
and  east  of  the  Colorado  (Durrant  and  Dean 
1959).  Benson  (1935)  reports  nine  collections 
from  south  of  Navajo  Mountain.  Our  records 
(BYU  1971-1972)  are  from  Sites  19  and  33. 

Castoridae  (Beaver) 

Castor   canadensis    repetitintis    Goldman. 

Beaver.  Prior  to  the  coming  of  the  white 
trappers,  beavers  were  widespread  through- 
out the  length  of  the  Colorado  River  and  its 
tributaries.  Tliis  abundance  of  beaver  enticed 
the  famous  French  trapper  and  fur  trader 
Denis  Julien  to  navigate  the  Colorado  in 
1836  as  far  south  as  Cataract  Canyon.  The 
best-known  trapper  in  the  Canyon  was  Nath- 
an Galloway;  in  1895,  1896,  and  on  other  oc- 
casions he  traveled  from  (ireen  River  to 
Lee's  Ferry  with  William  Richmond  (Cramp- 
ton  1959).  Durrant  and  Dean  (1959)  reported 
that  the  populations  in  Glen  Canyon  were 


among  the  largest  in  the  state.  During  the 
course  of  their  expedition,  30  June  to  9  .Au- 
gust 1958,  they  observed  beaver  or  their  signs 
daily.  They  indicated  that  these  bank 
dwellers  fed  almost  entirely  on  willows  and 
only  on  one  occasion  was  Tamarix  utilized. 
The  creation  of  Lake  Powell  has  nearly  elim- 
inated their  main  source  of  food  and  greatly 
reduced  their  numbers.  Observations  by  BYU 
field  crews  (1971-1974)  indicate  that  beavers 
utilize  representatives  of  nearlv  every  woody 
species  in  the  drainages  of  the  Colorado. 
They  have,  of  necessity,  been  pushed  into  the 
side  canyons,  particularlv  where  permanent 
springs  or  seeps  exist.  It  appears  that  they 
have  attempted  to  build  dams  in  some  of 
these  regions  to  create  a  more  suitable  habi- 
tat. 


Cricetidae  (Voles,  Rats,  Mice) 

Reithrodontomys  megalotis  (Baird).  West- 
ern Harvest  Mouse.  The  subspecies  R.  m. 
megalotis  (Baird)  is  restricted  to  the  west  side 
of  the  Colorado  River  in  the  Glen  Canyon 
area.  Cockrum  (1960)  collected  it  at  Lee's 
Ferry;  Durrant  and  Dean  (1959)  collected 
specimens  at  river  mile  43  and  93;  and  BYU 
(1971-1974)  collected  it  at  Sites  2,  3,  and  10. 
Durrant  and  Dean  (1959)  collected  R.  m.  az- 
tecus  J.  A.  Allen  on  the  east  side  of  the  Colo- 
rado at  river  miles  43  and  83. 

Peromyscus  eremicus  eremicus  (Baird). 
Cactus  Mouse.  Cockrum  (1960)  reported  it 
from  the  south  side  of  the  Colorado  at  Lee's 
Ferry. 

Peromyscus  tnaniculatus  (Wagner).  Deer 
Mouse.  Durrant  and  Dean  (1959)  have  estab- 
lished that  P.  m.  sonoriensis  (LeConte)  is  the 
onlv  subspecies  of  Deer  Mouse  in  Glen  Can- 
von  and  that  it  was  the  most  common  mam- 
mal in  Glen  Canyon  during  their  study.  They 
collected  specimens  from  Lee's  Ferry  to  river 
mile  78  on  the  Colorado.  Presnall  (1934)  re- 
ported this  subspecies  from  Bryce  Canyon; 
Benson  (1935)  indicated  that  it  was  the  most 
abundant  mammal  in  the  region  of  Navajo 
Mountain;  and  BYU  (1971-1973)  collected  it 
at  Sites  1,  2,  3,  6,  7,  8,  15,  19,  22,  27,  28,  and 
30,  and  Navajo  Mountain. 

Peromyscus  crinitus  Merriam.  Canyon 
Mous.  Two  subspecies  of  the  Canyon  Mouse 
occur    in   the   Kaiparowits   Basin,   viz.   P.   c. 


344 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


auripectus  (Osgood)  and  P.  c.  stephensi 
(Meam.s).  Durrant  and  Dean  (1959)  noted 
that  subspecies  auripectus  occured  only  on 
the  east  side  of  Glen  Canyon  and  collected 
specimens  at  river  miles  21,  28,  69,  78,  and 
83.  Benson  (1935)  reported  one  from  Rain- 
bow Bridge  and  Navajo  Mountain  Trading 
Post.  Additional  records  are:  BYU  (1972)  Site 
19,  BYU  (1971-1974)  Three  Garden  above 
confluence  of  the  San  Juan  and  Lake  Powell, 
and  Cockrum  (1960)  at  Lee's  Ferry  and  Nav- 
ajo Mountain. 

Peromyscus  c.  stephensi,  as  reported  by 
Durrant  and  Dean  (1959),  occurs  throughout 
the  Kaiparowits  Basin  west  of  the  Colorado 
River.  They  made  collections  at  river  miles 
23,  41,  56,  and  88,  and  at  Lee's  Ferry;  BYU 
(1971)  collected  this  subspecies  at  Sites  5,  6, 
8,  and  12;  BYU  (1972)  also  collected  it  at 
Sites  23,  27,  and  30. 

Peromyscus  boylii  (Baird).  Brush  Mouse. 
Two  subspecies  occur  in  tlie  Kaiparowits  Ba- 
sin, viz.  P.  b.  roioleyi  (Allen)  and  P.  b.  uta- 
hensis  Durrant.  The  former  species  is  re- 
stricted to  the  east  side  of  the  Colorado  and 
has  been  collected  at  river  miles  28,  43,  69, 
78,  and  83  by  Durrant  and  Dean  (1959). 
Cockrum  (1960)  cites  a  specimen  from  Rain- 
bow Lodge  at  the  southwest  base  of  Navajo 
Mountain  and  BYU  (1972-1974)  cites  speci- 
mens from  Three  Garden  near  the  San  Juan 
confluence  and  at  Pool  Garden  in  Reflection 
Canyon. 

The  latter  subspecies,  Peromyscus  b.  uta- 
Jiensis,  occurs  only  on  the  west  side  of  the 
Colorado  and  has  been  collected  by  Durrant 
and  Dean  (1959)  from  river  miles  34,  41,  and 
56  and  by  BYU  (1971)  from  Site  8  and  from 
Reflection  Canyon.  Durrant  and  Dean  in- 
dicate that  this  subspecies  was  nearly  as  com- 
mon as  the  deer  mice.  They  are  reported  to 
be  excellent  climbers,  preferring  brushy  habi- 
tats among  cliffs  and  rockv  areas. 

Peromyscus  truei  (Shufeldt).  Pinyon 
Mouse.  Benson  (1935)  reported  a  collection 
from  the  mesa  top  near  Navajo  Mountain  and 
one  from  the  Trading  Post.  Presnall  (1934) 
reported  this  species  from  Bryce  Canyon. 
During  the  course  of  our  study  (BYU 
1971-1974),  we  have  records  for  this  species 
from  Sites  2,  3,  6,  1.3,  14,  19,  20  21,  23,  27, 
28,  and  30,  and  Navajo  Mountain. 


Peromyscus  difficilis  nasutus  (Allen)  ( = 
Peromyscus  nasutus  nasutus  Allen).  Long- 
nosed  Deer  Mouse.  Benson  (1935)  reported 
this  species  from  Rainbow  Bridge  in  rocky 
places  where  stands  oi  Juniperiis  occur. 

Onychomys  leucogaster  (Wied-Neuwied). 
Northern  Grasshopper  Mouse.  Durrant 
(1952)  indicates  that  O.  I.  melanophrys  Mer- 
riam  is  confined  to  the  west  side  of  the  Colo- 
rado River.  Presnall  (1934)  reported  it  from 
Bryce  Canyon  and  BYU  (1971-1974)  from 
Sites  1,  2,  3,  4,  6,  8,  14,  17,  18,  and  .30;  O.  /. 
pallescens  Merriam  is  confined  to  the  east 
side  of  the  Colorado  River  (Durrant  1952). 
Available  records  are:  Benson  (1935)  several 
specimens  from  5  and  7  miles  south  of  Nav- 
ajo Mountain  and  BYU  (1971-1972)  from 
Sites  19,  20,  and  22.  One  specimen  (No.  181 
from  Nipple  Bench)  collected  by  BYU  (1971) 
appears  to  be  an  intergrade  between  these 
two  subspecies. 

Neotoma  albigula  laplataensis  Miller, 
White-throated  Wood  Rat.  Durrant  and 
Dean  (1959)  indicate  that  this  species  is  con- 
fined to  the  east  side  of  the  Colorado  River  in 
Glen  Canyon.  Collections  were  made  at  river 
miles  69  and  78.  Benson  (1935)  reported  a 
collection  from  Rainbow  Bridge  and  the 
mesa  top  south  of  Navajo  Mountain.  Studies 
by  BYU  (1971-1974)  in  Glen  Canyon  and  vi- 
cinity confirm  the  observations  of  Durrant 
and  Dean.  In  addition,  collections  by  BYU 
(1971-1972)  were  made  at  Sites  19  and  21. 

Neotoma  lepida  monstabilis  Goldman. 
Desert  Wood  Rat.  Durrant  and  Dean  (1959) 
state  that,  "members  of  this  species  are  the 
counter  parts  of  N.  albigula  and  A',  mexicana 
of  western  side  of  the  Colorado  River."  Col- 
lections of  N.  /.  monstrabilis  were  made  at 
river  miles  41,  56,  88,  and  91.  Additional  re- 
cords for  the  Kaiparowits  Basin  are:  Presnall 
(1934)  Bryce;  Tanner  (1940a)  Kaiparowits 
Basin;  Pritchett  (1962)  Cockscomb;  and  BYU 
(1971)  Sites  1,  2,  3,  4,  6,  8,  12,  and  13,  Step 
Garden,  and  Reflection  Canyon. 

Neotoma  stephensi  relicta  Goldman. 
Stephen's  Wood  Rat.  Benson  (1935)  reported 
one  collection  from  Rainbow  Bridge,  four 
specimens  from  south  of  Navajo  Mountain  on 
tlie  mesa  top,  and  four  from  Navajo  Moun- 
tain Trading  Post.  We  have  not  collected  this 
species  at  any  of  our  sites.  ^ 


December  1980 


Atwood  et  al.:  Kaiparowits  Vertebrates 


345 


Neotoma    mexicana    inopinata    Goldman. 

Mexican  Wood  Rat.  This  species  is  restricted 
to  the  east  side  of  Glen  Canyon  (Durrant  and 
Dean  1959).  Specimens  were  obtained  bv 
them  from  river  miles  69,  78,  and  83.  Benson 
(1935)  reported  it  from  War  (iod  Spring  on 
Navajo  Moimtain.  BYU  (1972-1974)  record 
are  from  Three  Garden  just  above  the  con- 
fluence of  the  San  Juan  on  the  east  side  of  the 
Colorado  River  and  from  Ribbon  Garden  in 
Ribbon  Can  von. 

Neotoma  ciyierea  arizonae  Merriam.  Bush- 
tailed  Wood  Rat.  According  to  Durrant  and 
Dean  (1959),  N.  cinerea  is  the  onlv  species  of 
wood  rat  that  occiu-s  on  both  sides  of  Glen 
Canyon.  Neotoma  c.  arizonae  occurs  only  on 
the  east  side  and  N.  c.  acraia  on  the  west  side. 
However,  no  specimens  of  the  latter  sub- 
species were  reported  bv  Durrant  and  Dean. 
They  did  collect  specimens  of  A\  c.  arizonae 
from  river  miles  21,  43,  and  83.  Additional 
records  for  the  Kaiparowits  Basin  are:  BYU 
(1971-1972)  Navajo  Mountain,  Site  27,  and 
Site  33. 

Microtus  mexicanus  navajo  Benson.  Mexi- 
can Vole.  Benson  (1934a  and  1935)  described 
this  species  from  specimens  obtained  from 
Soldier  and  War  God  Springs  on  the  east 
slope  of  Navajo  Mountain.  Thev  were  col- 
lected in  a  Ceanothus,  Symplioricarpos, 
Arctostaphylos,  and  Rosa  vegetation  type. 
This  taxon  is  known  only  from  the  type  col- 
lections. 

Lagurus  curtatiis  intermedins  (Taylor), 
Sagebrush  Vole.  Presnall  (1934)  reported  a 
specimen  from  Bryce  Canyon,  but  was 
unable  to  assign  a  species  name.  Durrant 
(1952)  cites  a  specimen  of  this  species  from 
David  Hollow  in  Bryce  Canyon  that  may  be 
the  one  cited  by  Presnall.  We  did  not  en- 
counter this  species  at  any  of  our  trap  sites. 

Erethizontidae  (Porcupine) 

Erethizon  dorsatum  (Linnaeus).  Porcu- 
pine. According  to  Durrant  (1952)  two  sub- 
species occur  in  the  Kaiparowits  Basin,  viz. 
£.  d.  epixanthnm  Brandt  on  the  west  side  of 
the  Colorado  River  and  E.  d.  consei  Mearns 
on  the  east  side.  Durrant  and  Dean  (1959)  re- 
ported that  expedition  members  observed 
one  animal  in  Navajo  Canyon.  Weight  (1932) 
and  Presnall  (1934)  reported  the  former  sub- 


species from  Bryce  Canyon  and  BYU 
(1971-1972)  reported  it  from  Sites  2,  3,  and 
4,  and  Reflection  Canyon.  Their  signs  are  not 
uncommon  in  stands  of  Finns  ednlis. 


Canidae  (Coyotes,  Foxes,  and  Wolves) 

Canis  latrans  Say.  Coyote.  Two  subspecies 
occur  in  the  Kaiparowits  Basin,  i.e.,  C.  /.  estor 
Merriam  and  C.  /.  lestes  Merriam.  The  for- 
mer is  by  far  the  most  conunon  reported  bv 
Tanner  (1940a)  for  the  Kaiparowits  Plateau; 
bv  BYU  (1971)  for  Sites  3,  10,  11,  12,  15,  and 
Wahweap  Creek;  by  BYU  (1972)  for  Site  14, 
Butler  Valley,  Cottonwood  Wash,  Covote 
Creek,  near  Glen  Canvon,  and  Warm  Creek; 
and  by  BYU  (1973)  for  Sites  3,  6,  8,  23,  27, 
and  28,  and  Smoky  Mountain.  The  first  re- 
cord of  this  species  in  Glen  Canyon  was 
made  by  members  of  Powell's  first  Colorado 
River  expedition  in  1869.  While  in  the  vicin- 
ity of  the  Escalante  River,  Sumner  (Powell 
1875)  records  as  follows,  "Dunn  killed  a  half- 
stayed  coyote  near  camp,  the  only  sign  of  an- 
imal life  we  have  seen  for  three  davs."  Dur 
rant  and  Dean  (1959)  observed  numerous 
signs  of  coyotes  in  Glen  Canyon  proper  and 
in  several  side  canvons.  The  senior  author,  in 
company  with  S.  L.  Welsh  and  J.  R.  Mur- 
dock,  witnessed  in  1971  the  fall  and  death  of 
a  coyote  from  a  200-foot  cliff  in  Last  Chance 
Creek. 

Canis  I.  lestes  is  known  only  from  the 
northern  portion  of  the  Kaiparowits  Basin  as 
reported  bv  Presnall  (1934)  from  Bryce  Can- 
yon. BYU  (1973)  observed  coyotes  north  of 
Canaan  Mountain  along  High\\a\  12  and 
these  may  be  C.  /.  lestes. 

Canis  lupus  youngi  Goldman.  Wolf.  .\c- 
cording  to  Young  and  Goldman  (1944)  and 
Cockrum  (1960),  the  wolf  formerly  occurred 
throughout  this  region,  but  no  specimens 
were  cited.  It  is  doubtful  that  it  occurs  in  the 
Basin  at  the  present  time. 

Vulpes  vulpes  maeroura  Baird  (  =  Vidpes 
fulva  maeroura  Baird).  Red  Fox.  .\t  present 
the  red  fox  is  rare  in  Utah  due  to  hunting, 
trapping,  and  use  of  poison  bait  stations.  Sev- 
eral records  were  reported  prior  to  1940. 
Presnall  (1934)  reported  that  this  animal  was 
occasionally  seen  at  Bryce  Canyon;  Benson 
(1935)  reported  one  from  the  north  side  of 
Navajo  Mountain;  Durrant  and  Dean  (1959) 


346 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


indicate  that  one  was  seen  two  miles  east  of 
Rainbow  Bridge  by  a  member  of  their  expe- 
dition. Durrant  (1952)  cites  one  from  Wah- 
weap  Creek  and  states,  "possibly  the  red  fox 
of  the  Colorado  River  is  an  undescribed  sub- 
species . .  .  and  I  tentatively  refer  them  to  V. 
/.  macroiira." 

Urocyon  cinereoargenteus  scottii  Mearns. 
Gray  Fox.  The  gray  fox  occurs  throughout 
the  Basin  and  has  been  reported  by  Presnall 
(1934)  for  Bryce  Canyon  and  Benson  (1935) 
for  Navajo  Mountain;  and  Hayward  et  al. 
(1958)  reported  their  signs  from  nearly  all 
areas  studied  from  the  Paria  to  the  Escalante 
River.  BYU  (1971-1974)  observed  this  animal 
in  Cottonwood  Wash,  Cockscomb,  and  on 
Brigham  Plains.  Tliis  species  is  much  more 
common  than  the  Red  Fox. 

Ursidae  (Bears) 

Ursus  americanus  cinnamomum  Audubon 
&  Bachman.  Black  Bear.  Presnall  (1934)  in- 
dicated that  in  1922  a  black  bear  was  killed 
in  Bryce  Canyon;  and  "Mr.  Ruby  Syrett  and 
Ranger  Cope  both  say  that  black  and  grizzly 
bears  used  to  cross  the  Bryce  Canyon  region 
at  irregular  intervals,  apparently  traveling 
between  the  Parowan  Moimtains  on  the  west 
and  the  Escalante  Mountains  on  the  east. 
Bears  are  still  found  in  both  ranges,  although 
rarely."  One  of  us  (Atwood)  observed  in  1965 
tracks  of  a  bear  on  the  south  slope  of  the 
Boulder  Mountains.  On  11  June  1975,  tracks 
of  a  solitary  bear  were  observed  by  S.  L. 
Welsh  at  the  head  of  Paradise  Canyon  about 
11  miles  north  of  Horse  Mountain  junction. 
The  left  hind  foot  track  measured  10.2  X 
22.9  cm.  In  late  June  1975,  a  small  bear  was 
sighted  at  Canaan  Peak  by  BYU  personnel. 

Ursus  horribilis  Merriam.  Grizzly  Bear. 
Presnall  (1934)  indicates  that  they  have 
crossed  through  Bryce.  For  additional  infor- 
mation the  reader  is  referred  to  the  discussion 
of  the  preceding  species. 

Procyonidae  (Ring-tailed  Cat,  Raccoon) 

Bassarisciis  astutus  (Lichtenstein).  Ring- 
tailed  Cat.  According  to  Durrant  (1952)  the 
Colorado  River  serves  as  a  barrier  for  two 
subspecies.  A  specimen  of  subspecies  B.  a. 
arizonensis  Goldman  was  obtained  from  W. 


Wilson,  which  he  had  shot  in  a  chicken  coop 
at  Rainbow  Lodge  in  November  1932  (Ben- 
son 1935).  Wetherill,  Flattum,  and  Stearns 
(1961)  recorded  in  their  journal  on  15  Janu- 
ary 1931  that  a  ring-tailed  cat  came  into 
camp.  They  were  in  or  near  Bridge  Canyon. 
The  entry  by  John  Wetherill  on  17  January 
states,  "Lots  of  skunks  around,  some  ring- 
tailed  cats,  fox.  .  .  ."  Bassariscus  a.  nevadensis 
Miller  has  been  reported  by  trappers  and  lo- 
cal residents  of  Kaiparowits  Basin  to  be  fairly 
common.  Durrant  and  Dean  (1959)  observed 
small  five-toed  tracks  in  the  talus  dust  and 
along  the  base  of  ledges  throughout  Glen 
Canyon.  During  the  course  of  our  study  we 
have  not  met  directly  with  this  animal,  but 
have  observed  their  tracks  in  the  major  drain- 
ages. 

Procyon  lotor  pallidus  Merriam.  Raccoon. 
Hall  and  Kelson  (1959)  list  this  subspecies  as 
occurring  throughout  the  Colorado  and  its 
tributaries.  Durrant  and  Dean  (1959),  how- 
ever, were  unable  to  find  any  evidence  to 
verify  their  presence  in  the  Colorado  drain- 
age. Observations  by  BYU  (1971-1974)  con- 
firmed the  conclusions  of  Durrant  and  Dean. 


Mustelidae  (Weasels,  Skunks,  etc.) 

Mustela  erminea  muricus  (Bangs).  Ermine. 
Durrant  (1952)  reports  one  specimen  from 
Boulder  Mountain  north  of  Escalante.  This 
species  probably  occurred  throughout  the 
Boulder  Mountains  and  probably  in  the  re- 
gion of  Bryce  Canyon 

Mustela  frenata  nevadensis  Hall,  Long- 
tailed  Weasel.  Presnall  (1934)  reported  this 
species  was  common  at  Bryce  Canyon.  Ben- 
son (1935)  indicated  that  one  was  observed  at 
Rainbow  Lodge  in  1932.  The  Long-tailed 
Weasel  is  probably  the  most  widely  dis- 
tributed carnivore  in  Utah  (Durrant  1952). 

Taxidea  taxus  berlandieri  Baird.  Badger. 
Presnall  (1934)  indicated  that  they  were  com- 
mon on  the  rim  of  Brvce  Canvon.  Benson 
(1935)  observed  fragments  of  badger  taken  by 
trappers  in  the  Navajo  Mountain  region.  Tan- 
ner (1940a)  reported  them  from  the  Kaiparo- 
wits Plateau.  Durrant  and  Dean  (1959)  in- 
dicated that  this  species  was  one  of  the  most 
common  and  abundant  carnivores  in  Glen 
Canyon.  They  observed  their  tracks  and  bur- 
rows    everywhere.     Our     records     (BYU 


December  1980 


Atwood  et  al.:  Kaiparowits  Vertebrates 


347 


1971-1974)  indicate  their  presence  at  Sites  1, 
3,  13,  19,  27,  and  34,  and  in  every  drainage  in 
the  basin. 

Spilogale  gracilis  Merriam.  Spotted  Skunk. 
The  Colorado  River  is  a  barrier  for  kiteral 
movement  of  this  animal  with  S.  g.  gracilis 
Merriam  confined  to  the  east  side.  It  was  re- 
ported by  Benson  (1935)  for  the  Navajo 
Mountain  area.  S.  g.  saxatilis  (Merriam)  is  re- 
ported by  Presnall  (1935)  from  Bryce  Can- 
yon. BYU  (1972)  noted  their  occurrence  on 
Cedar  Mountain  south  of  Glen  Canyon  City 
and  in  Little  Valley  Creek  at  the  south  base 
of  the  Kaiparowits  Plateau.  Durrant  and 
Dean  (1959)  observed  signs  of  this  species  on 
both  sides  of  the  Colorado  River,  and  one 
was  observed  by  investigators  from  BYU  at 
Three  Garden  on  the  east  side  of  Glen  Can- 
yon. 

Mephitis  mephitis  estor  Merriam.  Striped 
Skunk.  Presnall  (19.34)  indicates  that  this  spe- 
cies was  rare  at  Bryce  Canyon.  Durrant 
(1952)  reports  that  the  subspecies  M.  m.  estor 
is  limited  to  the  Colorado  River  drainage  in 
.southern  and  eastern  Utah.  Specimens  are 
cited  for  both  San  Juan  and  Washington 
Counties,  but  are  not  cited  for  the  Kaiparo- 
wits Basin. 

Lontra  canadensis  nexa  Goldman.  River 
Otter.  Gregory  (1938)  reports  otter  from 
Glen  Canyon.  Durrant  and  Dean  (1959)  ques- 
tion his  recording  of  the  species  in  Glen  Can- 
yon, but  do  not  discard  the  possibility  entire- 

iv. 


Felidae  (Cats) 

Felis  concolor  kaihabensis  Nelson  &  Gold- 
man. Moimtain  Lion.  Benson  (1935)  indicates 
that  an  animal  was  killed  near  Inscription 
House  by  Navajos.  Presnall  (1934)  reported 
that  an  occasional  lion  wanders  through 
Bryce  Canyon.  Observations  of  this  species 
were  made  by  BYU  (1971  and  1973)  in 
Nipple  Creek,  Tibbet  Canyon,  and  Willow 
Creek.  The  observation  in  Willow  Creek  was 
a  female  with  two  kittens.  In  addition,  a 
solid-black-colored  lion  was  observed  Octo- 
ber 1976  on  Cannon  Peak. 

Lynx  rufus  baileyi  Merriam.  Bobcat.  Ap- 
parently the  only  specimens  available  from 
the  Kaiparowits  Basin  are  those  collected  bv 
BYU  (1973)   in  Tibbet  Canyon,   Buckskin 


Gulch,  and  Cottonwood  Canyon.  Tracks  and 
scat  have  been  observed  throughout  the  re- 
gion. 

Cervidae  (Deer  and  Elk) 

Cervus  elaphus  nehoni  V.  Bailey.  Wapiti, 
Elk.  Cope  (1932)  reported  that  A.'w.  Ivins, 
an  early  resident  of  this  region,  informed  him 
that,  according  to  the  Indians,  elk  once  re- 
sided in  Bryce  Canyon  and  the  Paria  Valley. 
Mr.  Amnion  Davis,  of  Cannon ville,  informed 
him  that  he  had  found  part  of  the  head  and 
horns  of  a  bull  elk  on  Willis  Creek  just  south- 
east of  Bryce  Canyon  Rim.  This  material  is 
now  in  the  museum  collection  of  Brvce.  Pre- 
snall (1934)  reports  further  on  the  elk  of 
Bryce-Paria  Valley  as  follows:  "now  extinct 
in  this  region,  but  one  was  reported  killed  in 
Willis  Creek  many  years  ago  bv  a  Mr.  John- 
ston of  Cannonville." 

There  is  some  question  as  to  which  is  the 
correct  name,  i.e.,  Cervus  canadensis  nelsoni 
y.  Bailey  or  Cervus  elaphus  L.  Since  the 
question  has  yet  to  be  resolved,  this  treat- 
ment follows  Durrant  (1952). 

Odocoileus  hemionus  hemionus  (Rafi- 
nesque).  Mule  Deer.  No  records  of  deer  in 
the  Kaiparowits  were  made  by  early  explor- 
ers of  the  region.  Pre.snall  (1934)  indicated 
that  they  were  common  in  Bryce  Canyon. 
During  the  course  of  our  study  (BYU 
1971-1974)  frequent  sightings  have  been 
made,  especially  along  tributary  canyons 
along  Lake  Powell,  .\nimals  have  been  ob- 
served onlv  where  water  and  forage  are  ade- 
quate throughout  the  year  and  they  appear 
to  be  residents.  The  main  areas  of  concentra- 
tion (sens,  lat.)  are  on  Four  Mile  Bench, 
Cockscomb.  Canaan  Mountain,  Bryce  Can- 
yon, and  the  Kaiparow  its  Plateau. 

Antilocapridae  (Pronghom) 

Antilocapra  americana  americana   (Ord). 

Pronghorn.  Tanner  (1940a)  indicates  tliat 
pronghorn  antelope  were  common  in  the 
Kaiparowits  Basin  in  pioneer  days.  Presnall 
(1938)  also  reported  their  occurrence  at 
Brvce  Canyon.  Apparently  the  early  herds 
were  himted  too  severely  because  records 
since  the  beginning  of  this  century  are  lack- 
ing. 


348 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


In  November  1971  some  104  antelope 
were  transplanted  from  northeastern  Utah  to 
East  Clark  Bench,  10  miles  west  of  Glen  Can- 
yon City,  Utah.  Since  then,  40  separate  sight- 
ings of  these  animals  have  been  recorded  by 
personnel  of  the  U.S.  Bureau  of  Land  Man- 
agement and  BYU  (1971-1974).  These  sight- 
ings, for  the  most  part,  have  been  in  Cotton- 
wood Wash,  East  and  West  Clark  Benches, 
Coyote  Creek,  and  as  far  north  as  Butler  Val- 
ley and  on  Nipple  Bench.  Sightings  were  in- 
frequent in  1974  and  even  fewer  in  1975;  in 
1976  a  pair  was  known  to  range  in  the  vicin- 
ity of  Nipple  Spring. 

Bovidae  (Bovids) 

Ovis  canadensis  canadensis  Shaw,  Moun- 
tain Sheep.  Powell's  party  made  several  ref- 
erences to  these  animals  and  occasionally 
used  them  for  food.  On  27  July  1869,  Sumner 
recorded  (Powell  1875),  "Killed  two  moun- 
tain sheep  today— a  Godsend  to  us,  as  our 
bread  and  rotten  bacon  is  a  poor  diet  for  as 
hard  work  as  we  have  to  do."  Then  on  3  Au- 
gust below  Music  Temple,  Sumner  recorded, 
"pulled  out  early,  made  a  good  run.  Saw  two 
mountain  sheep  in  a  little  valley  on  the  south 
side.  How  they  got  there  I  will  leave  others 
to  judge,  as  there  is  no  outlet  to  the  valley 
that  a  man  can  climb.  Killed  one  and  chased 
the  other  through  the  natiual  pasture  for  an 
hour  and  pulled  out  again."  Gregory  and 
Moore  (1931)  reported  sheep  from  Dark  Can- 
yon at  the  southeast  base  of  the  Kaiparowits 
Plateau. 

Durrant  (1952)  indicates  that  the  species  is 
disappearing  rapidly,  and  if  adequate  speci- 
mens from  extreme  south  central  Utah  are 
available  for  study,  they  would  probably 
prove  to  be  O.  c.  mexicanus. 

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l^ESNALL,  C.  C.  1933.  Along  nature's  highwa\s.  Zion 
and  Bryce  Nat.  Notes  6(6):52-57. 

1934.  Animals  and  birds  of  Bryce  Canyon  Nation- 
al Park.  Zion  and  Brvce  Nat.  Notes  6(6):52-57. 

19.35.  The  reptiles  of  Zion  and  Brvce  Canvon  .Na- 
tional Parks.  Zion  and  Bryce  Nat.  Notes 
8(4):28-31. 

. 19.36.  Abundance  of  redbreasted  nutcatchers  in 

southeastern  Utah.  Condor  38(l):44-45. 

19.37.  Three  new  records  from  Bryce  Canyon, 

Utah.  Condor  39:259. 

1938.    Mammals    of   Zion,    Bryce,    and    Cedar 

Breaks.  Zion  and  Bryce  Mus.  Bull.  No.  2:1-20. 

Pritchett,  C.  L.  1962.  Vertebrate  distribution  in  rela- 
tion to  certain  habitats  in  central  Kane  County, 
Utah.  Unpublished  thesis,  Brigham  Young  Uni- 
versity, Provo,  Utah.  83  pp. 

Russell,  R.  W.,  and  H.  E.  Thompson.  1964.  .\  listing  of 
the  common  plants  and  vertebrate  animals  of 
Bryce  Canyon  National  Park.  Unpublished  manu- 
script. Bryce  Canyon,  Utah. 

Schmidt,  K.  P.  1953.  Check-list  of  North  American  am- 
phibians and  reptiles.  University  of  Chicago 
Press,  Chicago.  280  pp. 


S.Now.  I).  W,  1967a.  Pages  84-85  (Vol.  12)  in  R.  A.  Payn- 
ter,  Jr..  ed.,  Peter's  check-list  of  the  birds  of  the 
world.  Museum  of  Comparative  Zoology,  Har- 
vard University,  Cambridge,  .Massachusetts. 

19671).  Aegithalidae.  Page  59  (Vol.   12)  in  R.  A. 

Paynter,  Jr.,  ed.,  Peter's  check-list  of  the  birds  of 
the  world.  Museum  of  Comparative  Zoology, 
Harvard  University,  Cambridge,  Massachusetts. 

Tanner,  V.  M.  19.30.  The  amphibians  and  reptiles  of 
Brvce  Canvon  National  Park,  Utah.  Copeia  No. 
2:41-4.3. 

1940a.  .\  biotic  study  of  the  Kaiparowits  region  of 

Utah.  (;reat  Basin  Nat.  1:97-126. 

194()b.  The  flying  squirrel  collected  in  Garfield 

County,  Utah.  Great  Basin  .Nat.  1:126. 

Tanner,  W.  W.  1953.  Herpetological  notes.  Herp. 
9(3):  1.39-140. 

1954a.  A  new  species  Sceloponis  magistvr  from 

eastern  Utah.  Great  Basin  Nat.  1.5(1 -4):.32-.34. 

1954b.  .\dditional  note  on  the  genus  HypsioU-na 

with  a  description  of  a  new  subspecies.  Herp. 
10(l):54-56. 

1954c.   Herpetological  notes  concerning  some 

reptiles  in  Utah  and  .Arizona.  Herp.  10(2):92-96. 

19.54d.  In  the  field  notes  associated  with  data  and 

information  published  in  Herpetological  notes 
concerning  some  reptiles  in  Utah  and  .\rizona. 
Herp.  10(2):92-96. 

1957.  .New  Xanttisia  from  southeastern  Utah. 

Herp.  13(1):5-11. 

1958a.  Herpetology  of  Glen  Canyon  of  the  upper 

Colorado  River  basin.  Herpetologica  14(4): 
193-195. 

1958b.  Herpetological  range  extensions.  Herpe- 
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1964.  Notes.  Southwestern  Nat.  9(l):4.5-46. 

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Proc.  Utah  -Acad,  of  Sci.,  .\rts,  and  Letters 
.52(l):4-8. 

1980.  Personal  conununication  with  C.  L.  Prit- 
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Crotdplnjtus  wizlizenii  ptinctattts.  Brigham  Yotmg 
Uiii\ersity,  Provo,  Utah. 

Tanner,  W.  W.,  and  D.  F.  .\vehy.  1964.  .\  new  Scniro- 
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Tanner,  W.  W.,  and  B.  F.  Banta.  1963.  The  distribu- 
tion of  Tantilla  uUdiensis  Blanchard.  Cireat  Basin 
Nat.  22:116-118. 
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of  the  (ireat  Basin  and  adjoining  areas,  with  a  de- 
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Basin.  Great  Basin  Nat.  .37(2):22.5-240. 

ToKT,  C.  A.  1972.  Keys  to  the  lizards  (Sauria),  frogs,  and 
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Wetherill,  J.,  p.  M.  Flattum,  and  F.  A.  Stearns. 
1961.  Early  trip  up  the  Colorado  from  Lee's  Fer- 
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.34(3):.3.3-49. 


350 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


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munity of  passerine  birds.  Publication  and  manu- 
script in  lieu  of  Ph.D.  dissertation.  Brigham 
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7):  1-225. 

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birds  of  the  southeastern  Great  Basin,  Utah.  Un- 
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Young,  S.  P.,  and  E.  A.  Goldman.  1944.  The  wolves  of 
North  America.  American  Wildlife  Institute. 
Monumental  Printing  Company,  Baltimore.  686 
pp. 


A  NEW  SPECIES  OF  FOSSIL  CHRYSOTHAMNUS  (ASTERACEAE) 
FROM  NEW  MEXICO 


Loran  C.  Anderson' 


.\bstr.'\ct.—  The  new.  presmnahlx  extinct  species,  C'lt 
trated.  The  plant  materials  came  from  Holoeene  packrat 
species  to  extant  taxa  are  discussed. 

Study  of  Holoeene  environments  through 
packrat  {Neotoma  sp.)  middens  in  Chaco 
Canyon,  New  Mexico,  by  J.  L.  Betancourt 
and  T.  R.  Van  Devender  (1980)  provided 
many  samples  of  Chrysothamnus  plant  re- 
mains. Most  materials  proved  to  be  C.  nau- 
seosus  ssp.  bigelovii,  a  frequent  constituent  of 
the  present-day  flora.  Several  involucres  from 
a  midden  (referenced  as  Mockingbird  Canyon 
No.  2)  represent  an  undescribed,  extinct  spe- 
cies of  Clirysothamnus. 

The  plant  materials  of  the  new  species 
were  in  excellent  condition;  some  were  sub- 
jected to  anatomical  study.  One  might  ques- 
tion iLsing  the  term  fossil  for  such  material, 
but  precedent  has  been  set  by  Spilman  (1976) 
in  his  description  of  a  new  .species  of  beetle 
from  packrat  middens.  Spilman  defined  fossil 
as  "a  specimen,  a  replacement  of  a  specimen, 
or  the  work  or  evidence  of  a  specimen  that 
lived  in  the  past  and  was  naturally  preserved 
rather  than  biuied  by  man."  Since  this  new 
Chrysothamnus  is  apparently  extinct,  I 
choose  to  describe  it  as  a  fossil. 

Chrysothamnus  pulchelloides   L.   C.   Ander- 
son, sp.  nov. 

Vegetatively  unknown;  involucres  cylindri- 
cal, 7-8  mm  long,  phyllaries  .strongly  gradu- 
ated in  five  series  in  distinct  vertical  rows, 
chartaceous-coriaceous,  carinate  with  en- 
larged -subapical  co.sta,  hyaline  margined  be- 
low apex,  acimiinate-cuspidate;  disk  flowers 
4-6,  corollas  presumably  yellow  (tawny,  as  in 
dried  flowers  of  extant  yellow-flowered  spe- 
cies), 4.3-4.6  mm  long,  corolla  lobes  lanceo- 
late, erect,  ca  0.5  mm  long;  stigmatic  lines 
shorter  than  appendages  (ca  40  percent  of  to- 
tal style  branch   length);   achenes  cylindric. 


rysothiinnitts  pulchelloides,  is  tormally  described  and  illiis- 
middens.  .\natomical  and  phyletic  relation.ships  of  the  fossil 

1-2  mm  long  (probably  inunature),  glabrous, 
pappus  of  capillary  bristles,  nearlv  as  long  as 
corolla. 

TYPE:  New  Mexico,  San  Juan  Co..  (>haco 
Canyon  National  Monument,  shallow  lenticu- 
lar rock  shelter  in  sandstone  of  .small  alcove 
at  head  of  minor  tributarv  of  Mockingbird 
Canyon,  36°  3'  15"  N,  107°  55'  W,  elev. 
1927  m,  /.  L.  Betancourt  6  T.  R.  Van  Deven- 
der s.  n.  in  1979  (Fig.  1;  involucres  at  FSU!). 

Midden  material  from  Mockingbird  Can- 
yon No.  2  was  dated  at  1910  ±90  B.P.  on 
Juniperus  nionospeiina  twigs  (/\-2111);  other 
fossils  in  the  midden  included  Pinus  eduHs, 
Rhus  aromatica,  Cowania  mexicana,  and  Ar- 
temisia cf.  tridentata.  Present-day  plants 
growing  on  the  talus  immediatelv  below  the 
midden  include  Artemisia  htdoviciana,  Cir- 
sium  pulchellus,  and  Stanleya  pinnata;  only  a 
few  spindly  Juniperus  numosperma  are  found 
in  the  vicinity. 

Clirysothamnus  puhheUoides  is  particu- 
larly distinctive  in  its  phyllaries  that  are 
acuminate-cuspidate  with  subapical  thick- 
ened spots  (Fig.  1).  It  is  related  to  members 
of  .section  Pulchelli  (.see  Anderson  and  Fisher, 
1970,  for  sectional  composition  of  the  genus) 
with  its  strongly  ranked  phyllaries  and 
glabrous  achenes  with  long  pappus;  it  resem- 
bles C.  pulcheUus  in  its  short  corolla  lobes 
and  C.  molestus  in  its  hyaline  margined  phyl- 
laries. 

Two  intact  heads  of  C.  pulchelloides  were 
revived  and  sectioned  as  in  Anderson  (1970). 
The  phvllaries  have  prominent  secretory 
canals  and  sclercnchyma  distribution  as  in 
section  Pulchelli.  .Vpical  portions  of  the  ph>  1- 
laries  are  covered  adaxially  with  glandniar 
trichomes.   Ovarian  vasculature   is  abundant 


'Department  of  Biological  Science.  Florida  State  University'.  Tallahassee.  Florida  32306. 


351 


352 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


lO 


Fig.  1.  Camera  lucida  drawings  of  involucre  and  flow- 
er of  C.  ])tilchcUoidcs.  Note  style  branches  are  largely  in- 
cluded in  corolla  (stamens  withered).  Pappus  bristles  av- 
erage 97  percent  of  corolla  length,  shortened  here  to 
.show  corolla  lobes  more  clearly. 

with  8-10(12)  bundles.  The  vascular  pattern 
of  the  achene-coroUa  transition  follows  pat- 
terns "a"  and  "c"  (Anderson,  1970)  wherein 
the  additional  ovarian  bundles  end  blindly 
distally  and  the  style  bundles  are  derived 
from  the  ventral  and  dorsal  corolla  bundles. 
Secretory  canals  a.s.sociated  with  the  bundles 
are  abundant  in  the  corolla,  less  frequent  in 
the  achene,  and  ab.sent  in  the  .style. 

Anatomically,  the  fo.ssil  species  relates  well 
to  section  Pulchelli,  but  it  is  less  specialized 
in  vasculature  and  secretory  canal  abundance 
and  also  has  less  pronounced  stigniatic  lines. 
A   phylogenetic  index  of  specialization  was 


developed  for  Chrysothamnus  taxa  from  flor- 
al data  in  Anderson  and  Fisher  (1970).  Extant 
members  of  section  Pulchelli  have  indices  of 
4.2-7.1;  C.  piilchelloides  would  have  an  index 
of  specialization  of  3.5-lower  than  all  extant 
Chrysothamni  except  the  least  specialized 
subspecies  of  C.  parnji  and  C.  nauseosus  of 
section  Nau-seosi  (which  is  considered  the  bas- 
al section  of  the  genus).  The  fossil  species 
may  well  be  ancestral  to  some  of  the  present 
members  of  section  Pulchelli. 

To  my  knowledge,  this  report  represents 
the  only  record  of  a  plant  species  extinction 
documented  for  the  Holocene  in  this  region. 
Drastic  reduction  of  the  tree  species,  Pinus 
edulis  and  Juniperus  monosperma,  in  the 
Chaco  Canyon  area  occurred  during  the 
Holocene  in  relation  to  fuel  demands  of  the 
Anasazi  culture  (Van  Devender  and 
Betancourt,  pers.  comm.).  Desertification 
probably  continued  with  Navajo  grazing  ac- 
tivities. These  phenomena  may  have  contrib- 
uted toward  the  extinction  of  C.  piilchel- 
loides. 

Acknowledgments 

Special  thanks  are  due  Julio  Betancourt 
and  Tom  Van  Devender  for  giving  me  these 
and  other  fossil  materials  of  Chrysothamnus. 
They  also  kindly  supplied  radiocarbon  dates 
and  descriptive  data  for  the  collection  site. 

Literature  Cited 

.\nderson,  L.  C.  1970.  Floral  anatomy  of  Chryso- 
thamnus (Astereae,  Conipositae).  Sida  3:466-503. 

Anderson,  L.  C,  and  P.  S.  Fisher.  1970.  Phylogenetic 
indicators  from  floral  anatomx'  in  Chri/sothanmus 
(Astereae,  Conipositae).  Phytomorphology 
20:112-118. 

Betancourt,  J.  L.,  and  T.  R.  Van  Devender.  1980. 
Holocene  environments  in  Chaco  Canyon,  New 
Mexico:  the  packrat  midden  record.  Natl.  Park 
Serv.  Report,  Albuquerque,  New  Mexico. 

.SpiL.viAN,  T.  J.  1976.  A  new  species  of  fossil  Ptiniis  from 
fossil  wood  rat  nests  in  California  and  Arizona 
(Coleoptera,  Ptinidae),  with  a  postscript  on  the 
definition  of  a  fossil.  Coleopt.  Bull.  30:239-244. 


NEW  AMERICAN  BARK  BEETLES  (COLEOPTEKA:  SCOLYTIDAE), 
WITH  TWO  RECENTLY  INTRODUCED  SPECIES 

Stoplicii  I..  Wood 

.\hstrac;t.-  Species  luiiiied  as  new  to  science  include:  Coiioplitlioriis  iiiirlioacciiKic.  ('.  Icocultiiit,  //i//c.si;u/.s  (iztccii\ 
(Mexico).  Fhloeocleptus  punctdtits  (Closta  Rica),  I'sctidotliijsanocs  (ilonins  iN'enezuela).  P.  lecchi  (California).  Pitiio'^- 
cm-s  mcxiaiiuis.  Aivpttis  spcriosiis,  Amplticnmits  spcrtiis  (Mexico),  and  Xijlchonis  pnwstans  (Panama).  Also  reported 
,ire  the  first  records  of  the  notorions  Xijlosiinilnis  rompactus  (Eichhoff)  from  .South  America  (Brazil),  and  the  first 
American  records  oi  Xijlebomn  fomicatus  Eichhoff  (Panama.)  and  .V.  laliclus  Eichhoff  (New  York  and  Pennsylvania). 


As  indicated  in  the  above  abstract,  the  fol- 
lowing pages  report  the  first  .\nierican  re- 
cords of  two  species  of  Xylclwrus,  the  exten- 
sion of  the  range  of  Xylosandrus  compactus 
(Eichhoff)  into  South  America,  and  the  de- 
scription of  10  species  of  American  Scolv- 
tidae  new  to  science.  The  species  new  to  sci- 
ence represent  the  genera  Conoplithorus  (2), 
Hyh  sinus  (1),  Fhloeocleptus  (1),  Pseudo- 
thysanoes  (2),  Pityogenes  (1),  Xylehorus  (1), 
Araptus  (1),  and  Amphicranus  (1),  and  were 
taken  in  California  (1),  Mexico  (6),  Costa 
Rica  (1),  Panama  (1),  and  Venezuela  (1). 

New  Introductions 
Xt/lehorus  fomicatus  Eichhoff 

Xylchonis    fornittittis    Eichhoff,    1868.    ISeriiner    Ent. 
Zeitschr.  12:151  (.Syntypes:  Ceylon;  not  located) 

This  species  has  caused  extensive  economic 
damage  in  southern  Asia,  Sri  Lanka,  In- 
donesia, Micronesia,  Africa,  and  Hawaii  for  a 
half  century  or  more  in  a  verv  large  luunber 
of  host  species. 

A  living  specimen  of  this  species  was  taken 
in  July  1979  from  a  small  branch  of  a  tree  in 
the  Canal  Zone,  Panama.  This  constitutes  the 
first  record  of  this  species  in  any  American 
country.  Since  it  is  moderatelv  aggressive,  it 
is  expected  to  be  of  some  economic  impor- 
tance in  the  tropical  and  subtropical  areas 
into  which  it  spreads. 


Xylehorus  itilidus  Eichhoff 

Xillclxniis  vdlidiis  Eichhoff.  1875.  Ann.  .Soc.  Ent.  Bel- 
Liicjue  18:202  (S\ntyi)es.  female;  Japan;  apparent- 
ly at  Brussels  Mus. 

A  breeding  population  of  this  Japanese 
species  was  first  found  on  Long  Island  near 
New  York  City  (Nassau  Co.)  in  1976.  Several 
additional  collections  were  taken  at  Du 
Pont's  Farm  near  Newtown  Square,  Dela- 
ware Co.,  Pennsvlvania,  on  2-VII-1980.  from 
Quercus  velutina,  bv  Ceorge  Stevens. 

This  species  breeds  in  the  stumps  and  logs 
or  in  the  boles  of  a  broad  spectrum  of  trees. 
Host  genera  in  japan  include:  Abies,  Acan- 
thopanax,  Acer,  Betida,  Curpinus,  Castanea, 
Cletha,  Cryptomeria,  Fagus.  Julians,  Mal- 
lotus,  Phellodcndron,  Piuus,  Prunus,  Quercus, 
and  y.elkowd.  It  is  apparenth'  more  aggresive 
than  native  ambrosia  beetles  and  should  be  of 
economic  concern. 


Xylosandrus  compactus  (Eichhoff) 

Xi/lchonis  coiiipdcliis  l'"-ichhoti.  1875,  .\mi.  Ent.  Soc.  Bel- 
<4i(l\ie  18:201  iS\ntvpes.  female.  Japan:  one  syn- 
t\pe  in  Schedl  C'oll.  at  Vienna) 

This  destructive  .species  apparently  origi- 
nated in  southern  Asia  and  .spread  to  .Africa. 
Micronesia,  and  adjacent  areas  more  than  a 
half  centurv  ago.  More  recently  it  reached 
Hawaii,  Cuba,  Florida,  Georgia,  and  Loui- 
siana. 


'Life  .Science  Maseum  and  Department  of  Ztwloi^v.  Bn^hain  Young  University.  Provo.  Utah  84602.  Scolytidae  contrihiition  No.  70. 


353 


354 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


On  7  December  1979  numerous  collections 
of  this  species  were  made  in  the  vicinity  of 
Manaus,  Amazonas,  Brazil,  from  a  variety  of 
hosts.  This  is  the  first  documented  record  of 
this  species  in  South  America.  Its  occurrence 
at  this  remote  locality  in  virgin  forest  sug- 
gests that  it  is  widespread  in  South  America 
and  that  it  has  been  there  for  quite  some 
time.  Those  concerned  with  plant  protection 
should  be  advised  of  the  range  expansion  of 
this  notorious  pest  species. 


New  Taxa 
Conopthonis  midioacanae,  n.  sp. 

This  species  is  distinguished  from  apach- 
ecae  Hopkins  by  the  more  slender  body,  by 
the  less  densely  punctured,  smoother  basal 
half  of  the  elytral  disc,  by  the  broader,  more 
gradual  elytral  declivity,  and  by  other  char- 
acters cited  below. 

Male.—  Length  3.9  mm  (paratypes  3.0-4.2 
mm),  2.4  times  as  long  as  wide;  color  very 
dark  reddish  brown. 

Frons  as  in  apachecae  except  never  with  a 
median  crest  or  tubercle,  a  weak,  transverse 
impression  usually  present  on  upper  half  of 
median  half  of  area  below  upper  level  of 
eyes. 

Pronotum  essentially  as  in  apachecae  ex- 
cept slightly  more  slender. 

Elytra  resembling  apachecae  except  1.48 
times  as  long  as  wide;  discal  striae  with  punc- 
tures not  as  close,  mostly  in  rows,  interstriae 
sparsely  punctured,  punctures  only  slightly 
confused  on  basal  fifth,  surface  smooth,  not 
wrinkled;  declivity  not  as  steep,  less  strongly 
arched,  sulcus  deeper  and  much  wider,  tu- 
bercles on  interstriae  3  very  small  (less 
strongly  arched  and  more  broadly  sulcate 
than  in  ponderosae);  vestiture  less  abundant, 
slightly  coarser. 

Female.—  Similar  to  male  in  all  respects. 

Type  locality.—  Uniapan,  Michoacan, 
Mexico. 

Type  material.—  The  male  holotype,  fe- 
male allotype,  and  24  paratypes  were  taken 
at  the  type  locality  in  Febniary  1980,  from 
Pinus  michoacana  cones  by  Adolfo  A.  del  Rio 
Mora. 

The  holotype,  allotype,  and  paratypes  are 
in  my  collection. 


Conophthorus  teocotum,  n.  sp. 

This  species  is  distinguished  from  ponde- 
rosae Hopkins  by  the  subacutely  elevated 
median  carina  on  the  lower  half  of  the  frons 
in  both  sexes,  by  the  totally  obsolete  punc- 
tures on  declivital  striae  2  except  near  base, 
and  by  other  characters  cited  below. 

Male.—  Length  3.4  mm  (paratypes  3.1-3.7 
mm),  2.3  times  as  long  as  wide;  color  very 
dark  brown. 

Front  weakly,  transversely  impressed  as  in 
ponderosae;  median  line  on  more  than  lower 
half  with  a  conspicuous,  subacutely  elevated 
carina,  end  of  carina  somewhat  tuberculate 
at  epistomal  margin. 

Pronotum  as  in  ponderosae  except  asper- 
ities averaging  smaller,  serrations  on  anterior 
margin  usually  reduced,  impressed  points 
rather  numerous  and  sharply,  distinctly  im- 
pressed. 

Elytra  as  in  ponderosae  except  punctures 
on  declivital  striae  2  obsolete  except  on  less 
than  basal  fourth,  declivity  more  broadly, 
slightly  less  strongly  impressed,  tubercles  on 
declivital  interstriae  3  slightly  larger. 

Female.—  Similar  to  male  except  trans- 
verse frontal  impression  more  extensive, 
slightly  more  conspicuous,  carina  slightly 
shorter. 

Type  locality.—  LIrupan  Michoacan, 
Mexico. 

Type  material.—  The  male  holotype,  fe- 
male allotype,  and  two  female  paratypes 
were  taken  at  the  type  locality  in  March 
1980,  from  Pinus  teocote  cones,  by  Adolfo  A. 
del  Rio  Mora. 

The  holotype,  allotype,  and  two  paratypes 
are  in  my  collection. 


Hylesinus  azteciis,  n.  sp. 

This  species  is  distinguished  from  califor- 
nicus  (Swaine)  by  the  larger  size,  by  the  pres- 
ence of  a  fine,  low,  median,  frontal  carina  in 
both  sexes,  by  the  less  strongly  concave  male 
frons  and  less  strongly  convex  female  frons, 
and  by  differences  in  the  declivital  interstrial 
setae  described  below. 

Male.—  Length  3.8  mm  (paratypes  3.8-4.2 
mm),  1.8  times  as  long  as  wide;  vestiture  of 
dark  brown  and  tan  scales  in  a  pattern  similar 
to  calif ornicus. 


December  1980 


Wood:  American  Bark  Beetles 


355 


Frons  similar  to  californicus  except  very 
shallowly  concave  from  epistoma  to  upper 
level  of  eyes,  a  low,  median  carina  on  lower 
half,  and  granules  on  upper  and  lateral  areas 
of  head  conspicuously  larger. 

Pronotum  similar  to  californicus  except  as- 
perities smaller,  piuictures  smaller,  less  defi- 
nite, and  scales  averaging  much  more  slen- 
der. 

Elytra  similar  to  californicus  except  inter- 
strial  crenulations  more  numerous,  smaller, 
confused  (a  median  row  not  predominating); 
declivital  interstriae  1  less  strongly  elevated, 
2  wider;  groimd  setae  on  declivital  interstriae 
2  in  two  indefinite  ranks  (never  uniseriate); 
erect  setae  always  absent  on  2,  present  on  1 
and  3,  each  four  to  eight  times  as  long  as 
wide,  spaced  within  a  row  by  distances  great- 
er (one  to  four  times)  than  length  of  a  seta. 

Female.—  Similar  to  male  except  frons  less 
strongly,  more  broadly  impressed  (irregularly 
flattened);  declivital  interstriae  1  less  strongly 
elevated  (vestiture  not  clearly  sexually  dimor- 
phic). 

Type  locality.—  Chapingo,  Mexico,  Mex- 
ico. 

Type  material.—  The  male  holotype  and 
seven  paratypes  were  taken  at  the  type  local- 
ity on  12-XII-1979,  from  Fraxinus  uhdei,  by 
T.  H.  Atkinson.  The  female  allotype  and  five 
paratypes  bear  similar  data  except  they  were 
taken  on  17-VIII-1979. 

The  holotype,  allotype,  and  paratypes  are 
in  mv  collection. 


Phloeocleptus  punctatus,  n.  sp. 

This  species  is  distinguished  from  trcs- 
mariae  (Schedl)  by  the  slightly  smaller  size, 
by  the  fringe  of  long  setae  at  the  upper  mar- 
gin of  the  female  frontal  concavity,  by  the 
coarser  strial  punctures,  and  by  other  charac- 
ters cited  below. 

Male.— Length  1.6-1.7  mm  (females  both 
1.8  mm),  2.4  (female  2.6)  times  as  long  as 
wide;  color  very  dark  brown. 

Frons  convex,  a  slight  transverse  impres- 
sion just  above  epistoma;  surface  rugose-reti- 
culate on  lower  half,  more  irregularly  rugose 
above,  pimctures  moderately  coarse,  rather 
close.  Antennal  scape  slender,  elongate,  orna- 
mented by  less  than  a  dozen  long  setae. 


Pronotum  about  as  in  tresmariae  except  an- 
terior margin  finely  serrate  and  sparse  vesti- 
ture on  posterior  half  of  mixture  of  fine,  slen- 
der hair  and  stout  scales. 

Elytral  outline  about  as  in  tresmariae; 
striae  distinctly  impressed  on  posterior  third 
of  disc,  punctures  at  base  rather  small,  gradu- 
ally increasing  to  twice  as  large  and  very 
deep  at  base  of  declivity;  interstriae  slightlv 
wider  than  striae  at  base,  narrower  than 
striae  at  base  of  declivity,  punctures  unise- 
riate, fine  at  base,  becoming  replaced  by 
rounded  granules  near  base  of  declivity.  De- 
clivity broadly,  strongly  convex,  steep;  strial 
punctures  decrease  in  size  from  base,  moder- 
ately coarse  at  apex;  interstriae  as  wide  as 
striae  on  lower  half,  all  imiseriately  granulate 
to  apex.  Vestiture  of  erect,  uniseriate,  inter- 
strial  scales,  each  three  to  four  times  as  long 
as  wide,  almost  as  long  as  distance  between 
rows,  spaced  within  a  row  by  about  two- 
thirds  length  of  a  scale. 

Female.—  Similar  to  male  except  more 
slender;  frons  moderately  concave  almost 
from  eye  to  eye  from  epistoma  to  vertex;  its 
surface  minutely  irregular,  punctures  fine, 
obscure,  its  upper  margin  ornamented  by  a 
dense  fringe  of  long  hair,  these  setae  equal  in 
length  to  about  one-third  diameter  of  con- 
cave area;  scape  with  a  larger  tuft  of  long 
setae;  anterior  margin  of  pronotum  imarmed; 
elytral  punctures  and  granules  distinctly 
smaller,  interstrial  scales  each  four  to  five 
times  as  long  as  wide. 

Type  locality.—  Santa  Rosa  National 
Park,  Guanacaste  Province,  Costa  Rica. 

Type  material.—  The  female  holotype, 
male  allotype,  and  three  paratypes  were 
taken  at  the  tvpe  locality  between  15  De- 
cember 1979  and  6  January  1980,  from  the 
phloem  of  an  unidentified  tree,  by  George 
Stevens. 

The  holotype,  allotype,  and  paratypes  are 
in  my  collection. 


Pseudothysanoes  atoinus,  n.  sp. 

This  species  is  distinguished  from  coluni- 
hianus  (Blackman)  and  other  representatives 
of  the  genus  by  the  very  small  size  and  b\  the 
apparent  replacement  of  most  strial  punc- 
tures by  granules. 


356 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


Male.—  Length  0.8  mm  (paratypes  0.7-0.8 
mm),  2.3  times  as  long  as  wide;  color  yellow- 
ish brown. 

Frons  convex;  surface  shining  and  almost 
smooth  in  central  area,  becoming  reticulate 
toward  margins,  pimctures  fine,  sparse,  some 
replaced  by  fine  granules;  vestiture  sparse, 
inconspicuous.  Antennal  scape  elongate,  or- 
namented by  several  hairs;  club  without  su- 
tures, small,  rather  slender. 

Pronotum  as  long  as  wide;  outline  typical 
of  genus;  anterior  margin  armed  by  four 
coarse,  closely  set  denticles;  posterior  areas 
shining,  subreticulate  in  some  areas,  sparse 
punctures  obscure,  replaced  by  fine  granules 
behind  summit.  Vestiture  of  inconspicuous 
fine  hair. 

Elytra  1.4  times  as  long  as  wide,  1.5  times 
as  long  as  pronotum;  striae  not  impressed, 
punctures  fine,  distinct,  those  on  posterior 
two-thirds  with  a  tubercle  between  punc- 
tures; interstriae  as  wide  as  striae,  shining, 
surface  irregular,  indistinct,  fine  pimctures 
replaced  by  coarse  tubercles  before  declivity. 
Declivity  steep,  convex;  strial  punctures 
scarcely  evident,  all  striae  and  interstriae 
with  rows  of  rather  large,  rounded  tubercles. 
Vestiture  mostly  confined  to  declivity,  of 
erect  rows  of  slender  strial  and  stout  inter- 
strial  setae,  all  of  uniformly  rather  short 
length,  setae  on  interstriae  2  apparently  ab- 
sent except  at  base  of  declivity. 

Female.—  Similar  to  male  except  slightly 
more  slender;  anterior  margin  of  pronotum 
unarmed;  strial  and  interstrial  tubercles  much 
smaller  (but  present);  all  declivital  setae  slen- 
der. 

Type  locality.—  Finca  Monasterios,  Cau- 
cagua,  Miranda,  Venezuela. 

Type  material.—  The  male  holotype,  fe- 
male allotype,  and  11  paratypes  were  taken 
at  the  type  locality  in  1971,  from  Theobroma 
cacao  branches. 

The  holotype,  allotype,  and  paratypes  are 
in  my  collection. 


Pseudothysanoes  leechi,  n.  sp. 

This  species  is  distinguished  from  pJiora- 
clendri  Blackman  of  the  southwestern  USA  by 
the  larger  size,  by  the  much  shorter,  stouter 
male  declivital  scales,  and  by  the  much  long- 
er setae  on  the  female   vertex.   It   is   much 


more  closely  related  to  the  Mexican  verdicus 
Wood  but  is  distinguished  by  the  stouter 
scales  on  the  male  declivity,  by  the  much  less 
strongly  impressed  female  frons,  with  the 
setae  on  the  vertex  distinctly  shorter  and  less 
abundant,  and  by  other  characters  cited  be- 
low. 

Male.—  Length  1.6  mm  (paratypes,  males 
1.4-1.7  mm,  females  1.6-1.8  mm),  2.4  (fe- 
male 2.5)  times  as  long  as  wide;  color  very 
dark  brown,  vestiture  pale. 

Frons  as  in  verdictis  except  median  third 
on  lower  half  of  frons  more  distinctly,  con- 
cavely  impressed. 

Elytra  1.45  times  as  long  as  wide,  1.7  times 
as  long  as  pronotum;  as  in  verdicus  except 
strial  punctures  on  disc  slightly  larger,  inter- 
strial granules  distinctly  larger,  extending  to 
base,  interstrial  scales  closer,  shorter,  those 
on  declivity  about  twice  as  long  as  wide, 
each  half  to  two-thirds  as  long  as  distance  be- 
tween rows. 

Female.—  Similar  to  female  verdicus  ex- 
cept frons  shallowly  concave  on  median  two- 
thirds  of  lower  two-thirds,  setae  on  vertex 
shorter,  less  abundant,  tips  of  longest  attain- 
ing middle  of  frons;  posterior  areas  of  pro- 
notum without  reticulation;  strial  punctures 
slightly  deeper,  interstrial  punctures  and 
scales  closer,  scales  distinctly  shorter.  A  very 
small  tuft  of  hair  on  scape. 

Type  locality.—  North  side  of  Howell 
Mountain,  3  km  NNE  Angwin,  Napa  Co., 
California. 

Type  material.—  The  male  holotype,  fe- 
male allotype,  and  19  paratypes  were  taken 
at  the  type  locality  (reared)  on  27-VIII-1980, 
from  Phoradendron  flavescens  var.  villosum 
(taken  from  Quercus  kelloggii),  by  H.  B. 
Leech.  Other  paratypes  emerged  or  were  cut 
from  the  same  sample  on  the  following  1980 
dates:  (3)  18- VI,  (2)  21-VII,  (4)  25-VII,  (1)  6- 
VIII,  (1)  13-VIII,  (1)  10-VIII. 

The  holotype,  allotype,  and  part  of  the 
paratypes  are  in  the  California  Academy  of 
Sciences;  the  remaining  paratypes  are  in  my 
collection. 


Pityogenes  mexicanus,  n.  sp. 

This  species  is  distinguished  from  merid- 
ianus  Blackman  by  the  much  larger  size,  by 
the  shorter,  more  strongly  hooked  upper 


December  1980 


Wood:  American  Bark  Beetles 


357 


spines  on  the  male  elytral  declivity,  bv  the 
more  regularly  punctured  discal  interstriae, 
and  by  the  much  more  strongly  convex  fe- 
male elytral  declivity.  The  female  frons  of 
this  species  and  nwridiamis  differs  from  all 
other  American  Pityogencs  in  lacking  a  deep- 
ly excavated  central  area. 

Male.—  Length  3.2  mm  (paratypes  3.2-3.4 
mm),  2.6  times  as  long  as  wide;  color  verv 
dark  brown. 

Frons  broadly  granulate,  a  few  fine  punc- 
tures interspersed;  vestiture  of  fine,  long, 
moderately  abimdant  hair. 

Pronotum  essetially  as  in  mcridianus  ex- 
cept minute,  impressed  points  nuich  more 
numerous. 

Elytra  essentially  as  in  meridianus  except 
interstrial  punctures  regular,  about  equal  in 
size  to  those  of  striae;  upper  declivital  spines 
slightly  shorter,  more  strongly  hooked,  series 
of  tubercles  on  lower  fourth  of  lateral  margin 
much  lower  and  romided  except  lowest  one 
larger  and  pointed  (male  meridianus  not  at 
hand;  comparison  based  on  Blackman's  draw- 
ing). 

Female.—  Similar  to  male  except  median 
line  on  upper  half  of  frons  shallowly  concave; 
epistomal  area  on  median  third  slightly  pro- 
tuberant, granulate,  and  ornamented  by  mod- 
erately abundant,  fine,  short  hair;  frontal  tu- 
bercles smaller;  declivity  shallowly,  narrowly 
sulcate  (more  shallowly  impressed  than  any 
other  American  Pihjogenes),  declivity  with 
stouter,  more  abundant  vestiture  than  in 
meridianus. 

Type  locality.—  Parque  Nacional  Zoquia- 
pan,  Mexico,  Mexico. 

Type  material.—  The  female  holotype, 
male  allotype,  and  six  paratypes  were  taken 
at  the  type  locality  in  August  1979,  from  a 
Pinus  luirtwegii  branch  (shaded  out?),  by  T. 
H.  Atkinson. 

The  holotype,  allotype,  and  paratypes  are 
in  mv  collection. 


Araptus  speciosus,  n.  sp. 

A  specimen  of  this  species  in  the  U.S.  Na- 
tional Museum,  which  had  been  incorrectly 
identified  as  Neodryocoetes  exquisitus  Black- 
man,  led  me  to  apply  Blackman's  name  in- 
correctly. His  exquisitus  (  =  F.  inceptis  Wood) 
must  be   referred  to  PityophtJiorus  and   the 


very  similar,  misidentified  specimens,  here 
named  speciosus,  to  Araptus.  In  all  probabili- 
ty, both  species  should  be  in  Araptus,  al- 
though the  paucity  of  material  for  studv 
makes  resolution  of  the  problem  difficult. 

This  species  is  distinguished  from  exquis- 
itus by  the  more  broadly  flattened  female 
frons,  with  longer,  more  broadly  distributed 
frontal  vestiture;  by  the  less  distinctlv  reticu- 
late, more  finely  punctured  pronotum;  and 
by  the  slightly  shorter  elytral  vestiture. 

Female.—  Length  1.7  mm  (paratvpes 
1.4-1.7  mm),  2.7  times  as  long  as  wide;  color 
dark  reddish  brown. 

Frons  broadly  flattened  from  epistoma  to 
vertex,  shining,  finely,  closely  punctured, 
sparsely  pubescent  at  center,  densely  orna- 
mented by  long  yellow  hair  at  sides  and 
above,  longest  setae  on  vertex  extend  two- 
thirds  distance  to  epistoma. 

Pronotum  1.1  times  as  long  as  wide;  widest 
slightly  behind  middle;  sides  on  posterior  half 
weakly  arcuate,  feebly  constricted  on  ante- 
rior half,  then  rather  narrowly  rovmded  in 
front;  anterior  margin  armed  bv  four  to  ten 
serrations;  summit  poorly  developed,  slightly 
in  front  of  middle;  asperities  on  anterior 
slope  moderately  large,  arranged  into  dis- 
continuous, irregular,  subconcentric  rows, 
posterior  areas  mostly  smooth,  shining,  some 
specimens  with  very  obscure  indications  of 
reticulation,  punctures  moderately  coarse, 
deep,  rather  close.  Glabrous. 

Elytra  1.7  times  as  long  as  wide,  1.5  times 
as  long  as  pronotum;  sides  straight  and  paral- 
lel on  basal  two  thirds,  rather  broadl) 
rounded  behind;  striae  not  impressed,  pimc- 
tures  rather  small,  moderately  deep;  inter- 
striae about  twice  as  wide  as  striae,  almost 
smooth,  shining,  impunctate,  impressed 
points  not  clearly  visible.  Declivity  steep, 
rather  broadly  convex;  sutural  interstriae 
feebly  elevated;  strial  punctures  smaller  than 
on  disc,  a  few  verv  small  interstrial  punctures 
also  present.  Vestiture  confined  to  declivity, 
consisting  of  a  few  interstrial  bristles,  each  al- 
most as  long  as  distance  between  rows. 

Type  locality.—  Five  miles  or  8  km  south 
of  La  Huerta,  Jalisco,  Mexico. 

Type  material.—  The  female  holotype 
and  eight  female  paratypes  were  taken  at  the 
type  locality  on  l-VII-1965,  No.  168,  from  a 
Ficus  twig,  by  me. 


358 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


The  holotype  and  paratypes  are  in  my  col- 
lection. 


Amphicranus  spectiis,  n.  sp. 

This  species  is  distinguished  from  spec- 
tabilis  Wood  by  the  larger  size,  by  the  more 
elongate  antennal  club,  with  more  strongly 
arcuate  sutures,  by  the  more  shallowly  im- 
pressed elytral  punctures,  and  by  the  more 
strongly,  more  acutely  elevated  lateral  mar- 
gin of  the  elytral  declivity  from  base  to  apex. 

Male.—  Length  2.9  mm,  3.2  times  as  long 
as  wide;  color  reddish  brown  (fully  mature?). 

Frons  about  as  in  spectahilis  except  surface 
more  finely  pimctvired,  raised  median  gran- 
ular area  slightly  larger  (occupying  almost 
meridian  third),  much  more  sharply  defined. 
Antennal  club  more  slender,  1.5  times  as  long 
as  wide,  sutures  more  strongly  arcuate  than 
in  spectahilis. 

Pronotiun  1.6  times  as  long  as  wide;  about 
as  in  spectahilis  except  punctures  on  posteri- 
or areas  slightly  smaller. 

Elytra  1.8  times  as  long  as  wide,  1.13  times 
as  long  as  pronotum;  similar  to  spectahilis  ex- 
cept punctures  on  disc  very  shallow, 
obscurely  impressed,  declivity  more  deeply 
excavated,  lateral  margin  more  acutely,  more 
strongly  elevated,  more  strongly  explanate 
below,  basal  area  of  spine  1  protruding 
slightly. 


Type  locality.—  Pichucalco,  Chiapas, 
Mexico. 

Type  material.—  The  male  holotype  was 
taken  at  the  type  locality  on  26-III-1980, 
from  Theobroma  cacao. 

The  holotype  is  in  my  collection. 

Xyleborus  praestans,  n.  sp. 

This  species  is  distinguished  from  meritus 
Wood  by  the  larger  size,  by  the  different  de- 
clivity, and  by  other  characters  cited  below. 

Female.—  Length  3.9  mm,  2.9  times  as 
long  as  wide;  color  dark  brown. 

Frons  and  pronotum  as  in  meritus  except 
pronotal  summit  more  subacutely  elevated. 

Elytra  about  as  in  meritus  except  declivity 
slightly  steeper,  more  nearly  convex,  more 
broadly  rounded  behind;  interstrial  punctures 
on  disc  more  nearly  obsolete,  irregular  inter- 
strial lines  on  disc  more  conspicuous;  strial 
punctures  on  declivity  more  distinct,  not 
larger  than  those  on  disc,  interstrial  tubercles 
on  1  and  2  not  as  close,  very  slightly  larger. 

Type  locality.—  Cerro  Punta,  Chiriqui, 
Panama. 

Type  material.—  The  female  holotype 
was  taken  in  the  vicinity  of  the  type  locality 
on  31-V-1972,  6000-8000  ft,  by  f .  L.  and  L. 
J.  Erwin. 

The  holotype  is  in  the  Canadian  National 
Collection. 


HELD  OBSERVATIONS  ON  THE  RESPONSE  OF  THE  RAH.HOAD  VALLEY 
SPRINGFISH  {CRENICHTHYS  XEVADAE)  TO  TEMPERATURE 

Thomas  M.  Baus^li'  and  Briicf  G.  Browir 

Abstract.—  The  presence  of  Crcniclithi/s  ncvadac  Hiihbs  is  verified  from  37.8  to  18.3  C  in  the  Bi^  Sprint^s  aquatic 
system. 

The  Railroad  Valley  springfish  {Crenichthijs  of  Railroad  Valley,  Nye  County,  Nevada. 
nevadae  Hubbs)  is  a  small  (ca  5  cm)  cvprino-  One  location  in  this  valley  is  Big  Springs  on 
dont  found  naturally  only  within  the  confines      Lockes  Ranch  closely  adjacent  to  Highway  6. 


N 


Flu.  1.  Map  of  Lockes  Ranch  spring-stream-pond  complex.  .\  to  B.  spring  and  segment  of  stream  to  first  down- 
stream culxert;  B  to  C,  stream  on  west  side  of  highway  between  two  culverts;  C  to  F,  stream  between  second  down- 
stream culvert  and  pond. 


'1020  Custer  Avenue,  Ogden.  Ltah  H4-1(I4. 
'142  Eccles.  Ogden,  Utah  84401. 


359 


360 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


On  21  March  1980,  the  authors,  while  in- 
volved in  other  work,  had  the  opportunity  to 
make  a  cursory  survey  of  water  temperatures 
at  several  locations  from  the  spring  to  the 
terminus  of  the  stream  in  a  pond  on  Lockes 
Ranch  and  to  relate  these  temperatures  to 
the  presence  of  C.  nevadae.  (The  length  of 
the  total  aquatic  system  from  spring  to  pond 
is  estimated  to  be  about  500  yards.) 

Temperatures  were  taken  and  observations 
were  made  from  1459  to  1606  hours  Pacific 
Standard  Time.  The  air  temperature  was 
about  7.3  C.  There  were  scattered  clouds  and 
wind  gusts  accompanied  by  snow  flurries. 

All  temperatures  were  taken  with  a  West- 
on thermometer  (Model  2265)  and  the  pres- 
ence of  C.  nevadae  in  waters  of  various  tem- 
peratures was  verified  by  both  authors. 

The  spring-stream-pond  complex  (Fig.  1) 
can  be  divided  into  three  unequal  segments: 
Segment  1  (A-B)  encompasses  the  spring  and 
that  segment  of  the  stream  to  the  first  down- 
stream culvert.  Segment  2  (B-C)  is  composed 
of  that  segment  of  the  stream  on  the  west 
side  of  the  highway  between  the  two  cul- 
verts. Segment  3  (C-F)  is  composed  of  the 
segment  of  stream  between  the  second  down- 
stream culvert  and  the  pond. 


The  temperature  gradient  from  spring  to 
pond  was  37.8  to  17.8  C.  The  temperatures 
at  various  points  along  the  system,  under  the 
climatic  conditions  described  above,  were  as 
follows: 


Point Temperature  (C) 


Fish 


A 

37.8 

Yes 

B 

32.2 

Yes 

C 

26.7 

Yes 

D 

20.0 

Yes 

E 

17.8 

No 

F 

17.8 

No 

The  last  fish  noted  by  the  authors  were  be- 
tween points  D  and  E  at  a  temperature  of 
18.3  C.  No  fish  were  found  below  that  tem- 
perature nor  were  any  seen  in  the  shallow 
water  along  the  perimeter  of  the  pond, 
where  temperatures  were  a  uniform  17.8  C. 
The  presence  of  fish  was  verified  visually  or, 
where  vegetation  made  viewing  impossible, 
by  capture  in  a  fine  mesh  net.  Those  fish 
noted  at  18.3  C  were  taken  in  a  net. 

This  cursory  survey  establishes  a  tempera- 
ture profile  for  the  Big  Springs  system  of  37.8 
to  17.8  C  and  a  temperature  tolerance  range 
for  C.  nevadae  of  37.8  to  18.3  C. 


WOODRAT  NEST  FLEA  AXOMIOrSYLIA'S  AMPIIIBOLUS 
IN  SOUTHEASTERN  ORE(X)N 

llaroki  |,  K<j;()sciie' 


Ahsthact.-  The  lira  .\/i()»nii/)si///i/v  (iiiipliilxihis  is  reported  from  soutlieastern  Oieuon,  a  ian<4e  extension  of  aliont 
173  km  from  tlie  nearest  reported  totalities  in  northw  esteiii  I'tali. 


To  m\  kno\vlcd<j;e  fleas  in  the  yenus  Ano- 
miopsyllus  have  not  been  previonslv  reeord- 
ed  in  Oregon  (Hubbard  1947,  Barnes,  Tipton 
and  Wildie  1977),  although  one  or  more 
kinds  of  woodrats  {Neotoma)  that  normally 
host  these  interesting  nest  fleas  are  found 
throughout  the  state.  This  note  reports  the 
presence  oi  Anomiopsi/Uus  amphihohis  Wag- 
ner, 1936,  in  southeastern  Oregon,  a  range 
extension  of  about  475  km  from  the  nearest 
localities  mapped  by  Barnes  et  al.  (1977)  in 
northwestern  Utah. 

My  first  specimen,  a  female  (H.J.E. 
No.6641),  was  collected  25  November  1968 
from  a  bushy-tailed  woodrat,  Neotoma  cine- 
tea  alticola,  captured  11.2  km  south  of 
Crane,  Harney  County,  elevation  1290  m.  A 
male  A.  amphibolus  (H.J.E.  No. 8026)  was 
collected  21  April  1980  from  a  deer-mouse, 
Pewnnjscus  manicukiius  ssp.  caught  less  than 
4  m  from  the  woodrat  den  where  the  1968 
host  was  trapped. 

This  locality  is  near  the  northeastern  limits 
of  the  Great  Basin  in  Oregon,  in  arid  waste- 
land characterized  bv  low  annual  precipi- 
tation and  vegetation  dominated  by  desert 
shrubs.  Traps  were  set  along  a  steep,  dry 
talus-  and  boulder-strewn  hillside  that  includ- 
ed several  prominent  lava  outcrops.  Spiny 
hopsage,  Grai/ia  spinosa,  is  the  dominant 
plant.  The  area  is  treeless,  the  nearest  scat- 


tered stands  of  juniper  being  some  miles  dis- 
tant. Other  small  mammals  trapped  here  in- 
cluded desert  woodrats,  Xeotoma  lepida 
nevadensis,  canyon  mice,  Peromyscus  crinitus 
crinitu.s,  and  Great  Basin  pocket  mice,  Per- 
ognathiis  parvus  parvus.  Other  fleas  in  the 
subfamily  Anomiopsvllinae  found  here  were 
Stenistomera  huhhardi,  S.  (dpina.  and  Callis- 
topsyllus  terinus  terinus. 

The  sympatric  occurrence  at  this  place  of 
the  bushy-tailed  woodrat  and  desert  woodrat 
was  unexpected.  Zonally,  the  habitat  seemed 
ideal  for  the  latter  species  but  too  low  and 
arid  for  .V.  cinerea,  although  Finley  (1958) 
found  that  lack  of  suitable  den  sites  more 
than  type  of  vegetation  limited  the  distribu- 
tion of  bushy-tailed  woodrats  in  Colorado. 

Field  work  in  1980  was  done  under  Scien- 
tific Taking  Permit  No.  06.3  courtesv  of  the 
Oregon  Department  of  Fish  and  W  ilillife. 

LiTKHATL  RK  (hTKl) 

Bahms,  a.  M..  \'.  J.  Tipton,  and  J.  A.  Wiinn:.  1977. 
The  suhtamilv  Anomiopsvllinae  (H\s- 
tiieliopsyllidae:  Siphonaptcra).  I.  A  revision  of 
the  s^enns  Anoniiopstilliis  Raker.  Creat  Basin  Nat. 
.37: 138-206. 

FiM.i:-^.  K.  B..  Jr.  19.5S.  The  woodrats  of  Colorado:  dis- 
tribution and  ecologv.  Univ.  Kansas  Publ..  Mns. 
Nat.  Hist.  l()(6):213-.552. 

Ill  lUiAHi).  C  .\.  1947.  Siphonaptera  of  western  North 
.\nieriea.  Iowa  State  Colle'^e  Press.  .533  pp. 


'297  West  Durfee  Street,  Granlsvi 


361 


POSTEMERGENCE  DEVELOPMENT  AND  INTERYEAR  RESIDENCE  OF 
JUVENILE  COLUMBIAN  GROUND  SQUIRRELS  IN  THE  IDAHO  PRIMITIVE  AREA 

(lliarlcs  L.  Elliott'  and  |erran  T.  FIjikIlms' 

Abstract.—  A  colonv  of  Clohinihiaii  giouiid  squirrels  in  the  Idaho  Primitive  Area  was  observed  from  1976  to  1978. 
Seven  hodv  measurements  were  recorded  tor  juveniles  obtained  in  1978.  There  was  a  lack  of  sexual  dimorphism 
among  developing  juveniles.  The  hind  foot  was  the  fastest  developing  feature.  Juveniles  obtain  adult  size  their  sec- 
ond vear.  Juvenile  males  exhibited  the  lowest  interyear  residency  of  either  sex  or  age  group  examined. 


Pengelley  (1966),  in  a  comparison  of  devel- 
opmental patterns  of  four  species  of  ground 
squrrels  (Genus:  SpennopJiihis),  noted  that 
developmental  rates  appeared  to  have  an 
adaptive  value  for  the  particular  habitats  oc- 
cupied by  each  species.  Thus  an  under- 
.standing  of  the  developmental  pattern  of  a 
particular  species  may  provide  insight  into 
the  basic  ecology  of  that  .species.  Information 
on  various  aspects  of  growth  and  devel- 
opment for  several  species  of  Spewiophilus 
has  been  reported  (Svihla  1939,  Blair  1942, 
Maver  and  Roche  1954,  Tomich  1962, 
Mckeever  1964,  Neal  1965,  Clark  1970,  Iver- 
son  and  Turner  1972,  Zimmerman  1972, 
Michener  1974,  Turner  et  al.  1976),  but  little 
data  have  been  compiled  for  the  Columbian 
groimd  squirrel  (Spermopliiliis  columhianus). 
Levenson  (1979)  presented  a  growth  rate 
constant  from  birth  to  50  days  for  S.  colum- 
hianus and  Shaw  (1925)  traced  the  devel- 
opment of  Columbian  ground  squirrels  from 
birth  to  emergence  from  the  den,  but  infor- 
mation concerning  postemergence  devel- 
opment is  lacking. 


Methods 

A  description  of  the  Cold  Meadows  study 
area  and  trapping  procedure  used  have  been 
described  elsewhere  (Elliott  and  Flinders 
1980).  In  addition  to  ground  squirrels  live 
trapped,  specimens  were  collected  from  out- 
side the  trap  grid  using  a  small  caliber  rifle. 
Postemergence  development  reported  here  is 
based  on  data  acquired  during  the  1978  field 
season. 


Results  and  Discussion 

Seven  body  measurements  were  taken  on 
each  sex  of  juvenile  (juvenile  =  young  of  the 
year)  Columbian  ground  squirrel  live  trapped 
or  shot  (Table  1).  No  significant  difference 
(unpaired  t-test)  was  found  between  measure- 
ments of  males  and  females  (excluding  body 
weights  for  July  or  August).  This  lack  of  sex- 
ual dimorphism  in  developing  ground  squir- 
rels has  been  observed  for  other  spermophiles 
(Kiell  and  Millar  1978). 


Table  1. 

.\U 

•an 

measurements 

1  ±Sl^)  of  juvenile  Co' 

lumbiau  gi 

ound  s([uirrels  eollectec; 

1  at  Cold  M 

eadous.  Idaiio 

Primitixe  Ai 

ea. 

19' 

78. 

Foot 

Ear 

Tail 

Total 

Bodv 

Zvgomatic 

Condvlobasal 

Date 

Sex 

N 

length 

length 

length 

length 

weight 

breadth 

length 

Julv  17-24 

M 

4 

47  ±  1  mm 

17±1 

82  ±.5 

2.58  ±14 

200±.34g 

27±2 

49  ±3 

F 

8 

46  ±2 

17±] 

82  ±7 

2.58  ±15 

173  ±34 

27±1 

51  ±3 

c: 

omliiued 

12 

46  ±  1 

17±1 

82  ±  6 

2,58  ±14 

182  ±.35 

27±1 

50  ±3 

August  14-2 

,1 

M 

9 

.50  ±  1 

18±  1 

82  ±4 

280  ±  7 

311  ±43 

.30  ±2 

.52  ±3 

1-^ 

9 

49  ±  1 

17±  1 

81  ±.3 

276±11 

264  ±31 

29  ±  3 

49  ±1 

c 

ombined 

18 

49  ±1 

18±1 

82  ±.3 

278  ±9 

288  ±  44 

29  ±2 

50±2 

'Dc'partiiR'nl  of  Botany  and  Rani^e  Science,  Brighain  Yonng  University,  Provo,  lUali  84fi()2 


362 


December  1980         Elliott,  Flinders:  Columbian  Ground  Squirrels 


363 


T.\BLE  2.  Size  of  juvenile  Columbian  ground  scjuirrels  expressed  as  a  percent  of  the  size  of  adults  taken  during  the 
same  collection  period  in  Cold  Meadows,  Idaho  Primitive  Area,  1978. 


Foot 

Ear 

Tail 

Total 

Body 

Zygomatic 

Condvlobasal 

Date 

Sex 

length 

length 

length 

length 

weight 

breadth 

length 

Julv  17-24 

M 

92 

85 

87 

78 

33 

80 

84 

F 

93 

86 

92 

81 

34 

83 

9() 

Combined 

92 

85 

89 

79 

32 

82 

88 

August  14-21 

M 

98 

89 

87 

84 

53 

88 

89 

F 

99 

89 

91 

87 

51 

89 

87 

Combined 

98 

89 

89 

86 

52 

89 

88 

The  body  dimensions  were  compared 
(Table  2)  to  corresponding  measurements  of 
76  adult  squirrels  taken  in  the  same  collec- 
tion periods.  The  hind  foot  was  the  fastest  de- 
veloping item  measured,  a  feature  also  noted 
in  S.  riduirdsonii  (Clark  1970),  S.  lateralis 
(Clark  and  Skrvja  1969),  S.  pamjii  (Kiell  and 
Millar  1978),  s'  tereticaiidus  (Neal  1965),  and 
S.  Jiamsii  (Neal  1965).  All  mea.surements  ex- 
cept tail  length  and  condylobasal  length  were 
significantly  larger  (unpaired  t-test,  P<0.01) 
during  August  than  July.  Shaw  (1925)  noted 
Columbian  ground  squirrels  did  not  complete 
their  growth  cycle  until  the  second  season. 
Based  on  12  juveniles  captured  in  1977  and 
recaptured  in  1978,  ground  squirrels  at  Cold 
Meadows  also  obtain  adult  size  their  second 
year  of  life.  The  combined  proportions  for 
both  sexes  in  August  (Table  2)  indicate  ap- 
proximately 90  percent  of  the  adult  dimen- 
sions (excluding  body  weight)  are  obtained  by 
the  end  of  the  first  season.  This  delaying  of 
maturity  has  been  observed  in  other  species 
of  Sperrnophilus  (Bridgwater  1966,  Morton 
and  Tung  1970).  Morton  et  al.  (1974)  noted 
that  in  S.  heldingi  fattening  and  overall 
growth  were  concurrent  at  first  but  that  ca- 
loric intake  was  then  diverted  primarily  to- 
ward lipid  synthesis  and  storage  for  catabo- 
lism  during  hibernation.  This  caloric 
diversion  resulted  in  a  late  season  slowing  of 
increase  in  linear  dimensions.  The  Columbian 
ground  squirrels  at  Cold  Meadows  are  active 
four  months  out  of  the  year,  hibernating  for 
the  remaining  period.  The  necessity  to  "trade 
off"  calories  for  body  growth  to  develop 
greater  body  reserves  for  hibernation  may  ac- 
count for  the  inability  of  juveniles  to  attain 
adult  size  their  first  season. 

Utilizing  capture-recapture  data  acquired 
during  1976-1978,  the  percent  interyear  resi- 
dence  for  each   sex  and  age  group   of  S. 


Table  3.  Percent  interyear  residence  of  Columbian 
ground  squirrels  at  Cold  .Meadows,  Idaho  Primitive 
Area,  1976-1978. 


1977 

1978 

recaptures/ 

recaptures/ 

1976 

1977 

captures 

Percent 

captures 

Percent 

Adult 

males 

10/16 

62.5 

5/13 

38.4 

Adult 

females 

16/22 

72.7 

7/20 

35.0 

Juvenile 

males 

3/14 

21.4 

2/12 

16.6 

Juvenile 

females 

4/9 

44.4 

3/9 

33.3 

columbianus  was  calculated  (Table  3).  Juve- 
nile males  exhibited  the  lowest  rate  of  fide- 
litv.  Michener  and  Michener  (1971)  observed 
the  .same  residency  pattern  for  S.  richard- 
sonii.  Reasons  for  the  absence  of  juvenile 
male  ground  scjuirrels  has  been  postulated  to 
be  the  result  of  dispersal  (Evans  and  Hold- 
enried  1943,  Fitch  1948,  McCarley  1966. 
Quanstrom  1971,  Ycaton  1972),  exceptional 
vulnerabilitv  to  predation  (Schmutz  1979), 
and/or  overwinter  mortality  (Michener  and 
Michener  1977).  We  did  not  determine 
which  specific  factor(s)  resulted  in  the  ob- 
served low  juvenile  male  interyear  residency 
for  the  Cold  Meadows  colony. 

We  thank  the  I'niversity  of  Idaho  for  per- 
mission to  use  the  facilities  at  the  Ta\lor 
Ranch  Field  Station,  Idaho  Primitive  Area. 


Blai 


LiTER-\TURE  Cited 

F.  \V,  1942.  Rate  of  development  of  young  spotted 
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364 


Great  Basin  Naturalist 


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Clark,  T.  M.,  and  D.  D.  Skkyja.  1969.  Postnatal  devel- 
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Morton,  M.  L.,  C.  S.  M.vxwell,  and  C.  E.  Wade.  1974. 
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87:314-325. 


FLOOD  FREQUENCY  AND  THE  ASSEMBLAGE  OF  DISPERSAL  TYPES  LN 
HANGLNG  GARDENS  OF  THE  NARROWS,  ZION  NATIONAL  PARK,  UTAH 

George  P.  Malanson'  and  Jeaiiiie  Kay- 

.\bstr.\ct.—  Hanging  gardens  of  the  Narrows.  Zion  National  Park,  Utah,  are  plant  commnnities  growing  at  per- 
manent seeps  on  the  canyon  walls.  The  gardens  are  isolated  from  each  other  and  from  other  plant  connnunities  by 
\ertical  expanses  of  sandstone.  Gardens  consist  mostly  of  herbaceous  species  less  than  1  m  tall.  Though  not  individ- 
ually species-rich,  the  hanging  gardens  are  diverse  as  a  group,  and  verv  di.ssimilar. 

This  study  considers  two  explanations  of  the  heterogeneous  distribution  of  species  in  hanging  gardens.  The  assem- 
i)lages  of  dispersal  types  in  gardens  of  different  spatial  attributes  and  disturbance  frequencies  are  examined.  The  C.^^ 
statistic,  a  log  likelihood  ratio  test,  analyzes  the  incidence  of  dispersal  types  among  classes  of  three  spatial  and  two 
disturbance  variables. 

Tlie  disturbance  variables  of  expected  flood  frequency  and  soil  depth  segregate  dispersal  t\pes;  and  the  spatial 
\ariables  of  area,  distance  to  possible  seed  sources,  and  relative  isolation  do  not.  Ferns  and  mosses,  dispersing 
through  spores,  dominate  a  heterogeneous  higitive  guild  in  the  flood-prone  gardens.  Infre(|ucntlv  flooded  gardens 
support  more  large-seeded  species. 


Zion  National  Park  of  .southwestern  Utah  is 
renowned  for  the  sheer  canyons  of  the  North 
Fork  of  the  Virgin  River,  which  dissect  over 
600  m  of  Navaho  sandstone.  Compared  to 
the  arid  and  semiarid  environments  of  the 
Colorado  Plateau,  the  climate  of  the  narrow 
canyons  is  cool  and  moist,  and  direct  sunlight 
may  penetrate  for  only  a  few  hours  per  day. 
Large  expanses  of  the  canyon  walls  support 
no  vegetation.  Vascular  plants,  including 
shrubs  and  trees  such  as  Finns  ponderosa,  in- 
habit the  occasional  crevices.  Perhaps  the 
most  beautiful  plant  commmiities  of  the  can- 
von  faces  are  the  haniiino;  gardens,  which  in- 
elude  ferns  and  wildflowers  among  their  spe- 
cies. Hanging  gardens  are  defined  here  as 
insular  plant  communities  growing  at  per- 
manent seeps  on  canyon  walls  (Fig.  1). 

Seeps  occur  where  precipitation  has  per- 
colated downward  through  a  porous  forma- 
tion imtil  meeting  less  permeable  strata,  and 
it  then  flows  laterally  until  a  canyon  inter- 
sects this  plane.  The  volume  of  water  dis- 
charged from  permanent  seeps  in  Zion  Can- 
yon varies  from  barely  perceptible  trickles  to 
gushing  springs.  The  seeps,  and  consequently 
the  hanging  gardens,  assume  a  variety  of 
shapes.  Seeps  usually  create  a  less  steep,  rela- 
tivelv  narrow  ledge  on  the  canyon  wall.  At 


other  .sites,  vertical  jointing  concentrates  the 
seepage,  or  travertine  deposits  from  calcium 
carbonate  in  the  water  create  bulging  forms. 
Relict  potholes  containing  permanent  .seep- 
age water  supplies  also  support  hanging  gar- 
dens. 

Hanging  gardens  illustrate  several  bio- 
geographic  problems.  The  effects  of  the  un- 
u.sual  environments  of  hanging  gardens  on 
species  composition  have  not  been  adequate- 
ly explained.  Also,  the  i.solation  of  gardens  by 
steep  rock  surfaces  provides  another  testing 
groimd  for  concepts  of  island  biogeography. 
Most  studies  of  island  biogeography  treat 
broad  areas  where  pathways  of  dispersal  be- 
tween sites  are  not  restricted  (Simberloff 
1974).  The  linear  course  of  Zion  Canyon, 
however,  is  an  additional  constraint  on  the 
movement  of  propagules  between  hanging 
gardens. 

Earlier  studies  of  hanging  gardens  (Welsh 
and  Toft  1976,  Nebeker  et  al.  1977),  island 
biogeography  (MacArthur  and  Wilson  1967, 
Diamond  1975),  and  plant  distributions  (Piatt 
1975,  Levin  1976a,  1976b)  suggest  several  ex- 
planations of  the  assemblages  of  hanging  gar- 
den species.  Species  distributions  are  in- 
fluenced by  (1)  habitat  requirements  and 
tolerances,  (2)  species'  abilities  to  disperse  to 


Department  of  Geography.  University  of  California,  l.os  Angeles,  California  90024. 
Department  of  Geography.  University  of  Utah.  Salt  Lake  City,  Utah  841 12. 


365 


366 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


Fig.  1.  A  hanging  garden  in  the  Narrows,  Zion  National  Park,  Utah. 


sites  of  different  sizes  and  degrees  of  isola- 
tion, and  (3)  time  elapsed  since  sites  were 
opened  by  disturbance. 

Few  authors  have  investigated  the  vegeta- 
tion of  hanging  gardens,  and  their  con- 
clusions emphasize  the  importance  of  habitat 
and  isolation  as  controls  of  plant  distribution. 
Woodbury  (1933)  outlined  primary  succes- 
sion at  seeps  from  algae  to  mosses  and  vascu- 
lar plants.  Welsh  and  Toft  (1976)  disclosed 
the  geographical  affinities  of  species  they 
found  at  seeps  of  different  morphology,  call- 
ing hanging  gardens  "relictual  refugia"  of 
species  not  native  to  the  region.  Welsh  and 
Wood  (1976)  studied  structure,  and  Wood 
and  Welsh  (1976)  measured  productivity  of 
hanging  gardens,  finding  stability  of  both. 
Nebeker  et  al.  (1977)  examined  floristic  sim- 
ilarity, flower  size,  and  dispersal  type  in 
hanging  gardens,  and  concluded  that  the  as- 
semblages were  "random  assortments  of  indi- 
viduals from  the  species  pool  capable  of  ex- 
ploiting the  environments  of  individual  sites." 

Malanson  (1980)  recently  explored  the 
relationships  of  .species  and  habitat  in  hang- 


ing gardens  of  Zion  Canyon.  Although  the 
gardens  were  floristically  dissimilar,  species 
assemblages  did  not  clearly  vary  according  to 
perceptible  differences  in  physical  environ- 
ment. Tests  of  species  presence  across  a 
range  of  habitat  and  spatial  variables  pro- 
duced few  significant  relationships.  A  few 
gardens  had  relatively  high  levels  of  solar 
radiation  that  might  exclude  several  species. 
He  also  found  a  species-area  relationship 
characteristic  of  small  islands  (Whitehead 
and  Jones  1969). 

Malanson  (1980)  concluded  that  other  fac- 
tors must  influence  assemblages  of  hanging 
gardens.  This  study  examines  the  ideas  that 
spatial  characteristics  and  disturbance  his- 
tories of  the  gardens  affect  the  incidence  of 
dispersal  types,  and  thereby  the  plant  assem- 
blages. 

Methods 

We  sampled  29  of  the  60  hanging  gardens 
observed  in  an  8  km  section  of  the  Narrows 
and   in   0.75   km   of  a   tributary   (Orderville 


December  1980 


Malanson,  Kay:  Hanging  Gardens 


367 


'^      >^>^.    V 


Fit^.  2.  Tlie  location  ot  sampled  han^iiit;  t^arcloiis  and  the  t()pf)<4rai)li\  of  the  Narrows. 

Canyon)  between  June  and  September  1977  of  the  i;arden.s,  spaced  at  2  m  intervals  and 

(Fig.   2).   Sites  were  selected   for  approach-  perpendicular  to  the  long  axis.  Species  were 

ability,    though    five    were    reached    only  identified  in  the  field  or  at  the  Garrett  Her- 

through  technical  climbing,  and  for  variety  barium.   University  of  Utah,  and  were  sub- 

of  garden  sizes.  Species  presence  was  record-  sequently    classified    according    to    dispersal 

ed  along  line  transects  spanning  the  lireadth  tvpe,   according   to   Dansereau   and   Lems 


368 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


(1957).  Seed  descriptions  in  floras  of  the 
western  United  States  (Arnow  and  Wyckoff 
1977,  Davis  1952,  Flowers  1973,  Hitchcock 
and  Cronquist  1973,  Munz  and  Keck  1970) 
facilitated  this  classification.  Malanson  (1980) 
provides  additional  information  on  sample 
design. 

We  measured  three  spatial  variables  to  test 
the  applicability  of  island  biogeographic  hy- 
potheses to  the  distribution  of  hanging  gar- 
den species.  These  variables  are  (1)  area  of 
the  gardens,  (2)  relative  isolation  from  other 
gardens,  and  (3)  distance  of  gardens  from  the 
terminus  of  the  Gateway  to  the  Narrows 
Trail.  Area  was  derived  from  vegetation  sur- 
vey transects.  Isolation  is  defined  as  the  sum 
of  the  distances  from  each  garden  to  its  three 
nearest  neighbors,  as  determined  from  a  top- 
ographic map  (ZNHA  1977).  This  arbitrary 
measure  was  suggested  by  the  stepping-stone 
effect  whereby  species  colonize  one  "island" 
from  another  (MacArthur  and  Wilson  1967). 
The  terminus  of  the  Gateway  to  the  Narrows 
Trail  coincides  with  the  entrance  to  the  Nar- 
rows (Fig.  2).  The  trail  receives  much  pedes- 
trian tourist  traffic,  but  use  declines  markedly 
beyond  it.  The  distance  provides  a  crude  in- 
dex of  gardens'  accessibility  to  animal- 
dispersed  or  riparian  species  originating  out- 
side the  Narrows  environment. 

Because  flash  floods  are  a  common  envi- 
ronmental disturbance  of  Zion  Canyon,  we 
evaluated  the  susceptibility  of  gardens  to  in- 
undation. The  discharge  of  the  North  fork  of 
the  Virgin  River  is  recorded  at  a  USGS  gauge 
10  km  downstream  from  the  Narrows.  Mad- 
dox.  Hart,  and  Hawkins  (1977)  calculated  the 
expected  return  periods  for  instantaneous 
peak  flows  from  magnitude  and  frequency 
data  recorded  at  this  gauge.  After  measuring 
the  various  canyon  widths  and  elevations 
where  hanging  gardens  are  located,  we  esti- 
mated the  probable  frequencies  of  flooding 
for  each  garden.  Two  classes  of  flood  fre- 
quency were  used:  less  than  7.5  yr  and  great- 
er than  15  yr.  These  classes  should  be  distinct 
and  allow  some  margin  of  error  without 
overlap.  We  also  measured  soil  depth,  assum- 
ing that  high-velocity  floods  would  scour  soil 
from  affected  gardens. 

The  Gh  statistic  for  heterogeneity  (Sokal 
and  Rohlf  1969)  was  used  to  disclose  signifi- 
cant differences  in  the  incidence  of  the  four 


Table  1.  Incidence  of  tested  dispersal  types  in  classes 
of  the  variables  and  relationship  of  soil  depth  and  flood 
frequency  in  hanging  gardens. 


Class 

Spore 

D 
Wind 

ispersal  Ty 
Plume 

pe 
Fleshy 

Area 

0-10 

21 

10 

4 

3 

(m2) 

10-25 

19 

11 

5 

2 

25-50 

14 

8 

4 

0 

<50 

19 

28 

12 

12 

Isolation 

0-80 

10 

13 

7 

5 

(ni) 

81-160 

26 

24 

12 

19 

161-240 

14 

5 

3 

1 

241-320 

11 

9 

1 

0 

<320 

12 

/ 

2 

2 

Distance 

500-1630 

11 

25 

9 

5 

(m) 

1631-2760 

8 

6 

1 

0 

2761-3890 

23 

11 

5 

0 

3891-5020 

21 

12 

9 

8 

5021-6150 

10 

4 

1 

4 

Soil  deptl 

1      0-1 

19 

12 

1 

1 

(cm) 

1-2 

29 

18 

10 

10 

2-4 

8 

1 

0 

2 

4-8 

1 

1 

0 

3 

<8 

12 

25 

14 

11 

Flood 

<7.5 

51 

23 

12 

14 

(yr) 

>15 

20 

25 

11 

12 

Flood  (yr 

) 

Soil  depth  (cm) 
0-1        1-2       2-4 

4-8 

<8 

<7.5 

7 

9 

2 

0 

1 

>15 

1 

1 

0 

1 

4 

most  common  dispersal  types  between  classes 
of  the  spatial  variables  and  soil  depth  (Table 
1).  Onlv  spores,  wind-blown,  plumed,  and 
fleshy  types  were  abundant  enough  to  pro- 
vide meaningful  tests.  We  used  a  probability 
of  p  =  .05  for  significance.  Because  we  made 
multiple  comparisons,  the  .05  chance  of  Type 
I  error  applies  to  the  individual  tests  where 
df  =  3,  but  the  probability  of  "experiment 
wide  error"  is  much  higher.  To  limit  experi 
ment  wide  error  to  .05  we  judged  individual 
tests  at  p  =  .001  (Gabriel  1966).'^Single  tests  of 
dispersal  type  and  of  soil  depth  between  the 
two  flood  frequency  categories  were  made  at 
p  =  .05. 

Results 

Forty-eight  species  were  counted  in  the  29 
hanging  gardens  (Table  2).  The  frequency  of 
species  occurrence  ranges  from  1  to  17.  Only 
13  species  were  found  in  more  than  four  gar- 
dens. The  number  of  species  per  garden 
ranges  from  2  to  20.  The  average  richness  is 


December  1980 


Mal.\nson,  Kay:  Hanging  Gardens 


369 


Table  2.  Hanging  garden  plant  species. 


Species 

Frequency 

Diaspora  type 

Abies  concolor 

1 

heavy 

Acer  ncfiimdo 

3 

winged 

Adiantum  capiUus-venehs 

15 

spore 

Acluintuin  pcdattini 

6 

spore 

Aiiuirarilhiis  (dhiis 

1 

heavy 

Anopludis  ituirficiritaccd 

4 

plumed 

ApocynuDi  ciinnubinuin 

1 

plumed 

Acjuileoiu  spp. 

13 

wind-blown 

Aralia  raccmosa 

12 

tlesh) 

Artemisia  litdovicidrui 

1 

u  iiul-hlown 

Aster  eatonii 

8 

jiluined 

Berheris  repcns 

3 

flesln 

Brickelia  grandifhna 
Bromus  ciliatus 

1 
3 

spins 
plumed 

Calainagrostis  seopii  lorn  in 

5 

plumed 

Cirsiutn  ariz^niicinit 

1 

spin) 

Ctjstopteris  frd'^ili.s 
Dodecdtheon  pulcltcllum 

17 
9 

spore 
wind-blown 

Drijopteris  filix-inas 
Ek'ocharis  sp. 

2 
3 

spore 
wind-blown 

Epipactis  ^i^igantea 
Equisetum  hijemcde 

3 

1 

heavy 
heavy 

Fraxinus  velutinu 

3 

winged 

Galium  aparine 

4 

glandular 

Hepaticae 
Hcuchera  versicolor 

10 

2 

spore 
wind-blow  n 

Jumiis  sp. 

2 

windblown 

Lobelia  cardinalis 

3 

wind-blovMi 

Mimulus  cardinalis 

13 

wind-blown 

Mimulus  ^uttatus 

1 

heavy 

Muhlenben^ia  andina 

1 

wind-blown 

Mtddeid>erf:^iu  mexicana 

2 

wind-biouii 

Sasttirtiitm  officinale 

2 

witid-blowii 

Poa  nevadensis 

2 

wind-blown 

Rhus  radicans 

2 

fleshy 

Ritbtis  leucodermis 

3 

fleshy 

Rumex  sp. 

Salix  sp. 

1 
1 

wind-blown 
wind-blown 

Smilacina  steUata 

~ 

flesh  \ 

Sphagnum  sp. 
Sphagnaceae 
Taraxacum  officimdis 

14 

9 
4 

spore 
spore 
plumed 

Thalictrum  fendleri 

2 

wind-blown 

Viola  spp. 

3 

expulsi\e 

unidentified  #  1 

1 

unidentified  #2 

1 

unidentified  #3 

2 

unidentified  #4 

1 

7.3  species,  but  in  the  seven  gardens  found  to 
be  infrequently  flooded  the  average  is  12. 
These  seven  gardens  contain  80  percent  of 
the  rare  species.  All  but  one  species  identi- 
fied are  perennial. 

The  sizes  of  the  hanging  gardens  vary 
greatly,  from  2  to  100  m  among  samples. 
Most  values  of  isolation  are  low.  Twenty-four 
gardens  are  less  than  300  m  from  the  nearest 
three  neighbors.  All  but  four  distances  from 


sampled  gardens  to  the  Gateway  to  the  Nar- 
rows trail  are  clustered  between  500-2000  m 
and  300()-5()00  m. 

Nineteen  sampled  hanging  gardens  are 
within  the  range  of  flash  floods  with  an  ex- 
pected recurrence  interval  of  7.5  yr.  Only 
seven  gardens  are  high  enough  on  the  canyon 
walls  to  escape  flood  crests  with  a  15  yr  ex- 
pected recurrence  interval,  and  gardens  28 
and  29  probably  never  have  been  inundated. 
Three  gardens  could  not  be  put  unequivo- 
callv  in  either  cla.ss. 

Individual  tests  at  p  =  .05,  df  =  3  indicate  a 
higher  incidence  of  spore  dispersal  types  in 
the  smaller  and  more  isolated  gardens  and  a 
larger  proportion  of  the  heavier,  plumed,  and 
fleshy  types  in  the  larger  and  less  isolated 
gardens.  However,  when  applving  the 
p  =  .001  level  to  limit  the  probability  of  Type 
I  error  within  the  groups,  the  statistic  re- 
vealed no  significant  differences  in  the  26 
spatial  tests  (Table  3). 

Among  the  10  soil  depth  tests,  the  Gh  sta- 
tistic indicated  a  significant  difference  in  dis- 
persal type  between  the  shallowest  (0-1  cm) 
and  the  deepest  (8  cm)  classes.  The  incidence 
of  dispersal  types  and  soil  depths  significantly 
differed  between  the  two  classes  of  flood  sus- 
ceptibility. The  frequently  flooded  gardens 
usually  have  thin  soils  and  a  high  incidence 
of  spore  dispersal  t\  pes,  and  the  heavier, 
plumed,  and  fleshy  dispersal  types  and  deep- 
er soils  are  more  common  in  the  infrequently 
flooded  eardens. 


Discussio.N  AM)  Conclusions 

The  length  of  time  seeps  are  available  for 
colonization  between  disturbances  seems  an 
important  control  of  plant  assemblages.  The 
mosses  and  ferns  disperse  by  microscopic 
wind-blown  spores  and  can  establish  them- 
selves rapidly  in  recently  flooded  gardens. 
However,  at  least  seven  species  disperse 
through  spores,  and  a  few  individuals  of  other 
dispersal  tvpes  do  establish  themselves  in  fre 
quentlv  flooded  gardens,  so  garden  commu- 
nities are  not  necessarily  similar.  The  postu- 
lated susceptibility  of  65  percent  of  this 
sample  to  frequent  floods  may  explain  wh\ 
spore-dispersed  plants  were  the  mo.st  com- 
mon tvpes.  Nebeker  et  al.  (1977)  found  bird- 
dispersed  types  to  be  mo.st  common  in  hang- 


370 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


Table  3.  Significant  differences  disclosed  by  Gh  statistic  (p  =  .05  or  less)  for  incidence  of  dispersal  types  in  cate- 
eories  of  spatial  variables,  soil  depth,  and  flood  susceptibility  and  incidence  of  soil  depth  classes  in  categories  of  flood 
susceptibility. 

Probability  of  a  Type  I 
error  among  tests 


.40 


.80 


.80 


.80 


.0,5 


.01 


Area  (m^) 

10-25 

25-50 

<50 

0-10 

- 

.05 

.05 

10-25 

- 

— 

- 

25-50 

— 

— 

— 

Isolation  (m) 

81-160 

161-240 

241 -.320 

<.320 

0-80 

— 

- 

.05 

— 

81-160 

.05 

.01 

- 

161-240 

- 

— 

241-320 

— 

Distance  (m) 

1631-2760 

2761-3890 

3891-5020 

5020-6150 

500-1630 

.01 

.05 

.05 

1631-2760 

— 

— 

— 

2761-3890 

— 

— 

3891-5020 

— 

Soil  depth  (cm) 

1-2 

2-4 

4-8 

<8 

0-1 

.05 

- 

.02 

.001 

1-2 

— 

— 

.05 

2-4 

- 

.01 

4-8 

— 

Flood  (yr) 

<15 

7.5 

.05 

Soil  depth  (cm) 

Flood 

15 

Flood  <7.5 

.01 

ing  gardens  of  Arches  and  Canyonlands  Na- 
tional Parks.  Where  garden  habitats  are  sel- 
dom disturbed,  large-seeded  species  are  prob- 
ably more  successful  competitors  in  plant 
succession,  because  their  larger  propagules 
provide  more  energy  to  their  seedlings. 

Tlie  term  hanging  gardens  is  useful  from 
the  standpoint  of  vegetation  physiognomy, 
but  it  has  little  relevance  to  floristic  composi- 
tion and  ecology.  The  species  compositions  of 
29  hanging  gardens  in  the  Narrows  of  Zion 
Canyon  were  quite  di.ssimilar,  and  variables 
of  the  gardens'  physical  environment  do  not 
afford  a  convincing  explanation  of  the  differ- 
ences (Malanson  1980).  Dispersal  seems  to  be 
more  important  in  influencing  plant  assem- 
blages than  the  early  succcssional  environ- 
ment represented  by  .soil  depth,  because  Ma- 
lanson (1980)  did  not  find  many  species 
limited  by  that  variable. 

The  spatial  variables  of  area,  distance  to  a 
likely  seed  .source,  and  relative  isolation  com- 
monly are  used  by  island  biogeographers  to 
predict   species   distributions.    Hanging   gar- 


dens in  the  Narrows,  however,  are  not  de- 
monstrably in  equilibrium,  and,  without  fur- 
ther investigation,  we  cannot  support  a  spa- 
tial explanation  of  their  plant  assemblages. 

Mo.st  species  of  the  Zion  Narrows  could  be 
considered  "fugitive  species"  sensu  Piatt 
(1975),  in  being  both  perennial  and  com- 
paratively vagile  (annual  colonizers  are  clas- 
sified as  "ruderals").  According  to  this  meth- 
od, following  flash  floods,  the  able  dispersers 
would  quickly  occupy  open  spaces.  During 
later  stages  of  colonization,  perennials  would 
be  more  successful  than  annuals.  Dispersal  of 
annuals  to  a  new  site  already  occupied  by 
perennials  would  have  a  low  probability  of 
success  because  of  the  scarcity  of  favorable 
spots.  There  may  also  be  a  high  ri.sk  of  seed 
loss  from  any  established  annuals,  given  the 
restricted  habitat  spaces  available  and  the 
vertical  nature  of  the  Narrows  environment. 
At  the  gardens  situated  above  the  flood 
crests,  the  fugitive  species  would  give  way  to 
species  with  larger  seeds. 


December  1980 


Malanson,  Kay:  Hanging  Gardens 


371 


A  minority  of  seven  gardens  support  the 
conclusion  by  Nebeker  et  al.  (1977)  that  gar- 
den assemblages  are  "random"  collections  of 
plants.  The  majority  of  gardens  in  the  Nar- 
rows apparently  maintain  their  dissimilar, 
fugitive  assemblages  through  response  of  dis- 
persal types  to  frequent  disturbance. 

Acknowledgments 

We  appreciate  the  assistance  of  Kimball  T. 
Harper,  Paul  A.  Kay,  Kezia  M.  Nielsen,  Gar- 
ry F.  Rogers,  Lois  A.  Arnow,  Robert  W.  Aus- 
tin, Walter  E.  Westman  and  Jonathan  D. 
Sauer.  This  research  was  fimded  in  part  by 
grants  from  the  University  of  Utah  Student 
Research  Grants  in  Geography,  the  Zion  Nat- 
ural History  Association,  and  Sigma  Xi,  the 
Scientific  Research  Society  of  North  Ameri- 
ca. 


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D.\NSEREAU,  P.,  AND  K.  Lems.  1957.  The  grading  of  dis- 
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Flowers,  S.  1973.  Mo.sses  of  Utah  and  the  West.  Brig- 
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Hitchcock,  C.  L.,  and  A.  Cronquist.  1973.  Flora  of  the 
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Levin,  S.  .\.  1976a.  Population  dynamic  models  in 
heterogeneous  environments.  Anna.  Rev.  Ecol.  & 
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1976b.   Spatial  patterning  and  the  structure  of 

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Mac.\rtiilr,  R.  .,  and  E.  O.  Wilson.  1967.  The  theo- 
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.Maddo.x,  J.  D.,  G.  E.  Hart,  and  R.  H.  Hawkins.  1977.  A 
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the  Virgin  River,  Zion  National  Park.  Depart- 
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State  University,  Logan. 

Malanson,  G.  P.  1980.  Habitat  and  plant  distributions 
in  hanging  gardens  of  the  Narrows,  Zion  National 
Park.  Great  Basin  Nat.  40:178-182. 

Mu.NZ,  P.  A.,  and  D.  D.  Keck.  1970.  A  California  flora. 
University  of  California  Press,  Berkelev. 

Nebeker,  G.  T.,  K.  T.  Harper,  J.  D.  Brothehson,  and  S. 
L.  Welsh.  1977.  Characteristics  of  plants  of  com- 
mon occurrence  in  hanging  gardens  of  the  Colo- 
rado Plateau,  Utah.  Unpublished  manuscript.  De- 
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Young  University.  Provo,  Utah. 

Plait,  W.  J.  1975.  The  colonization  and  formation  of 
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disturbances  in  a  tall  grass  prairie.  Ecol.  .Monogr. 
45:28.5-.3()5. 

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biogeographv  and  ecologv.  .\nnu.  Rev.  Ecol.  & 
Syst.  5:161-182. 

SoKAL,  R.  R.,  AND  F.  J.  RoHLF.  1969.  Biometry.  W.  H. 
Freeman,  San  Francisco. 

Welsh,  S.  L.,  and  C.  .\.  Toft.  1976.  Biotic  comnnmities 
of  hanging  gardens  in  .southeastern  Utah.  Pages 
468-498  in  J.  R.  Murdock,  W.  L.  Welsh,  and  B. 
W.  Wood,  eds.,  Navaho-Kaiparowits  environmen- 
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and  Environmental  Studies,  Brigham  Young  Uni- 
versity, Provo,  Utah. 

Welsh,  S.  L.,  and  B.  W.  Wood.  1976.  Structure  of  a  se- 
lected hanging  garden.  Pages  499-511  in  J.  R. 
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Navaho-Kaiparowits  environmental  baseline 
studies  1971-1974.  Center  for  Health  and  Envi- 
romnental  Studies,  Brigham  Young  L'niversitv, 
Provo,  Utah. 

Whitehead,  D.  R.,  and  C.  E.  Jones.  1969.  Small  islands 
and  the  equilibrium  theorv  of  insular  bio- 
geography. Evolution  23:171-179. 

Wood,  B.  W.,  and  S.  L.  Welsh.  1976.  Productivity  of 
hanging  gardens.  Pages  512-.521  in  J.  R.  .Mur- 
dock, W.'  L.  Welsh,  and  B.  W.  Wood,  eds.,  Nav- 
aho-Kaiparowits environmental  baseline  studies 
1971-1974.  Center  for  Health  and  Environmental 
Studies,  Brigham  Young  University,  Provo,  Utah. 

Woodbury,  A.  M.  1933.  Biotic  relationships  of  Zion 
Canyon,  Utah,  with  special  reference  to  succes- 
sion. Ecol.  Monogr.  3:147-245. 

Zion  Natural  History  .\ssociation.  1977.  Topograph- 
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Springdale,  Utah. 


ZONATION  PATTERNS  IN  THE  POTHOLES  OF  KALSOW  PRAIRIE,  IOWA 

Jack  D.  Brotherson' 

,\bstract.—  Kalsow  Prairie,  a  mesic  prairie  remnant  in  central  Iowa,  was  acquired  by  the  state  in  1949  and  later 
established  as  a  botanical  reserve.  This  study  concerns  a  complex  of  14  potholes  and  adjacent  drainage  areas  within 
the  prairie.  The  potholes  varied  in  depth  and  degree  of  fill  and  are  thus  usefid  in  studies  of  plant  succession.  All  14 
potholes  exliibit  zonation  patterns.  A  total  of  36  transects  were  taken  in  the  various  comnuuiity  zones.  Canopy  cover 
data  were  taken  in  every  zone.  The  zones  are  ordinated  into  three-dimensional  space  as  well  as  clustered.  Inter- 
specific association  patterns  are  elucidated.  A  successional  sequence  is  proposed. 

There  are  prairies  three,  six,  ten,  and  twenty  leagues 
in  length,  and  three  in  width,  surrounded  by  forests  of 
the  .same  extent;  beyond  these,  the  prairies  begin  again, 
so  that  there  is  as  much  of  one  sort  of  land  as  of  the 
other.  Sometimes  we  saw  the  grass  very  short,  and,  at 
other  times  five  or  six  feet  high;  hemp,  which  grows  nat- 
urally there,  reaches  a  height  of  eight  feet. 

A  settler  would  not  there  spend  ten  years  in  cutting 
down  and  burning  trees;  on  the  day  of  his  arrival,  he 
could  put  his  plough  into  the  ground.  (Louis  Jolliet) 

These  are  the  finest  and  most  fertile  countries  in  the 
world.  .  .  .  From  time  to  time  there  are  vast  prairies 
where  the  gra.ss  is  ten  or  twelve  feet  high  at  all  sea- 
sons. .  .  .  No  settler  arriving  in  the  country  will  not  find 
at  first  enough  to  support  plenteously  a  large  family,  or 
will  not,  in  less  than  two  years"  time  be  as  comfortably 
settled  as  in  any  place  in  Europe.  (Douay) 


Such  are  the  early  accounts  from  explorers 
and  settlers  of  presettlement  lowan  vegeta- 
tion (Dondore  1926).  A  government  survey 
begun  in  March  1832,  when  Iowa  was  still  a 
territory,  and  completed  in  August  1859,  first 
documented  the  original  extent  of  this  vast 
grassland  area.  That  survey  indicated  that  in 
the  1850s  about  85  percent  of  Iowa  was  cov- 
ered by  grassland  (U.S.  Government  1868, 
Hayden  1945,  Hewes  1950,  Dick-Peddie 
1955). 

Accounts  by  the  first  explorers,  surveyors, 
and  settlers  in  the  area  describe  three  major 
types  of  landscape  in  Iowa:  (1)  woodland,  (2) 
well-drained  prairie,  and  (3)  poorly  drained 
prairie  and  marsh  (Dondore  1926,  Berry 
1927,  Hewes  1950).  The  woodlands  were 
confined  to  stream  valleys  and  adjacent 
slopes,  and  the  prairie  was  .said  to  occupv  the 
remaining  portions  of  the  land  (Berry  1927, 
Hewes  1950).  The  well-drained  prairie  areas 
were  the  most  extensive  except  in  portions  of 


the  state  subjected  to  late  Wisconsinan  gla- 
ciation;  there  the  poorly  drained  prairie  and 
marsh  were  more  common  (Hewes  1950, 
Hewes  1951,  Hewes  and  Frandson  1952).  The 
well-drained  prairie  was  described  as  highly 
fertile  (Dondore  1926,  Berry  1927),  whereas 
the  poorly  drained  prairie  was  filled  with  wa- 
ter most  of  the  season  and  "capable  of  pro- 
ducing nothing  but  wild  rice,  frogs,  and  mos- 
quitoes" (Hewes  and  Frandson  1952). 

The  characteristics  of  Iowa  prairie  in  terms 
of  vegetation  types,  structure,  and  general 
ecology  of  the  dominant  species  were  the 
subjects  of  several  papers  during  the  1930s 
and  1940s  (Steiger  1930,  Rydberg  1931, 
Weaver  and  Fitzpatrick  1934,  Hayden  1943). 
These  authors  recognized  the  existence  of  six 
grassland  communities  and  generally  con 
eluded  that  water  relations,  as  affected  by 
climate,  soil,  and  topography,  were  respon- 
sible for  local  variations  in  the  structure  and 
distribution  of  prairie  vegetation.  "In  varying 
the  water  relations  of  soil  and  air  they  merely 
bring  about  changes  in  the  groupings  of  the 
dominant  grasses  and  accompanying  segrega- 
tions and  rearrangements  of  the  forbs" 
(Weaver  and  Fitzpatrick,  1934). 

The  major  grassland  types  as  alluded  to  in 
the  above  studies  were  labeled  "con- 
sociations" after  Weaver  and  Clements 
(1938)  and  were  designated  as  follows: 

1.  Big  Bluestem  type  {Andropogon  ge- 
rardi)—iound  on  the  lower  moist  slopes 
and  well-aerated  lowlands. 

2.  Slough  Grass  type  [Sportino  pectinata)— 
found  on  poorly  aerated  and  wet  soils  of 
sloughs  and  natural  drainage  systems. 


'Department  of  Botany  and  Range  Science.  Brighani  Young  University.  Provo,  Utah  84602. 


372 


December  1980 


Brotherson:  Kalsow  Prairie,  Iowa 


373 


3.  Tall  panic  grass-wildrve  type  [Panicum 
virgatum  and  Elymus  canadensis)— 
found  on  soils  intermediate  between 
Slough  Grass  and  Big  Bluestein  tvpes. 

4.  Little  Bluestem  type  (Schizocherium 
scoparius)—moiii  important  upland  tvpe 
on  well-drained  soils. 

5.  Needle  Grass  type  {Stipa  spartea)— 
found  on  the  uplands,  often  occurring 
as  a  narrow  zone  following  the  shoul- 
ders of  ridges. 

6.  Prairie  dropseed  type  (Spowbohis 
hcterolepis)—ionn(\  locally  on  the  driest 
upland  sites. 

Mover  (1953),  Aikman  and  Thorne  (1956), 
Ehrenreich  (1957),  and  Kennedy  (1969)  have 
presented  ecological  and  taxonomic  descrip- 
tions of  four  state-owned  native  prairie 
tracts.  The  vegetation  complexes  studied 
were  basically  limited  to  upland  prairie. 
Those  studies  include  information  on  soils, 
microclimate,  topography,  and  management, 
as  well  as  extensive  literature  reviews. 

These  prairies  are  presentlv  owned  bv 
state  agencies  and  were  purchased  and  set 
aside  as  natural  areas  with  the  intent  that  the 
various  typical  landscapes,  wild  flowers,  and 
wildlife  of  the  native  tall-grass  prairie  region 
be  preserved  for  posterity.  It  was  also  in- 
tended that  these  areas  serve  as  game  and 
wildlife  sanctuaries;  as  examples  of  the  native 
prairie  soil  types,  where  comparisons  could 
be  made  with  cultivated  soils  of  the  same  soil 
association;  and  as  prairie  reserves  where  sci- 
entific investigations  could  be  undertaken  on 
problems  concerning  the  native  flora  and 
fauna  of  the  various  topographic,  climatic, 
and  prairie  districts  throughout  Iowa.  The 
prairies  were  also  meant  to  serve  as  reference 
points  by  which  future  generations  could 
compare  the  postsettlement  influences  of 
man  on  Iowa  (Hayden  1946,  Moyer  1953, 
Aikman  1959,  Landers  1966). 

One  such  area  is  Kalsow  Prairie,  160  acres 
of  unplowed  grassland  in  Pocahontas  Coimty, 
Iowa.  Criteria  for  its  purchase  dictated  that 
this  area  satisfy  the  requirements  of  a  game 
preserve,  contain  one  or  more  soil  types  of  an 
association,  and  include  several  regional  veg- 
etation types  (Hayden  1946).  The  prairie  was 
purchased  in  1949  by  the  Iowa  State  Con- 
servation Commi.ssion  and  since  its  purchase 
has  been  the  object  of  several  studies  dealing 


with  its  vegetation,  soils,  management,  in- 
sects, response  to  fire,  mammals,  and  nema- 
todes (Mover  1953,  Ehrenreich  1957,  Esau 
1968,  Richards  1969.  Brennan  1969,  Norton 
and  Ponchillia  1968,  Schmitt  1969). 

The  present  investigation  was  undertaken 
to  provide  information  on  the  phvtosociolog\' 
and  ecological  relationships  of  poorl\-  drained 
prairie  and  marsh  areas  of  Kalsow  Prairie.  It 
includes  information  on  species  composition 
and  distribution,  zonation  patterns,  and  inter- 
relationships within  and  between  these  zones. 

Methods 

General 

This  study  was  begun  in  the  spring  of  1967 
and  continued  through  the  following  vear 
(1968)  and  into  the  summer  of  1969.  Kalsow 
Prairie  is  one  of  four  state-owned  Iowa 
prairies.  It  is  located  five  miles  northwest  of 
Manson,  Iowa,  and  comprises  the  \E  '4  of 
Section  36,  Belleville  Township,  T  90N,  R 
32W,  Pocahontas  County.  It  occurs  in  a  part 
of  north  central  Iowa  that  was  glaciated  dur- 
ing the  most  recent  advances  of  the  W'is- 
consinan  glaciation  and  within  the  Clarion- 
Nicollet-Web.ster  soil  association  area  (Ruhe 
1969).  The  area  was  chosen  for  study  on  the 
basis  of  its  vegetational  composition,  i.e.,  flo- 
ristic  richness  and  the  presence  of  several 
pothole  areas  (poorly  drained  prairie  and 
marsh). 

Taxonoiu)' 

Voucher  specimens  were  collected  in  du- 
plicate throughout  the  growing  seasons  of 
1967  and  1968.  All  specimens  were  identi- 
fied, and  identical  sets  have  been  deposited 
in  the  herbaria  of  Iowa  State  University, 
Ames,  Iowa,  and  Brigham  Young  University, 
Provo,  Utah.  Nomenclature  follows  Pohl 
(1966)  for  the  grasses,  Gilly  (1946)  for  the 
sedges,  and  Glea.son  (1952)  for  the  forbs. 

Community  Types 

Kalsow  Prairie  contains  within  its  bound- 
aries a  complex  of  potholes  and  correspond- 
ing drainage  ways.  The  vegetation  of  these 
communitv    types   was   analyzed    itsing   two 


374 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


separate  approaches.  The  first  involved  the 
identification  and  listing  of  all  species,  and 
the  second  utilized  random  sample  plots  to 
determine  percent  cover,  composition,  and 
interspecific  relationships  of  species  within 
these  subcommunities. 


Quadrat  Analysis 

The  vegetation  of  each  area  was  sampled 
by  using  a  20  X  50  cm  (1000  cm^)  quadrat. 
Twenty  quadrats  were  located  along  tran- 
sects on  a  restricted  basis  to  reduce  bias  and 


GRAZED    AREA 


Map  of   the  KALSOW  PRAIRIE 

A   areas  affected  by  soil  drift  from   adjacent  fields 
yCi^  potholes   and    drainage 

—  location  of  1000  foot  bisect 


Fis;.  1.   Map  of  Kalsow  Fraiiic  showing  locations  ot  potholes  and  drainagf  ways. 


December  1980 


Brotherson:  Kalsow  Prairie,  Iowa 


375 


to  keep  adjacent  quadrats  at  fairlv  e(jual  dis- 
tances apart.  The  number  of  samples  varied 
with  the  subcommunity  or  zone,  but  a  total 
of  720  quadrats  were  taken  within  the  com- 
munity. Sampling  was  done  between  1  Au- 
gust and  15  September  each  vear  when  most 
species  had  reached  their  maximum  growth. 
Cover  estimates  were  made  for  each  quadrat 
through  use  of  Daubenmire's  (1959)  method. 

Coverage  was  determined  separately  for 
all  species  overlapping  the  plot  regardless  of 
where  the  individuals  were  rooted.  Coverage 
was  projected  to  include  the  perimeter  of 
overlap  of  each  species  regardless  of  super- 
imposed canopies  of  other  species.  The  cano- 
pies of  different  species  were  commonly  in- 
terlaced or  superimposed  over  the  same  area; 
therefore  coverage  percents  often  total  great- 
er than  100  percent. 

Commiuiity  Analysis 

Analysis  of  these  areas  was  accomplished 
by  dividing  the  sites  into  subimits  or  zones 
(Fig.  1)  based  upon  location  and  dominant 
species.  Each  subvmit  was  then  sampled  by 
randomly  locating  a  starting  point  and  then 
placing  a  quadrat  every  3  m  along  a  transect. 
Twenty  quadrats  were  taken  for  each  zone. 

Data  Analysis 

General  descriptive  data.—  Data  collected 
from  quadrat,  mapping,  and  topographic 
studies  were  used  to  describe  the  vegetation 
of  each  zone.  Frequency  values  and  average 
cover  values  were  determined  for  all  species 
in  every  stand.  Cover  values  were  deter- 
mined by  summing  the  midpoints  of  the 
cover-class  ranges  and  dividing  by  the  num- 
ber of  sample  quadrats  in  each  zone. 

Ordination  analysis.—  An  ordination  tech- 
nique proposed  by  Orloci  (1966)  was  em- 
ployed to  ordinate  vegetation  units  within 
the  different  subcommimities  listed  above. 
Raw  data  were  first  summarized  by  hand  cal- 
culation and  then  transferred  to  punch  cards. 
The  entities  to  be  ordinated  (i.e.,  plant  spe- 
cies or  stands  of  vegetation)  were  projected 
as  points  into  n-dimensional  space.  Such 
points  were  positioned  by  attribute  scores 
through  the  application  of  the  R  and  Q-tech- 
niques  of  factor  analysis  (Orloci  1967).  Once 


established,  this  multidimensional  array  of 
points  was  then  reduced  to  a  three-dimen- 
sional system.  This  was  accomplished  by  se- 
lecting the  two  most  different  stands  or  spe- 
cies and  placing  one  at  zero  and  the  other  at 
some  distance  along  the  abscissa.  All  other 
stands  or  species  under  consideration  were 
then  positioned  linearly  in  relationship  to 
these  two  extremes.  This  action  thus  estab- 
lished the  X-axis.  The  above  process  was  re- 
peated until  all  points  had  been  established  in 
three-dimensional  space  (i.e.,  Y  and  Z  axes 
had  been  added).  Coordinate  values  for  the 
X,  Y,  and  Z  axes  were  given  as  output  from 
the  computer. 

Cluster  analifsis.-  Cluster  analyses  were 
performed  by  calculating  a  similarit\'  index 
(SI)  (Ruzicka  1958)  in  percent  from  the  for- 
mula: 


SI  = 


2  min  (Xi,  Yi) 


X  max  (Xi,  Yi) 

and  then  clustering  the  indices  using  un- 
weighted pair-group  clustering  techniques 
(UPGMA)  (Sneath  and  Sokal  1973).  The  UP- 
GMA  method  computes  the  average  sim- 
ilarity of  each  unit  to  the  clu.ster,  using  arith- 
metic averages.  It  is  widelv  used  and  has 
been  found  to  introduce  less  distortion  than 
other  methods  (Kaesler  and  Cairns  1972). 

Interspecific  association  analysis.—  Expres- 
sions of  interspecific  association  were  at- 
tempted using  Cole's  Index  (1949).  Step  one 
in  the  computation  of  the  index  involved  the 
accumulation  of  2  X  2  contingency  tables 
(Fig.  7).  Actual  calculation  of  the  index  in- 
volved the  following  three  sets  of  formulas: 

when  ad  <  be: 


C.  ±  Sdc  = 


ad-bc 


(a  +  c)(c  +  d) 


(a  +  b)(b  +  d)        n(a  +  b)(l)  +  c) 
when  be  >  ad  and  d  <  a: 


C,  ±  Sdc  = 


ad-bc 


(b  +  d)(c  +  d) 


(a  +  b)(a  +  c)        n(a  +  b)(a  +  c) 

when  be  >  ad  and  a  >  d: 

ad-bc  (a  +  b)(a-Hc) 

C,  ±  Sdc  =      ^^^^j^^^._^j^    ±  n(b-l-d)(c  +  d) 

where  C;  =  Cole's  Index  of  Interspecific 
Association 

Sdc  =  standard  deviation  of  Cole's  in- 
dex 

n  =  total  number  of  samples 


376 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


and  a,  b,  c,  and  d  represented  the  four  cells  of 
the  2  X  2  contingency  table. 

Tests  of  statistical  significance  were  per- 
formed by  means  of  the  Chi-square  test.  The 
Chi-squares  were  computed  by  the  formula: 

(ad-bc)2n 

^'  "     (a  +  b)(a  +  c)(c  +  d)(b  +  d) 

where  X^  =  Chi-square  value 
n  =  number  of  samples 
and  a,  b,  c,  and  d  represented  the  different 
cells  of  the  2x2  contingency  table. 

In  all  cases,  a  single  degree  of  freedom  was 
used.  Chi-quare  values  greater  than  3.84 
were  significant  at  the  5  percent  level,  while 
values  greater  than  6.63  were  significant  at 
the  1  percent  level. 

Data  representation.—  The  three-dimen- 
sional graphic  representation  of  data  ob- 
tained from  ordination  analysis  was  drawn  by 
the  computer.  Such  representation  was  ac- 
complished through  the  use  of  a  plotting 
technique  developed  and  programmed  by 
Mr.  Howard  Jesperson,  Agricultural  Experi- 
ment Station,  Iowa  State  University. 


Results  and  Discussion 

Marean  and  Jones  (1903)  gave  the  follow- 
ing description  of  the  landscape  in  central 
Iowa: 

Low  knolls  are  separated  by  saucerlike  depressions  in 
which  impounded  water  often  stands  the  year  around. 
In  many  cases  these  low-lying  areas  have  been  reclaimed 
by  artificial  drainage,  but  in  the  main  rainwater  which 
falls  upon  the  upland  has  to  escape  by  seepage  or  evapo- 
ration. Little  ponds  and  marshes  are  foimd  in  almost  in- 
numerable places  scattered  all  over  the  coimtry. 

These  saucerlike  depressions  have  been  es- 
timated as  covering  more  than  50  percent  of 
that  part  of  Iowa  subjected  to  late 
Wisconsinan  glaciation  (Hewes  1950).  They 
were  early  recognized  as  supporting  a  dis- 
tinct vegetation  from  that  of  the  adjacent  up- 
land prairie  (Yapp  1909,  Sherff  1912,  Shimek 
1915,  and  Berry  1927).  The  grasses  of  these 
areas  were  described  as  being  "ten  to  twelve 
feet  tall  all  season"  (Dondore  1926).  These 
and  later  descriptions  indicate  that  the  pot- 
holes and  drainage  ways  were  often  charac- 
terized by  very  discrete  zones  of  vegetation 
(Sherff  1912,  Shimek   1915,  Schaffner  1926, 


Fig.  2.  Three-dimensional  ordination  of  pothole  and  drainage  zones  found  in  Kalsow  Prairie,  with  numbers  corre- 
sponding to  pothole  and  drainage  mimbers  from  Figure  1  except  3.3-36,  which  are  prairie  edge  areas. 


December  1980 


Brotherson:  Kalsow  Prairie,  Iowa 


377 


Hayden  1943,  Trauger  1967).  Three  to  four 
zones  were  generally  recognized,  yet  in  all 
cases  little  information  was  given  on  the  rela- 
tionships of  these  zones  to  one  another  either 
floristicallv  or  spatially.  Some  authors  (Sherff 
1912,  Schaffner  1926,  Hayden  1943),  how- 
ever, indicated  that  succession  was  taking 
place  in  these  areas  and  proposed  the  follow- 
ing successional  scheme: 
I  Pond  center 
II      Sedge  zone 

III  Slough  grass  zone 

IV  Dry  margin  of  slough  grass 
V      Andropogon  gerardi 

VI  Upland  prairie 
Within  the  boundaries  of  Kalsow  Prairie 
there  exists  a  complex  of  14  potholes  and  cor- 
responding drainage  ways  (Fig.  1).  These 
areas  are  found  scattered  throughout  the  160 
acres  at  different  elevations.  They  also  vary 
in  depth  and  degree  of  fill.  These  character- 
istics make  them  extremely  useful  in  studies 
of  plant  succession  and  zonation.  All  14  pot- 


holes studied  exhibited  strong  zonation.  Each 
zone  was  subsampled  20  times  for  cover  and 
then  averaged  to  obtain  a  characteristic  veg- 
etation for  each  zone.  The  36  zones  were  or- 
dinated  into  three-dimensional  space  using 
Orloci's  (1966)  method  (Figs.  2  and  3)  and 
then  clustered  according  to  Sneath  and  Sokal 
(1973)  (Fig.  4). 

Following  the  ordination  and  cluster  analy- 
ses, the  zones  were  then  grouped  into  six 
units  as  .shown  in  Figures  3  and  4.  TTiis  pro- 
cedure seemed  justified  since  each  zone  rep- 
resented a  rather  discrete  vegetational  unit. 
After  grouping,  the  data  from  all  zones  in- 
cluded in  each  new  unit  were  averaged  and 
placed  in  Table  1.  These  six  groups  (with  one 
exception,  Group  5)  correpond  in  reality  to 
the  suggested  successional  sequence  shown  in 
Figure  5.  Table  1  is  so  designed  that  columns 
1  through  6  represent  values  from  the  center 
of  each  pothole  through  a  transition  to  up- 
land prairie,  respectively.  The  positioning  of 
each   species  within   Table    1    was  done  by 


50- 


IV 


,.\ 


y 


\  -36 


32/         -<ll./ 


0  02< 


024  O         0  = 

323        Q28 


Oio 


ill 


/^\T 


V&VI 


\    I 


Y 


50 


100 


Fiu.  3.   Two-dimensional  ordination  of  pothole  and  drainage  zones  of  Kalsow  Praine.  grouped  as  shown  in  Table  1 
with  minor  exceptions,  llie  factors  responsible  for  the  ordination  are  nnknown. 


378 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


Table  1.  Average  percentage  cover  values  in  the  six  groups  according  to  Orloci  ordination  for  the  pothole  and 
drainage  communities. 


Species 


1 


3 


Polygonum  coccineum* 
Lysimachia  hybrida^ 
Scirpus  fluviatilis^ 
C^arex  atherodes^ 
Spartina  pectinata^ 
Carex  aquatilis* 
Carex  retrorsa 
Phalaris  anmdinacea 
Sagittaria  latifolia 
Eleocharis  sp. 
Calamagrostis  canadensis^ 
Apocvnuni  sibiricum 
Lvcopus  americanus 
Convolvulus  sepium 
Teucrimn  canadense 
Carex  meadii 
Iris  virginica 
.\sclepias  incarnata 
Hordeum  jubatum 
Runiex  crispus 
Panicum  capillare 
Cirsium  altissimum 
Asclepias  sidlivantii 
Zizia  aurea 

Pycnanthemum  virginanum 
Elymus  canadensis 
Thalictrum  dasycarpuni 
Helenium  autumnale 
Helianthus  laetiflorus 
Anemone  cylindrica 
Solidago  rigida 
Gentiana  andrewsii 
Agrostis  hiemalis 
Heliopsis  helianthoides 
Ciciita  maculata 
Lythnuu  alatum 
Aster  ericoides 
Panicum  virgatuni 
Lathynmi  palustris 
Silphium  laciniatum 
Eryngiimi  yuccifolium 
Desmodium  canadense 
Liatris  pycnostachya 
Vernonia  fasciculata 
Rosa  sufhdta 
Fragaria  virginiana 
Senecio  pauperculus 
Solidago  gymnospermoides 
Audropogon  gerardi 
Poa  pratensis 
Solidago  canadensis 
Sporobolus  heterolepis 
.\ster  simplex 
Galium  obtusum 
Carex  lasiocarpa* 
Helianthus  grosseserratus 


66.15 

19.50 

5.55 

2.45 


.05 


26.74 

..39 

22.94 

10.56 

1.89 

1.44 

.80 

.39 

.20 

2.60 

.40 

.24 

.41 


31.15 

1.55 

.53.45 

.80 

.75 
.20 


.05 
3.15 


.40 


5.27 

.55 

.11 

.06 

17.54 

14.51 

3.27 

2.64 

1.21 

.73 

26.73 

.98 

.18 

.16 

.10 

.01 

.01 

.07 

.01 

.01 


.01 
.01 
.01 


.55 


1.15 


.01 


.01 


.02 


.24 
.32 


.58 
.01 

.86 

.35 

5.55 

.65 


2.58 
.36 

.12 

12.53 

3.42 

2.10 

2.21 

.01 

.29 

57.10 

2.08 

1..36 

.14 

.76 

.26 

.14 

.01 


.02 


.21 

.07 
.06 


.53 
.10 
.01 

.17 

.60 
.01 

2.01 
1.56 

7.43 
1.98 


.57 
.07 


2.20 

6.05 

.02 


6.42 
.75 

1.07 
.12 
.60 


.02 

.02 

.05 

.07 

.10 

.15 

.15 

.20 

.20 

.20 

.20 

.20 

.22 

.22 

.22 

.22 

.25 

.25 

.35 

.37 

.40 

.40 

.55 

.55 

.72 

1.32 

1.52 

1.70 

2.20 

2.35 

3.05 

4.07 

5.22 

5.65 

10.15 

13.10 


^Species  picked  by  the  three-dimensional  ordination  as  indicator  species. 


December  1980 


Brotherson:  Kalsow  Prairie,  Iowa 


379 


assigning  those  species  with  the  highest  val- 
ues for  Group  1  at  the  top  and  those  species 
with  the  highest  values  for  Group  6  at  the 
bottom  of  the  list.  It  was  then  possible  to  de- 
termine from  tlie  table  the  characteristic  dis- 
tributional patterns  of  man\'  of  the  species  as 
well  as  their  positions  of  importance  within 
each  zone  (i.e.,  Helianthus  grosseserratus  is 
mainly  restricted  to  Groups  5  and  6  and  is 
the  dominant  species  of  Group  6). 

The  species  of  these  different  zones  were 
also  ordinated  into  three-dimensional  space 
(Fig.  6).  The  ordination  did  not  yield  groups 
of  stronglv  associated  taxa  but  rather  picked 
out  eight  species  exhibiting  distinct  and  dif- 
ferent distributional  patterns.  It  placed  all 
other  species  within  the  areas  covered  by  the 
circles  A,  B,  and  C.  When  the  results  of  Fig- 
ure 6  are  compared  with  those  of  Table  1,  it 


can  be  seen  that  the  species  picked  bv  this 
method  as  indicator  species  are  those  taxa 
which  represent  the  dominants  or  sub- 
dominants  c.f  Groups  1  through  6. 

A  lOOO-foot  bisect  of  the  area  noted  in  Fig- 
ure 1  was  taken  in  an  effort  to  correlate  the 
distribution  of  the  dominants  of  each  zone 
with  elevation  and  topography.  This  informa- 
tion has  been  summarized  in  Figure  7.  The 
data  show  that  elevation  changes  of  6  to  12 
inches  altered  the  distribution  patterns  of  the 
zone  dominants. 

Attempts  to  pick  groups  of  associated  spe- 
cies through  the  application  of  Cole's  Index 
are  shown  in  Table  2  and  in  Figures  8  and  9. 
Figure  8  represents  a  clearly  definable  cluster 
and  includes  the  dominant  species  of  Groups 
1  through  3  of  Table  1.  These  species  are 
Carex  atfwrodes,   LijswiacJxia   lu/hrida.   Poh/- 


PERCENT    SIMILARITY 


-c 


^ 


Hi  polygonum    COCCINEUM      LTSMACHIA    HTBtdDA 

'lai  POLVCONUM  COCCINEUM 

14'  POLYGONUW   COCCINEUM  -SCIRPUS    FLUVIATIliS 

1 15  I  POLTCONUM    COCCINEUM   -  SClW^US    FLUVIATIttS 

9,  POLYGONUM   COCCINEUM   -  SOflPus    FLUVUTIks 

i6i  POLYGONUM   COCCINEUM  -  SCIRPUS    FLUVIAflLIS  -CAREX   ATHEBOOES 

1121  POLYGONUM    COCCINEUM  -SCtRPUS   FLUVIATILIS  -CA«I   ATMEROOCS 

1171  POLYGONUM   COCCINEUM  -  SCIflPUS   FLUVIATILIS  -CAREX   ATHEROOES 

171  POLYGONUM   COCCINEUM -CAREX     ATHERODES 

(191  POLYGONUM    COCCINEUM  -  CAREX     ATMEROOtS 

^13i  POLYGONUM   COCCINEUM  -CAREX     ATHEROOES 

II  POLYGONUM   COCCINEUM       CAREx    AOUATiLIS 

131  CAREX     AOUATILIS 

|29)  POLYGONUM  COCCINEUM   -SPARTIna    PECT 


]" 


LAMOCnOSTIS    CANADENSIS 


l32»      SPARTI 


PECT 


-CALAMOGROSTIS  CANADENSIS 

'  CALAMOGROSTIS  CANADENSIS 

-CALAMQOROSTIS  CANADENSIS 

-  CALAMOGROSTIS  CANADCNSIS 
-CALAMOGROSTIS  CANADENSIS 
-CALAMOGROSTIS  CANADENSIS 

CALAMOGROSTIS  CANADCNSIS 

-  CALAMOGROSTIS  CANADENSIS 

-  CALAMOGROSTIS  CANAOCNilS 

-CALAMOGROSTIS 

-  CALAMOGROSTIS 


SPARTINA     PECTtMATA-CALAMOG«OSTl 
CALAMOGROSTIS   CANADENSIS 
CALAMOGROSTIS    CANADENSIS 
CANADCNSIS 


28  CALAMOGROSTIS     CANADCNSIS 

—  3Si  PRAiW'l    COGC 

—  36  PRAWK    EDUC 
\3T'  PR»>"it    EDGB 


i34l     PRAIRIE    I 


Fig.  4.   Phenogram  of  36  potholo  zones  as  developed  from  cluster  analysis  tSneath  and  Sokai  1973).  Croups  are  a.s 
shown  in  Table  1  with  minor  exceptions. 


380 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


gonum  coccineum,  and  Scirpus  fluviatilis. 
Figures  8b  and  9  show  several  definable  clus- 
ters and  include  taxa  found  in  Groups  4 
through  6  of  Table  1.  The  cluster  of  species 

POLYGONUM  COCCINEUM-LYSIMACHIA   HYBRlDA  I 

i 

POLYGONUM  COCCINEUM- SCIRPUS   FLUVIATILIS  II 

I 

POLYGONUM  COCCINEUM -CARE X  ATHERODES  m 

1/  ^ 

SPARTINA  PECTINATA  ^CAREX    AQUATILIS  IV&V 

CALAMOGROSTIS  CANADENSIS 

CALAMOGROSIIS  CANADENSIS  VI 

I 

PRAIRIE    EDGE  VII 

Fig.  5.  Suggested  successional  sequence  for  potholes 
of  Kalsow  Prairie. 


designated  by  A  in  Figure  9  contains  species 
found  entirely  in  Groups  4  and  5  of  Table  1. 
Those  clusters  identified  by  the  letters  B  and 
C  of  this  same  figure  contain  only  plants 
found  in  Group  6  of  Table  1  and  correspond 
in  reality  to  the  prairie  edge.  Cluster  A  and 
Clusters  B  and  C  are  bridged  by  a  single  spe- 
cies {Aster  simplex)  that  is  found  growing 
mainly  along  the  border  between  Groups  5 
and  6  of  Table  1. 

The  vegetation  of  the  potholes  and  drain- 
age ways  of  Kalsow  Prairie  can  best  be  de- 
scribed as  a  series  of  five  zones  (Fig.  5),  each 
of  which  exhibits  different  spatial  and  floris- 
tic  properties.  This  characteristic  zonation 
can  be  expected  to  repeat  itself  from  pothole 
to  pothole  when  controlling  environmental 
factors  are  similar.  The  zones  themselves  are 
best  described  by  starting  at  the  center  of  the 
potholes  and  moving  toward  the  prairie  edge. 
Zone  1  (Group  1  of  Table  1,  etc.)  is  found  at 
the   center   of   the   deepest   potholes   and   is 


lOO— 


SO— 


X 


o  o 


09  O' 


-7f- 


Y 


50 


100 


220 


Fig.  6.  Two-dimensional  ordination  of  species  found  in  pothole  and  drainage  areas  of  Kalsow  Prairie;  A,  B,  and  C 
=  clusters  of  species  not  showing  distinct  distribution  patterns,  d  =  Polyg,onu)n  coccincuui  (usually  in  center  of 
pothole),  e  =  Sci>/;(/.s  fluviatilis.  f  =  Q/re.v  atherodes.  g  =  Lijsiutachia  hyhrida,  h  =  Spartina  pectinata.  i  =  Carcx 
(Ujuatilis,  i  =  Carex  la.siocurfm,  k  =  Calamagrostis  canadensis  (usually  in  outer  zone  of  pothole  complex). 


December  1980 


Brothersox:  Kalsow  Prairie,  Iowa 


381 


Table  2.  Cole's  Index  values  expressing  positive  interspecific  association  in  pothole  and  drainage  communities. 


Species 


Species 


X2a 


Ct" 


sde, 


Andropogon  gerardi 


Anemone  cvlindrica 


Apocvnuni  sibiricum 


.Aster  simplex 


Calamasrrostis  canadensis 


Carex  atherodes 

Carex  meadii 
Carex  aquatilis 

Carex  lasiocarpa 

Carex  retrorsa 
Convolvulus  sepium 
Desmodium  canadense 


Eleocharis  sp. 
Elvmus  canadensis 


Eryngium  ynecifoli\im 
Fragaria  virginiana 


Galium  ohtusum 


Helenium  autunniale 
Heliantlnis  grosseserratus 


Iris  virginica 


Eryngium  \aiccifolium 
Liatris  pycnostaclna 
Sporobolus  heterolepis 
Fragaria  virginiana 
Panicum  virgatum 
Thalictrum  das\carpum 
Calamagrostis  canadensis 
Carex  lasiocarpa 
Carex  retrorsa 
Spartina  pectinata 
Calamagrostis  canadensis 
Carex  lasiocarpa 
C.alium  ohtusum 
Heliantlius  grosseserratus 
Poa  pratensis 
Spartina  pectinata 
Carex  acjuatilis 
Carex  lasiocarpa 
Carex  retrorsa 
Spartina  pectinata 
Polygonum  coccincum 
Scirpus  fluviatilis 
Rosa  suffulta 
Carex  lasiocarpa 
Carex  retrorsa 
Spartina  pectinata 
Carex  retrorsa 
Spartina  pectinata 
Spartina  pectinata 
Galium  ohtusum 
.\ster  ericoides 
Liatris  pycnostachya 
Ratihida  columnifera 
Senecio  pauperculus 
Phalaris  anmdinacea 
.\ster  ericoides 
Galium  ohtusum 
Helianthus  grosseserratus 
Lathvrus  palustris 
Desmodimn  canadense 
Liatris  pycnostachya 
Senecio  pauperculus 
Thalictrum  dasycarpum 
Sporoholus  heterolepis 
Andropogon  gerardi 
Ciaiium  ohtusum 
Helianthus  grosseserratus 
Poa  pratensis 
Senecio  pauperculus 
Solidago  canadensis 
Sporoholus  heterolepis 
Helianthus  grosseserratus 
Poa  pratensis 
Spartina  pectinata 
Helianthus  grosseserratus 
Poa  pratensis 
Solidago  canaclciisis 
Poa  pratensis 
Rosa  sufhilta 
Vemonia  fasiculata 


119.80 

.30 

.02 

130.35 

.38 

.03 

65.90 

.22 

.02 

89.71 

.19 

.02 

84.61 

.24 

.02 

91.73 

..33 

.03 

41.22 

.68 

.10 

30.42 

.57 

.10 

9.44 

.18 

.05 

19.63 

..37 

.08 

.36.47 

.56 

.09 

31.79 

.51 

.09 

212.39 

.51 

.03 

231.49 

.51 

.03 

85.80 

.18 

.01 

19.10 

.31 

.07 

49..30 

.17 

.02 

338.21 

.68 

.03 

81. .33 

.19 

.02 

200.44 

.41 

.02 

153.47 

.93 

.07 

120..34 

.47 

.04 

28.47 

..38 

.07 

73..33 

.49 

.05 

53.85 

.24 

.03 

19.61 

.20 

.04 

111.72 

.70 

.06 

18.3.90 

.40 

.02 

43.76 

.35 

.05 

8.76 

..39 

.13 

.54.86 

.28 

.03 

87.08 

.42 

.04 

200.56 

.28 

.02 

189.69 

.71 

.05 

28.62 

.25 

.04 

65.57 

..33 

.04 

4..34 

..39 

.18 

.30.83 

1.00 

.18 

62.34 

.49 

.06 

64.57 

..33 

.04 

44.21 

.32 

.04 

.33.25 

..32 

.05 

91.73 

.33 

.03 

.54.86 

.28 

.03 

.37.61 

.19 

.03 

46.46 

.70 

.10 

104.86 

1.00 

.09 

19.29 

.2.5 

.05 

37.61 

.19 

.a3 

.30.90 

.27 

.04 

75.87 

.19 

.02 

242.41 

.60 

.03 

86.58 

.20 

.02 

13.73 

.29 

.07 

5..54 

.60 

.2.5 

131.97 

.2.5 

.02 

112.89 

.20 

.01 

7..38 

.29 

.10 

23. 1 1 

..32 

.06 

8.39 

.29 

.10 

382 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


Table  2  continued. 


Species 


Species 


Lathyrus  paliistris 

I  .iatris  pycnostachya 

I.Nsiniachia  hybrida 

Lvthnnn  alatuin 
Panicuni  virgatum 

Poa  pratensis 
Polygonum  coccineum 
Rosa  sufftilta 

Solidago  canadensis 
Zizia  aurea 


Poa  pratensis 

Solidago  gymnospermoides 
Thalictruni  dasycarpiun 
Senecio  pauperculus 
Sporobolus  heterolepis 
Polvgoniun  coccineum 
Scirpus  fluviatilis 
Vernonia  fasciculata 
Poa  pratensis 
Teucrium  canadense 
Solidago  canadensis 
Scirpus  fluviatilis 
Andropogon  gerardi 
Solidago  canadensis 
Senecio  paupercidus 
Desm  odium  canadense 


^hi-square 

"Cole's  Index 

'^Standard  dexiation  Cole's  Index 


-rNNW 


FEET 


X^a 


1000 


Sd&, 


9.53 

.20 

.06 

62.18 

.30 

.03 

75.89 

.18 

.02 

44. .56 

.29 

.04 

248..50 

..50 

.03 

16.21 

.61 

.15 

16.11 

..34 

.08 

5.46 

.20 

.08 

14.89 

.29 

.07 

45.6.3 

..39 

.05 

92.29 

.30 

.03 

223.30 

.32 

.02 

42.51 

.21 

.03 

.35.46 

..30 

.05 

160.63 

.30 

.02 

1.32.56 

.66 

.05 

Fig.  7.  Correlation  of  dominant  species  of  each  zone  from  potholes  and  drainages  with  changes  in  elevation  along 
1,000  ft  (925  m)  bisect;  1  =  Sporobolus  heterolepis,  2  =  Helianthus  ^rosseserrottis.  3  =  Calarnagrostis  canadensis.  4 
=  Carex  atherodes,  5  =  Scirpus  fluviatilis,  6  =  Polygonum  coccineum. 


dominated  chiefly  by  Polygonum  coccineum 
and  Lijsimachia  hijhrida.  Zone  2  is  found  to 
conipletelv  encircle  Zone  1  and  is  character- 
ized by  the  dominants  Polygonum  coccineum 
and  Scirpus  fluviatilis.  Zone  3  is  found  as  a 
very  narrow  band  that  encircles  Zone  2  or 
occurs  as  rather  wide  patches  in  areas  of 
equivalent  elevation.  It  is  characterized 
chiefly  by  Carex  atherodes.  Zones  4  and  5  are 
best  distinguished  in  potholes  and  drainage 
ways  that  are  somewhat  shallow.  Zone  4 
most  often  occupies  the  center  of  these  shal- 
low depre.ssions  surrounded  by  Zone  6.  The 
dominant  species  {Carex  aquatilis)  of  Zone  5 
is  u.sually  found  as  a  subdominant  of  Zone  4 


but  at  times  appears  as  a  dominant  zone  sur- 
rounded bv  Zone  6.  Whether  this  relationship 
is  due  to  competition  and/ or  environmental 
influences  is  unknown.  Zone  4  is  character- 
ized by  the  species  Spartina  pectinata,  Carex 
aquatilis,  and  Calamogrostis  canadensis. 
Zone  6  is  distinguished  by  the  dominant  spe- 
cies Calamagrostis  canadeiisis  and  a  few 
other  participating  species  (i.e.,  Apocynum 
sibiricum,  Lycopus  americanus,  Teucrium 
canadense,  Carex  meadii,  and  Iris  virginica). 
Zone  7  and  column  6  of  Table  1  represent 
the  prairie  edge  and  are  characterized  pri- 
marilv  by  the  presence  of  Helianthus  grosse- 
serratus. 


December  1980 


Brotherson:  Kalsow  Prairie,  Iowa 


383 


These  zones  appear  to  represent  a  succes- 
sional  sequence  that  is  controlled  basically  by 
the  degree  of  pothole  fill  and  consequently 
by  related  moisture  regimes.  The  successional 
scheme  (Fig.  5)  parallels  in  many  respects  a 
scheme  proposed  bv  earlier  authors  (Sherff 
1912,  Schaffner  1926,  Hayden  1943). 

The  actuality  of  this  scheme  is  based  on 
the  repeatability  of  the  zonation  pattern  as 
found  within  the  potholes  of  Kalsow  Prairie. 
Evidence  for  change  of  fluctuations  in  pot- 
hole vegetation  paralleling  this  sequence  will 
depend  on  the  results  obtained  from  long- 
term  studies. 

Literature  Cited 

AiKMAN,  J.  M.  1959.  Prairie  research  in  Iowa.  American 
Biology  Teacher  21:7-8. 

.\iKMAN.  J.  M.,  AND  R.  F.  Thorne.  1956.  Tlie  (Javier 
Prairie:  an  ecologic  and  taxonomic  stuck'  of  a 
northwest  Iowa  Prairie.  Iowa  Acad.  .Sci. 
ft3: 177-200. 

Berry,  W.  J.  1927.  The  influence  of  natural  environment 
in  north  central  Iowa.  Iowa  J.  Historv  and  Poli- 
tics 25:277-298. 


Fig.  8.  .\ssociated  species  of  potholes  and  drainage, 
Groups  I  and  2  (Table  1)  as  determined  by  Cole's  (1949) 
Index;  (a)  Ca  at  =  Carex  athcrodes.  Ly  hy  =  Lysi- 
machia  liyhrida,  Po  co  =  Polygonum  coccineum,  Sc  fl 
=  Scirpns  fluviatilis\  (b)  El  sp  =  Eleocharis  sp.,  Ph  ar 
=  Phalaris  unindinacea. 


Bhen.nan,  K.  M.  1969.  Vertebrate  fauna  of  Kalsow 
Prairie.  I'npublished  thesis.  Iowa  State  Univer- 
sity. 

Cole,  L.  C.  1949.  The  measurement  of  interspecific  as- 
sociation. Ecology  .30:411-424. 

Daibe.N'mire,  R.  1959.  A  canopy-coverage  method  of 
vegetational  analvsis.  Northwest  Science 
.33:4.3-66. 

Dic:k-Peddie,  VV.  .\.  1955.  Presettlement  forest  types  in 
Iowa.  Unpublished  thesis.  Iowa  State  Univcrsitv. 

Do.NDORE,  D.  \.  1926.  The  prairie  and  the  making  of 
middle  .America:  four  centuries  of  description. 
Cedar  Rapids,  Iowa,  Torch  Press.  472  pp. 

Ehrenreich,  J.  H.  1957.  Management  practices  for 
maintenance  of  native  prairies  in  Iowa.  Unpub- 
lished dissertation,  Iowa  State  l'niversit\'. 

Esau,  K.  L.  1968.  Carabidae  (Coleoptcra)  and  other  ar- 
thropods collected  in  pitfall  traps  in  Iowa  corn- 
fields, fencerows  and  prairies.  Unpublished  dis- 
sertation, Iowa  State  University. 

GiLLY,  C.  L.  1946.  The  Cvperacae  of  Iowa.  Iowa  State 
Coll.  J.Sci.  21:.5.5-151. 

(Ileason,  H.  \.  1952.  The  new  Britton  and  Brown  illus- 
trated flora  of  the  northeastern  United  States  and 
adjacent  Canada.  Lancaster,  Penns\lvania.  Lan- 
caster Press,  Inc.  3  vols. 

H.\yde.\,  .\.  1943.  .\  botanical  survey  in  the  Iowa  lake 
region  of  Clav  and  Palo  .\lto  Counties.  Iowa 
State  Coll.  }.  17:277-416. 

1945.   The  selection   of  prairie  areas   in   Iowa 

which  should  be  preserved.  Proc.  Iowa  .\cad.  Sci. 
52:127-148. 

1946.  .\  progress  report  on  the  preservation  of 

prairie.  Proc.  Iowa  .\cad.  Sci.  .53:45-82. 

Hewes,  L.  19.50.  Some  features  of  earlv  woodland  and 
prairie  settlement  in  a  central  Iowa  count)'.  .An- 
nals .\ssoc.  .\mer.  Geog.  40:40-.57. 

1951.   The  northern  wet   prairie  of  the   United 

States:  nature,  sources  of  information,  and  extent. 
.\nnals  .\ssoc.  .\mer.  Geog.  41:.307-.32.3. 

Hewes,  L.,  a.nh  P.  E.  Fra.ndson.  19.52.  Occupying  the 
wet  prairie:  the  role  of  artificial  drainage  in  Story 
Countv,  Iowa.  .\nn.  .\ssoc.  .\mer.  Geog. 
42:24-.50. 

Kaesler,  R.  L.,  a.nd  J.  Cair.ns.  1972.  Cluster  analysis  of 
data  from  limnological  surveys  of  the  upper  Poto- 
mac River.  .\m.  Midi.  Nat.  88:56-67. 

Kennedy,  R.  K.  1969.  \x\  analysis  of  tall-grass  prairie 
vegetation  relative  to  slope  position,  Shccder 
Prairie,  Iowa.  Unpublished  thesis,  Iowa  State 
Universitv. 

Landers,  R.  Q.  1966.  Visit  the  \irgin  prairie.  Iowa  Farm 
Science  21:418-419. 

Marean,  H.  VV.,  AND  G.  B.  Jones.  1903.  Soil  survey  of 
Storv  Countv,  Iowa.  Field  Operations  of  the  Bu- 
reau of  Soils  836. 

MoYER.  J.  F.  1953.  Ecologv'  of  native  prairie  in  Iowa. 
I'npublished  dissertation,  Iowa  State  University. 

Norton,  D.  C..  and  P.  E.  Ponchilla.  1968.  Stylet-bear- 
ing nematodes  associated  with  plants  in  Iowa 
prairies.  J.  Iowa  .\cad.  Sci.  75:32-35. 

Oreoci.  L.  1966.  Geometric  models  in  ccolog).  I.  The 
theorv  and  application  of  some  ordination  meth- 
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PoHL,  R.  \\'.  1966.  The  grasses  of  Iowa.  Iowa  State  J. 
Science  40:341 -.566. 


384 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


Fig.  9.  Associated  species  of  potholes  and  drainage.  Groups  3-6  (Table  1)  as  determined  by  Cole's  (1949)  Index, 
the  more  lines  between  species,  the  greater  the  association;  groups  A,  B,  and  C  are  basic  clusters;  .\n  ge  =  Andwpo- 
gon  gerardi.  An  cv  =  Anemone  eijlindnca,  Ap  si  =  Apoci/ntim  sihiiicuni.  As  er  =  Aster  ericoides.  As  si  =  Aster 
simplex.  Ca  aq  =  ('arex  cujuatilis,  Ca  ca  =  Cakimagrostis  eanadensis,  Ca  la  =  Carex  hisiocarpa,  Ca  me  =  Carex 
meadii.  Ca  re  =  Carex  retrorsa,  De  ca  =  Desmodiiim  canadensis.  El  ca  =  Elymits  canadensis,  Er  yu  =  Enjngium 
i/iicci folium.  Fr  vi  =  Fragaria  virginiana,  Ga  ob  =  Galium  obtusum.  He  gr  =  Helianthus  grosseserratus,  Ir  vr  =  Iris 
virginica.  La  pa  =  Lathyrus  paUistris,  Li  pii  =  Liatris  pycnostachya,  Ly  al  =  Lythrum  alatum.  Pa  vi  =  Panicum 
virgatum.  Po  pr  =  Poa  pratensis,  Ra  co  =  Ratibida  columnifera,  Ro  su  =  Rosa  suffulta,  Se  pa  =  Serteeio  pauper- 
culus.  So  ca  =  Solidago  canadensis.  So  gy  =  Solidago  gymnospermoides.  Sp  he  =  Sporobohis  heterolepis,  Sp  pe  = 
Spaiiina  pectinata.  Te  ca  =  Teucrium  canadense,  Tha  da  =  Tlialictrum  dasycarpum.  Ve  fa  =  Vernonia  fasicukita. 
Zi  au  =  7.iziii  aurea. 


Richards,  M.  S.  1969.  Observations  on  responses  of 
prairie  vegetation  to  an  .\pril  fire  in  central 
Iowa.  Unpublished  thesis,  Iowa  State  University. 

RiHE,  R.  V.  1969.  Quaternary  landscapes  in  Iowa.  .Ames, 
Iowa,  Iowa  State  University  Press.  249  pp. 

RuziCKA,  M.  1958.  Anwendung  mathematish-statisticher 
methoden  in  der  Geobotanik  (Synthetische  bear- 
bietung  von  aufnahmen).  Biologia,  Bratisl. 
1.3:649-661. 

Rydberg,  p.  \.  1931.  A  short  phvtogeographv  of  the 
prairies  and  great  plants  of  Central  North  .\nieri- 
ca.  Brittonia  1:57-66. 

ScHAFFNER,  J.  H.  1926.  Grasslands  of  the  Central  United 
States.  Ohio  State  Universitv  Studies  Contr.  Bot. 
178:5-56. 

Sc:hmitt,  D.  p.  1969.  Plant  parasitic  nematodes  and 
nematode  populations  in  the  Kalsow  Prairie.  Un- 
published thesis,  Iowa  State  Universitv. 

Sherff,  E.  E.  1912.  Tlie  vegetation  of  Skokie  marsh, 
with  special  reference  to  subterranean  organs 
and  their  interrelationships.  Bot.  Gaz. 
53:415-435. 


Shi.mek,  B.  1915.  The  plant  geography  of  the  lake  Oko- 
boji  region.  Bull.  Lab.  Nat.  Hist.,  State  Univ. 
Iowa  7:1-90. 

Sne.\th,  p.  H.  a.,  and  R.  R.  Sokal.  1973.  Numerical  ta.x- 
onomy.  W.  H.  Freeman.  San  Francisco,  Califor- 
nia. 573  pp. 

Steiger,  T.  L.  1930.  Structure  of  prairie  vegetation. 
Ecology  11:170-217. 

Traucer,  D.  L.  1967.  Habitat  factors  influencing  duck 
brood  use  of  semi-permanent  and  permanent 
prairie  potholes  in  North  Dakota.  Unpublished 
thesis,  Iowa  State  University. 

U.S.  Government.  1868.  First  survey  of  the  state  of 
Iowa.  Plants  deposited  in  the  State  House,  Des 
Moines. 

Weaver,  J.  E.,  and  F.  E.  Clements.  1938.  Plant  ecolo- 
gy. 2d  ed.  McGraw-Hill,  New  York. 

Weaver,  J.  E.,  and  T.  J.  Fitzp.atrick.  1934.  The  prairie. 
Ecol.  Monogr.  4:109-295. 

Yapp,  R.  H.  1909.  On  stratification  in  the  vegetation  of  a 
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perature. Ann.  Bot.  23:275-320. 


PLANTS  OF  ANGEL  ISLAND,  MAIUN  COUNTY,  CALlFORNL\ 

J.  D.  Ripley' 

Abstract.—  The  floristic  composition  of  .\nge\  Island,  Marin  County,  California,  is  presented.  The  vascular  flora 
consists  of  416  native  and  naturalized  plants,  representing  2.52  genera  and  7.J  families.  Eight)  species  of  ornamental 
plants  persisting  from  cultivation,  representing  64  genera  and  '39  families,  are  also  identified.  The  nonvascular  flora 
consists  of  42  species  of  fvuigi,  7  species  of  green  algae,  8  species  of  brown  algae,  and  41  species  of  red  algae.  The 
\ascular  flora  occurs  in  seven  plant  communities  and  is  composed  mainly  of  the  following  plant  types:  forbs  (69 
percent),  grasses  (1.3  percent),  shnibs  (8  percent),  and  trees  (4  percent),  .\pproximately  25  percent  of  the  1605  native 
and  naturalized  vascular  plants  of  Marin  ('ounty  occur  on  .Xngel  Island. 


Angel  Island  is  situated  in  San  Francisco  Tiburon  Peninsula,  hv  Raccoon  Strait,  which 

Bav  off  the  southern  coast  of  Marin  Countv,  is  about   a   mile   in   width   at   the   narrowest 

California.  The  island  lies  approximately  3 '2  point. 

miles  north  of  the  city  of  San  Francisco  and  The  shape  of  the  island,  as  shown  in  Figure 

is  separated  from  the  nearest  mainland,  the  1.  roughly  resembles  an  equilateral  triangle 


10 


12 


13 


15 


16 


17 


19 


20 


CAMPBELL    POINT 

North    Garrison 


SIMPTON    POINT 


POINT    lONE 


STUART    POINT 


West    Garrison 
Camp     Reynolds 

KNOX    POINT 


LEGEND 

aacz:^     Fire     Road 
=  =  =  ==      Lower     Road 

Sunset    Trail 

North     Ridge     Trail 


FT.    Mc    DOWELL 


East    Gorrison 


Quorry    Beach 


SAN    FRANCISCO    BAY 


BLUNT    POINT 


10 


12 


13 


15 


16 


19 


20 


Fig.  I.    Mapot  Augcl  MaiKJ. 
■Department  of  Biology,  US.\F  .\cademy.  Colorado  80840.       Present  address:  Departn.cnt  of  Botanv  and  PUnt  Pathology.  OreRon  Stale  University. 


Corvallis,  Oregon  973.30. 


385 


386 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


with  the  sides  measuring  about  VA  miles  in 
length.  Hence,  the  area  of  the  island  is 
sliglitly  larger  than  one  square  mile  and  equal 
to  approximately  740  acres. 

The  topography  of  the  island  is  roughly 
pyramidal  in  form,  with  the  central  peak,  Mt. 
Livermore  (elevation  781  ft),  forming  the 
central  axis  for  various  spurs  that  radiate  out 
to  the  perimeter  of  the  island. 

Numerous  water  seepages  occur  in  the 
canyons  formed  by  the  radiating  ridges,  but  it 
is  only  in  the  canyon  extending  from  the 
summit  of  Mt.  Livermore  to  Perle's  Beach 
that  any  significant  water  flow  occurs.  The 
damming  of  this  small  stream  in  two  loca- 
tions has  resulted  in  the  formation  of  two 
small  ponds. 

With  the  exception  of  several  miles  of  nar- 
row, sandy  beaches,  the  island  is  generally 
hilly.  In  some  areas,  particularly  near  Simp- 
son Point  and  Blunt  Point,  the  cliffs  are  ex- 
tremely steep. 

Climate 

Due  to  the  steady  sweep  of  air  from  the 
Pacific  Ocean,  the  climate  of  Angel  Island  is 
characterized  by  few  extremes  of  heat  or 
cold.  Like  other  central  coastal  regions  of 
California,  the  winters  are  cool  and  wet  and 
the  summers  are  warmer  and  dry.  Until  re- 
cently, only  intermittent  precipitation  and 
temperature  records  were  maintained  on  the 
island.  The  limited  data  available  indicate  a 
total  annual  precipitation  ranging  from  a  low 
of  2L34  inches  to  a  high  of  34.53  inches, 
with  the  average  being  26.62  inches.  These 
data  further  reveal  that  84  percent  of  the  to- 
tal average  precipitation  occurs  during  the 
five-month  period  between  November  and 
March  and  that  measurable  amounts  of  pre- 
cipitation occur  on  the  average  less  than  70 
days  a  year.  As  in  many  coastal  regions  of 
California,  summer  fog  and  low  overcast  play 
an  important  role  in  tlie  island's  climate. 
Summer  mornings  are  frequently  foggy,  but 
clearing  usually  begins  early  in  the  forenoon. 
These  low  summer  fogs  no  doubt  contribute 
soil  moisture  as  condensation  occurs  on 
leaves  of  trees  and  drips  to  the  ground  (Ober- 
lander  1956). 

The  average  annual  temperature  on  Angel 
Island  is  56  F.  The  average  monthly  mini- 


mum temperature  occurs  in  January  and  is  42 
F,  and  the  average  monthly  maximum  tem- 
perature occurs  in  September  and  is  73  F. 
The  average  monthly  temperature  ranges 
from  a  low  of  46  F  in  January  to  a  high  of  65 
F  in  September. 

Geology 

Ransome  (1894)  conducted  the  first  and 
most  extensive  study  of  the  island's  geology. 
This  study  has  been  supplemented  and  up- 
dated by  Bloxam  (1960)  and  by  Schlocker, 
Bonilla,  and  Radbruch  (1958),  who  compiled 
an  extensive  geological  map  of  the  island. 
The  geological  features  of  the  island  have 
also  provided  source  material  for  other  stud- 
ies of  rock  formations  in  the  San  Francisco 
Bay  area  such  as  those  of  Bloxam  (1956), 
Schlocker  (1961),  and  Bailey,  Irwin,  and 
Jones  (1964).  Ransome  (1894)  describes  the 
structure  of  the  island  as  consisting  essen- 
tially of  a  syncline  trough  with  its  axis  plung- 
ing to  the  northeast.  The  rocks  of  Angel  Is- 
land all  belong  to  the  Franciscan  Formation 
and  consequently  were  formed  during  the  Ju- 
rassic approximately  100  million  years  ago. 
Bloxam  (1960)  identified  the  following  six 
major  rock  groups  on  the  island:  (1)  Francis- 
can graywackes,  conglomerates,  and  radio- 
larian  cherts;  (2)  jadeite-bearing  metagray- 
wackes  derived  from  the  Franciscan 
graywackes;  (3)  glaucophane-schists  derived 
from  graywackes,  cherts,  and  basic  igneous 
rocks;  (4)  a  large  and  apparently  concordant 
sheet  of  altered  diabase  intrusive  into  the 
Franciscan  sediments;  (5)  pillow-lavas  at 
Blunt  Point;  and  (6)  a  vertical  dikelike  body 
of  serpentinite  and  pyroxenite  near  the  west- 
ern end  of  the  island.  Of  these  rock  groups, 
the  near  vertical  serpentine  dike  extending 
across  the  western  portion  of  the  island  is  the 
predominant  geological  feature  of  the  area. 
Bloxam  (1960)  further  points  out  that  all  the 
rocks  of  the  island  are  metamorphosed  to 
various  degrees,  with  the  exception  of  the 
graywacke  and  shale  at  Quarry  Point,  which 
are  unmetamorphosed. 

Vegetation  Communities 

Seven  categories  of  vegetation  cover  are 
recognized  as  occurring  on  Angel  Island. 


December  1980 


Ripley:  Angel  Island  Plants 


387 


S  niXED  EVERGRtEM  FOREST 
I      1  GRASSLAND 
\M  CHAPARRAL 
H  COASTAL  SCRUB 
H  COASTAL  5TRAND 
^  DISTURBED  AREAS 
FRESHWATER  POND 


Fi<i.  2.   Map  ot  Angel  Island  showing  location  ot  its  se\("n  tatcgories  of  vegetation  cover. 
Table  1.  Tabulation  of  the  native  and  naturalized  vacular  flora. 


Division 
Class 

Family 

Genus 

Specific 

and  infraspec 

ific  taxa 

Subclass 

Native 

Naturalized 

Total 

Calamophyta 
Traeht'ophvta 

I 

1 

2 

0 

2 

Filicinae 

3 

7 

7 

0 

7 

Cyinnospermae 
.\ngiosperiTiae 

Monocotvledonae 

2 
10 

2 
45 

0 
60 

2 
30 

2 

90 

Dicotyledoneae 

57 

197 

213 

102 

315 

Totals 

73 

252 

282 

134 

lib 

These  include  mixed  evergreen  forest  (33 
percent),  grassland  (28  percent),  altered  or 
disturbed  areas  (20  percent),  coastal  scrub  (10 
percent),  chaparral  (4  percent),  coa.stal  strand 
(4  percent),  and  freshwater  ponds  (less  than 
one  percent).  The  location  of  each  vegetation 
type  is  indicated  in  Figure  2. 


Flora  of  the  Island 

The  native  and  naturalized  vascular  flora 
of  Angel  Island  consists  of  73  families.  252 
genera,  and  416  species  and  infraspecific  taxa 
(Table  1).  In  addition,  80  species  of  culti- 
vated plants  have  been  identified  (Table  2)  as 


388 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


Table  2.  Tabulation  of  the  nonvascular  flora  identi- 
fied. 


Division 


Genus        Species 


Eumycophyta 
Chlorophyta 
Phaeophyta 
Rhodophyta 

Totals 


28 
5 

8 
33 

74 


42 
7 
8 

41 


well  as  42  species  of  higher  fungi  and  56  spe- 
cies of  marine  algae  (Table  3).  The  10  largest 
families  are  listed  in  Table  5  and  contain  138 
genera  and  265  species,  or  64  percent  of  the 
total  native  and  naturalized  vascular  flora. 
Comparing  the  416  species  and  infraspecific 
taxa  or  native  and  naturalized  vascular  plants 
with  the  1605  species  and  infraspecific  taxa 
recognized  by  Howell  (1970)  for  all  Marin 
County  indicates  that  the  flora  of  Angel  Is- 
land represents  approximately  25  percent  of 
the  total  Marin  County  flora.  No  endemic 
species  occur  on  Angel  Island  nor  is  the  is- 
land the  type  locality  for  any  plants. 


Table  3.   Tabulation  of  the  cultivated  plants  identi 
fied. 


Class 
Subclass 


Family       Genera       Species 


Gymnospermae  5  11  12 

Angiospermae 

Monocotyledoneae  5  8  10 

Dicotyledoneae  29  45  58 

Totals  39  64  80 


Checklist 

The  relative  inaccessibility  and  long  mili- 
tary history  of  Angel  Island  precluded  a  de- 
tailed floristic  examination  of  the  area  prior 
to  the  study  upon  which  the  checklist  below 
is  largely  based  (Ripley  1969).  Early  collec- 
tors included  Albert  Kellogg  (no  date), 
George  Vasey  (1876),  T.  S.  Brandegee  (no 
date),  Volney  Rattan  (1877),  and  Joseph 
Burtt-Davy  (no  date).  Only  one  or  two  Angel 
Island  collections  exist  from  each  of  these 
collectors.  Collectors  in  the  first  half  of  the 


Table  4.  Plant  characteristics  of  Angel  Island  native,  introduced,  and  cultivated  plants. 


Percent  of 

Percent  of 

total  flora 

total  native 

(native. 

Generalized  life  for 

ni 

Total 

Percent  of 
individual 
category 

and 

introduced 

flora 

introduced, 
and 

Growth  habit 

Annual 

Biennial 

Perennial 

cultivated) 

I.  Native  plants 

Fems 

0 

0 

7 

7 

3 

1,7 

1.3 

Sedges 
Rushes 

1 
1 

0 
0 

5 

5 

6 
6 

2 
2 

1.5 
1.5 

1,2 
1,2 

Grasses 

5 

1 

24 

30 

11 

7.3 

6,0 

Forbs 

73 

8 

110 

191 

67 

46.0 

.39.0 

Shrubs 

0 

0 

.30 

30 

11 

7,2 

6.0 

Trees 

0 

0 

10 

10 

4 

2.4 

2.0 

Subtotals 

80 

9 

191 

280 

100 

67.3 

56.4 

II.  Introduced  plants 

Grasses 

17 

0 

8 

25 

19 

6,0 

5,0 

Forbs 

61 

8 

28 

97 

71 

23,0 

19,9 

Shnibs 

0 

0 

7 

7 

5 

1,7 

1,3 

Trees 

0 

0 

7 

7 

5 

1,7 

1,3 

Subtotals 

78 

8 

50 

136 

100 

32.7 

27,4 

III.  Cultivated 

(ira.sses 

Forbs 

Shnibs 

Trees 

PLANTS 

0 
0 
0 
0 

0 

0 
0 
0 
0 

0 

1 

6 
22 
51 

80 

1 

6 

22 

51 

80 

1,3 

7.5 

26.5 

64.7 

,20 
1,2 
4.4 
10.0 

Subtotals 

100.0 

16.2 

Grand  totals 

158 

17 

320 

493 

December  1980 


Ripley:  Angel  Island  Plants 


389 


Table  5.  The  10  largest  families  of  vascular  plants. 


Species 

and 

infraspecific 

Family 

Genera 

taxa 

Compositae 

47 

76 

Gramineae 

25 

61 

Letjuniinosae 

12 

45 

ScTophiilariaceae 

7 

16 

Cniciterac 

14 

16 

Rosaccac 

10 

12 

I'mhelliferae 

10 

12 

Car\()phyllaceae 

6 

11 

Onagraceae 

4 

9 

Cyperaceae 

3 

7 

Totals 

138 

265 

twentieth  centurv  included  E.  C.  Sntliffe 
(1920),  Alice  Eastwood  (1925),  and  John 
Thomas  Howell  (1946  and  1949).  The  51 
specimens  collected  by  Howell  are  the  most 
important  of  these  collections.  In  addition, 
numerous  plants  were  observed  but  not  col- 
lected by  Howell  on  his  two  trips  that 
formed  the  basis  for  many  citations  in  Marin 
Flora  (1970).  The  most  recent  collectors  have 
included  Otto  T.  Solbrig  (1956),  Peter  H. 
Raven  and  Michael  P.  Johnson  (1967),  J. 
Douglas  Riplev  (1967-1969),  Gordon  True 
(1978),  Alva  Dav  (1979-1980),  and  Catherine 
G.  Burke  (1979-1980).  Of  these  collections, 
the  most  extensive  are  those  of  Ripley  with 
1074  and  Raven  and  Johnson  with  337. 

The   following   abbreviations   are   used   in 
this  checklist   to   indicate   individual   collec- 


tors 
B 

B-D 
D 


(Catherine  G.  Burke 

Joseph  Bvirtt-Davy 

Alva  Dav 
E  Alice  Eastwood 

H  John  Thomas  Howell 

K  Albert  Keilogt; 

R  V'olnev  Rattan 

JDR     J.  Douglas  Ripley 

R  &  J    Peter  H.  Raven  and  Michael  P.  Johnson 
S  E.  G.  Sutlifte 

OS        Otto  T.  Solbrig 
T  Gordon  True 

TB        T.  S.  Brandegee 
\'  George  Vasey 

Certain  plant  characteristics  are  also  in- 
dicated in  the  checklist  following  the  scien- 
tific name  for  all  vascular  plants.  The  follow- 
ing abbreviations  are  used  for  this  purpose: 


P  perennial 

B  biennial 

A  annual 

N  native 

I  introduced 

('  ornamental  plant  persisting  from  cultivation 

T  tree 

S  shrub 

F  fori) 

Fr  fern 

G  grass 

R  rush 

Sd  sedge 

Finally,  the  approximate  collection  loca- 
tion of  each  specimen  is  indicated  bv  map 
grid  coordinates  from  Figure  1 . 

The  checklist  is  arranged  in  alphabetical 
order  by  families  for  ease  of  reference.  Iden- 
tification of  the  algal  flora  follows  that  of 
Smith  (1944)  and  Hollenberg  and  .\bbott 
(1966).  The  vascular  plant  identification  fol- 
lows that  of  Munz  (1973)  and  Howell  (1970). 

Division  Eumycophyta— Higher  Fimgi 

Agrocybe  pediades  (Fr.)  Favod. 

JDR  1008,  2  Mar  1968, 'L-6;  JDR  1048,  31 
Mar  1968,  G-12. 

Aleuria  atirantia  (Fr.)  Fuckel. 
JDR  1000,2  Mar  1968,  F- 10. 

Amanita  gemmata  (Fr.)  Gill. 
JDR  1005,  2  Mar  1968,  F- 10. 

Amanita  pantherina  (DC.  ex  Fr.)  Seer. 
JDR  1656,  30  Dec  1968,  F-10. 

Amanita  rubescens  (Pers.  ex  Fr.)  Gray 
JDR  1704,  20  Apr  1969,  E-Il. 

Amanita  vaginata  (Fr.)  Vitt. 
JDR  998,  2  Mar  1968,  F-10. 

Bolbitius  vitcllinus  (Fr.)  Fr. 

JDR  975,  6  Feb  1968,  HI  3;  JDR  1002,  2 
Mar  1968,  D-11. 

Boletus  chrysenteron  (Bull,  ex  Fr.)  Q)uel. 
JDR  1003,  2  Mar  1968,  F-12. 

Boletus  subtomentosus  (L.  ex  Fr.)  Quel. 
JDR  1662,  30  Dec  1968,  D-10. 

Chroogomphus  rutilus  (Fr.)  Miller 
JDR  974,  6  Feb  1968,  J-9. 

Clavariadelphus  ligula  (Fr.)  Donk. 
JDR  963,  6  Feb  1968,  F-10. 

Clavulina  cristata  (Fr.)  Schroet. 
JDR  1660,  30  Dec  1968.  F-10. 


390 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


Laccaria  striatula  (Peck)  Peck 
JDR  1659,  30  Dec  1968,  F-10. 

Lactarius  chrysorheus  Fr. 
JDR  964,  6  Feb  1968,  F-10. 

Lactarius  suhdulcis  (Bull,  ex  Fr.)  Gray 

JDR  965,  6  Feb  1968,  F-10;  JDR  1669,  30 
Dec  1968,  E-12. 

Lepiota  rachodes  (Vitt.)  Quel. 
JDR  944,  27  Dec  1967,  J-4. 

Marasmius  Candidas  Fr. 
JDR  999,  2  Mar  1968,  F-10. 

Marasmius  plicatulus  Peck 

JDR  968,  6  Feb  1968,  F-10;  JDR  1004,  2 
Mar  1968,  F-12. 

Nematolonia  fascicidare  (Fr.)  Karst. 

JDR  945,  27  Dec  1967,  F-7;  JDR  976,  6 
Feb  1968, 1-13. 

Peziza  badia  Merat. 

JDR  480,  28  Apr  1967,  J-9. 

Pleuteits  cervinus  (Schaeff  ex  Seer.)  Kummer 
JDR  1667,  30  Dec  1968,  F-12. 

Ramariopsis  kiinzei  (Fr.)  Donk. 
JDR  1010,  2  Mar  1968,  K-9. 

Schizophylhim  commune  Fr. 
JDR  1047,  31  Mar  1968,  F-10. 

Suillus  pungens  Thiers 

JDR  938,  27  Dec  1967,  L-5;  JDR  939,  27 
Dec  1967,  J-9;  JDR  1657,  30  Dec  1968,  F-10. 

Trernella  mesenterica  Fr. 
JDR  970,  6  Feb  1968,  E-11. 

Clitocybe  inversa  (Fr.)  Quelet 
JDR  971,  6  Feb  1968,  K-14. 

Clitocybe  ntida  (Fr.)  Cooke 

JDR  969,  6  Feb  1968,  F-10;  JDR  1049,  31 
Mar  1968,  G-12. 

Clitocybe  robusta  Peck 

JDR  1661,  30  Dec  1968,  F-12. 

Cortinarius  cinnamonieus  Fr. 
JDR  976,  6  Feb  1968,  F-10. 

Cortinarius  croceofolius  Peck 
JDR  1007,  2  Mar  1968,  J-9. 

Entolonia  serriceus  (Fr.)  Kuhn.  &  Romagn. 
JDR  966,  6  Feb  1968,  F-10. 

Gyroniitra  hicunosa  Afx.  ex  Fr. 
JDR  475,  28  Apr  1967,  J-9. 


Hebeloma  crustiliniforme  Quelet 
JDR  476,  28  Apr  1967,  J-9. 

Hygrophorus  ehurneus  Fr. 

JDR  1672,  30  Dec  1968,  H-12. 

Hygrophorus  flavescens  (Kauffmann)  Smith  & 
Hesler 

JDR  1009,  2  Mar  1968,  K-9. 

Hygrophorus  hypothejus  (Fr.)  Fr. 
JDR  1666,  30  Dec  1968,  E-12. 

Hygrophorus  ponderatus  Britz. 
JDR  479,  28  Apr  1967,  J-9. 

Laccaris  amethystina  (Bolt,  ex  Fr.)  B.  &  Br. 

JDR  940,  27  Dec  1967,  J-9;  JDR  1671,  30 
Dec  1968,  H-12. 

Tricholoma  flavovirens  (Fr.)  Lundell 

JDR  942,  27  Dec  1967,  J-9;  JDR  1644,  30 
Dec  1968,  F-12. 

Tricholoma  virgatum  (Fr.  ex  Fr.)  Kummer 
JDR  1359,  29  Apr  1968,  F-10. 

Volvariella  speciosa  (Fr.)  Sing. 
JDR  1673,  30  Dec  1968,1-11. 

Xylaria  hypoxylon  (Linn.)  DuMortier 
JDR  1673,  30  Dec  1968,  Ml. 

Division  Chlorophyta— Green  Algae 

Bryopsis  corticulans  Setchell 
JDR  1492,  10  Jun  1968,  E-9. 

Cladophora  columbiana  Coll. 
JDR  1475,  10  Jun  1968,  J-3. 

Enteromorpha  intestinalis  (L.)  Link. 
JDR  1466,  10  Jun  1968,  J-4. 

Spongomorpha  coalita  (Ruprecht)  Collins 
JDR  1449,  10  Jun  1968,  L-16. 

Ulva  angusta  Setchell  &  Gardner 
JDR  1471,  10  Jun  1968,  J-3. 

Ulva  linza  L. 
JDR  1488,  10  Jun  1968,  J-3. 

Vha  lobata  (Kutzing)  Setchell  &  Gardner 
JDR  1472,  10  Jun  1968,  J-3. 

Division  Phaeophyta— Brown  Algae 

Cystoseira     osmundacea     (Menzies)     C.A. 
Agardh 

JDR  1531,  14  Jun  1968,  C-13. 

Desmarestia  ligulata  (Lightf.)  Lamour 
JDR  1456,  10  Jun  1968,  N- 17. 


December  1980 


Ripley:  Angel  Island  Plants 


391 


Ectocarpus  acutus  Setchell  &  Gardner 
JDR  1456,  10  Jun  1968,  J-3. 

Egregia  menziesii  (Turn.)  Aresch. 
JDR  1483,  10  Jun  1968,  J-3. 

Fucus  distichus  L.  ssp.  cdcntatus  (De  la  Py- 
laise)  Powell 

JDR  1470,  10  Jun  1968,  J-4. 

Laminaria  sinchiirii  (Harvev)  Farlow 
JDR  1431,  10  Jun  1968,6-17. 

Xcrcocystis  luctkeana  (Mertens)  Ruprecht  & 
Postels 
JDR  1532,  14  Jun  1968,  K- 16. 

Pelvctia  fastigiata  (J.  G.  A^ardli)  De  Toni 
JDR  1477,  10  Jun  1968,^K-4. 

Division  Rhodophyta— Red  Algae 

Ahnfcltia  pliaita  (Hud.son)  Fries. 
JDR  1439a,  10  Jun  1968,  0-17. 

AnfitluininioneUa  glandulifera  (Kvl.)  Woll. 
PR  1431,  10  Jun  1968,0-17.  ' 

CaUithamnion  pikcanum  Harvey 
JDR  1443,  10  Jun  1968,  0-17^ 

CaUoplu/Ilis  obtusifolia  J.  G.  Agardh 
PR  i437,  10  Jun  1968,0-17. 

Ceramium  eatonianum  (Farlow)  De  Toni 
JDR  1430,  10  Jun  1968,  0-17. 

Ceramium  gardneri  Kvlin 

JDR  1451,  10  Jun  1968,0-17. 

Cnjptopleura  violacea  (J.  G.  Agardh)  Kylin 
JDR  1434,  10  Jun  1968,  0-17. 

Cryptosiphonia  woodii  J.  G.  Agardh 
JDR  1438,  10  Jun  1968,  0-17. 

Endockidia  murkata  (Postels  &  Ruprecht)  A. 
G.  Agardli 

JDR  1480,  10  Jun  1968,  L-5. 

GelidUim  coulteri  Harvey 
JDR  1440,  10  Jun  1968,0-17. 

Gigartina  agardhii  Setchell  &  Gardner 
JDR  1468,  10  Jun  1968,  J-3. 

Gigartina  exasperata  Harv.  &  Bail. 
JDR  1478,  10  Jun  1968,  J-3. 

Gigartina  canaliculata  Harvey 
JDR  1447,  10  Jun  1968,  0-i7. 

Gigartina  papillata  (C.  Ag.)  J.  Ag. 
JDR  1479,  10  Jun  1968,  J-4. 


Gracihiriopsis  sjocstcdtii  (K\lin)  Dawson 
JDR  1460,  10  Jun  1968,  k-16. 

Grateloupia  dorypfwra  (Montagne)  Howe 
JDR  1491.  lOJun  1968,  E-8. 

Gi/mnogongru.s  Icptopln/llus  J.  G.  .\gardh 
JDR  1436,  10  Jun  1968,0-17. 

Gymnogongnis  linearis  (Turner)  J.  G.  Agardh 
JDR  1446,  10  Jun  1968,  N- 16. 

Gymnogongnis  plati/pliyllus  Gardner 
JDR  1432,  10  Jun  1968,  O- 17. 

Ihdosaccion  glandiforme  (Gmelin)  Ruprecht 
JDR  1530,  14  Jun  1968,  K-4. 

Leptocladia  conferata  Setchell 
JDR  1464,  10  Jun  1968,  N-17. 

IJthophyllum  decipiens  (Foslie)  Foslie 
JDR  1490,  10  Jun  1968,  J-4. 

Iridaea  heterocarpa  Postels  &  Ruprecht 
JDR  1445,  10  Jun  1968,  0-17. 

Iridaea  eordata  S.  &  G.  var.  splendens  (S.  & 
G.)  Abb. 

PR  1485,  10  Jun  1968.  J-4. 

Melobesia  mediocris  (Foslie)  Setchell  &:  Ma- 
zon 

JDR  1428,  10  Jun  1968,0-17. 

Microeladia  borealis  Ruprecht 
JDR  1444,  10  Jun  1968,  0-17. 

Microckidia  coulteri  Harvey 
JDR  1453.  10  Jun  1968,0-17. 

Odonthalia  florcosa  (Esper)  Falkenberg 
JDR  1433,  10  Jun  1968,0-17. 

Petrocelis  franciscana  Setchell  &  Gardner 
JDR  1487,  10  Jun  1968,0-17. 

Pikea  californica  Harve\' 

JDR  1441,  10  Jun  1968.  O- 17. 

Platyihamnion  pectinatum  Kylin 
JDR  1462,  10  Jun  1968,  J-3. 

Polyneura  latissima  (Harvev)  Kylin 
JDR  1435,  10  Jun  1968,6-17. 

Polysi})honia  hcndn/i 

JDR  1463,  10  Jun  1968,  J-3. 

Porphyra  lanceolata  (Setchell  &  Hus)  G.  .\1. 
Smith 

JDR  1482,  10  Jun  1968,  J-4. 

Pterosiphonia  dcndroidca  (Montagne)  Falken- 
berg 

JDR  1439,  10  Jun  1968.  O- 17. 


392 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


Ptilota  filicina  (Farlow)  J.  G.  Agardh 
JDR  1465,  10  Jun  1968,  0-17. 

Rliodomeki  larix  (Turner)  G.  A.  Agardli 
JDR  1481,  10  Jun  1968,  J-4. 

Scfiizymenia  pacifica  Kylin 
JDR  1457,  10  Jun  1968,  0-17. 

Smithora  naiadum  (Anderson)  Hollenberg 
JDR  1427,  10  Jun  1968,  0-17. 

Stenogramme  interrupta  (C.  A.  Agardli)  Mon- 
tague 
JDR  1429,  10  Jun  1968,  0-17. 

Division  Calamophyta 

Class  Equisetinae 

Equisetaceae 

Equisetum  hyemale  L.  var.  affine  (Engelm.) 
A.  A.  Eat.    PNF 
JDR  703,  25  Jun  1967,  L-16. 

Equisetum    telmateia    Ehrh.    var.    braunii 
Milde.    PNF 

JDR  386,  20  Apr  1967,  L-7;  JDR  625,  7 
Jun  1967,  K-8. 

Division  Pterophyta— Ferns 

Class  Filicinae 

Dryopteridaceae 

Dryopteris  arguta  (Kaulf.)  Maxon.    PNFr 
PR  953,  27  Dec  1967,  G-9. 

Polystichum  mimittim  (Kaulf.)  Presl.    PNFr 

JDR  563,  7  Jun  1967,  1-6;  JDR  982,  6  Feb 
1968,  F-10. 

Polypodiaceae 

Polypodiiim  californicum  Kaulf.    PNFr 

JDR  564,  7  Jun  1967,  E-7;  JDR  981,  6  Feb 
19681,  F-10. 

Pteridaceae 

Adiantum  jordanii  C.  Muell.    PNFr 

JDR  559,  2  May  1967,  G-7;  JDR  1191a,  11 
Apr  1968,  H-13;  JDR  980,  6  Feb  1968,  K-8. 

Pellaea  andromedifolia  (Kaulf.)  Fee.    PNFr 

JDR  1069,  31  Mar  1968,  E-6;  JDR  1176, 
11  Apr  1968,  K-9. 


Pityrogramma    triangularis    (Kaulf.)    Maxon. 
PNFr 

JDR  457,  28  Apr  1967,  F-13;  JDR  511,  2 
May  1967,  C-14;  JDR  587,  7  Jun  1967,  F-11. 

Pteridiiim   oquilimim   (L.)  Kuhn  var.  pub- 
escens  Underw.    PNFr 

JDR  565,  7  Jun  1967,  F-7;  JDR  588,  7  Jun 
1967,  E-12;  JDR  1320,  12  Apr  1968,  1-9. 

Division  Coniferophyta 

Araucariaceae 

Araucaria  bidwillii  Hook.    PCT 
JDR  497,  2  May  1967,  D-13. 

Araucaria   heterophylla   (Salisb.)  Franco. 
PCT 
JDR  499,  2  May  1967,  D-13. 

Cupressaceae 

Chamaecyparis  laivsoniana  (A.  Murr.)  Pari. 
PCT 
JDR  749,  22  Sep  1967,  F-14. 

Cupressus  macrocarpa  Hartw.  ex  Gord.    PIT 
JDR  469,  28  Apr  1967,  K-14. 

Calocedrus  dccurrens  (Torr.)  Florin.    PCT 
JDR  513,  2  May  1967,  H-14. 

Pinaceae 

Abies  concolor  (Gord.  &  Glend.)  Lindl.    PCT 
JDR  486,  2  May  1967,  D-13. 

Cedrus  deodara  (D.  Don)  G.  Don.    PCT 

JDR  485,  2  May  1967,  D-13;  JDR  991,  6 
Feb  1968,  H-15. 

Pinus  monophxjUa  Torr.  &  Frein.    PCT 
Observed  by  B,  Jan  1980,  F-8. 

Pinus  ponderosa  Doiigl.  ex  P.  &  C.  Lawson. 
PCT 

JDR  598,  7  Jun  1967,  G-13. 

Pinus  radiata  D.  Don.    PIT 
JDR  470,  28  Apr  1967,  J-9. 

Pscudotsuga  nienziesii  (Mirbel)  Franco.    PCT 
JDR  1365,  29  Apr  1968,  J-13. 

Taxaceae 

Taxus  baccata  L.    PCT 

JDR  697,  25  Jun  1967,  D13. 

Taxodiaceae 

Cryptonieria  japonica  D.  Don.    PCT 
JDR  516,  2  May  1967,  H-15. 


December  1980 


Ripley:  Angel  Island  Plants 


393 


Seciiioia  senipenirens  (D.  Don)  Endl.    PCT 

JDR  568,  7  Jun  1967,  1-4;  JDR  1251a,  12 
Apr  1968,  K-13. 

Scijuoiadcndnm   gigfintcum   (Lindl.)   Buchh. 
PCT 
JDR  514,  2  May  1967,  J- 16. 

Division  Anthrophyta— Flowering  Plants 

Class  Dicottjh'don cae 

Aceraceae 

Acer  negundo  L.  ssp.  ralifomicuni  (T.  &  C.) 
W'esniael.    PCT 

JDR  491,  2  Mav  1967,  E-13;  JDR  726,  25 
Jun  1967,  K-6. 

Acer  macrophijlhim  Piirsh.    PNT 
JDR  597,  7  Jun  1967,0-12. 

Aizoaceae 

Mescmhrijanthemum  chdense  Mol.    PNF 
JDR  1533,  14  Jun  1968,  J-4. 

Mesemhnjandxeinwn  ediile  L.    PIF 
JDR  660,  20  Jun  1967,  L- 16. 

Mcscinhrijanthcmum  floribiinduni  Haw.    PIF 
JDR  535,  2  May  1967,1-17. 

Tctragonia  tetragonioides  (Pall.)  O.  Kuntze. 
PIF 
JDR  715,  25  Jun  1967,  K-4. 

Anacardiaceae 

Rhus  divcrsdoha  T.  &  C.    PNS 

JDR  410,  20  Apr  1967,  1-7;  JDR  1224,  12 
Apr  1968,  L-6. 

ScJiinus  mollc  L.    PCT 
JDR  515,  2  May  1967,  H- 15. 

Apocynaceae 

Neriutn  oleander  L.    PCS 
JDR  756,  22  Sep  1967,  E-14. 

Vincd  major  L.    PIF 

JDR  441,  20  Apr  1967,  J-5;  JDR  1231,  12 
Apr  1968, 1-4. 

Aristolochiaeeae 

Aristolochia  californica  Torr.    PNF 

K  without  number  or  date;  R  without  num- 
ber, 1877;  JDR  812,  22  Sep  1967,  F-9;  JDR 
1014,  2  Mav  1968,  E-10. 


Basellaceae 

Andrcdera  cordi folia  (Ter.)  Stennis.    PCF 
JDR  717,  22  Sep  1967,1-14. 

Berberidaceae 

Bcrbcris  daruiiiii  Hook.     PCS 
JDR  526,  2  May  1967,  H-15. 

Betulaceae 

Corylus  cornuta   Marsh,  var.  californica   (A. 
DC.)  Sharp.    PNT 

JDR  637,  20  Jun  1967,  K-15;  JDR  670a,  20 
Jmi  1967,  F-12;  JDR  1364,  29  Apr  1968,  H- 
10. 

Boraginaceae 

Amsinckia  intermedia  F.  &  M.    ANF 

JDR  418,  20  Apr  1967.  1-5:  JDR  1071.  31 
Mar  1968,  F-7. 

Cynoglosstim  grandc  Dougl.  ex  Lehm.    PNF 

JDR  560,  2  Mav  1967,  E-7;  JDR  1013.  2 
Mar  1968,  F-11. 

Echiiim  fastuosum  Jacq.    PIS 

JDR  385,  20  Apr  1967,  J-4:  JDR  1180.  11 
Apr  1968,  C- 11. 

Myosotis  latifolia  Poir.    PIF 
JDR  524,  2  May  1967,  E-14. 

Callitrichaceae 

Callitrichc  marginata  Torr.    PNF 
JDR  632,  20  Jun  1967,  J-9. 

Caprifoliaceae 

Loniccra   hispidula    Dougl.    var.   lacillans 
Crav.    PNS 

JDR  561,  7  Jun  1967,  F-6. 

Lonicera  japonica  Thunb.    PIS 
JDR  709,  25  Jun  1967,  D-14. 

Samhucus  coerulca  Raf.    PNS 
JDR  569,  7  Jun  1967, 1-6. 

Symphoricarpos  rivularis  Suksdort.    P.\S 
JDR  763,  22  Sep  1967,  E-Il. 

Symphoricarpos  mollis  Nutt.     PNS 
R.  6c  J.  21177,2  May  1967,  J-12. 

Car\'ophyIlaceae 

Arcnaria  douglasii  Fen/.l  ex  T.  6i  C.    ANF" 

JDR  401,  20  Apr  1967,  Ml  6:  JDH  595.  7 
Jim  1967,  E-12. 


394 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


Cerastium  ghmeratum  Thuill.    AIF 

JDR  1103,  11  Apr  1968,  F-7;  JDR  1271,  12 
Apr  1968,  E-9. 

Pohjcarpon  tetraphiiUum  (L.)  L.    AIF 
JDR  1409,  10  Jun  1968, 1-16. 

Sagina  apetala  Ard.  var.  harbata  Fenzl.    AIF 
R.  &  J.  21158,  2  May  1967,  K-8. 

Silene  califomica  Durand.    PNF 
JDR  1389,  8  Jun  1968,  C-11. 

Silene  gallica  L.    AIF 

JDR  423,  20  Apr  1967,  1-4;  JDR  1149,  11 
Apr  1968,  J-14;  JDR  1378,  29  Apr  1968,  K- 
15. 

Spergularia  bocconii  (Scheele)  Foucaud.    AIF 
JDR  1358,  29  Apr  1968,  J-14. 

Spergularia   macrotheca   (Hornem.)   Heynh. 
PNF 

JDR  390,  20  Apr  1967,  M-15;  JDR  1311, 
12  Apr  1968,  K-9. 

Spergularia  rubra  (L.)  J.  &  C.  Presl.    PIF 
JDR  647,  20  Jun  1967,  H-13. 

Spergularia  villosa  (Pers.)  Canib.    PIF 

JDR  531,  2  May  1967,  H-15;  JDR  1108,  11 
Apr  1968,  F-7;  JDR,  1051,  31  Mar  1968,  G-8; 
JDR  1221,  12  Apr  1968,  K-6. 

Stellaria  media  (L.)  Vill.    AIF 

JDR  996,  6  Feb  1968,  E-14;  JDR  1017,  2 
May  1968,  F-10. 

Casuarinaceae 

Casuarina  striata  Dry.    PCT 
JDR  487,  2  May  1967,  D-14. 

Celastraceae 

Maytenus  boaria  Mol.    PCT 
JDR  1407,  10  Jun  1968,  G-14. 

Chenopodiaceae 

Atriplex  leucophylla  (Moq.)  Dietr.    PNF 
Observed  by  H  on  16  May  1946. 

Chenopodium  album  L.    AIF 
JDR  667,  20  Jun  1967,  L-6. 

Chenopodium   ambrosioides    L.    var.    vagans 
(Standi.)  J.  T.  Howell.    PIF 

Observed  by  H  on  16  May  1946. 

Chenopodium    californicum    (Wats.)    Wats. 
PNF 

JDR  1411,  10  Jun  1968,  0-16. 


Chenopodium  multifidum  L.    AIF 
JDR  1395,  8  Jun  1968,  N-16. 

Salicornia  virginica  L.    PNF 
JDR  12.32,  12  Apr  1968,  L-5. 

Cistaceae 

Helianthemum  scoparium  Nutt.  var.  vulgare 
Jeps.    PNF 

JDR  1054,  31  Mar  1968,  G-12. 

Compositae 

Achillea  borealis  Bong.  ssp.  califomica  (Pol- 
lard) Keck.    PNF 

JDR  408,  20  Apr  1967,  1-5;  JDR  1115,  11 
Apr  1968,  J-5. 

Achijrachaena  mollis  Schauer.    ANF 
Observed  by  H  on  16  May  1946. 

Agoseris  apargioides  (Less.)  Greene.    PNF 
JDR  1297,  12  Apr  1968,  H-8. 

Agoseris  grandiflora  (Nutt.)  Greene.    PNF 
JDR  580a,  7  Jun  1967,  E-11. 

Ambrosia  chamissonis  (Less.)  Greene  var.  bi- 
pinnatisecta  J.  T.  Howell.    PNF 

JDR  664,  20  Jun  1967,  K-5;  JDR  720,  25 
Jun  1967,  N-15. 

Anthemis  cotula  L.    AIF 

JDR  1706,  16  Jun  1969,  E-12. 

Artemisia  califomica  Less,  in  Hook.    PNS 

JDR  725,  25  Jun  1967,  F-13;  JDR  1505,  14 
Jun  1968,  L-5. 

Artemisia  douglasiana  Bess.    PNF 
JDR  713,  25  Jun  1967,  J-14. 

Artemisia  pijcnocephala  DC.    PNF 
JDR  950,  27  Dec  1967,  1-9. 

Aster  chilensis  Nees.    PNF 
JDR  814,  22  Sep  1967,  J-9. 

Baccharis  pilularis  DC.    PNS 
JDR  822,  22  Sep  1967,  J-9. 

Baccharis   pilularis    DC.    var.    consanguinea 
(DC.)  C.  B.  Wolf.    PNS 
JDR  708,  25  Jun  1967,  H-7. 

Bcllis  perennis  L.    PNF 
JDR  558,  2  May  1967,  F-8. 

Carduus  pycnocephalus  L.    AIF 
JDR  1373,  29  Apr  1968,  J-12. 

Carduus  tenuiflorus  Curt.    AIF 

JDR  521,  2  Mav  1967,  L-16;  JDR  438,  20 
Apr  1967,  J-8. 


December  1980 


Ripley:  Angel  Island  Plants 


395 


Centaurea  meUtensis  L.    AIF 

JDR  656,  20  Jun  1967,  J- 15;  JDR  694,  25 
fun  1967,  D-14;  JDR  1525,  14  Jun  1968,  K-9. 

Cirsiiun  hrcvisti/lmu  Cronq.     PNF 
JDR  623,  7  Jun  1967,  K-9. 

Cirsium  occidentale  (Nutt.)  Jeps.    P\F 
JDR  696,  25  Jun  1967,  D-13. 

Cirsiimi  piotcanuni  J.  T.  Howell.    P\F 
R.  &  J.  21304,  2  May  1967,  D-10. 

Cirsium  qucrcctorum  (Grav)  Jeps.    P\F 
JDR  616,  7  Jun  1967,  K-6. 

Cirsium  rcmotifolium  (Hook.)  DC.    P\F 
H  21878,  16  May  1946,  without  location. 

Cirsium  vulgare  (Savi)  Ten.    BIF 
JDR  454,  28  Apr  1967,  K-8. 

Conijza  bonariensis  (L.)  Cronq.    AIF 
Observed  by  H,  16  May  1946. 

Comjza  canadensis  (L.)  Cronq.    AIF 
Observed  by  H,  16  May  1946. 

Cotula  australis  (Less.)  Hook.    AIF 

JDR  435,  20  Apr  1967,  1-4;  JDR  1074,  1 1 
Apr  1968,  F-8;  JDR  1097,  11  Apr  1968,  F-9. 

Cotula  coronopifulia  L.    PIF 
JDR  432,  20  Apr  1967, 1-4. 

Ci/nara  scoh/mus  L.    PCS 
JDR  807,22  Sep  1967,  F-8. 

Erechtites  arguta  (A.  Rich.)  DC.    AIF 
JDR  595a,  7  Jim  1967,  G-13. 

Erechtites  prenanthoides  (A.  Rich.)  DC.    AIF 
JDR  737,  22  Sep  1967,  E-I4. 

Erigeron  glaucus  Ker.    PNF 
JDR629,  7  Jun  1967,  N- 17. 

EriophiiUum   confertiflorum   (DC.)  Grav. 
PiNS 

PR  589,  7  Jun  1967,  J- 14. 

EriopJu/lhim    staechadifolium    Lag.    var.    ar- 
temisiifolium  (Less.)  Macbr.    PNS 
JDR  504,  2  May  1967,  J-6. 

Evax  sparsiflora  (Grav)  Jeps.    ANF 
JDR  1353,  12  Apr  1968,  K-9. 

Fikigo  gallica  (L.)  L.    AIF 

JDR  643,  20  Jun  1967,  K-16;  JDR  692,  25 
Jun  1967,  D-14. 

Gnaphalium  califurnicum  D(>.    BNF 

JDR  574,  7  Jun  1967,  H-6;  JDR  1194,  12 
Apr  1968,  E-11. 


Gnaphalium  chilcnsc  Spreng.    B\F 

JDR  673,  20  Jun  1967,  K-15;  JDR  1191,  11 
Apr  1968, 1-I4;  JDR  1399,  8  Jun  1968,  H-16. 

Gnaphalium  luteo-alhum  L.    AIF 

JDR  1192,  11  Apr  1968,  1-14;  JDR  1094, 
12  Apr  1968,  G-7. 

Gnaphalium  purpureum  L.    BNF 
JDR  1192a.  11  Apr  1968,1-14. 

Grindclia    campurum    Cireene    var.    Daii/i 
(Jeps.)  Steverm.    PNF 

JDR588a,  7  Jun  1967,  F- 13. 

Grindelia  hirsutula  H.  (be  A.  var.  hrcvisquama 
Steyerm.    PNF 

JDR  1337,  12  Apr  1968,  1-9. 

Gutierrczia  californica  (DC.)  T.  &  G.    PNS 

E  without  number,  3  Apr  1925;  OS  2681, 
Oct  1957;  JDR  636,  20  Jun  1967,  1-6;  JDR 
716,  25  Jun  1967,  J-7. 

Haplopappus  ericoides  (Less.)  Hook.  6c  .Arn. 
PNS 

JDR  721,  25  Jun  1967,  M-10. 

Heterotheca  bolanderi  Grav.    PNF 

JDR  669,  20  Jun  1967, 'K-15;  JDR  699,  25 
Jun  1967,  E-12. 

Hijpochoeris  glabra  L.    PIF 

JDR  380,  20  Apr  1967,  M-15;  JDR  1233, 
12  Apr  1968,  L-5;  JDR  1269,  12  Apr  1968,  E- 
9;  JDR  1281,  12  Apr  1968,  K-9. 

Hijpochoeris  radicata  L.    PIF 

JDR  809,  22  Sep  1967,  F-7;  JDR  580,  7  Jun 
1967,  F-11. 

Iva  axillaris  Pursh.    PNF 

JDR  661,  20  Jun  1967,  L-15;  JDR  765,  22 
Sep  1967,  K-9. 

iMctuca  saligna  L.    AIF 
JDR  813,  22  Sep  1967,  F-7. 

Lasthenia   chrysostoma   (F.   &   W.)  Greene. 
ANF 

JDR  1295,  12  Apr  1968.  K-8. 

Madia  gracilis  (Smith)  Keck.    PNF 
JDR  553,  2  .May  1967,  K-13. 

Madia  sativa  .Mol.    PNF 
JDR  621,  7  Jun  1967,  K-8. 

Matricaria  matricarioidcs  (Less.)  Porter.     \\Y 
JDR  435a,  20  April  1967,  E-13;  JDK  1067, 
11  Apr  1968,  G-10;  IDH  1.357.  29  Apr  1968, 
1-8. 


396 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


Micropus  califomiciis  F.  &  M.    ANF 

JDR  1225,  12  Apr  1968,  J-5;  JDR  1344,  12 
Apr  1968,  K-9. 

Microseris  higelovii  (Gray)  Sch.-Bip.    ANF 
H  21859,  16  May  1946. 

Microseris  douglasii  (DC.)  Sch.-Bip.    ANF 
JDR  1347,  12  Apr  1968,  K-9. 

Microseris  heierocarpa  (Niitt.)  Chamb.    ANF 
JDR  1349,  12  Apr  1968,  K-9. 

Microseris  linearifolia  (Nutt.)  Sch.-Bip.    ANF 
JDR  1310,  12  Apr  1968,  K-9. 

Pentachaeta  alsinoides  Greene.    ANF 
H  21865,  16  May  1946. 

Picris  echioides  L.    BIF 

R.  &  J.  21032,  2  May  1967,  K-9. 

Psilocarphus  teneUiis  Nutt.    ANF 
Observed  by  H,  16  May  1946. 

Rafinesqiiia  californica  Nutt.    ANF 
Observed  by  H,  16  May  1946. 

Senecio  aronicoides  DC.    PNF 

E  without  number,  3  Apr  1925;  JDR  466, 
28  Apr  1967,  E-11;  JDR  1030,  2  Mar  1968, 
G-13. 

Senecio  rnikanioides  Otto.    PIF 
JDR  756a,  22  Sep  1968,  K-16. 

Senecio  sylvaticiis  L.    AIF 
JDR707,  25Jun  1967,  L- 15. 

Senecio  vulgaris  L.    AIF 

JDR  433,  20  Apr  1967,  1-4;  JDR  1336,  12 
Apr  1968, 1-8. 

Silybum  marianum  (L.)  Gaertn.    BIF 

JDR  581a,  7  Jun  1967,  F-11;  JDR  623,  7 
Jun  1967,  L-16;  JDR  949,  27  Dec  1967,  F-8. 

Solidago  californica  Nutt.    PNF 
JDR  752,  22  Sep  1967,  L-15. 

SoHva  daucifolia  Nutt.    ANF 
JDR  663,  20  Jun  1967,  L-7. 

Soliva  pterospertna  (Juss.)  Less.    ANF 

JDR  1096,  11  Apr  1968,  J-5;  JDR  1304,  12 
Apr  1968,  K-8. 

Soliva  sessilis  R.  &  P.    ANF 
D  80-39,  29  May  1980,  F-8. 

Sonchus  asper  (L.)  Hill.    AIF 

JDR  1.388,  29  Apr  1968,  1-6;  JDR  1414,  10 
Jun  1968,  N-17. 

Stephanonieria  virgata  Benth.    ANF 
Observed  by  H,  16  May  1946. 


Stylocline  amphihola   (Gray)  J.  T.   Howell. 
ANF 

R.  &  J.  21328,  2  May  1967,  K-16. 

Taraxacum    officinale    Weber    in    Wiggers. 
PIF 

JDR  508,  2  Mav  1967,  D-14;  JDR  1092,  11 
Apr  1968,  J-4. 

Tragopogon  porrifolius  L.    BIF 
JDR  1423,  10  Jun  1968,  H-16. 

Wyethia  angustifolia  Nutt.    PNF 

JDR  594,  7  Jun  1967,  H-7;  JDR  1287,  12 
Apr  1968,  K-8. 

Xanthium  spinosum  L.    AIF 
JDR  755,  22  Sep  1968,  K-16. 

Convolvulaceae 

Calystegia  polymorpha  (Greene)  Munz.    PNF 
Observed  by  H,  16  May  1946. 

Calystegia    purpurata    (Greene)    Bnnnmitt. 
PNF 

JDR  450,  28  Apr  1967,  G-13;  JDR  624,  7 
Jun  1967,  K-7;  JDR  1117,  11  Apr  1968,  J-9. 

Calystegia  suhacaulis  H.  &  A.    PNF 
JDR  1305,  12  Apr  1968,  K-8. 

Convolvulus  arvensis  L.    PIF 
Observed  by  H,  16  May  1946. 

Dichondra  donneUiana  Tharp  &  Johnston. 
PNF 

JDR  1103a,  11  Apr  1968,  F-7;  JDR  1187a, 
11  Apr  1968,1-14. 

Crassulaceae 

Dudleya    farinosa    (Lindl.)    Britt.    &    Rose. 
PNF 

R  without  number,  26  Mav  1877;  JDR  570, 
7  Jun  1967,  J-4. 

Sedum  spathulifolium  Hook.    PNF 
JDR  505,  2  May  1967,  J-3. 

Tillaea  erecta  H.  &  A.    ANF 

JDR  1037,  2  Mar  1968,  K-8;  JDR  1236,  12 
Apr  1968,  K-5. 

Cruciferae 

Arabis  glabra  (L.)  Bernh.    BNF 
JDR  1302,  12  Apr  1968.  K-8. 

Brassica  campestris  L.    AIF 
JDR  1333,  12  Apr  1968,  G-7. 

Brassica  geniculata  (Desf.)  J.  Ball.    BIF 

JDR  626,  7  Jun  1967,  1-7;  JDR  631,  20  Jun 
1967,  E-12. 


December  1980 


Ripley:  Angel  Island  Plants 


39- 


Cakile  maritima  Scop.    AIF 

JDR  431,  20  Apr  1967,  K-5;  JDR  634,  20 
Jun  1967,  L-16. 

CapscUa  hursa-pastoris  (L.)  Medic.    AIF 

JDR  1129,  11  Apr  1968,  F-8;  JDR  1173,  11 
Apr  1968,  G-7. 

Cardamine  oligospenna  Nutt.    BNF 

JDR  1012,  2  Mar  1968,  E-10;  JDR  1120, 
11  Apr  1968,  E-6. 

Coronopus  didymus  (L.)  Smith.    BIF 
Observed  by  H,  16  May  1946. 

Dentaria    integrifolia    Nutt.    var.    californica 
(Nutt.)Jeps.    PNF 

JDR  414,  20  Apr  1967,  J-4;  JDR  979,  6  Feb 
1968,  E-10. 

Enjshmim  capitatiun  (Dougl.)  Greene.    BNF 
JDR  502,  2  May  1967,  D- 14. 

Lepidium  nitidum  Nutt.    ANF 

JDR  1036,  2  May  1968,  J-7;  JDR  1518,  14 
Jun  1968,  K-9. 

Lepidium  strictum  (Wats.)  Rattan.    AIF 
D  80-44,  29  May  1980,  F-9. 

Lobidaria  maritima  (L.)  Desv.    PIF 
JDR  409,  20  Apr  1967,  J-5. 

Raplianus  sativus  L.    BIF 
JDR  510,  2  May  1967,  D-14. 

Rorippa  nasturtium-aquaticum  (L.)  Britt.  & 
Rendl.    PIF 

JDR  735,  22  Sep  1967,  L-9;  JDR  1415,  10 
Jun  1968,  N-16. 

Thehjpodium     lasiophyllum     (H.     &     A.) 
Greene.    ANF 

JDR  1503,  14  Jun  1968,  K-9;  JDR  1529,  14 
Jun  1968,  E-6. 

Thysanocarpus  curvipes  Hook.    ANF 
JDR  1207,  12  Apr  196,  E-6. 

Cucurbitaceae 

Marah  fabaceus  (Naud.)  Dunn.    PNF 
JDR  389,  20  Apr  1967,  L-6. 

Marah  oregonus  (T.  &  G.)  Howell.    PNF 
JDR  1330,  12  Apr  1968,  1-9. 

Ericaceae 

Arbutus  rtienziesii  Pursh.    PNT 

JDR  405,  20  Apr  1967,  D-12;  JDR  818,  22 
Sep  1967,  E-11. 


Arctostaphylo.s  glandulosa  Eastw.  var.  Cush- 
ingiana  (Eastw.)  .Adams  ex  McMinn.    PNS 
JDR  978,  6  Feb  1968,  E-11. 

Euphorbiaceae 

Euphorbia  pephis  L.    AIF 
JDR  731,  22  Sep  1967,  H-15. 

Euphorbia  spathulata  Lamk.    ANF 

JDR  1351,  12  Apr  1968,  K-9;  JDR  1.384,  29 
Apr  1968,  K-15;  JDR  1417.  10  Jun  1968,  \- 
17. 

Fagaceae 

Quercus  agrifolia  Nee.    PNT 
JDR  423,  20  Apr  1967, 1-4. 

Quercus  wislizeni  A.  DC.  var.  frutescens 
Engelm.    PNT 

JDR  977,  6  Feb  1968,  D- 11. 

Quercus  suber  L.    PCT 

Observed  by  B,  Jan  1980,  K-14  and  L-8. 

Garryaceae 

Garrya  eUiptica  Dougl.    PNS 
JE)R  566a,  7  Jun  1967,  E-7. 

Gentianaceae 

Centauriitm     muhlcnbergii     (Griseb.)     W. 
Wight.    ANF 

H  21857,  16  May  1946. 

Geraniaceae 

Erodium  botrys  (Gav.)  Bertol.    AIF 
JDR  1516,  14  Jim  1968,  K-9. 

Erodium  cicutarium  (L.)  L'Her.    .\IF 

JDR  .396,  20  Apr  1967,  N-16;  JDR  811,  22 
Sep  1967,  G-9;  JDR  984,  6  Feb  1968,  E-11. 

Erodium  moschatum  (L.)  L  Her.    .\IF 

JDR  1170,  11  Apr  1968,  G-7;  JDR  1413,  10 
Jun  1968,  N-16. 

Geranium  carolinianum  L.    ANF 
JDR  1106a,  11  Apr  1968,  G-7. 

Geranium  dissectum  L.    .-MF 
JDR  417,  20  Apr  1967,  1-5. 

Geranium  molle  L.    BIF 

JDR  520,  2  May  1967,  H-16;  JDR  537,  2 
May  1967,  C-11. 

Pelargonium  zonale  (L.)  L'Her.    PGS 
JDR  .530.  2  May  1967,1-16. 


398 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


Hippocastanaceae 

Aesculus  californica  (Spach)  Nutt.    PNT 
JDR602,  7Jiin  1967,  H- 14. 

Hydrophyllaceae 

Eucnjpta  chrifsanthetnifolia  (Benth.) 
Greene.    ANF 

TB  without  number  or  date. 

Nemophila  heterophyUa  F.  &  M.    ANF 
JDR  1217,  12  Apr  1968,  F-6. 

Phacelia  californica  Cham.    PNF 

JDR  437,  20  Apr  1967,  J-6;  JDR  1158,  11 
Apr  1968,  F-12. 

Phacelia  malvifolia  Cham.    ANF 
JDR  1393,  29  Apr  1968,  C-11. 

Phacelia  nenioralis  Greene.    BNF 
H  21892,  16  May  1946. 

Pholistoma  aiiritum  (Lindl.)  Lilja.    ANF 
Observed  by  H,  16  May  1946. 

Hyperieaceae 

Hypericum  caneriensis  L.    PCS 
JDR  1408,  10  Jun  1968,  K-15. 

Juglandaceae 

Juglans  hindsii  (Jeps.)  Jeps.    PCX 
JDR  730,  25  Jun  1967,  K-15. 

Labiatae 

Lepechinia  cahjcina  (Benth.)  Epl.    PNS 

JDR  472,  28  Apr  1967,  H-8;  JDR  583,  7 
Jun  1967,  K-7. 

Mentha  pulegium  L.    PIF 
Observed  by  B,  1980,  E- 11. 

Monardella   villosa    Benth.    var.    franciscana 
(Ehner)Jeps.    PNF 
JDR  615,  7  Jun  1967,  K-7. 

Rosmarinus  officinalis  L.    PCS 
JDR  1140,  11  Apr  1968,  F-8. 

Salvia  leucantha  Cav.    PCS 
JDR  962,  27  Dec  1967,  J-6. 

Stachys  rigida  Nutt.  var.  quercetorum  (Hel.) 
Epl.    ANF 

JDR  416,  20  Apr  1967,  J-3;  JDR  545,  2 
May  1967,  D-12;  JDR  1109,  11  Apr  1968,  G- 

7. 


Lauraceae 

Umbellularia    californica    (H.    &   A.)    Nutt. 
PNT 

JDR  413,  20  Apr  1967,  1-4;  JDR  1019,  2 
May  1968,  F-10. 

Leguminosae 

Acacia  deciirrens  (Wendl.)  Willd.    PIT 

JDR  494,  2  May  1967,  E-13;  JDR  1043,  2 
May  1968,  F-7. 

Acacia  longifolia  Willd.    PIT 
Observed  iDy  B,  1980,  H-15. 

Albizia  lophatha  (Willd.)  Benth.    PIT 
JDR  960,  6  Feb  1968,  L-6. 

Acacia  melanoxylon  R.  Br.    PIT 
JDR  442,  20  Apr  1967,  K-6. 

Acacia  verticillata  (L'Her.)  Willd.    PCT 
JDR  523,  2  May  1967,  E-14. 

Astragalus  gambellianus  Sheld.    ANF 

JDR  1511,  14  Jun  1968,  K-9;  JDR  1315,  12 
Apr  1968,  M-16. 

Astragalus  nuttalli  (T.  &  G.)  J.  T.  Howell  var. 
virgatus  (Gray)  Barneby.    PNF 

S  without  number,  May  1920;  H  21869,  16 
May  1946. 

Cytisus  monspessulanus  L.    PIS 
JDR  961,  27  Dec  1967,  K-14. 

Cytisus  multiflorus  (L'Her.)  Sweet.    PCS 
JDR  1142,  11  Apr  1968,1-15. 

Cytisus  scoparius  (L.)  Link.    PIS 

JDR  542,  2  May  1967,  C-10;  JDR  1214,  12 
Apr  1968,  J-7;  JDR  1334,  12  Apr  1968,  MO. 

Cytisus  scoparius   (L.)   Link.   var.  andrianus 
(Puiss.)  Dippel.    PIS 

D  80-42,  29  May  1980,  F-7. 

Lathyrus  latifolius  L.    PIF 
JDR  1392,  8  Jun  1968,  L-15. 

Lathyrus  vestitus  Nutt.  ex  Torr.  &  Grav  ssp. 
Bolanderi  (Wats.)  C.  L.  Hitchc.    PNF 

JDR  612,  7  Jun  1967,  F-12;  JDR  1059,  31 
Mar  1968,  D-15;  JDR  1133,  11  Apr  1968,  E- 
11. 

Lotus  scoparius  (Nutt.)  Ottlev.    PNF 
JDR  578,  7  Jun  1967,  E-12. 

Lotus  strigosus  (Nutt.)  Greene.    ANF 
H  21869,  16  May  1946. 


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399 


Lotus  siihpinnatus  Lag.    ANF 

JDR  556,  2  Mav  1967,  K-13;  JDR  1223,  12 
Apr  1968,  L-6;  JDR  1299,  12  Apr  1968,  K-9; 
JDR  1340,  12  Apr  1968,  G-12. 

Lupinus    albifwns    Benth.    var.    collinu.'i 
Greene.    PNF" 

H  21873,  16  May  1946. 

Lupinus  arborcus  Sims.    PNS 

JDR  603,  7  Jmi  1967,  E-12;  JDR  536,  2 
May  1967,  C-11. 

Lupinus    hicolor    Lindl.    var.    umhcllatus 
(Greene)  C.  P.  Sm.    ANF 

JDR  379,  20  Apr  1967,  N-15;  JDR  1370,  29 
Apr  1968,  J- 11. 

Lupinus  densiflorus  Benth.    ANF 
H  21863,  16  May  1946. 

Lupinus  fonnosus  Greene.    PNF 
JDR  459,  28  Apr  1967, 1-9. 

Lupinus  latif alius  Agardh.    PNF 
JDR  575,  7  Jun  1967,  J-8. 

Lupinus  poll/corpus  Greene.    ANF 
JDR  1510",  14  Jun  1968.  K-9. 

Lupinus  rivularis  Dougl.    PNF 
JDR  604,  7  Jun  1967, 1-7. 

Lupinus  succulentus  Dougl.    ANF 
Observed  by  H,  16  May  1946. 

Medicago  arahica  (L.)  All.    AIF 

JDR  393,  20  Apr  1967,  M-16;  JDR  1376, 
29  Apr  1968,  H-14. 

Medicago  pohjmorpha  L.    AIF 

JDR  1179,  11  Apr  1968,  F-15;  JDR  1335, 
12  Apr  1968,  I-IO;  JDR  1521,  14  Jun  1968. 
K-9. 

Medicago  poli/inorpha   L.   var.   l)revispina 
(Benth.)  Hevn."  AIF 

JDR  1175,  11  Apr  1968,  G-7;  JDR  1247,  12 
Apr  1968,  E-10;  JDR  1332,  12  Apr  1968,  1-8. 

Melilotus  indicus  (L.)  All.    BIF 

JDR  704,  25  Jun  1967,  K-14;  JDR  1401,  10 
Jun  1968,  J-16;  JDR  1416,  10  Jun  1968,  M-16. 

Rohinid  pseudo-acacia  L.    PCT 
JDR  567,  7  Jun  1967,  1-6. 

Sophora  micropIu/Ua  Aiton.     P(>T 
JDR  496,  2  May  1967,  D- 14. 

Thennopsis  macrophijUa  H.  &  A.    PNF 
JDR  1.387,  29  Apr"  1968,  1-6. 

Trifoliwn  amplectens  T.  &  G.    ANF 
JDR  1296,  12  Apr  1968,  K-9. 


TrifoHum  alhopurpurcum  T.  &  G.     .\NF 
JDR  1293,  12  Apr  1968,  K-9. 

Trifoliutn  hifidum  Grav.    .\NF 
JDR  1151,  11  Apr  1968,  H-13. 

Trifoliuni  duhiuin  Sibth.     .\IF 

JDR  1080,  11  Apr  1968,  G-13;  JDR  1377, 
29  Apr  1968,  J-15;  JDR  1418,  10  Jun  1968, 
N-16. 

Trifoliuni  fucatuni  Lindl.    .\.\F 
Observed  by  H,  16  May  1946. 

TrifoHum  gracilentum  T.  &  G.    .\NF 

JDR  1135,  11  Apr  1968,  F-11;  JDR  1200, 
12  Apr  1968,  1-14;  JDR  1354,  12  Apr  1968, 
K-9. 

Trifoliuni  macraci  H.  &  .\.    ANF 

JDR  461,  28  Apr  1967,  KIO;  JDR  1316,  12 
Apr  1968,  K-8. 

Trifoliuni  niicroccphc.hnn  Pursh.    .ANF 

JDR  1216,  12  Apr  1968,  J-7;  JDR  1265,  12 
Apr  1968,  J-5;  JDR  1276.  12  Apr  1968.  K-9; 
JDR  1339,  12  Apr  1968,  1-9. 

TrifoHum  microdon  H.  &  A.    ANF 

JDR  552,  2  May  1967,  K-14;  JDK  1121.  II 
Apr  1968,  F-12. 

TrifoHum  repcns  L.    PIF 
JDR  671,  20  Jun  1967,  K-14. 

TrifoHum  tridentatum  Lindl.    .\NF 

JDR  423a,  20  Apr  1967,  J-5;  JDR  1 166.  11 
Apr  1968,  1-9;  JDR  1254,  12  Apr  1968,  D-IO. 

TrifoHum  variegatum  Nutt.    .\NF 

JDR  391,  20  Apr  1967.  M-15:  JDR  555.  2 
May  1967,  C-11. 

Vicia  americana   Muhl.  var.  orcgana  (Nutt.) 
Nels.    PNF 

R.  &  J.  21333,  2  May  1967,  E-12. 

Vicia  angustifolia  L.    .\IF 
JDR  1215.  12  Apr  1968.  J-7. 

Vicia  cxigua  Nutt.    ANF 

JDR  11.34,  11  Apr  1968,  F-13;  JDR  1250, 
12  Apr  1968,  D-10. 

V'irjV;  sativa  L.    .\IF 
JDR  546,  2  May  1967.  C-11. 

Lvthraceae 

Lijtlirum  hyssopifoHa  L.    PNF 

JDR  768,  22  Sep  1967,  H-9;  JDR  1523,  14 
Jun  1968,  K-9. 


400 


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Vol.  40,  No.  4 


Magnoliaceae 

Magnolia  grandi flora  L.    PCT 
JDR  805,  22  Sep  1967,  J-6. 

Malvaceae 

Lavatera  cretica  L.    AIF 

H  21899,  16  May  1946,  H-16. 

Malva  parviflora  L.    AIF 
JDR  668,  20  Jun  1967, 1-6. 

Sidalcea  malviflora  (DC.)  Gray.    PNF 

JDR  421,  20  Apr  1967,  1-4;  JDR  1270,  12 
Apr  1968,  C-12. 

Moraceae 

Ficus  carica  L.    PCT 

PR  742,  22  Sep  1967,  D-14. 

Myoporaceae 

Mijoporwn  laetum  Forst.    PCS 
Observed  by  B,  1980,  E-9. 

Myricaceae 

Myrica  califomica  C.  &  S.    PNT 
JDR  1679,  16  May  1969,  G-15. 

Myrtaceae 

Eucalyptus  amygdalina  Labill.    PCT 

JDR  488,  2  May  1967,  D-13;  JDR  702,  25 
Jun  1967, 1-14. 

Eucalyptus  cornuta  Labill.    PCT 
JDR  754,  22  Sep  1967,  K-13. 

Eucalyptus  eugenioides  Sieb.    PCT 
JDR  638,  20  Jun  1967,1-14. 

Eucalyptus  ficifolia  F.  J.  Muell.    PCT 
JDR  744,  22  Sep  1967,  D-13. 

Eucalyptus  globulus  Labill.    PIT 
JDR  610,  7  Jun  1967,  1-15. 

Eucalyptus  goniocalyx  F.  J.  Muell.    PCT 
JDR  525,  2  May  1967,  D-13. 

Eucalyptus  leucoxylon  F.  J.  Muell.    PCT 
JDR  819,  22  Sep  1967, 1-15. 

Eucalyptus  polyanthemos  Schau.    PCT 
JDR  823,  22  Sep  1967, 1-14. 

Eucalyptus  pulverulenta  Sims.    PCT 
JDR  714,  25  Jun  1967,1-16. 

Melaleuca  nesophila  F.  J.  Muell.    PCS 
JDR  748,  22  Sep  1967,  D-14. 

Syzygium  paniculatum  Gaertner.    PCT 
JDR  729,  25  Jun  1967,  G-15. 


Nyctaginaceae 

Ahronia  htifolia  Esch.    PNF 
JDR  534,  2  May  1967,  J- 16. 

Abronia  umbellata  Lamk.    PNF 
JDR  533,  2  May  1967,  J-16. 

Bougainvillea  spectabilis  Willd.    PCS 
JDR  723,  25  Jun  1967,  1-15. 

Oleaceae 

Ligustrum  ovalifolium  Hassk.    PCS 
JDR  609,  7  Jun  1967,1-16. 

Syringa  vulgaris  L.    PCS 
JDR  495,  2  May  1967,  E-14. 

Onagraceae 

Clarkia    amoena    (Lehm.)    Nels.    &    Macbr. 
ANF 

JDR605,  7  Jun  1967,  G- 14. 

Clarkia  concinna  (F.  &  M.)  Greene.    ANF 
JDR586,  7  Jun  1967,  E-11. 

Clarkia  unguiculata  Lindl.    ANF 
JDR  760,  22  Sep  1967,  J-15. 

Epilobium  adenocaulon  Hausskn.  var.  Parishii 
(Trel.)  Munz.    PNF 

JDR  1524,  14  Jun  1968,  K-9. 

Epilobium  brachycarpum  Presl.    ANF 
R.  &  J.  21064,  2  May  1967,  K-9. 

Epilobium  minutum  Lindl.    ANF 
JDR  1356,  12  Apr  1968,  K-9. 

Epilobium    watsonii    Barbey    var.  francisca- 
num  (Barbey)  Jeps.    PNF 

Observed  by  H,  16  May  1946. 

Fuchsia  magellanica  Lam.    PCS 
JDR  600,  20  Jun  1967,  H-15. 

Oenothera  ovata  Nutt.    PNF 
JDR  458,  28  Apr  1967,  J-9. 

Zauschneria  califomica  Presl.    PNF 

JDR  718,  22  Sep  1967,  J-6;  JDR  946,  27 
Dec  1967,  K-14. 

Orobanchaceae 

Orobanche  fasciculata  Nutt.  var.  franciscana 
Achey.    ANF 

JDR  1703,  20  Apr  1969,  K-8. 

Oxalidaceae 

Oxalis  pes-caprae  L.    PIF 

JDR  403,  20  Apr  1967,  J-16;  JDR  1169,  11 
Apr  1968,  F-8. 


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401 


Oxalis  pilosa  Nutt.    PNF 
JDR995,  6Feb  1968,  D-13. 

Oxalis  rubra  St.  Hil.    PIF 
JDR  404,  20  Apr  1967,  J- 16. 

Papaveraceae 

Eschscholzia  caUfornica  Cham.    ANF 

JDR  .383,  20  Apr  1967.  M-14;  JDR  601,  7 
Jun  1967,  H-14;  JDR  1078,  11  Apr  1968,  F- 
14;  JDR  1182,  11  Apr  1968,  J-15. 

Stylomecon  heterophylla   (Benth.)  G.  Tavl. 
ANF 

JDR  1068,  11  Apr  1968,  E-7. 

Pittosporaceae 

Pittosponini    crassifoUiim    Banks   &    Soland. 
PCX 
JDR  408,  20  Apr  1967,  D-14. 

Pittosponini  eugenioidcs  A.  Cunn.    PCT 
JDR  490,  2  May  1967,  J-6. 

Pittosporum  rhomhifolium  A.  Cunn.    PCT 
JDR  743,  22  Sep  1967,  D-14. 

Pittosporum  undulatwn  Vent.    PCT 
JDR  449,  20  Apr  1967.  K-6. 

Pittosporum  viridiflorum  Sims.    PCT 
JDR  712.  25  Jmi  1967,  K-6. 

Plantaginaceae 

Plantago  hookeriana  F.  &  M.  var.  caUfornica 
(Greene)  Poe.    ANF 

JDR  1034,  31  Mar  1968,  H-12;  JDR  1161. 
11  Apr  1968,  G- 13. 

Plantago  lanceolata  L.    PIF 

JDR  607,  7  ]nn  1967,  G-I3;  JDR  1193,  11 
Apr  1968,  F-14;  JDR  1227,  12  Apr  1968,  K-6. 

Polemoniaceae 

Cilia  achilleijolia  Benth.  ssp.  multicaulis 
(Benth.)  V.  &  A.  Grant.    ANF 
JDR  1089,  11  Apr  1968,  F-6. 

Cilia  capitata  Dougl.    ANF 
JDR  382,  20  Apr  1967,  M-14. 

Cilia  clivorum  (Jeps.)  V.  Grant.    ANF 
JDR  1355,  12  Apr  1968,  K-9. 

Navarretia  squarrosa  (Esch.)  H.  &  A.    ANF 
JDR  624,  7  Jun  1967,  K-14. 

Polemonium  carneum  Gray.    PNF 
V  without  number,  1876. 


Folygalaceae 

Poll/gala  \  dalmaisina  Baile\ .     PCS 
JDR  1406,  10  Jun  1968,  E-13. 

Polygonaceae 

Eriogonum  latifoliuin  Smith  in  Rees.    PNF 
JDR  654,  2  May  1967,  F-7. 

Poli/gonum  avicularc  L.    ,\IF 
JDR  764,  22  Sep  1967,  F- 12. 

Pterostegia  drymarioides  F.  6c  .M.    .\NF 
JDR  1038,  2  May  1968,  K-9. 

Rumex  acetosella  L.    PIF 

JDR  635,  20  Jun  1967,  L-16;  JDR  666.  20 
Jun  1967,  M-16;  JDR  1199,  12  Apr  1968,  H- 
14. 

Rumex  crispus  L.    PIF 

JDR  665,  20  Jun  1967,  L-16. 

Rumex  pulclwr  L.    PIF 

JDR  766,  22  Sep  1967,  E-12. 

Portulacaceae 

Calandrinia  ciliata  (R.  6c  P.)  DC.  var.  men- 
ziesii  (Hook.)  Macbr.    ANF 
JDR  1055,  31  Mar  1968,  E-12. 

Montia  perfoliata  (Donn)  Howell.    .\NF 

JDR  388,  20  Apr  1967,  L-5;  JDR  1015,  2 
Mar  1968.  E-10. 

Primulaceae 

Anagallis  arvensis  L.    .\IF 
JDR  462.  28  Apr  1967,  F-7. 

Anagallis     arvensis     L.     forma     caerulea 
(Schreb.)  Baiung.    .\IF 

Observed  by  R.  &  J.,  2  May  1967. 

Dodecatheon  hendersonii  Grav.    PNF 

JDR  1024,  2  Mav  1968,  E-'lO;  JDR  1031.  2 
May  1968,  G-I2. 

Proteaceae 

Grevillea  robusta  \.  Cunn.    PCT 
JDR  7.39.  22  .Sep  1967,  D-14. 

Ranunculaceae 

Delphinium  califoniicum  T.  &  G.    PNF 
JDR  613,  7  Jun  1967,  J-9. 

Ranunculus  califomicus  Benth.    PNF 

JDR  415,  20  Apr  1967.  1-4:  JDR  10.32,  2 
Mav  1968,  1-9. 


402 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


Ranunculus  niuricatus  L.    AIF 

JDR  658,  20  Jim  1967,  J-9;  JDR  1705,  20 
Apr  1969,  F-7. 

Thalictrum  pohjcarpum  (Torr.)  Wats.    PNF 

JDR  1057,  31  Mar  1968,  E-10;  JDR  1184, 
11  Apr  1968,  G-12. 

Rhamnaceae 

Ceanothus  thyrsiflorus  Esch.    PNS 

JDR  406,  20  Apr  1967,  D-13;  JDR  706,  25 
Jun  1967, 1-4. 

Rhamnus  crocea  Nutt.    PNS 

JDR  948,  6  Feb  1968,  J-8;  JDR  983,  6  Feb 
1968,  E-12. 

Rosaceae 

Acaena  californica  Bitt.    PNF 
R.  &  J.  21187,  2  May  1967,  J-11. 

Adenostoma  fascicuhtum  H.  &  A.    PNS 
JDR  569,  7  Jun  1967,  E-12. 

Cotoneaster  franchetti  Bois.    PCS 
JDR  483,  2  May  1967,  E-14. 

Cotoneaster  pannosus  Franch.    PIS 
D  79-40,  6  July  1979,  G-8. 

Crataegus  monogyna  Jacq.    PCT 
JDR  492,  2  May  1967,  E-14. 

Eriobotnja  japonica  (Thunb.)  Lindl.    PCT 
JDR  7.38,  22  Sep  1967,  E-14. 

Heteromeles  arhutifoUa  (Ait.)  M.  Roem.    PNS 

JDR  492a,  2  May  1967,  G-14;  JDR  611,  7 

Jun  1967,  K-15;  JDR  951,  27  Dec  1967,  L-15. 

Holodiscus  discolor  (Pursh)  Maxim.    PNS 

JDR  562,  7  Jun  1967,  G-8;  JDR  701,  25  Jun 
1967,  D-10. 

Horkelia  californica  Cham.  &  Schlecht.    PNF 
R.  &  J.  21194,  2  May  1967,  H-9. 

Mains  sijlvestris  Mill.    PCT 
JDR  493,  2  May  1967,  E-14. 

Oernleria   cerasifortnis   (T.    &   G.)    Landon. 
PNS 

JDR  412,  20  Apr  1967,  G-7;  JDR  1683,  30 
Dec  1968,  L-7. 

Potentilla  glandulosa  Lindl.    PNF 
Observed  by  H,  16  May  1946. 

Prunus  amieniaca  L.    PCT 
JDR  754,  22  Sep  1967,  E- 13. 

Prunus  avium  L.    PCT 
JDR  566,  7  Jun  1967,  F-8. 


Prunus  cerasifera  Ehrh.  'atropurpurea'.    PCT 
JDR  484,  2  May  1967,  E-13. 

Pyracantha  angustifolia   (Franch.)  Schneid. 
PCS 
JDR  946,  27  Dec  1967,  K-15. 

Pyrus  communis  L.    PCT 

JDR  1374,  29  Apr  1968,  K-15. 

Rosa  californica  C.  &  S.    PNS 
JDR  1390,  8  Jun  1968, 1-16. 

Rosa  gymnocarpa  Nutt.    PNS 
JDR  576,  7  Jun  1967,  J-4. 

Rubus  parviflorus  Nutt.  var.  velutinus  (H.  & 
A.)  Greene.    PNS 

JDR  590,  7  Jun  1967,  F-U;  JDR  1114,  11 
Apr  1968,  1-8. 

Rubus  procerus  P.  J.  Muell.    PNS 
JDR  1070,  11  Apr  1968,  F-7. 

Rubus  ursinus  C.  &  S.    PNS 

JDR  453,  28  Apr  1967,  L-7;  JDR  1072,  11 
Apr  1968,  G-7. 

Rubiaceae 

Coprosma  repens  A.  Rich.    PCS 
JDR  740,  22  Sep  1967,  D-13. 

Galium  aparine  L.    AIF 

JDR  1264,  12  Apr  1968,  C-12;  JDR  1286, 
12  Apr  1968,  K-9. 

Galium  nuttallii  Grav.    PNF 

JDR  463.  28  Apr  1967,  F-11;  JDR  1267,  12 
Apr  1968,  D-12;  JDR  1298,  12  Apr  1968,  K- 
9;  JDR  1329,  12  Apr  1968,  G-7. 

Sherardia  arvensis  L.    AIF 

JDR  1421,  10  Jun  1968,  F-8;  JDR  1106,  11 
Apr  1968,  G-7. 

Salicaceae 

Populus  alba  L.    PCT 

JDR  473,  28  Apr  1967,  J-6. 

Populus  nigra  L.  var.  italica  Du  Roi.    PCT 
JDR  734,  22  Sep  1967,  J-5. 

Salix  babylonica  L.    PCT 
JDR  557,  2  May  1967,  F-8. 

Salix  lasiolepis  Benth.    PNT 

JDR  719,  22  Sep  1967,  L-11;  JDR  987,  6 
Feb  1968,  J-15;  JDR  988.  6  Feb  1968,  G-15. 

Salix  lasiolepis   Benth.   var.   bigelovii   (Torr.) 
Bebb.    PNT 

JDR  989,  6  Feb  1968,  L-8. 


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Ripley:  Angel  Island  Plants 


403 


Saxifragaceae 

Heuchera  niicranlha  Doiigl.    PNF 

JDR  577,  7  Jun  1967,  H-7;  JDR  1528,  14 
Jun  1968,  E-7. 

Lithophra^nia  hctvwplujUum  (H.  &  A.)  T.  & 
G.    PNF 

JDR  547,  2  Mav  1967,  D-11;  JDR  1058,  31 
Mar  1968,  F-IO. 

Rihes  cahfornicuin  (H.  6c  A.)    PNS 
JDR  1243,  12  Apr  1968,  L-6. 

Saxifrag^a  califomica  Greene.    PNS 
JDR  1016,2  May  1968,  F- 10. 

Scrophulariaceae 

Castilleja  affinis  H.  &  A.    PNF 
JDR593",  7  Jun  1967,  F-10. 

Castilleja  foliolosa  H.  &  A.    PNF 
JDR  1052,31  Mar  1968,  E-11. 

Castilleja  franciscana  Penn.    PNF 
R.  &  J.  21310,  2  May  1967,  D-10. 

Castilleja  latifolia  H.  &  A.    PNF 

JDR  1150,  11  Apr  1968,  J-15;  JDR  1258, 
12  Apr  1968,  D-11;  JDR  1322,  12  Apr  1968, 
G-12. 

CoUinsia  heterophylla  Buist  ex  Grah.    ANF 

JDR  538,  2  May  1967,  D-11;  JDR  1212,  12 
Apr  1968,  G-7;  JDR  1346,  12  Apr  1968,  K-9. 

CoUinsia  multicolor  Lindl.  &  Paxt.    ANF 
R.  &  J.  21312,  2  May  1967,  D-10. 

Linaria  canadensis  (L.)  Dum. -Coins,  var.  tex- 
ana  (Scheele)  Penn.    BNF 
JDR  381,  20  Apr  1967,  M-12. 

Linaria  cipnbalaria  (L.)  Mill.    PIF 
JDR  1502,  14  Jun  1968,  L-5. 

Mimuhts  aurantiacus  Curt.    PNS 

JDR  468,  28  Apr  1967,  K-15;  JDR  461,  28 
Apr  1967,  C- 11. 

Mimnlus  guttatus  Fisch.  ex  DC.    PNF 
JDR  655,  20  Jun  1967,  K-16. 

Mitnulus  guttatus  Fisch.  ex  DC.  var.  arvensis 
(Greene)  Munz.    PNF 

JDR  1400,  10  Jun  1968,  N- 15. 

Orthocarf)us  attenuatus  Gray.    ANF 
JDR  1322,  12  Apr  1968, 1-8. 

Otihocarpus  densiflorns  Benth.    ANF 

JDR  456,  28  Apr  1967,  F-12;  JDR  584,  7 
Jun  1967,  J-14;  JDR  1244,  12  Apr  1968,  D- 
13. 


Orthocarpus  pusillus  Benth.    .\NF 

JDR  426,  20  Apr  1967,  1-4;  JDR  1147,  11 
Apr  1968,  F-8;  JDR  1245,  12  Apr  1968,  D- 
10. 

Scropliularia  califomica  ChauL    PNF 
JDR  591,  7  Jun  1967,  E-11. 

Veronica  arvensis  L.    .AIF 
JDR  1098,  11  Apr  1968,  F-8. 

Simarubaceae 

Ailanthus  altissinia  (Mill.)  Swingle.    PCX 
JDR  648,  20  Jun  1967,  L-16. 

Solanaceae 

Nicotiana  glauca  Grah.    PCS 
Observed  by  B,  1980,1-15. 

Solanum  nigriwi  L.    AIF 

Observed  by  H,  16  .May  1946. 

Tropaeolaceae 

Tropaeoluui  niajiis  L.    .\IF 
JDR  1424.  10  Jun  1968,  N-16. 

Umbelliferae 

Angelica  tomentosa  Wats.    PNF 
Observed  by  H,  16  May  1946. 

Daucus  ptisilhis  Michx.    ANF 
JDR  644,  7  Jun  1967,  F-7. 

Foeniculum  vulgare  Mill.    PNF 
JDR  722,  25  Jun  1967.  E- 13. 

Ileracleum  lanatum  Michx.    PNF 

JDR  503,  2  May  1967,  D-12;  JDR  1209,  12 
Apr  1968,  F-7. 

Lomatiiim  dusycarpuni   (T.  ik  G.)  C.  &  R. 
PNF 

JDR  467,  28  Apr  1967,  1-8. 

Osniorhiza  chilensis  H.  &  A.    PNF 
JDR  617,  7  Jun  1967,  J-8. 

Perideridia  kelloggii  (Grav)  Mathias.    PNF 
R.  &  J.  21 123,  2  May  i967,  K-9. 

Sanicula  hipinnatifida  Dougl.    P.NF 

JDR  420,  20  Apr  1967,  1-9;  JDH  670.  7  Jun 
1967,  K-15;  JDR  1273,  12  Apr  196S,  F  in 
JDR  1212,  12  Apr  1968,  G-7. 

Sanicida  crassicaulis  Poepp.    PNF 

JDR  436,  20  Apr  1967,  1-4;  JDR  118.3.  11 
Apr  1968,  1-15;  JDR  1282,  12  Apr  1968,  K-9. 

Scandix  pecten-veneris  L.     \\F 
JDR  518,  2  May  1967,  H-15. 


404 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


Torilis  arvensis   (Huds.)   Link   ssp.  purpurea 
(Ten.)  Hayek.    AIF 

D  80-39,  29  May  1980,  F-8. 

Torilis  nodosa  L.    AIF 

JDR  663,  20  Jun  1967,  L-6;  JDR  1160,  11 
Apr  1968,  F-14;  JDR  1348,  12  Apr  1968,  K-9. 

Urticaceae 

Urtica  holosericea  Nutt.    PNF 
JDR  736,  22  Sep  1967,  G-15. 

Valerianaceae 

Centranthus  ruber  (L.)  DC.    PIF 
JDR  528,  2  May  1967,  H-17. 

Plectritis  macrocera  T.  &  G.    ANF 
JDR  434,  20  Apr  1967,  E-14. 

Verbenaceae 

Verbena  robusta  Greene.    PNF 
Observed  by  H,  16  May  1946. 

Violaceae 

Viola  pedunculata  T.  &  G.    PNF 

JDR  419,  20  Apr  1967,  1-4;  JDR  1268,  12 
Apr  1968,  D-12. 

Vitaceae 

Parthenocissus   tricuspidata    (Sieb.   &   Zucc.) 
Planch.    PCF 
JDR  724,  25  Jun  1967,  G-14. 

Class  Monocotyledoneae 

Agavaceae 

Agave  americana  L.    PCS 
JDR  803,  22  Sep  1967,  K-4. 

Agave  americana  L.  'variegata\    PCS 
JDR  959,  27  Dec  1967,  K-6. 

Cordyline  striata  Endl.    PCS 
JDR  990,  6  Feb  1968,  H-15. 

Amaryllidaceae 

Allium  dichlamijdeum  Greene.    PNF 
JDR  622,  7  Jun  1967,  K-8. 

Amaryllis  belUidonna  L.    PCF 
JDR  725,  25  Jun  1967,  J-6. 

Brodiaea  lam  (Benth.)  Wats.    PNF 
JDR  592,  7  Jun  1967,  F-11. 

Brodiaea  pulchella  (Salisb.)  Greene.    PNF 

JDR  440,  20  Apr  1967,  J-6;  JDR  1190,  11 
Apr  1968, 1-14;  JDR  1237,  12  Apr  1968,  K-5. 


Narcissus  poetaz  Hort.    PCF 
JDR  992,  6  Feb  1968,  E-13. 

Narcissus  pseudo-narcissus  L.    PCF 
JDR  1042,  2  Mar  1968,  1-7. 

Araceae 

Zantedeschia  aethiopica  (L.)  Spreng.    PIF 
JDR  806,  22  Sep  1967,  1-6. 

Cyperaceae 

Carex  barbarae  Dewey.    PNSd 

JDR  700,  25  Jun  1967,  F-12;  JDR  1189a, 
11  Apr  1968,  G-14;  JDR  1367,  29  Apr  1968, 
D-14. 

Carex  brevicaulis  Mkze.    PNSd 
H  21850,  16  May  1946. 

Carex  gracilior  Mkze.    PNSd 
JDR  1288,  12  Apr  1968,  K-9. 

Carex  tumulicola  Mkze.    PNSd 

JDR  400,  20  Apr  1967,  M-15;  JDR  1093, 
11  Apr  1968,  M-16;  JDR  1347,  12  Apr  1968, 
K-9. 

Cyperus  eragrostis  Lanik.    PNSd 
JDR  750,  22  Sep  1967,  G-15. 

Scirpus  cernuus  Vahl.  var.  californicus  (Torr.) 
Beetle.    ANSd 

JDR  1402,  10  Jun  1968,  N-15. 

Gramineae 

Agrostis  diegoensis  Vasey.    PNG 
JDR  619,  7  Jun  1967, 'K-8. 

Agrostis  exarata  Trin.  var.  pacifica  Vasev. 
PNG 

JDR  672,  20  Jun  1967,  L-6. 

Agrostis  hall  a  Vasey.    PNG 
Observed  by  H,  16  May  1946. 

Agrostis    setniverticillata    (Forsk.)    C.    Chr. 
PIG 

JDR  769,  22  Sep  1967,  M-15;  JDR  747,  22 
Sep  1967,  D-11. 

Aira  caryophyllea  L.    AIG 

JDR  628,  7  Jun  1967,  K-8;  JDR  1164,  11 
Apr  1968,  F-14;  JDR  1314,  12  Apr  1968,  K-9. 

Ammophila  breviligulata  Fernald.    PIG 
JDR  947,  27  Dec  1967,  K- 16. 

Avena  barbata  Brot.    AIG 

JDR  395,  20  Apr  1967,  M-I4;  JDR  1145, 
11  Apr  1968,  1-8;  JDR  1263,  12  Apr  1968,  C- 
12. 


December  1980 


Ripley:  Angel  Island  Plants 


405 


Aiena  fotua  L.    AIG 

JDR  572,  7  Jun  1967,  K-8;  JDR  1382,  29 
Apr  1968,  K-15. 

Briza  maxima  L.    AIG 

JDR  1701,  20  Apr  1969,  1-7;  JDR  1707,  20 
Apr  1969,  E-U. 

Briza  minor  L.    AIG 

JDR  572,  7  Jun  1967,  H-7;  JDR  1382,  29 
Apr  1968,  J-15. 

Brumus  carinattis  H.  &  A.    BNG 

JDR  447a,  20  Apr  1967,  1-7;  JDR  1171,  11 
Apr  1968,  G-7;  JDR  1284,  12  Apr  1968,  K-9. 

BroDius  diandrus  Roth.    AIG 

JDR  394,  20  Apr  1967,  M-15;  JDR  447,  20 
Apr  1967,  J-7;  JDR  509,  2  May  1967,  D-14; 
JDR  527,  2  May  1967,  1-16:  JDR  1086.  11 
Apr  1968,  F-14. ' 

Bromus  madritensis  L.    AIG 

JDR  446,  20  Apr  1967,  J-6:  JDR  646,  20 
Jun  1967,  L-5. 

Bromus  marginatus  Nees  +  PNG 
JDR  1324,  12  Apr  1968, 1-8. 

Bromus  mollis  L.    AIG 

JDR  427,  20  Apr  1967,  J-5;  JDR  1372,  29 
Apr  1968,  J-13;  JDR  1404,  10  Jun  1968,  N- 
16. 

Bromus  racemosus  L.    .\IG 

R.  &  J.  21330,  2  May  1967,  D-10. 

Bromus  ruhens  L.    AIG 

JDR  1124,  11  Apr  1968,  F-12;  JDR  1338, 
29  Apr  1968,  1-9. 

Cortaderia    selloana    (Schult.)    Aschers.    & 
Graebn.    PCG 

JDR  532,  2  May  1967,  K-16;  JDR  633,  20 
Jun  1967,  H-16;  JDR  957,  27  Dec  1967,  L-15. 

Cynodon  dactijlon  (L.)  Pers.    PIG 
JDR  1522,  14  Jun  1968,  K-9. 

Cynosurus  erhinatus  L.    AIG 
D  79-41,  6  Jul  1979,  E- 12. 

Dactylis  glomerata  L.    PIG 
Observed  by  H,  16  May  1946. 

Danthonia  californica  Roland,  var.  amcricana 
(Scribn.)  Hitchc.    PNG 
JDR  1188,  11  Apr  1968,  F-14. 

Danthonia  pilosa  R.  Br.    PIG 
T8468,  16  Jun  1978,  H-7. 

Distichlh  spicata  (L.)  Greene.    PNG 
JDR  728,  25  Jun  1967,  K-5. 


Elymus  glaucuso  Buckl.    PNG 

JDR  460,  28  Apr  1967,  1-9;  JDR  548,  2 
May  1967,  K-8;  JDR  1280,  12  Apr  1968,  K-9. 

Ehjmus  triticoidcs  Buckl.    PNG 
Observed  by  H,  16  May  1946. 

Elymus  triticoides  Buckl.  ssp.  multif torus 
Gould.    ANG 

JDR  705,  25  Jun  1967,  G-8. 

Festuca  californica  Vasey.    PNG 

JDR  551,  2  May  1967,  K-8;  JDR  1023,  2 
Mar  1968,  1-9;  JDR  1077,  11  Apr  1968,  E-9; 
JDR  1123,  11  Apr  1968,0-12. 

Festuca     drrtonensis     (All.)     .\schers.     & 
Graebn.    AIG 

JDR  541,  2  May  1967,  G-11;  JDR  1104,  11 
Apr  1968,  G-7;  JDR  1285.  12  Apr  1968.  K-9: 
JDR  1231,  12  Apr  1968,  K-6. 

Festuca  idahoensis  Elmer.    PNG 
JDR  1146,  11  Apr  1968,1-9. 

Festuca  megalura  Nutt.    .ANG 

JDR  1082,  11  Apr  1968,  G-14;  JDR  1022,  2 
Mar  1968,  E-11;  JDR  1260,  12  Apr  1968.  D- 
10;  JDR  1303,  12  Apr  1968,  K-9. 

Festuca  myuros  L.    AIG 

JDR  507,  2  May  1967,  G-13;  JDR  550,  2 
May  1967,  D-11;  JDR  1262,  12  Apr  1968,  D- 
10.' 

Festuca  pacifica  Piper.    ANG 
JDR  1187,  11  Apr  1968,1-14. 

Festuca  reflexa  Buckl.    ANG 
JDR  1420,  10  Jun  1968,  K-9. 

Festuca  rubra  L.    PNG 

Observed  by  H,  16  May  1946. 

Gastridium    ventricosum    iCiouan)    Schin/   6c 
Thell.    ANG 

JDR  652,  20  Jun  1967,  K-15;  JDR  762,  22 
Sep  1967,  M-15;  JDR  1506.  14  Jun  1968.  L-6. 

Holcus  lanatus  L.    PIG 

JDR  1708,  16  Jun  1969,  F-7. 

Hordeum  hrachyantherum  Nevski.    PNG 

JDR  582,  7  jun  1967,  E-U;  JDR  693,  25 
Jun  1967,  D-13;  JDR  986,  6  Feb  1968,  D-13. 

Hordeum  glaucum  Steud.    AIG 

JDR  1095,  11  Apr  1968,  G-7;  JDR  1128,  11 
Apr  1968,  F-12. 

Hordeum  leporinum  Link.    MC 

JDR  443,  20  Apr  1967,  H-6:  JDH  1101.  11 
Apr  1968,  J-7;  JDR  1363.  29  Apr  1968,  1-9; 
JDR  1383,  29  Apr  1968.  J- 16. 


406 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


Hordeum  vulgare  L.    AIG 
Observed  by  H,  16  May  1946. 

Koeleria  macrantha  (Ledeb.)  Spreng.    PNG 

JDR  554,  2  May  1967,  K-14;  JDR  1307,  12 
Apr  1968,  K-9;  JDR  1324a,  12  Apr  1968, 1-9. 

Lolium  multiflorum  Lamk.  var.  muticum 
DC.    PIG 

JDR  651,  20  Jun  1967,  J-15;  JDR  1369,  29 
Apr  1968,  J- 13. 

Lolium  perenne  L.    PIG 

R.  &  J.  21040,  2  May  1967,  K-9. 

Melica  californica  Scribn.    PNG 

JDR  1203,  12  Apr  1968,  G-15;  JDR  1381, 
29  Apr  1968,  K-15. 

Melica  torreijana  Scribn.    PNG 

JDR  549,  2  May  1967,  K-8;  JDR  1249,  12 
Apr  1968,  D-12;  JDR  1163,  11  Apr  1968,  G- 
15;  JDR  1157,  11  Apr  1968,  G-12. 

Monenna  cylindrica  (Willd.)  Coss.  &  Dur. 
AIG 

Observed  by  H,  16  May  1946. 

Phahris  californica  H.  &  A.    PNG 

JDR  464,  28  Apr  1967,  1-9;  JDR  1143,  11 
Apr  1968,  H-10;  JDR  1331,  12  Apr  1968,  G- 
12. 

Plialaris  minor  Retz.    AIG 
JDR  618a,  7  Jun  1967,  K-8. 

Poa  annua  L.    AIG 

JDR  985,  6  Feb  1968,  D-11;  JDR  1020,  2 
Mar  1968,  F-13;  JDR  1100,  11  Apr  1968,  F-7. 

Poa  bolanderi  Vasey  ssp.  howellii  (Vasey  & 
Scribn.)  Keck.    ANG 
JDR  1186,  11  Apr  1968,  F-14. 

Poa  scahrella  (Thiirb.)  Benth.    PNG 

JDR  1085,  11  Apr  1968,  F-14;  JDR  1148a, 

11  Apr  1968,  1-14;  JDR  1240,  12  Apr  1968, 
D-12. 

Poa  unilateralis  Scribn.    PNG 

JDR  1087,  11  Apr  1968,  F-14;  JDR  1317, 

12  Apr  1968,  K-9;  JDR  1321,  12  Apr  1968,  I- 
9. 

Polypogpn  interrnptiis  H.  B.  K.  PIG 

JDR  1396,  8  Jun  1968,  N-17;  JDR  1403,  10 
Jun  1968,  M-14;  JDR  1526,  14  Jun  1968,  N- 
16. 

Pohjpogon  monspeliensis  (L.)  Desf.    AIG 

JDR  653,  20  Jun  1967,  L-15;  JDR  758,  22 
Sep  1967,  N-16. 


Sitanion   x  Jianscnii   (Scribn.)  J.   G.   Smith. 
PNG 

JDR  1371a,  29  Apr  1968,  D-14. 

Sitanion  jubatum  J.  G.  Smith.    PNG 

JDR  767,  22  Sep  1967,  J-15;  JDR  1381a,  29 
Apr  1968,  L-15. 

Stipa  lepida  Hitchc.    PNG 
JDR  1198,  12  Apr  1968,  F-13. 

Stipa  pulchra  Hitchc.    PNG 

JDR  539,  2  May  1967,  D-11;  JDR  1127,  11 
Apr  1968,  H-11;  JDR  1131,  11  Apr  1968,  E- 
11;  JDR  1371,  29  Apr  1968,  C-14. 

Iridaceae 

Iris  longipetala  Herbert.    PNG 
JDR  429,  20  Apr  1967,  J-7. 

Sisijrinchium  helium  Wats.    PNG 

JDR  387,  20  Apr  1967,  L-6;  JDR  1118,  11 
Apr  1968,  G-9;  JDR  1246,  12  Apr  1968,  C- 
11. 

Juncaceae 

Juncus  balticus  Willd.    PNR 
JDR  1292,  12  Apr  1968,  K-9. 

Juncus  bufonis  L.    ANR 
JDR  640,  20  Jun  1967,  J-8. 

Juncus  effusus  L.  var.  pacificus  Fern.  & 
Wieg.    PNR 

JDR  751,  22  Sep  1967,  G-15. 

Juncus  patens  E.  Mey.    PNR 

JDR  397,  20  Apr  1967,  M-16;  JDR  579,  7 
Jun  1967,  F-12. 

Juncus  tenuis  Willd.  var.  congestus  Engelm. 
PNR 

JDR  657a,  20  Jun  1967,  K-16;  JDR  1291, 
12  Apr  1967,  K-9. 

Luzula  subsesilis  (Wats.)  Buch.    PNR 

B-D  6912  without  date;  JDR  540,  2  May 
1967,  D-11;  JDR  1035,  2  Mar  1968,  F-12; 
JDR  1075,  11  Apr  1968,  J-7. 

Liliaceae 

Aloe  saponaria  (Ait.)  Haw.    PIS 
JDR  517,  2  May  1967,  G-14. 

Calochortus  luteus  Dougl.  ex  Lindl.    PNF 
D  80-47,  18  Jun  1980,  1-12. 

Chlorogalum  pomcridianum  (DC.)  Kunth  var. 
divaricatum  (Lindl.)  Hoover.    PNF 
JDR  630,  20  Jun  1967,  1-8. 


December  1980 


Ripley:  An(;el  Island  Plants 


407 


Fritillarid  hmceohta  Pursh.    PNF 

JDR  1206,  12  Apr  1968,  E-9;  JDR  1681,  16 
Mar  1969,  L-7. 

Knaphofia  uvaria  (L.)  Okeii.    PCS 
JDR  439,  20  Apr  1967,  J-5. 

S7nilacina  raccmosa  (L.)  Desf.    PNF 
JDR  1116,  11  Apr  1968,  G-9. 

Smilacina  stellata  (L.)  Desf.  var.  sessilifolia 
(Baker)  Renders.    PNF 

R.  &J.  21111,  2  May  1967,  L-9. 

Trilliutu  chloropctdluiu  (Torr.)  Howell.    PNF 
JDR  1680,  16  Mar  1969,  K-7. 

Zigadcnus  fremontii  (Torr.)  Torr.    PNF 

JDR  465,  28  Apr  1967,  1-9;  JDR  1033,  2 
Mar  1968,  G-12. 

Orchidaceae 

Hahernaria  elegans  (Lindl.)  Roland.    PNF 
JDR698,  25Jnn  1967,  F-12. 

Palmae 

Phoenix  canariensis  Chaboud.    PCT 
JDR  512,  2  May  1967,  C- 13. 

Typhaceae 

Typlia  angustifoUa  L.    PNF 
JDR  824,  22  Sep  1967,  J-9. 

Zosteraceae 

Zostcra    marina    L.    var.    latifolia    Morong. 
PNF 

JDR  1391,  8  Jun  1968,  F-8;  JDR  1426,  10 
Jun  1968,  G-6. 

PhijUospadix  torrciji  Wats.    PNF 
JDR  1425,  10  Jun  1968,  N-17. 


ACKNOWLEDG.VIENTS 

It  is  my  pleasure  to  acknowledge  the  in- 
valuable a.ssistance  of  the  following  individ- 
uals who  generously  assited  in  the  prepara- 
tion of  this  paper:  J.  T.  Howell,  H.  D.  Thiers, 
M.  Josselvn,  C.  Best,  C.  H.  True,  A.  Day,  and 
CO.  Burke. 


LlTEHATl  HE  ClTEl) 

Bailey,  K.  li.,  W  .  I',  liuviv.  and  D.  L.  Jones.  Ujf>4. 
Franciscan  and  related  rocks  and  their  signifi- 
cance in  the  neolojfy  of  western  California.  Cali- 
fornia Division  of  .Mines  and  Geologv,  Bull.  18.3. 

Blo.vvm,  T.  W.  19.56.  Jadeite-hearing  metagravwackes  in 
California,  .\merican  .Mineralogist  41:488-496. 

1960.  Jadite-rocks  and  glaucophane-schists  from 

.\ngel  Island,  San  Francisco  Bav.  California.  Am. 
Jour.  Sci.  258:.55.5-.57.3. 

HouLE.NBERG,  C.  S.,  A.ND  I.  .\.  .\bbott.  1966.  Supplement 
to  Smith's  marine  algae  of  the  Monterev  Penin- 
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HowEi.i.,  J.  T.  1970.  .Marin  flora  and  supplement.  Uni- 
versity of  California  Press.  Berkelev. 

-Mc.NZ,  P.  A.  1973.  .\  California  flora  and  supplement. 
University  of  California  Press,  BerkeleN'. 

Obkki.ander,  C  T.  19.56.  Simimer  fog  precipitation  on 
the  San  Francisco  Peninsula.  Ecology 
37(4):851-852. 

Ra.nso.me,  F.  L.  1894.  The  geolog\  of  .\ngel  Island.  Uni- 
versity of  California  Pub.  Dept.  Geology 
1:19.3-240. 

Ripley,  J.  D.  1969.  .\  floristic  and  ecological  study  of 
.\ngel  Island  State  Park,  .Marin  County,  Califor- 
nia. Unpublished  thesis,  San  Francisco  Slate  Uni- 
versity, San  Francisco.  California. 

ScHLOCKEH.  J.  M.  1961.  Podingite  from  .\ngel  Island. 
San  Francisco  Bav.  California.  U.S.  Geological 
Survey  Prof.  Paper  400-B,  p.  B311-B312. 

Scni.ocKER,  J.  M.,  M.  G.  Bomlla.  ano  D.  H.  Radhiuch. 
1958.  CJeology  of  the  San  Francisco  north  quad- 
rangle. California.  U.S.  Geological  Survey  Mi.sc. 
(;eol.  Inv.  .Map  1-272. 

Smith.  G.  .M.  1944.  .Marine  algae  of  the  Monterey  Penin- 
sula, (California.  Stanford  Uni\ersit\'  Press.  Stan- 
ford, California. 


ADDITIONS  TO  THE  VASCULAR  FLORA  OF  TETON  COUNTY,  WYOMING 

Ronald  L.  Hartman'  and  Roliert  W.  Lichvar^-^ 


Abstract.—  An  annotated  list  of  125  taxa  new  to  the  flora  of  Teton  County  is  presented,  increasing  the  number  of 


known  species  to  1043. 

Recently,  Shaw  (1976)  published  a  "Field 
Guide  to  the  Vascular  Plants  of  Grand  Teton 
National  Park  and  Teton  County,  Wyom- 
ing," which  was  said  to  be  based  in  part  on 
the  holdings  of  the  Rocky  Mountain  Herbar- 
ium. Unfortruiately,  that  herbarium  was  con- 
sulted mainly  for  the  taxonomic  groups 
treated  by  Dr.  Robert  D.  Dorn,  including  the 
Poaceae  and  the  genera  Artemisia  and  Salix. 
Consequently,  many  of  the  125  taxa  listed  be- 
low were  omitted.  Additionally,  recent  in- 
tensive collecting  by  the  junior  author  in  the 
Gros  Ventre  Mountains  resulted  in  the  dis- 
covery of  20  species  apparently  new  to  the 
flora  of  Teton  County.  This  paper  updates 
the  list  of  known  taxa  for  this  much-visited 
area  and  emphasizes  the  richness  of  its  flora 
(1043  species).  The  nomenclature  and  tax- 
onomy follow  that  of  Hitchcock  and 
Cronquist  (1973),  unless  otherwise  indicated 
by  synonymy.  The  senior  author  has  checked 
determinations  on  those  specimens  not  anno- 
tated by  specialists  in  their  respective  groups. 
Dr.  Dorn  is  acknowledged  for  assistance  with 
the  manuscript. 

Apiaceae 

Angelica  roseana  Hand.  Teton  Mts.,  16 
Aug  1899,  A.  Nelson  6  E.  Nelson  6500,  anno- 
tation by  M.  E.  Mathias  &  L.  Constance, 
1940. 

Cicuta  douglasii  (DC.)  Coult.  &  Rose. 
Kent's  Corner  (SI  T41N  R115W),  elev.  6700 
ft,  10  Aug  1977,  R.  Lichvar  1235. 

Cymopterus  terebinthinus  (Hook.)  T.  &  G. 
var.  calcareus  (M.  E.  Jones)  Cronq.  Teton 
Pass  Mts.,  elev.  7500  and  9000  ft,  22  Jul 
1920,  E.  B.  6  L.  B.  Paijson  2073,  annotation 
by  M.  E.  Mathias,  1930,  as  Pteryxia  t.  var.  c. 


Ligusticum  porteri  Coult.  &  Rose.  7.7  mi 

W  of  Flagg  Ranch,  elev.  7600  ft,  14  Jul  1956, 
W.  G.  Solheim  4571;  Whetstone  Creek,  24 
Jul  1929,  O.  /.  Murie  86. 

Apocynaceae 

Apocynum  cannabinum  L.  var.  glaberri- 
muni  DC.  Hot  Springs  Bar,  20  mi  S  of  Jack- 
son, 19  Jul  1901,  E.  D.  Merrill  6  E.  N.  Wilcox 
1042,  cited  by  Woodson  (1930). 

ASTERACEAE 

Antennaria  anaphaloides  Rydb.  Gros 
Ventre  River,  Jackson  Hole,  14  Jul  1901,  E. 
D.  Merrill  6  E.  N.  Wilcox  986;  Sheep  Creek 
(S16  T41N  R115W),  elev.  6900  ft,  5  Jun 
1977,  R.  Lichvar  242. 

Arnica  sororia  Greene.  Near  Moose  (SWi/4 
S16  T43N  R115W),  4  Jul  1970,  D.  W.  Sa- 
binske  13C. 

Aster  eatonii  (Gray)  Howell.  Along  Tag- 
gert  Creek,  elev.  7500  ft,  14  Aug  1932,  L. 
Williams  1041,  annotation  by  M.  L.  Dean, 
1963;  Gros  Ventre  River  (SI  T42N  R115W), 
elev.  6900  ft,  21  Aug  1977,  R.  Lichvar  1285. 

Chrysopsis  horrida  Rydb.  Gros  Ventre 
River,  16  Aug  1894,  A.  Nelson  1084,  annota- 
tion by  V.  L.  Harms,  1963,  as  Heterotheca 
horrida  (Rydb.)  Harms. 

Crepis  runcinata  (James)  T.  &  G.  ssp. 
glauca  (Nutt.)  Babe.  &  Stebb.  Gros  Ventre 
Mts.,  E  of  Gros  Ventre  Slide,  elev.  7300  ft,  28 
Aug  1949,  W.  G.  Solheim  2733. 

Erigeron  corymbosus  Nutt.  Jackson  Hole, 
10  Jul  1931,  L.  Williams  290,  annotation  by 
A.  Cronquist,  1943-44;  Crvstal  Creek  (S28 
T42N  R113W),  elev.  7100  ft,  7  Jul  1977,  R. 
Lichvar  655. 


'Rocky  Mountain  Herbarium,  Department  of  Botany.  University  of  Wyoming.  Laramie,  Wyoming  82071. 

■Present  address:  Wyoming  Natural  Heritage  Program,  The  Nature  Conservancy,  1603  Capitol  Avenue,  No.  .325,  Cheyenne,  Wyoming  82001. 


408 


December  1980 


Hartman,  Lichvau:  V asci  lah  Flora 


409 


Erigeron  flagellaris  Gray.  Glacier  Creek, 
Jackson  Hole,  elev.  7500  ft,  9  Aug  1920,  E.  B. 
6  L.  B.  Paijson  2258. 

Erigcron  rydbergii  Cronq.  Sheep  Mt.  (S34 
T42N  H114\V),  elev.  9800  ft,  1  Autr  1977,  R. 
Liclnar  1015. 

Erigeron  simplex  Greene.  Cirque  basin 
near  Amphitheater  and  Surprise  Lakes,  elev. 
9750  ft,  31  Jul  1962,  /.  Merklc  62-36-  Snake 
River,  22  Aug  1894,  A.  Xel.son  96.9,  annota- 
tions by  S.  Spongberg,  1970;  Sheep  Mt.  (S34 
T42N  R114W),  elev.  9800  ft,  13  Jul  1977,  R. 
Lichvar  804. 

Erigeron  tirsiniis  D.C.  Eat.  Two-gwo-tee 
Pass,  elev.  10,500  ft,  27  Jul  1932  L.  Williams 
947;  Alaska  Ba.sin,  W  slope  Teton  Range, 
elev.  9500-10.000  ft,  14  Aug  1965.  /.  Merkle 
65-44. 

Gaillarida  aristata  Pursh.  Kellv  Post  Of- 
fice (SI  T42N  R115W),  elev.  6900'ft,  18  Aug 
1977,  R.  Lichvar  1278. 

Gnaphalium  chilense  Spreng.  Along  Snake 
River,  20  mi  S  of  Jackson,  19  Jul  1901,  E.  D. 
Merrill  ir  E.  N.  Wilcox  949. 

Gnaphalium  viscosiim  H.B.K.  Cascade 
Canyon,  elev.  7500  ft,  30  Jul  1934,  L.  Wil- 
liams 1684. 

Haplopappus  lyallii  Gray.  Two  Ocean 
Mt.,  Two-gwo-tee  Pass,  elev.  10,000  ft,  1  Aug 
1933,  L.  Williams  1366. 

Microseris  nigrescens  Hend.  Near  Two- 
gwo-tee  Pass,  elev.  9650  ft,  W.  G.  6  R.  Sol- 
Iwim  4031. 

Senecio  debilis  Nutt.  Gros  Ventre  Mts., 
elev.  7100  ft,  28  Aug  1949,  T.  F.  6  M.  S. 
Reed  2702,  annotation  bv  T.  M.  Barkley, 
1960. 

Senecio  fremontii  T.  &  G.  var.  fremontii. 
Near  Leighs  Lake,  Teton  Mts.,  26  Jul  1901, 
E.  D.  Merrill  6  E.  N.  Wilcox  1047;  Gros 
Ventre  Mts.,  10  mi  N  of  Bondurant,  15  Aug 
1922,  £.  B.  6  L.  B.  Payson  .3022. 

Senecio  werneriaefolius  Gray.  Teton  Mts., 
elev.  9000  ft,  21  Aug  1894,  A.  Xelson  979. 

Solidago  gigantea  Ait.  var.  serotina 
(Kuntze)  Cronq.  Double  Diamond  Ranch, 
elev.  7000  ft,  31  Jul  1932,  L.  Williams  966. 

Townsendia  florifer  (Hook.)  Gray.  Snake 
River,  29  May  1892,  F.  McCullough  s.n.,  an- 
notation by  J.  H.  Beaman,  1956. 

Townsendia  hookeri  Beaman  rather  than 
T.  exscapa  as  reported  in  Shaw  (1976)  ac- 
cording to  R.  Dorn  (pers.  conun.). 


Townsendia  leptotes  (Gray)  Osterh.  Sheep 
Mt.,  ca.  14  mi  NE  of  Jackson,  elev.  11,190  ft, 
30  Jul  1957,  J.  //.  Beaman  6  K.  J.  Stone  1487. 

Toivnsendia  parryi  D.C.  Eat.  Road  above 
Upper  Slide  Lake,  (iros  Ventre  River,  elev. 
9300  ft,  27  Jul  1955,  W.  G.  6  R.  Solheim 
4143;  Sheep  Mt.  (S33  T42N  R115W),  elev. 
11,100  ft,  13  Jul  1977,  R.  Lichvar  824. 

Boraginaceae 

Cryptantha  amhigua  (Grav)  Greene.  Near 

Moose  iSE'4,  S27  T43N  R115\V),  1  Jul  1971, 
D.  W.  Sahinske  B3. 

Brassicaceae 

Arabis  lemmonii  Wats.  Teton  Pass  .\lts., 
elev.  10,100  ft,  25  Jul  1920,  E.  B.  6  L.  B. 
Payson  2135. 

Arabis  microphylla  Nutt.  var.  micro- 
phylla.  Treasure  Mt.  Scout  Camp,  Teton 
Can) on,  25  Jun  1955,  L.  C.  Anderson  109. 

Barbarea  vulgaris  R.  Br.  Elk  Ranch,  elev. 
6900  ft,  13  Jun  1948.  /.  F.  6  M.  S.  Reed  2318. 

Berteroa  incana  (L.)  DC.  Alta.  Jun  1969. 
//.  P.  Alley  s.n. 

Descurainia  californica  (Gray)  Schultz. 
Snow  King  Mt.  (S7  T41\  R116W)',  elev.  7700 
ft.  22  Jun  1977, /^.  Lichvar  407. 

Draba  lanceolata  Rovle.  Sheep  Mt.  (S16 
T41N  R114W),  elev.  lO.'oOO  ft,  23  Jun  1977. 
R.  Lichvar  462. 

Draba  praealta  Greene.  Hoback  Canvon, 
elev.  7500  ft,  23  Jun  1933,  L.  Williams  1143. 
annotation  by  C.  L.  Hitchcock,  1939. 

Erysimum  inconspicuum  (Wats.)  MacM. 
Bacon  Creek,  15  Aug  1894,  A.  Xelson  916. 

Hesperis  matronalis  L.  Snow  King  Mt.  (S7 
T41N  RI 16W),  elev.  6700  ft.  22  Jun  1977.  R. 
Lichvar  392. 

Physaria  didymocarpa  (Hook.)  Gray  var. 
integrifolia  Rollins.  Near  .\dams  Ranch, 
Jackson,  15  Jul  1901,  E.  D.  Merrill  6  E.  N. 
Wilcox  965;  Gros  Ventre  River,  16  Aug  1894, 
A.  Selson  927. 

Rorippa  curvipes  Greene  var.  curvipes  {R. 
obtusa  (Nutt.)  Britt.  var.  obtusa).  Lizard 
Point,  N  end  Jackson  Lake,  elev.  66(K)  ft.  15 
Aug  1961,  C.  L.  6  M.  Porter  8829a.  duplicate 
annotated  by  R.  L.  Stuckey,  s.d. 

Rorippa  curvipes  Greene  var.  alpina 
(Wats.)  Stuckey  {R.  obtusa  (Nutt.)  Britt.  \ar. 


410 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


alpina  (Wats.)  Britt.).  E  slope  Grand  Teton, 
elev.  8500  ft,  6  Aug  1920,  E.  B  ^  L.  B.  Pay- 
son  2210,  duplicate  annotated  by  R.  L. 
Stuckey,  s.d. 

Sisymbrium  altissimum  L.  Gros  Ventre 
River  (SI  T42N  R115W),  elev.  6800  ft,  27 
Jun  1977,  R.  Lichvar  497. 

Subularia  aquatica  L.  Near  Togwotee 
Pass,  elev.  9600  ft,  28  Aug  1952,  C.  L.  Porter 
6211. 

Thelypodium  paniculatum  A.  Nels.  Bacon 
Creek,  15  Aug  1894,  A.  Nelson  922. 

Callitrichaceae 

Callitriche  hermaphroditica  L.  Near  Tog- 
wotee Pass,  elev.  9000  ft,  4  Aug  1953,  C.  L. 
Porter  6373. 

Caryophyllaceae         • 

Arenaria  stricta  Michx.  ssp.  dawsonensis 
(Britt.)  Maguire.  Vicinity  of  Hoback  Canyon, 
elev.  7500  ft,  24  Jun  1932,  L.  Williams  6  R. 
Pierson  718. 

Dianthus  armeria  L.  Near  Pilgrim  Creek, 
17  Aug  1965,  A.  A.  Beetle  s.n. 

Stellaria  jamesiana  Torr.  Hoback  Canyon, 
6  mi  SE  of  Hoback  Junction,  elev.  ca.  7000 
ft,  7  Jun  1969,  R.  L.  Hartman  2845. 

Chenopodiaceae 

Chenopodium  glaucum  L.  var.  salinum 
(Standi.)  Boiv.  Near  Gros  Ventre  River,  9  or 
10  mi  E  of  the  slide,  elev.  7100  ft,  28  Aug 
1949,  /.  F.  ir  M.  S.  Reed  2700;  Jackson  Lake, 
24  Aug  1922,  E.  B.  ir  L.  B.  Payson  3087,  an- 
notations by  H.  A.  Wahl,  1960. 

Chenopodium  leptophyllum  (Moq.)  Wats. 
Gros  Ventre  Road  (S4  T42N  R115W),  elev. 
7100  ft,  18  Aug  1977,  R.  Lichvar  1281. 

Cyperaceae 

Carex  backii  Boott.  Treasure  Mt.  Scout 
Camp,  Teton  Canyon,  25  Jun  1956,  L.  C.  An- 
derson 359,  annotation  by  F.  J.  Hermann, 
1958,  as  C.  saximontana  Mack. 

Carex  buxbaumii  Wahl.  E  shore  String 
Lake,  elev.  ca.  6000  ft,  29  Jul  1964,  W.  M. 
Johnson  465,  annotation  by  F.  J.  Hermann, 
1964. 


Carex  illota  Bailey.  Alaska  Basin,  W  slope 
Teton  Range,  elev.  9500-10,000  ft,  14  Aug 
1965,  /.  Merkle  65-62. 

Carex  muricata  L.  Jackson  Hole,  24  Jul 
1901,  E.  D.  Merrill  6  E.  N.  Wilcox  907. 

Carex  narditia  Fries.  Summit  of  Table  Mt., 
elev.  11,100  ft,  17  Jul  1956,  L.  C.  Anderson 
532;  Head  of  S  Fork  Cascade  Canyon,  elev. 
10,500  ft,  1  Aug  1949,  H.  b  V.  Bailey  4936, 
annotations  by  F.  J.  Hermann,  1958  as  C. 
hepburnii  Boott. 

Carex  norvegica  Retz.  Just  E  of  Treasure 
Lake,  Teton  Canyon,  22  Jun  1956,  L.  C.  An- 
derson 347,  annotation  by  F.  J.  Hermann, 
1958,  as  C.  media  R.  Br. 

Carex  nova  Bailey.  Teton  Mts.,  16  Aug 
1899,  A.  Nelson  <b  E.  Nelson  6527,  annotation 
by  F.  J.  Hermann,  1958,  as  C.  pelocarpa  Her- 
mann. 

Carex  praticola  Rydb.  Grassy  Lake  road 
W  of  Flagg  Ranch,  elev.  6600  ft,  9  Jul  1959, 
C.  L.  6  M.  W.  Porter  7876,  annotation  by  F. 
J.  Hermann,  1959. 

Eleocharis  flavescens  (Poir.)  Urban,  Flagg 
Ranch,  Jul  1954,  A.  A.  Beetle  16409. 

Eleocharis  rostellata  Torr.  Huckleberry 
Hot  Springs,  12  Aug  1974,  A.  A.  Beetle  s.n. 

Eriophorum  polystachion  L.  Grand  Teton 
National  Park,  elev.  7000  ft,  1  Aug  1932,  L. 
Williams  989. 

Dipsacaceae 

Knautia  arvensis  (L.)  Coult.  Jackson,  Jun 
1969,  H.  P.  Alley  s.n. 

Ericaceae 

Vaccinium  globulure  Rvdb.  Along  Taggart 
Creek,  elev.  7500  ft,  15 'jun  1933,  L.  Wil- 
liams 1123,  cited  by  Camp  (1942). 

Fabaceae 

Astragalus  argophyllus  Nutt.  Near  Snake 
River  at  Simon's,  E  of  Moran,  elev.  6750  ft,  2 
Jun  1948,  /.  F.  6  M.  S.  Reed  1833,  annotation 
by  R.  C.  Banieby,  1960. 

Astragalus  diversifolius  Gray.  Gros  Ventre 
River,  16  Aug  1894,  A.  Nelson  1086. 

Astragalus  eucosmus  Robins.  Spread 
Creek,  E  side  of  Jackson  Hole,  elev.  7200  ft, 
13  Jun  1948,  /.  F.  6  M.  S.  Reed  2261,  annota- 
tion by  R.  C.  Barneby,  1960. 


December  1980 


Hartmax,  Lichvar:  Vascular  Flora 


411 


Astragalus  miser  Dougl.  ex  Hook.  var. 
tenuifolius  (Nutt.)  Banieby.  V^icinitv  of  Ho- 
back  Canyon,  elev.  8000  ft,  24  Jun'l932,  L. 
Williams  ir  R.  Pierson  721,  annotation  bv  D. 
Isely,  1976. 

Astragalus  teiwUus  Pursh.  Near  Adams 
Ranch,  Jackson  Hole,  15  Jnl  1901,  E.  D.  Mer- 
rill 6  E.  N.  Wileox  964. 

Lupinus  tiyethii  Wats.  Grand  Teton  Na- 
tional Park.  elev.  7000  ft,  21  Jun  1932,  L. 
Willidtiis  6801);  Two-gwo-tee  Pass,  elev. 
10,500  ft,  27  Jul  1932,  L.  Williams  940,  anno- 
tations by  B.  J.  Cox,  1969. 

Oxytropis  sericea  Nutt.  var.  spicata 
(Hook.)  Barneby.  Near  mouth  of  Fish  Creek, 
Gros  Ventre  drainage,  28  Jun  1958,  D.  E. 
Wilbeii  5. 

Oxytropis  viscida  Nutt.  Gros  Ventre  Riv- 
er, 16  Aug  1894,  A.  Nelson  928;  Sportsmans 
Ridge  (SIO  T40N  R112W),  elev.  9200  ft,  21 
Jul  1977,  R.  Liehvar  980. 


Juncus  torreyi  Gov.  Teton  Mts.,  21  .\ug 
1895,  A.  Selwn  956,  annotation  by  F.  J.  Her- 
mann, 1957. 


Juncaginaceae 

Triglochin  palustris  L.  2  mi  SW  of  Jack- 
son, elev.  2000  m,  30  Aug  1949,  J.  F.  Reed 
2726;  Near  Gros  Ventre  River,  ca.  9  mi 
above  the  slide,  elev.  7100  ft,  28  Aug  1949.  /. 
F.  6  M.  S.  Reed  2697. 


Lamlaceae 

Nepeta  cataria  L,  Kelh  Warm  Springs  (SI 
T42N  R115W),  elev.  6800  ft,  10  Aug  1977, 
R.  Lichvar  1240. 

Stachys  palustris  L.  var.  pilosa  (Nutt.) 
Fern.  20  mi  S  of  Jackson,  19  Julv  1901,  E.  D. 
Merrill  6  E.  N.  Wilcox  953. 


Gentianaceae 

Gentiana  barbellata  Engelm.  Crvstal 
Creek  divide,  25  Aug  1933,  O.  J.  Murie  1052. 

Gentiana  prostrata  Haenke.  E  of  the  slide, 
Ciros  Ventre  Mts.,  elev.  7300  ft,  28  Aug  1949, 
/.  F.  6  A/.  S.  Reed  2701. 

Grossul.ariaceae 

Ribes  aureum  Pursh.  Snake  River  bottom, 
6  mi  SW  of  Jackson,  elev.  6000  ft,  29  Jun 
1933,  L.  Williams  1185. 

Hydroch.aritaceae 

Elodea  longivaginata  St.  John.  Two-Ocean 
Lake,  elev.  6900  ft,  4  Aug  1953,  C.  L.  Porter 
6365. 


Hydrophyllaceae 

Phacelia  linearis  (Pursh)  Holz.  Snake  Riv- 
er, 29  May  1892,  F.  McCoullough  s.n.,  anno- 
tation bv  G.  W.  Gillett,  1958. 


JUNCACEAE 

Juncus  nodosus  L.  2  mi  SW  of  Jackson, 
elev.  2000  m,  30  Aug  1949,  /.  F.  Reed  2727. 


Lemnaceae 

Lemna  minuta  H.B.K.  Third  Creek  near 
Swan  Lake,  10  Jun  1959,/.  Wetherell  40. 

LiLIACEAE 

Allium  cernuum  Roth.  Big  Cow  Creek  (S3 
T41N  R112W),  elev.  8100  ft,  30  Jul  1977,  R. 
Lichvar  993. 

Asparagus  officinalis  L.  Gros  Ventre  Ris- 
er at  Kellv  (SI  T42\  R115W).  elev.  6700  ft. 
27  Jun  1977,  R.  Lichvar  496. 

Calochortus  nuttallii  T.  &  G.  Teton  Pass 
Mts.,  elev.  7000  ft,  22  Jul  1920.  E.  B.  6  L.  B. 
Payson  2093,  annotation  by  M.  Ownbey, 
1939. 

Malvaceae 

Sidalcea  oregana  (Nutt.)  Gray  var.  orc- 
gana.  Granite  Creek,  vicinity  of  Hoback 
C:anyon,  elev.  8000  ft,  10  Jul  1932,  /..  Wil- 
liams 836. 


Najaoaceae 

Najas  guadalupensis  (Spreng.)  Morong. 
Kellv  Warm  Springs,  elev.  6700  ft.  17  Aug 
1971.  K.  D.  Dorn  1424. 


412 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


Onagraceae 

Clarkia  pulchella  Pursh.  Snake  River,  29 
May  1892,  F.  McCouUough  s.n.,  annotation 
by  M.  &  H.  Lewis,  1951. 

Orchidaceae 

Habenaria  saccata  Greene.  Jackson  Hole, 
elev.  6700  ft,  8  Aug  1920,  E.  B.  6  L.  B.  Pay- 
son  2248,  annotation  by  D.  S.  Correll,  1946; 
Granite  Creek  (S5  T39N  R113W),  elev.  8200 
ft,  10  Jul  1977,  R.  Liclwar  721. 

Orobanchaceae 

Orobanche  corymbosa    (Rydb.)    Ferris. 

Near  Cliff  Creek,  Hoback  Canyon,  19  Aug 
1922,  E.  B.  ir  L.  B.  Paijson  3078,  annotation 
by  L.  R.  Heckard,  s.d. 

Plant  AGiNACEAE 

Plantago  tweedyi  Gray.  Trail  Creek,  vicin- 
ity of  Teton  Pass,  elev.  7500  ft,  1  Jul  1932,  L. 
Williams  790. 


POLEMONIACEAE 

Gilia  spicata  Nutt.  Sportsmans  Ridge  (SIO 
T40N  R112W),  elev.  9200  ft,  21  Jul  1977,  R. 
Lichvar  ,977;  Curtis  Canyon  Campground 
(S16  T41N  R115W),  elev.  8000  ft,  5  Jun 
1977,  R.  Lichvar  263. 

Gilia  tenerrima  Gray.  Near  Moose  (NWVi, 
S12  T44N  R115W),  3  Jul  1970,  D.  W.  Sa- 
binske  12A;  Teton  Canyon,  11  mi  E  of 
Driggs,  Idaho,  2  Jun  1956,  L.  C.  Anderson 
264. 

Navarretia  breweri  (Gray)  Greene.  Gros 
Ventre  River,  18  Aug  1894,  A.  Nekon  1094. 

POLYGONACEAE 

Erigonum  brevicaule  Nutt.  var.  laxifolium 
(T.  &  G.)  Reveal  (£.  chrysocephalum  Gray). 
Bacon  Creek,  15  Aug  1894,  A.  Nelson  903; 
Gros  Ventre  slide  area,  elev.  7000  ft,  10  Jul 
1959,  C.  L.  6  M.  W.  Porter  7892,  annotations 
by  J.  L.  Reveal,  1971;  Soda  Creek  (S14  T41N 
R112W),  elev.  7200  ft,  21  Jul  1977,  R.  Lich- 
var 941. 


Polygonum  achoreum  Blake.  Jackson  Hole 
Wildlife  Park,  elev.  6750  ft,  25  Aug  1949,  /. 
F.  6  M.  Reed  2674. 

Polygonum  confertiflorum  Nutt.  Near 
Jackson  Lake,  elev.  6750  ft,  21  Aug  1935,  M. 
Ownbey  976;  Snake  River,  15  Aug.  1899,  A. 
Nelson  b-  E.  Nelson  6463,  annotations  by  J. 
O.  Coolidge,  1963. 

Polygonum  saivatchense  Small.  Two-gwo- 
tee  Pass,  25  Jul  1939,  Miss  Gooding  39-139. 

Polygonum  watsonii  Small.  Jackson  Lake, 
21  Aug  1899,  A.  Nelson  6  E.  Nelson  6556; 
Hwy.  89,  2  mi  S  of  Moran,  27  Jul  1939,  /.  F. 
Brenckle  6-  O.  A.  Stevens  44,  annotations  by 
J.  O.  Coolidge,  1963. 

Primulaceae 

Primula  incana  Jones.  Hot  Spring  Bar,  20 
mi  S  of  Jackson,  19  Jul  1901,  E.  D.  Merrill  b 
E.  N.  Wilcox  1039;  Adams  Ranch,  Jackson 
Hole,  14  Jul  1901,  E.  D.  Merrill  6  E.  N.  Wil- 
cox 990;  Soda  Lake  (Sll  T41N  R112W),  elev. 
7100  ft,  21  Jul  1977,  R.  Lichvar  937. 

Potamogetonaceae 

Potamogeton  amplifolius  Tuckerman.  In 
Lake  of  the  Woods,  near  the  Continental  Di- 
vide, elev.  9400  ft,  8  Aug  1956,  C.  L.  Porter 
7196. 

Potamogeton  epihydrus  Raf.  Signal  Mt., 
elev.  6800  ft,  19  Jul  1963,  C.  L.  6  M.  W.  Por- 
ter 9401. 

Ranunculaceae 

Anemone  parviflora  Michx.  Teton  Pass 
Mts.,  elev.  9500  ft,  25  Jul  1920,  E.  B.  6  L.  B. 
Paijson  2134. 

Anemone  nuttalliana  DC.  Snow  King  Mt., 
(S35  T41N  R116W),  elev.  6500  ft,  25  Mav 
1978,  R.  Lichvar  1446. 

Delphinium  burkei  Greene.  N  drainage  of 
Pilgrim  Creek,  E  of  Colter  Bay,  20  Jul  1957, 
A.  A.  Beetle  13974,  annotation  bv  R.  J.  Tay- 
lor, 1960. 

Ratiuticulus  gmelinii  DC.  Lower  Slide 
Lake  (S5  T42N  Rl  14W),  elev.  6900  ft,  4  Jul 
1977,  fl.  Lichvar  572. 

Ranuticulus  uncinatus  D.  Don  var.  parvif- 
lorus  (Torr.)  Benson.  Snake  River  bottom, 
elev.  6500  ft,  28  Jul  1932,  L.  Williams  754, 
annotations  by  L.  Benson,  1945. 


December  1980 


Hartman,  Lichvar:  Vascular  Flora 


413 


Rosaceae 

Potentilla  ovina  Macoun.  Summit  of 
Table  Mt.,  elev.  11,100  ft,  7  Jul  1956,  L.  C. 
Anderson  429;  Togwotee  Pass,  8  Jul  1940,  L. 
E.  Wehmei/er  ct  al.  5322  (NY),  annotations  bv 
B.  C.Johnston,  1979. 

Potentilla  hookeriana  Lchm.  Hoback  Can- 
yon, elev.  7500  ft,  24  Jun  1933,  L.  Williams 
1164,  annotation  by  B.  C.  Johnston,  1978. 

Pyrtis  mains  L.  Kellv  Warm  Springs  (SI 
T42N  R115\V),  elev.  6800  ft,  15  Aug  1977, 
R.  Lichvar  1237. 

Salicaceae 

Poptilus  acuminata  Rvdb.  Gros  Ventre 
River  (SI  T42N  R115W),'  elev.  6800  ft,  27 
Jun  1977,  R.  Lichvar  524. 

Saxifragaceae 

Conimitella  williamsii  (D.C.  Eat.)  Rvdb. 

Hoback  Canyon,  elev.  7500  ft,  23  Jun  1933, 
L.  Williams  1144. 

Saxifraga  adscendens  L.  var.  oregonensis 
(Raf.)  Breit.  N  side  of  the  Grand  Teton,  elev. 
11,000  ft,  10  Aug  1932,  L.  Williams  1016. 

Saxifraga  caespitosa  L.  var.  minima 
Blake.  Single-Shot  Mt.,  4  Jul  1897,  R.  S.  Wil- 
liajns  s.n.,  cited  by  Hitchcock  et  al.  (1961). 

Saxifraga  cernua  L.  Sheep  Mt.  (S3  T41N 
R114W),  elev.  10,500  ft,  1  Aug  1977,  R. 
Lichvar  1028. 

Scrophularl\ceae 

Castilleja  gracillima  Rvdb.  Blackrock 
Creek,  Two-gwo-tee  Pass,  elev.  9500  ft,  29 
Jul  1932,  L.  Williams  ,960,  annotation  by  M. 
Ownbey. 

Mimulus  tilingii  Regel.  Grand  Teton  Na- 
tional Park,  elev.  7000  ft,  22  Jul  1933,  L. 
Williams  1322. 

Orthocarpus  tolmiei  H.  &  A.  Teton  Pass 
Mts.,  elev.  8000  ft,  22  Jul  1920,  E.  B.  b  L.  B. 
Payson  2072,  annotation  by  D.  D.  Keck, 
1926. 


Penstemon  glaher  Pursh  ssp.  glaber.  Gros 
Ventre  slide  area,  elev  7000  ft,  10  Jul  1959, 
C.  L.  6  .\/.  \V.  Porter  7S95,  annotation  bv  F. 
S.  Crosswhite,  1969. 

Penstemon  rydbergii  A.  Nels.  var.  ryd- 
bergii.  Grand  Teton  National  Park,  elev. 
8000  ft,  19  Jul  1932,  L.  Williams  S92,  annota- 
tion by  D.  V.Clark.  1971. 

Synthyris  pinnatifida  Wats.  Teton  Pass 
Mts.,  elev.  9200  ft,  22  Jul  1920,  £.  B.  6  /..  B. 
Payson  2079,  annotation  by  C.  G.  Schaack, 
1975;  Cache  Divide  (S14  T40N  R115W), 
elev.  9200  ft,  4  Aug  1977,  R.  Lichvar  1166. 

Veronica  scutellata  L.  Jackson  Lake,  12 
Aug  1899,  A.  Xelson  6  E.  Selson  6561,  anno- 
tation by  F.  W.  Pennell,  1920. 

Solanace.ae 

Solanum  dulcamara  L.  Spalding  Bav  (S34 
T44N  R114W),  elev  6400  ft,  1  Aug  1978,  R. 
Lichvar  1604. 

Violaceae 

Viola  canadensis  L.  var.  canadensis.  Vi- 
cinitv  of  Hoback  Canvon,  elev.  7000  ft.  25 
Jun  1932,  /..  Williams'{:r  R.  Pier.wn  731,  an- 
notation bv  N.  H.  Russell,  1967. 

Viola  nephrophylla  Greene.  Hot  Spring 
Bar,  20  mi  S  of  Jackson,  19  Jul  1901.  E.  D. 
Merrill  6  E.  X.  Wilcox  1040,  N'icinitv  of  .Mo- 
ran,  7500-8500  ft,  22-30  Jun  1935,  T.  G. 
Yuncker  5291,  annotations  bv  N.  H.  Russell, 
1967. 

Literature  Cited 

lIiTfncocK,  C.  L..  A.  Cronqiist.  \t.  Ownbev.  and  J. 
W.  Thompson.  1961.  Vascular  plants  of  the  Pacif- 
ic Northwest.  I'nivcrsitv  of  W'ashiiiiitoii  Press, 
Seattle,  vol.  .3.614  pp. 

HiT(  IK  ocK,  C.  L.,  A.ND  .\.  Cronqiist.  1973.  Flora  of  the 
Pacific  Northwest.  University  of  Washington 
Press,  Seattle,  xix  -f-  730  pp. 

Shaw,  R.  J.  (with  major  contributions  by  R.  D.  Dorn). 
1976.  Field  piide  of  the  vascular  plants  of  Grand 
Teton  National  Park  and  Teton  Co\inty,  Wyom- 
ing. Utah  State  University  Press.  Logan,  xvi  -1- 
301  pp. 


INDEX  TO  VOLUME  40 


The  genera  and  species  decrihed  as  new  to  science  in  this  vohiine  appear  in  hold  t\  pe  in 
this  index. 


Ahronia  argillosa,  p.  78. 

Additions  to  the  vascular  Oora  of  Teton 
County,  Wyoming,  p.  408. 

Allred,  Dorald  M.,  articles  by,  pp.  116.  165. 

Anipldcninus  spectus,  p.  358. 

Anaxylebonis,  p.  90. 

Andersen,  Ferron  L.,  and  Peter  M.  Schantz, 
article  by,  p.  216. 

Andersen,  William  R.,  and  Jack  D.  Broth- 
erson,  article  bv,  p.  121. 

Anderson,  Loran  C,  articles  bv,  pp.  73,  117, 
351. 

Andre,  John  B.,  and  James  A.  MacMahon,  ar- 
ticle bv,  p.  68. 

Androsace  alaskana,  var.  reedae,  p.  80. 

Apoxyleborus,  p.  90. 

Araptus  speciosus,  p.  357. 

Atwood,  N.  Duane,  Clyde  L.  Pritchett,  Rich- 
ard D.  Porter,  and  Benjamin  W.  Wood,  ar- 
ticle by,  p.  303. 

Bacterium  Thioploca  ingrica  on  wet  walls  in 
Zion  National  Park,  Utah,  The,  p.  98. 

Bateman,  Lucinda,  article  by,  p.  268. 

Baugh,  Thomas  M.,  article  by,  p.  139. 

Baugh,  Thomas  M.,  and  Bnice  C.  Brown,  ar- 
ticle by,  p.  359. 

Baumann,  Richard  W.,  and  Bill  P.  Stark,  ar- 
ticle by,  p.  63. 

Breinholt,  J.  Craig,  and  Richard  A.  Heck- 
mann,  article  by,  p.  149. 

Brotherson,  Jack  D.,  article  by,  p.  372. 

Brotherson,  Jack  D.,  and  William  R.  Ander- 
sen, article  bv,  p.  121. 

Brotherson,  Jack  D.,  William  E.  Evenson, 
and  Richard  B.  Wilcox,  article  by,  p.  167. 

Brotherson,  Jack  D.,  Lee  A.  Szyska,  and  Wil- 
liam E.  Evenson,  article  bv,  p.  229. 

Brown,  Bruce  C,  and  Thomas  M.  Baugh.  ar- 
ticle by,  p.  359. 

Bryan,  James  A.,  and  Ronald  \l.  Lanner,  ar- 
ticle by,  p.  190. 

Burton,  Sheril  D.,  Sanniel  R.  Rushforth,  Jef- 
frey R.  Johansen,  and  fuciith  A.  Grimes,  ar- 
ticle by,  p.  98. 

Campbell  III,  Thomas  M.,  and  Tim  W. 
Clark,  article  by,  p.  183. 


Chiracdnthiiini  sj^idcr  ])ite.  A.  p.  116. 

Cltnisi>th(iinnus  pulchelloides,  p.  351. 

Clark,  Tim  W.,  and  Thomas  M.  Campbell 
111,  article  by,  p.  183. 

Collin.s,  Nicholas  C,  and  Grav  Stirling,  ar- 
ticle by,  p.  131. 

Comparative  floral  biology  of  Pcnstrmon  ca- 
tonii  and  Pcnstemon  cyananthiis  in  central 
Utah:  a  preliminarv  studv,  p.  268. 

Comparison  of  epiph\  tic  diatom  assemblages 
on  living  and  dead  stems  of  the  conunon 
grass  Plinif^niitcs  (iiistnili.s,  .\,  p.  223. 

Cono))htlu)nis  michoacanae,  p.  354. 

Cou()j)lithoni.s  teocotum,  p.  354. 

Cryphalogenes,  p.  91. 

Ciyphal()<gcncs  cuphorbiae,  p.  91. 

CrypluiloigciU's  exiguus,  p.  92. 

Cushing,  Bruce  S.,  and  .\nne  Matherne.  ar- 
ticle by,  p.  193. 

Differential  habitat  utilization  l)\  the  sexes  of 
mule  deer,  p.  273. 

Dog  owners  and  hydatid  disease  in  Sanpete 
County,  Utah,  p.  216. 

Egoscue,  Harold  J.,  article  bv,  p.  361. 

Elliott,  Charles  L.,  and  Jerran  T.  Flinders,  ar- 
ticles by,  pp.  175,  362. 

Ernocladiu.s,  p.  93. 

Evenson,  William  E.,  Jack  D.  Brotherson, 
and  Lee  \.  Szvska,  article  by,  p.  229. 

Evenson,  \\'illiam  E.,  Jack  D.  Brotherson, 
and  Richard  B.  Wilcox,  article  by.  p.  167. 

Feeding  ecology  of  Gila  honixohitts  (Os- 
teichthves:  Cvprinidae)  endemic  to  a  ther- 
mal lake  in  southeastern  Oregon,  p.  101. 

Ferrington,  Leonard  C,  Jr.,  and  Bernard  C 
Swegman,  article  by,  p.  287. 

Field  observations  on  the  response  of  the 
Railroad  Valley  springfish  [Crcnichtliijs  nc- 
vadae)  to  temperature,  p.  359. 

First  record  of  the  pallid  bat  iAutroznits  piil- 
lidus)  from  .Montana,  p.  115. 

Flath,  Dennis  L.,  and  Jeff  Shryer,  article  l)\. 
p.  115. 

Flinders,  Jerran  T.,  and  Charles  L.  Elliott,  ar- 
ticles b\ .  pp.  175,  362. 


415 


416 


Great  Basin  Naturalist 


Vol.  40,  No.  4 


Flood  frequency  and  the  assemblage  of  dis- 
persal types  in  hanging  gardens  of  the 
Narrows,  Zion  National  Park,  Utah,  p. 
365. 

Garcia,  James  R.,  Clive  D.  Jorgensen,  and  H. 
Duane  Smith,  article  by,  p.  282. 

Genus  Eriogoniim  Michx.  (Polygonaceae)  and 
Michel  Gandoger,  The,  p.  143. 

Goodrich,  Sherel,  and  Stanley  L.  Welsh,  ar- 
ticle by,  p.  78. 

Grimes,  Judith  A.,  Samuel  R.  Rushforth,  She- 
ril  D.  Burton,  Jeffrey  R.  Johansen,  article 
by,  p.  98. 

Grimes,  Judith  A.,  Larry  L.  St.  Clair,  and 
Samuel  R.  Rushforth,  article  by,  p.  223. 

Habitat  and  plant  distributions  in  hanging 
gardens  of  the  Narrows,  Zion  National 
Park,  Utah,  p.  178. 

Hadrodemius,  p.  94. 

Haplopappits  alpinus,  p.  73. 

Haplopappus  alpinus  (Asteraceae):  a  new 
species  from  Nevada,  p.  73. 

Harper,  K.  T.,  and  L.  M.  Kunzler,  article  by, 
p.  127. 

Hartman,  Ronald  L.,  and  Robert  W.  Lichvar, 
article  by,  p.  408. 

Hatch,  Stephan  L.,  article  by,  p.  221. 

Heckmann,  Richard  A.,  and  J.  Craig  Brein- 
holt,  article  by,  p.  149. 

Henderson,  Jan  A.,  David  J.  Schimpf,  and 
James  A.  MacMahon,  article  by,  p.  1. 

Hesperoperla  hoguei,  p.  63. 

Hesperoperki  hoguei,  a  new  species  of  stone- 
fly  from  California  (Plecoptera:  Perlidae), 
p.  63. 

Hijlesinus  aztecus,  p.  354. 

Hijlurdrectonus  corticinus,  p.  94. 

Identity  of  narrow-leaved  CJinjsothamniis 
viscidiflonis  (Asteraceae),  p.  117. 

Impact  of  the  1975  Wallsburg  fire  on  ante- 
lope bitterbrush  {Purshia  tridentata),  p. 
299. 

Johansen,  Jeffrey  R.,  Samuel  R.  Rushforth, 
Sheril  D.  Burton,  and  Judith  A.  Grimes, 
article  by,  p.  98. 

Jorgensen,  Clive  D.,  H.  Duane  Smith,  and 
James  R.  Garcia,  article  by,  p.  282. 

Kay,  Jeanne,  and  George  P.  Malanson,  article 
by,  p.  365. 

Keiss,  Robert  E.,  Patrick  W.  Roberts,  and 
Donald  J.  Nash,  article  by,  p.  141. 

King,  Michael  M.,  and  H.  Duane  Smith,  ar- 
ticle by,  p.  273. 


Kunzler,  L.  M.,  and  K.  T.  Harper,  article  by, 
p.  127. 

Lanner,  Ronald  M.,  article  by,  p.  265. 

Lanner,  Ronald  M.,  and  James  A.  Bryan,  ar- 
ticle by,  p.  190. 

Lepidium  ostleri,  p.  80. 

Leptoxyleborus,  p.  94. 

Lichvar,  Robert  W.,  and  Ronald  L.  Hartman, 
article  by,  p.  408. 

Lygodesmia  entrada,  p.  83. 

MacMahon,  James  A.,  and  John  B.  Andre,  ar- 
ticle by,  p.  68. 

MacMahon,  James  A.,  David  J.  Schimpf,  and 
Jan  A.  Henderson,  article  by,  p.  1. 

Malanson,  George  P.,  article  by,  p.  178. 

Malanson,  George  P.,  and  Jeanne  Kay,  article 
by,  p.  365. 

Matherne,  Anne,  and  Bruce  S.  Gushing,  ar- 
ticle by,  p.  193. 

Microperus,  p.  94. 

Miscellaneous  plant  novelties  from  Alaska, 
Nevada,  and  Utah,  p.  78. 

Murphy,  Robert  W.,  John  R.  Ottley,  and 
Geoffrey  V.  Smith,  article  by,  p.  59. 

Nash,  Donald  J.,  Patrick  W.  Roberts,  and 
Robert  E.  Keiss,  article  by,  p.  141. 

New  American  bark  beetles  (Coleoptera: 
Scolytidae),  with  two  recently  introduced 
species,  p.  353. 

New  genera  and  new  generic  synonymy  in 
Scolytidae  (Coleoptera),  p.  89. 

New  grass  distribution  records  for  Arizona, 
New  Mexico,  and  Texas,  p.  221. 

New  records  of  western  Trichoptera  with 
notes  on  their  biology,  p.  287. 

New  species  of  fossil  Chrysothamnus  (Aste- 
raceae) from  New  Mexico,  A,  p.  351. 

Observations  on  seasonal  variation  in  desert 
arthropods  in  central  Nevada,  p.  292. 

Ottley,  John  R.,  Robert  W.  Murphy,  and 
Geoffrey  V.  Smith,  article  by,  p.  59. 

Parasites  from  two  species  of  suckers  (Ca- 
tostomidae)  from  southern  Utah,  p.  149. 

Parker,  Albert  J.,  article  by,  p.  254. 

Pediococtus  despainii,  p.  83. 

Phloeocleptus  punctatus,  p.  355. 

Pietruszka,  Robert  D.,  article  by,  p.  292. 

Pityogenes  mexicanus,  p.  356. 

Poisonous  plants  of  Utah,  p.  229. 

Porter,  Richard  D.,  N.  Duane  Atwood,  Clyde 
L.  Pritchett,  and  Benjamin  W.  Wood,  ar- 
ticle by,  p.  303. 


December  1980 


Index 


417 


Postemergence  development  and  interyear 
residence  of  juvenile  Columbian  groimd 
squirrels  in  the  Idaho  primitive  area,  p. 
362. 

Plants  of  Angel  Island,  Marin  ('ountv,  C^ali- 
foniia,  p.  385. 

Pritchett,  Clyde  L.,  N.  Duane  Atwood,  Rich- 
ard D.  Porter,  and  Benjamin  W.  Wood,  ar- 
ticle by,  p.  303. 

Psetidotliifsanocs  atonius,  p.  355. 

Pscudothi/sanocs  leechi,  p.  356. 

Recovery  of  Gambel  oak  after  fire  in  central 
Utah,' p.  127. 

Relationship  between  environmental  and 
vegetational  parameters  for  luiderstorv 
and  open-area  commimities,  p.  167. 

Relationships  among  total  dissolved  solids, 
conductivitv,  and  osmositv  for  five  Ar- 
temia  habitats  (Anostraca:  Artemiidae),  p. 
131. 

Reproduction  in  three  sympatric  lizard  spe- 
cies from  west-central  Utah,  p.  68. 

Reveal,  James  L.,  article  by,  p.  143. 

Ribulose  diphosphate  carboxylase  activities 
in  cold-resistant  common  mallow,  Malvo 
neglecta  Wallr.  and  a  cold-sensitive  to- 
mato, Lycopersicon  esculciituin  L.,  Ace  55 
var.,  p.  121. 

Ripley,  J.  D.,  article  by,  p.  385. 

Roberts,  Patrick  W.,  Donald  J.  Nash,  and 
Robert  E.  Keiss,  article  by,  p.  141. 

Rushforth,  Samuel  R.,  Sheril  D.  Burton,  Jef- 
frev  R.  Johansen,  and  Judith  A.  (wimes,  ar- 
ticle by,  p.  98. 

Rushforth,  Samuel  R.,  Judith  A.  Grimes,  and 
Larry  L.  St.  Clair,  article  by,  p.  223. 

St.  Clair,  Larry  L.,  Judith  A.  Crimes,  and 
Samuel  R.  Rushforth,  article  by,  p.  223. 

Schantz,  Peter  M.,  and  Perron  L.  Andersen, 
article  by,  p.  216. 

Schimpf,  David  J.,  Jan  A.  Henderson,  and 
James  A.  MacMahon,  article  by,  p.  1. 

Seasonal  activity  pattern  of  Columbian 
ground  squirrels  in  the  Idaho  primitive 
area,  p.  175. 

Self-pollination  experiment  in  Piiuis  ('dulis, 
A,  p.  265. 

Senecio  toiyabensis,  p.  86. 

Short-term  effects  of  logging  on  red-backed 
voles  and  deer  mice,  p.  183. 

Shryer,  Jeff,  and  Dennis  L.  Flath,  article  by, 
p.  115. 


Smith,  Geoffrey  V..  John  R.  Ottley,  and  Rob- 
ert W.  Murphy,  article  bv,  p.  59. 

Smith,  H.  Duane,  Clive  D.  Jorgensen.  and 
James  R.  Garcia,  article  by,  p.  282. 

Smith,  H.  Duane,  and  Michael  M.  King,  ar- 
ticle by,  p.  273. 

Soil  water  withdrawal  and  root  distribution 
under  grubbed,  sprayed,  and  undisturbed 
big  sagebrush  vegetation,  p.  157. 

Some  aspects  of  succession  in  the  spruce-fir 
forest  zone  of  northern  Utah,  p.  1. 

Spatiotemporal  variation  in  phenology  and 
abundance  of  floral  resources  on  .shortgrass 
prairie,  p.  197. 

Spawning  of  the  least  cinib  i/o/ir/i/Zii/.s 
pJiIegetJwntis),  p.  139. 

Sphaeralcea  grossularufoHa.  \'ar.  moorei,  p. 
35. 

Sphaeralcea  Icptopln/lla,  var.  janeae,  p.  36. 

Sphaeralcea  psoraloides,  p.  36. 

Stanton,  N.  L.,  and  V.  J.  Tepedino,  article  by, 
p.  197. 

Stark,  Bill  P.,  and  Richard  W.  Bauniann.  ar- 
ticle bv,  p.  63. 

Stinger  utilization  and  predation  in  the  scor- 
pion Paruroctonus  horeus,  p.  193. 

Stirling,  Gray,  and  Xicliolas  C.  Collins,  ar- 
ticle by,  p!  131. 

Sturges,  David  L.,  article  by,  p.  157. 

Successional  status  of  Cupressus  arizonica. 
The,  p.  254. 

Swarming  of  the  western  harvester  ant,  Pu- 
gonoDiijmiex  occidentalis,  p.  165. 

Swegman,  Bernard  G.,  and  Leonard  C. 
Ferrington,  Jr.,  article  by,  p.  287. 

Szyska,  Lee  A.,  Jack  D.  Brotherson,  and 
William  E.  Evenson,  article  by.  p.  229. 

Taphrodasus,  p.  95. 

Taurodemus,  p.  96. 

Taxonomic  status  of  the  rosy  boa  Lichonura 
roseofusca  (Serpentes:  Boidae),  The,  p.  59. 

Temporal  activity  patterns  of  a  Dipodonuis 
ordii  population,  p.  282. 

Tepedino,  V.  J.,  and  N.  L.  Stanton,  article  by, 
p.  197. 

Terminal  bud  formation  in  linilHT  piiK-.  p. 
190. 

Terrestrial  vertebrate  fauna  of  llu-  Kaiparo- 
wits  Basin,  p.  303. 

Transferrin  polymorphism  in  bighorn  sheep. 
Otis  canadensh,  in  Colorado,  p.  141. 

Utah  flora:  Malvaceae,  p.  27. 

Utah  flora:  miscellaneous  families,  p.  38. 


418                                                     Great  Basin  Naturalist                                    Vol.  40,  No.  4 

Waejstaff,  Fred  J.,  article  by,  p.  299.  Wood,  Benjamin  W.,   N.   Duane  Atwood, 

Wetsh,  Stanley  L.,  articles  by,  pp.  27,  38.  Clyde  L.  Pritchett,  and  Richard  D.  Porter, 

Welsh,  Stanley  L.,  and  Sherel  Goodrich,  ar-  article  by,  p.  303. 

tide  by,  p.  78.  Wood,  Stephen  L.,  articles  by,  pp.  89,  353. 

Wilcox,  Richard  B.,  William  E.  Evenson,  and  Woodrat  nest  flea  Anamiopsi/Uus  amphibolus 

Tack  D.  Brotherson,  article  by,  p.  167.  .          ^u      4-        r>,    ^           oa^ 

„           ^      ,       ^          -,  -r    1  V;   ^^r.ii.  in  southeastern  Oregon,  p.  361. 

Williams,  Cvnthia  D.,  and  Jack  E.  Williams,  ^,  ,  ,                              ^^'/ 

article  by,  p.  101.  Xyleborus  praestans,  p.  358. 

Williams,  Jack  E.,  and  Cynthia  D.  Williams,  Zonation  patterns  in  the  potholes  of  Kalsow 

article  by,  p.  101.  Prairie,  Iowa,  p.  372. 


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TABLE  OF  CONTENTS 

Impact  of  the  1975  Wallsburg  fire  on  antelope  Ijitterbnish  (Purshia  tiidcntata).  Fred 

J.  Wagstaff  299 

Terrestrial  vertebrate  fauna  of  the  Kaiparowits  Basin.  N.  Duane  Atwood,  Clyde  L. 

Pritchett,  Richard  D.  Porter,  and  Benjamin  W.  Wood 303 

A  new  species  of  fossil  ChrysotJiamnus  (Asteraceae)  from  New  Mexico.  Loran  C. 

Anderson  3.51 

New  American  bark  beetles  (Coleoptera:  Scolytidae),  with  two  recently  introduced 

species.  Stephen  L.  Wood  353 

Field  observations  on  the  respon.se  of  the  Railroad  Valley  springfish  (Crenichthijs 

nevadae)  to  temperature.  Thomas  M.  Baugh  and  Bruce  G.  Brown  359 

Woodrat  nest   flea  Anomiopsyllus  atnphibolus   in  southeastern  Oregon.   Harold  J. 

Egoscue  361 

Postemergence  development  and  interyear  residence  of  juvenile  Columbian  groimd 
squirrels  in  the  Idaho  primitive  area.  Charles  L.  Elliott  and  Jerran  T. 
Flinders  .362 

Flood  frequency  and  the  assemblage  of  dispersal  types  in  hanging  gardens  of  the 

Narrows,  Zion  National  Park,  Utah.  George  P.  Malanson  and  Jeanne  Kay  365 

Zonatioii  patterns  in  the  potholes  of  Kalsow  Prairie,  Iowa.  Jack  D.  Brotherson  372 

Plants  of  Angel  Island,  Marin  County,  California.  J.  D.  Ripley  .385 

Additions  to  the  vascular  flora  of  Teton  County,  Wyoming.  Ronald  L.  Hartman  and 

Robert  W.  Lichvar  .". '. '. 408 

Index  to  Volume  40  414 


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